Stability of the determined state

During development parts of plants become determined for specific fates which can persist in populations of dividing cells. Students of plant development have tended to favour the view that the determined state is stabilized at the supracellular level. We provide evidence for the alternative view that determination can be inherited at the cellular level. This conclusion is based on the observation that cultured cells derived from the leaf lamina and cortex of the stem of tobacco plants inherit different states of cytokinin requirement. Plant regeneration experiments show that the cytokinin-requiring (C-) state characteristic of leaf cells and the cytokinin autotrophic (C+) state characteristic of cortex cells are stable but not permanent. Progeny of both cell types can give rise to complete plants with tissues exhibiting the cytokinin requirement of comparable tissues of seed grown plants. Pith cells can shift from the C- to the C+ state by a process known as habituation. Evidence is presented that this epigenetic change is stabilized by a positive-feedback mechanism in which cytokinins appear to promote their own biosynthesis. A single dominant, Mendelian gene, H1-2, controls expression of the C- and C+ phenotypes. There are hints that this gene changes state in cultured cells at rates expected for epigenetic modifications. This suggests that the C- and C+ state can also be stabilized by genetic mechanisms.