Welcome to a new kind of tension: translating kinetochore mechanics into a wait-anaphase signal

Recent high-resolution studies of kinetochore structure have transformed the way researchers think about this crucial macro-molecular complex, which is essential for ensuring chromosome segregation occurs faithfully during cell division. Kinetochores mediate the interaction between chromosomes and the plus-ends of dynamic spindle microtubules and control the timing of anaphase onset by regulating the spindle assembly checkpoint (SAC). There is much debate in the SAC research community as to whether mitotic cells sense only microtubule attachment at the kinetochore, or both attachment and tension, before committing to anaphase. In this Commentary, we present a brief history of the tension-versus-attachment debate, summarize recent advances in our understanding of kinetochore structure and focus on the implications of a phenomenon known as intrakinetochore stretch for SAC regulation. We also hypothesize how intrakinetochore stretch might impact SAC function by regulating both microtubule attachment stability and the localization and activity of checkpoint components at the kinetochore.