分子进化与基因组多样性（张亚平）知识库

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	Mitochondrial genetic diversity, selection and recombination in a canine transmissible cancer.
	Strakova A1; Ní Leathlobhair M1; Allen JL4; Allum KM5; Bansse-Issa L6; Bisson JL1; Castillo Domracheva A7; Wang GD2; Yin TT2; Airikkala-Otter I3; Murchison EP*1; epm27@cam.ac.uk
	2016
发表期刊	ELIFE
卷号	5期号:X页码:e14552
摘要	Canine transmissible venereal tumour (CTVT) is a clonally transmissible cancer that originated approximately 11,000 years ago and affects dogs worldwide. Despite the clonal origin of the CTVT nuclear genome, CTVT mitochondrial genomes (mtDNAs) have been acquired by periodic capture from transient hosts. We sequenced 449 complete mtDNAs from a global population of CTVTs, and show that mtDNA horizontal transfer has occurred at least five times, delineating five tumour clades whose distributions track two millennia of dog global migration. Negative selection has operated to prevent accumulation of deleterious mutations in captured mtDNA, and recombination has caused occasional mtDNA re-assortment. These findings implicate functional mtDNA as a driver of CTVT global metastatic spread, further highlighting the important role of mtDNA in cancer evolution.
关键词	Dog Mitochondria Cancer Biology Transmissible Cancer Canine Transmissible Venereal Tumour Evolutionary Biology Genomics
资助者	This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support.
收录类别	SCI
语种	英语
资助者	This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support. ; This work was supported by grants from the Wellcome Trust (102942/Z/13/A) and the Royal Society (RG130615), and a Philip Leverhulme Prize from the Leverhulme Trust. We are grateful to Michael Stratton for his support.
文献类型	期刊论文
条目标识符	http://ir.kiz.ac.cn/handle/152453/9875
专题	科研部门_分子进化与基因组多样性（张亚平） 遗传资源与进化国家重点实验室
通讯作者	epm27@cam.ac.uk
作者单位	1.Department of Veterinary Medicine, University of Cambridge, Cambridge, United Kingdom. 2.State Key Laboratory of Genetic Resources and Evolution, Yunnan Laboratory of Molecular Biology of Domestic Animals, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China 3.International Training Center, Worldwide Veterinary Service, Aruvankadu, India. 4.Animal Management in Rural and Remote Indigenous Communities, Darwin, Australia 5.World Vets, Fargo, United States 6.Stichting Dierenbescherming Suriname, Paramaribo, Suriname 7.Corozal Veterinary Hospital, University of Panama, Panama City, Panama. 8.Department of Clinical and Veterinary Surgery, São Paulo State University, São Paulo, Brazil 9.St. George's University, True Blue, Grenada 10.The Nakuru District Veterinary Scheme Ltd, Nakuru, Kenya.
推荐引用方式 GB/T 7714	Strakova A,Ní Leathlobhair M,Allen JL,et al. Mitochondrial genetic diversity, selection and recombination in a canine transmissible cancer.[J]. ELIFE,2016,5(X):e14552.
APA	Strakova A.,Ní Leathlobhair M.,Allen JL.,Allum KM.,Bansse-Issa L.,...&epm27@cam.ac.uk.(2016).Mitochondrial genetic diversity, selection and recombination in a canine transmissible cancer..ELIFE,5(X),e14552.
MLA	Strakova A,et al."Mitochondrial genetic diversity, selection and recombination in a canine transmissible cancer.".ELIFE 5.X(2016):e14552.

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4444444.pdf（1043KB）	期刊论文	出版稿	开放获取	CC BY-NC-SA	请求全文

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