A Functionally Specialized Alpha-tubulin is Required for Oocyte Meiosis and Cleavage Mitoses in Drosophila

Three alpha-tubulin proteins contribute to microtubules during oogenesis and early embryogenesis in Drosophila melanogaster: alpha TUB84B, alpha TUB84D, and alpha TUB67C. alpha TUB67C is unique in two respects. It is a structurally divergent alpha-tubulin, sharing only 67% amino acid identity with the generic isotypes alpha TUB84B and alpha TUB84D, and its expression is exclusively maternal. The genetic analysis of the alpha Tub67C gene described here demonstrates that alpha TUB67C is required for nuclear division in the oocyte and early embryo. Both meiosis and cleavage-stage mitoses are severely affected by mutations that result in a substantial decrease in the ratio of alpha TUB67C/alpha TUB84B+alpha TUB84D. A large increase in this ratio, achieved by increasing the gene dosage of alpha Tub67C, has little or no effect on meiosis, but severely disrupts mitotic spindle function. Thus, both classes of alpha-tubulin isotype present in the mature oocyte, alpha TUB67C and alpha TUB84B/84D, are essential for normal spindle function in early Drosophila development. These alpha-tubulins provide the first example of tubulin isotypes known to be coexpressed in wild-type animals whose encoded variation is required for the normal function of a microtubule array.