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Chloride (Cl-) channels are transmembrane proteins in biological membranes, which form functional pores and allow the diffusion of negatively-charged Clions along the electrochemical gradients. These channels can also conduct other anions including I-, Br-, NO3-, aspartate, glutamate, etc. They are conventionally called Clchannels because Clis the most abundant and physiological anions in organisms. While cation (K+, Na+, and Ca2+) channels have received most of attentions in the past 50 years, the role of Clchannels in the pacemaker activity of the heart has been largely ignored and understudied (Duan, 2009). The possible contribution of Clcurrents to spontaneous cardiac action potentials was first described in multicellular preparations of Purkinje fibers by Carmeliet (Carmeliet, 1961) and Hutter & Noble in 1961 (Hutter & Noble, 1961). Later, investigators from several laboratories observed the Cldependence of the diastolic depolarization and action potential duration. They characterized a hyperpolarizationactivated Clcurrent in multicellular preparations of rabbit sinoatrial node (SAN) (De Mello, 1963;Noma & Irisawa, 1976;Seyama, 1979). Unfortunately, studies in the late 1970s raised serious doubts about the existence and functional role of any type of Clchannels in the heart. The potential functional role of the hyperpolarization-activated Clcurrents in cardiac pacemaker activity has therefore been completely disputed (Hume et al., 2000;Duan et al., 2005). Recent studies have characterized the functional and molecular expression of several Clchannels in cardiac SAN cells from different species (Duan, 2009). These include 1) a hyperpolarizationand cell swelling-activated inwardly rectifying Clcurrent (ICl.ir)(Duan et al., 2000), which may be encoded by ClC-2, a member of the ClC voltage-gated Clchannel gene superfamily (Thiemann et al., 1992;Britton et al., 2000;Duan et al., 2000;Britton et al., 2005;Huang et al., 2009); 2) a volume-regulated outwardly-rectifying Clcurrent (ICl,vol)(Hagiwara et al., 1992), which may be encoded by another member of the ClC superfamily, ClC-3 (Duan et al., 1997b;Duan et al., 1999a;Britton et al., 2000); and 3) a Ca2+activated Clcurrent (ICl.Ca) (Verkerk et al., 2002), which may be encoded by TEME16A (or Ano1) (Schroeder et al., 2008;Caputo et al., 2008;Yang et al., 2008;Hartzell et al., 2009). Therefore, experimental evidence are merging to support the potential role of Clchannels in

The Functional Role of Chloride Channels in Cardiac Pacemaker Activity