A bulb is a whole plant condensed into an underground organ. A geophyte's bulb comprises both food reserves and important developmental history that may affect its whole growth. In Easter lily (Lilium longiflorum), bulb size is associated with the plant's flowering pathwayvernalization or photoperiodand also affects sprouting, flower quality and abortion rate. The aim of this study was to investigate the reasons for the major physiological differences between large and small bulbs. Lily bulbs start their development from secondary meristems along the stem, with large bulbs being heavier and bear more scales than small ones. Peeling the outer scales of a large bulb converts its physiological responses into those of a small bulb, implying that the physiological discrepancies in plants developing from large or small bulbs are mediated by factors inherent to the bulb. We therefore performed broad analyses of the metabolite composition in the scales of bulbs subjected to temperature regimes affecting further plant development. We found a striking association between the level of glycerol, a primary metabolite mostly synthesized in the outer scales, and a delay in sprouting and flowering time, and reduction in abortion rate. Exogenous glycerol application to the bulbs before planting corroborated these results. Moreover, transcriptome analyses showed that flowering-promoting gene expression was downregulated in the bulb after glycerol treatment, while potential flowering inhibitor as well as a dormancy-related gene expressions were upregulated. Based on these studies, we postulate that glycerol is a major factor influencing both vegetative and reproductive development in lily.
At the reproductive stage, lily plants bear two morphological types of mature leaves, one at the lower and one at the upper part of the stem. At the vegetative stage, all the leaves are similar to each other and to the reproductive plant's lower leaves. This heterophylly has not yet been explored. In this study, we show that it is not a result of the plant's age but rather an outcome of floral induction. The induction appears as an on-going process, during which the meristem still produces leaves but progressively becomes committed to reproduction. This intermediate period lasts until the ultimate switch to flower primordia occurs. The leaves produced during floral induction, termed here as "inductive," appear at the upper part of the stem. Besides their typical higher stomata density, these leaves have a poly-layered palisade mesophyll, whose cells exhibit a unique morphology and contain more chlorophyll than leaves of vegetative plants. These leaves display higher carbon assimilation, soluble sugar production, and chloroplast-lipid accumulation. Accordingly, genes associated with stomata, chloroplast, and photosynthesis are upregulated in these leaves. Our results were obtained when floral induction was achieved either by vernalization or photoperiod signals, ruling out a mere environmental effect. We suggest that lily plants prepare themselves for the high-energy-demanding bloom by producing leaves with enhanced photosynthetic capacity, leading to an increase in soluble sugars. These novel findings introduce an adjacent affinity between photosynthesis and flowering and provide a nondestructive tool for identifying the plant's developmental stagevegetative or reproductive.
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