Shaping plastid stromules - principles of in vitro membrane tubulation applied in planta

Plastids undergo drastic shape changes under stress, including the formation of stroma-filled tubules, or `stromules'. Stromules are dynamic, and may extend, branch and retract within minutes. There are two prerequisites for stromule extension: excess plastid membrane and a force(s) that shapes the membrane into a tubule. In vitro studies provide insight into the basic molecular machinery for tubulation, and are often cited when discussing stromule formation. In this review, we evaluate in vitro modes of tubulation in the context of stromule dynamics, and find that most mechanisms fail to explain stromule morphology and behavior observed in planta. Current data support a model of stromule formation relying on pulling motors (myosins and kinesins) and cytoskeleton (actin and microtubules).