A genetic system for alternative stable characteristics in genomically identical homozygous clones

Paramecium tetraurelia, stock d113, although completely homozygous, produces two kinds of genomically identical clones: N (nondischarge) clones incapable of trichocyst exocytosis (discharge) from intact cells in response to picric acid; and D (discharge) clones that do respond. These alternatives are irreversibly determined (at 27°C) during a determination sensitive period the first day after fertilization (autogamy, conjugation, or cytogamy): D parents are always determined to produce D progeny; N parents produce mostly N progeny if kept in exhausted medium, but mostly D progeny if kept in bacterized nutrient medium, throughout the sensitive period. If connecting bridges between mates persist long after the time for pair separation, the N member of N×D conjugant and cytogamous pairs produces D progeny even if exposed to exhausted medium throughout the sensitive period, thus indicating the presence in D mates of a D‐determining cytoplasmic factor, δ, which overrides effects of external conditions. N and D determinations are brought about on newly developing somatic nuclei (macronuclear anlagen). After macronuclear development has been completed, determination is irreversible in it and its descendant macronuclei. M (mixed) clones produce N, D, and partial D cells; within these clones, diverse subclones can be selected. Crosses of d113 (N)×standard wild stock 51 (D) yield no segregation in F2, indicating no genomic difference between d113 (N) and wild type (D), δ may be a genic product regulating its own production. This results in “cytoplasmic inheritance” of D vs N in crosses of D×N followed by exhausted medium during the sensitive period. As with the only other well‐analyzed comparable example, mating types, neither a genetic nor an epigenetic interpretation has yet been excluded for this system of developmental differentiation.