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Membrane currents and development of tension in atrial trabeculae from elephant seal hearts were studied using a single sucrose-gap voltage-clamp technique. A transient outward current (Ito) was observed with kinetics, voltage and beat dependence, similar to those of tension. Ito had a bell-shaped voltage dependence similar to that of tension and the slow inward current (Isi). Ito, unlike Isi, showed beat dependence quite similar to developed tension. Increases in [Ca]o, frequency of stimulation, and addition of adrenaline enhanced Ito and developed tension. Ito was suppressed by addition of Mn2+, tetracaine, or by depolarizing pre-pulses (to -40 mV for 250 ms). Caffeine at low concentrations (1 mM) blocked beat dependence of Ito. At higher concentrations (greater than 5 mM) caffeine suppressed the activation of Ito, phasic tension, and the second component of the birefringence signal (related to Ca2+-releasing activity of the sarcoplasmic reticulum (s.r.]. Similar to Isi phasic tension and Ito, the voltage dependence of the second component of the birefringence signal was bell-shaped. Our studies suggest that activation of Ito is related to triggered release of Ca2+ from the s.r. which generates the phasic tension. An excitation-contraction coupling scheme is presented which incorporates these findings and suggests that Ito may be responsible for shorter action potentials found in atrial fibres.