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Brettanomyces also appears to metabolize ATHP into ETHP, the latter of which has a much higher taste and odor threshold than the former. This may explain how THP flavors and odors may "age out" of lambic over time <ref name=Joseph> Joseph, CML [http://slideplayer.com/slide/4381124/ | Aromatic Diversity of Brettanomyces]</ref>. ETHP is not believed to be further metabolized by Brettanomyces <ref name=Snowdon> Snowdon, EM; Bowyer, MC; Grbin, PR; and Bowyer, PK [http://pubs.acs.org/doi/pdf/10.1021/jf0528613 | Mousey Off-Flavor: A Review], 2006</ref>.
The second primary source of THP during fermentation are heterofermentive lactic acid bacteria (LAB), which various strains can produce all three major forms of THP found in lambic. While nearly every genus of LAB contains at least one strain known to produce THP, Lactobacillus appears to be the primary source of bacterial THP in beer<ref name=Costello> Costello, PJ [REFhttps://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=5&cad=rja&uact=8&ved=0ahUKEwi75dj-x7PXAhVG12MKHWo1DvQQFghJMAQ&url=https%3A%2F%2Fdigital.library.adelaide.edu.au%2Fdspace%2Fbitstream%2F2440%2F22432%2F2%2F02whole.pdf&usg=AOvVaw0Ekk50qgZfvBs3FVo_rLw2 | Formation of mousy off-flavour in wine by lactic acid bacteria], 1998</ref>. It is not known if oxygen plays a role in the production of THP by LAB as it does with Brettanomyces.
==References==
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