The consequences of a non-uniform tension across kinetochores: lessons from segregation of chromosomes in the permanent translocation heterozygote Oenothera.

The alternate (zigzag) configuration of the chromosome ring in oenotheras fulfills the requirement of high tension across kinetochores for stability of the configuration and the progression to anaphase. However, also semialternate configurations (two pairs of adjacent kinetochores interspaced among the zigzag) fulfill the requirement of high tension across kinetochores. If only the magnitude of tensile force acting on a kinetochore pair governs the stability of microtubule attachments, the probability of occurrence of the semialternate configurations would be higher than that of fully alternate configurations. Yet the percentage of irregularity in the zigzag configuration is surprisingly low, which means that the semialternate configurations are corrected. The only difference which distinguishes the fully alternate and the semialternate configurations with respect to the tension across kinetochores is that the tension across a kinetochore alternating with its neighbors is rather uniformly distributed over the kinetochore, while there is a gradient of the tension in the kinetochore having a non-alternating neighbor, with low tension on the side of this neighbor. Apparently, a low tension across a part of a kinetochore brings about correction of its attachment to microtubules. This hypothesis fits with the repeat subunit model of the kinetochore; apparently, each subunit can function autonomously in the tension-governed mechanisms, stabilizing its attachment and controlling the metaphase-to-anaphase transition.