A tubular system of sarcoplasmic reticulum that is not penetrated by extracellular markers is described in vertebrate smooth muscles. The sarcoplasmic reticulum forms fenestrations around the surface vesicles and also forms close appositions (an approximately 10 to 12 nm gap traversed by periodic electron dense materia) with the non-specialized surface membrane. The morphological couplings are considered to be the most probable sites of electromechanical coupling of the action potential to the twitch contraction. The relative volume of the sarcoplasmic reticulum varies in functionally different (tonic and phasic) smooth muscles, and correlates with the ability of the different smooth muscles to contract in the absence of extracellular calcium. Electron opaque deposits of strontium are accumulated by peripheral and central elements of the sarcoplasmic reticulum. The accumulation of strontium and barium by mitochondria raises the possibility that, in addition to the sarcoplasmic reticulum, mitochondria may play a role in the regulation of intracellular divalent cation levels in vertebrate smooth muscle.