Solid-state fermentation (SSF) is one of the oldest and most economical ways of producing and preserving foods which may improve the nutritional values, sensory properties, and functional qualities of raw materials (1). The SSF processes of most known natural and industrial fermented foods, such as grape wine (2) and cheese (3), are driven by complex communities of microorganisms. As such, studies on the formation and function of food microbiota, especially as-yet-uncultivated microorganisms, during SSF processes are becoming increasingly important.Traditional Chinese vinegars, referred to as cereal vinegars, are important seasoning and medicinal products in Chinese daily life (4). Solid-state acetic acid fermentation (AAF), an important step in producing the flavor compounds of cereal vinegar, is a repeated batch fermentation process that proceeds for many centuries without spoilage (4). In an open work environment, microbes that inhabit solid-state vinegar culture (termed Pei in Chinese) reproducibly metabolize nonautoclaved raw materials (e.g., sorghum, sticky rice, and wheat bran) and synthesize flavor compounds (5). Thus, the function of reproducible fermentation-based metabolism makes this acidic ecosystem (pH 3.0 to 3.5) amenable to adaptation for dissecting the formation and function of microbiota in food fermentation. Although many microbiological studies have been conducted to reveal the diversity and formation of microbial communities during the AAF of cereal vinegars (6-9), the gap between community assemblage and the function of this microbial ecosystem still exists.In vinegar microbiota, the utilization of ethanol and glucose originating from raw materials leads to diverse flavors and the formation of bioactive compounds. 2,3,5,6-Tetramethylpyrazine (TTMP), termed ligustrazine in traditional Chinese medicine, exists abundantly in the rhizome of Ligusticum wallichii. This alka-