What Makes Sourdough Bread Sour?

With sandwiches and toast, sourdough bread always adds an extra accented flavor to your meals. But where does the signature tartness come from?

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Sources:
http://www.npr.org/templates/story/story.php?storyId=6061648
http://www.npr.org/sections/thesalt/2012/03/07/148120591/into-the-wild-science-of-sourdough-bread-making
http://discovermagazine.com/2003/sep/featscienceof
https://www.exploratorium.edu/cooking/bread/recipe-sourdough.html
http://www.nytimes.com/2016/03/23/dining/sourdough-starter-bread-baking.html
http://viking.lkstevens.wednet.edu/cms/lib03/WA01001468/Centricity/Domain/824/htbchapter3%20yeast.pdf
http://comenius.susqu.edu/biol/312/lactobacillussanfranciscoakeysourdoughlacticacidbacteriumareview.pdf
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC377202/
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC377203/

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