Penicillium digitatum is the most important postharvest pathogen of citrus fruits. Along disease progression, the infected citrus peel tissue is acidified due to the accumulation of organic acids. So far, relatively little is known about the environmental factors that regulate pathogenicity in this fungus. In this study, the role of the pH signaling transcription factor PacC in the pathogenesis of P. digitatum was investigated. We identified the pacC ortholog (PdpacC) in P. digitatum and found that its transcript levels were elevated under alkaline conditions (pH a parts per thousand yenaEuro parts per thousand 7) in vitro, as well as during the infection of citrus fruits in spite of the low pH (about 3.0 to 3.5) of the macerated tissue. Na+ and pectin also induced the expression of PdpacC. Disruption of PdpacC resulted in impaired mycelial growth under neutral or alkaline pH conditions and on synthetic medium supplemented with pectin as the sole carbon source, and attenuated virulence towards citrus fruits. Introducing the full length of PdpacC into the Delta PdpacC mutant restored all these phenotypes. The expression of the polygalacturonase gene Pdpg2 and pectin lyase gene Pdpnl1 in P. digitatum was upregulated in the wild type strain but not or weakly upregulated in the Delta PdpacC mutant during infection. Disruption of Pdpg2 also resulted in attenuated virulence of P. digitatum towards citrus fruits. Collectively, we conclude that PdPacC plays an important role in pathogenesis of P. digitatum via regulation of the expression of cell wall degradation enzyme genes, such as Pdpg2 and Pdpnl1.