When plants are exposed to drastic environmental changes such as drought, salt or bacterial invasion, rapid stomatal movement confers tolerance to these stresses. This process involves a variety of guard cell expressed ion channels and their complex regulation network. Inward K+ channels mainly function in stomatal opening. On the other hand, guard cell anion channels play a crucial role in the closing of stomata, which is vital in terms of preventing water loss and bacterial entrance. Massive progress has been made on the research of these anion channels in the last decade. In this review, we focus on the function and regulation of Arabidopsis guard cell anion channels. Starting from SLAC1, a main contributor of stomatal closure, members of SLAHs (SLAC1 homologues), AtNRTs (Nitrate transporters), AtALMTs (Aluminum-activated malate transporters), ABC transporters, AtCLCs (Chloride channels), DTXs (Detoxification efflux carriers), SULTRs (Sulfate transporters), and their regulator components are reviewed. These membrane transport systems are the keys to maintaining cellular ion homeostasis against fluctuating external circumstances.