Tonic and phasic activity in smooth muscle

We have studied the electrical and mechanical behaviour of two very different smooth muscle preparations, mesenteric lymphatic ducts and proximal urethra. These tissues generate different patterns of spontaneous contraction adapted to fulfil their contrasting functions. While lymphatics undergo regular phasic contractions and relaxations, suited to their role in propelling lymph, the urethra remains in a state of contracture to maintain urinary continence. The challenge is to understand how both of these achieve their respective roles. Interestingly, electrical activity of lymphatics resembles that in the heart in having a one to one relationship between the action potential and phasic contraction. Patch clamp studies have shown that lymphatic cells express 3 ionic currents that are not present in urethral cells, but are shared with cardiac muscle. These are, i) fast Na+ current, ii) T-type Ca2+ current and iii) a hyperpolarization-activated cation current, Ir. The fast Na+ current is ideally suited to the propagation of the action potential over large distances, as required by a vessel capable of generating a rapid well co-ordinated contraction along its length. The T-current and Ir, on the other hand, appear to be involved in electrical pacemaking as they are in the heart. The urethra does not usually undergo regular phasic contractions and it lacks these currents. Instead, urethral tone may depend on an interaction between L-type Ca2+ current and a large Ca(2+)-activated Cl- current. Activation of Cl- channels (perhaps by spontaneous release of Ca2+ from intracellular stores) would depolarize the membrane potential to within the 'window current' range for L-type Ca2+ channels and result in Ca2+ influx and contraction. This process may be maintained for a time by positive feedback whereby the influx of Ca2+ continues to activate the Cl- channels.