Second thoughts on septation by the fission yeast, Schizosaccharomyces pombe: pull vs. push mechanisms with an appendix--dimensional modelling of the flat and variable septa.

The correlation of contraction by an actomyosin band with the closing of the septum of dividing cells of the fission yeast, Schizosaccharomyces pombe, cannot suggest cause-and-effect because contraction would be apparent whether the membrane enveloping the centripetally closing septum were pulled or were pushed. Thus the common observation of contraction is not critical. Diagrams of published electron micrographs of dividing wild-type fission yeasts illustrate variable (tilted) septal images that are counterintuitive to a pull model. Circumference calculations based on those images suggest that some variable forms might be only 6% closed even though their two-dimensional profiles would be 50% closed, if they were not tilted. Development of multiseptate forms of cdc4-8 and cdc4-377 temperature sensitive mutants incubated at their restrictive temperature was followed. These multiseptate forms are shown to have functional (functional in terms of generating divided uninucleate cytoplasts) but grotesque septa which are formed in the absence of actomyosin bands. By contrast, the myosin of the plant phragmoplast is not properly oriented for contractility, and Dictyostelium (attached cells) and Saccharomyces (mutants) have been shown to divide in the absence of myosin II, just as S. pombe does (above). Hence contractility, the essence of a pull model for septum closure, would seem to be non-essential. Other, non-contractile mechanisms of myosin are emphasized, and a push model becomes a rational default hypothesis. The essence of push models is that their synthesis/assembly mechanisms are driving force sufficient for septum closure.