In mmm^MB04232 homozygous males, mature sperm fail to be loaded into the seminal vesicles (in 14 days old adults). There fewer and more abnormal actin cones. Immediately after meiosis there is increased mitochondrial aggregation. Onion stage spermatids show nuclei that are enlarged and of heterogeneous size. Elongation stage spermatids show loss of mitochondrial integrity: the major mitochondrial derivative appears normal but the corresponding minor derivative is small and occasionally absent; while cells that lack the minor derivative only have the ~14 accessory microtubules surrounding the major derivative, cells with both derivatives have an increased number of accessory microtubules, leading to an overall increase in the number of accessory microtubules.