87% of embryos derived from homozygous female germline clones have nuclear division abnormalities during the syncytial blastoderm stage. The embryos often show asynchrony of nuclear division and irregular distribution of the nuclei. Chromosome defects are seen during mitosis, with a high frequency of lagging chromosomes and formation of chromosome bridges during anaphase. Interphase nuclei fused together or attached by chromosome bridges are also seen.

Brain neuroblasts of homozygous larvae have mitotic defects, showing chromosome congression defects during metaphase and chromosome lagging during anaphase.

Homozygous clones in the imaginal disc are significantly smaller than the twin-spot clones or are absent.

Females carrying homozygous germ-line clones lay a normal number of eggs for more than 15 days after eclosion. Egg chambers in these females (analysed 7-8 or 15 days after eclosion) show a normal determination of the oocyte, a normal condensed karyosome, a normal number of nurse cells, a normal fusome and produce eggs with a normal eggshell.

Homozygous clones in the follicle cells are significantly smaller and recovered less frequently than control clones.

san³/san⁴ animals pupate but die at the pupal stage.

san³/san⁴ third instar larvae have smaller brains than normal and extremely small imaginal discs.