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This work was supported by national funds through Fundação para a Ciência e a Tecnologia and by EUFEDER funding through the Operational Competitiveness Programme – COMPETE (grants FCOMP- 01-0124-FEDER-014321 [PTDC/BIA-PRO/110523/2009], FCOMP-01-0124-FEDER-028359 [PTDC/ BIA-MIC/2779/2012] and FCOMP-01-0124-FEDER-037276 [PEst-C/SAU/LA0001/2013-2014]), and operating grants to AJC from the Canadian Institutes of Health Research and the Natural Sciences and Engineering Research Council of Canada. We thank Dyanne Brewer, Armen Charchoglyan and David Sychantha for support with mass spectrometry. AM also acknowledges FCT for the PhD fellowship SFRH/BD/74845/2010