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ON THE NATURE OF THINGS: ESSAYS AMERICAN JOURNAL OF BOTANY expression of mitochondrial-and plastid-targeted genes in the nucleus, upregulated cytoplasmic genome copy number, elevated levels of organelle biogenesis) as well as over longer timescales (eg, subfunctionalization, pseudogenization)(Fig. 1). One indication that polyploidy has substantial stoichiometric consequences for cytonuclear interactions is that organelle-targeted genes appear to be one of the first and most overrepresented classes of nuclear genes to return to single copy following a WGD (De Smet et al., 2013), although there may be considerable variation in sensitivity to gene dosage imbalance across different functional pathways within the organelles (Coate et al., 2011). Additionally, the well-known positive relationship between nuclear genome size and cell size (Beaulieu et al., 2008) indicates that plants may be capable …