Filamentous overgrowth in aerobic granular sludge processes can cause reactor failure. In this work, aerobic granules were cultivated in five identical sequencing batch reactors with acetate or glucose as the carbon source with various values of influent pH (4.5-8). Microscopic observations revealed that acidic pH, rather than the species of carbon source, epistatically controls the aerobic granules with filamentous structure. An acidic pH shifted the structure of the microbial community in the granules, such that the fungus Geotrichum fragrans was the predominant filamentous microorganism therein. The acidic pH reduced the intracellular cyclic diguanylate (c-di-GMP) content for increasing the motility of the bacteria to washout and increase the growth rate of G. fragrans on glucose or acetate, together causing overgrowth of the fungus. Maintaining the suspension under alkaline condition is proposed as an effective way to suppress filamentous overgrowth and maintain granule stability.