The INDETERMINATE1 gene, ID1, encodes a putative transcription factor that plays an important role in regulating the transition to flowering in maize. Mutant id1 plants have a prolonged vegetative growth phase and fail to make normal flowers. The ID1 gene, which encodes a nuclear-localized zinc finger protein, is expressed exclusively in immature leaves, suggesting that ID1 regulates a leaf-derived floral inductive signal. It is shown by western analysis with anti-ID1-specific antibody that ID1 co-localizes with ID1 mRNA in developing, immature leaves and, similarly, is absent in mature, photosynthetically active leaf blades, as well as the shoot apical meristem. Immunolocalization with anti-ID1 antibody shows that ID1 protein is detected in the nuclei of all cell types in immature leaves. Examination of plants grown in different day/night cycles revealed that ID1 gene expression and protein levels are largely unaffected by variations in light and dark, and that mRNA and protein levels do not follow a circadian pattern. The absence of ID1 expression in greening leaf tips coincides with the sink-to-source transition of developing leaves. It was found that ID1 levels are down-regulated in mature albino leaves similarly as in normal green leaves, suggesting that ID1 activity is controlled developmentally and is not affected by the sink/source status of the leaf or the inability of a mature leaf to engage in photosynthesis. The finding that ID1 expression is developmentally regulated and is unperturbed by external stimuli such as photoperiod supports the supposition that ID1 acts through the autonomous floral inductive pathway in maize.
The INDETERMINATE1 (ID1) transcription factor is a key regulator of the transition to flowering in maize. ID1 is expressed in immature leaves where it controls the production or transmission of leaf-derived florigenic signals. Loss-of-function id1 mutants make many more leaves than normal plants and produce aberrant flowers; however, they exhibit no obvious developmental defects in early growth stages. A maize oligonucleotide microarray was used to assess the molecular differences between immature leaves of wild-type and id1 mutant plants prior to the floral transition. This analysis revealed 55 genes with a significant 2-fold difference in expression; 22 are down-regulated and 33 are up-regulated in id1 mutants. Most prominent is a novel family of three β-glucosidase genes that are most closely related to sorghum dhurrinases. These genes, termed Zmdhr1, Zmdhr2, and Zmdhr3, are undetectable in immature leaves of id1 mutants and are expressed exclusively in normal immature leaves in a pattern identical to the ID1 gene. Other down-regulated genes include a group of four zinc finger protein-encoding genes that are unrelated to ID1. A significant number of genes up-regulated in id1 mutant immature leaves have potential roles in photosynthesis and carbon fixation, substantiating a possible connection between floral induction and assimilate partitioning. Finally, expression of these genes was compared in florally induced versus uninduced teosinte, a photoperiod-sensitive progenitor of day-neutral maize. Only a few genes showed expression differences, suggesting that ID1 acts in a novel autonomous floral induction pathway that is distinct from the photoperiod induction pathway.
Little is known about the nature of floral inductive cues in day-neutral plants that are insensitive to photoperiod variations and, therefore, rely on endogenous signals to initiate reproductive growth. The INDETERMINATE1 (ID1) transcription factor is a key regulator of the transition to flowering in day-neutral maize. The ID1 gene is expressed exclusively in developing leaves, where it controls the production or transmission of leaf-derived florigenic signals. Florigen-producing source leaves were compared with mature leaves of late-flowering id1 plants, and metabolite and gene expression differences associated with the floral transition in maize were observed. While id1 mutants have a similar capacity for photosynthesis to wild-type siblings, id1 source leaves show quantitative differences in carbohydrate allocation prior to the floral transition stage, with a marked increase in sucrose and other soluble sugars, accompanied by a decrease in tricarboxylic acid (TCA) cycle organic acids. Importantly, source leaves of autonomous-flowering maize are typified by a higher transitory starch to sucrose ratio and a transcript profile enriched for sucrose synthesis and starch metabolism-related gene function. Finally, similar changes in transitory starch and sucrose are not observed in teosinte, the tropical progenitor of maize that requires short-day photoperiods to induce flowering. Together, these data define a transcript and metabolite signature associated with the autonomous floral transition in temperate maize leaves.
Agriculturally important grasses such as rice, maize and sugarcane are evolutionarily distant from Arabidopsis, yet some components of the floral induction process are highly conserved. Flowering in sugarcane is an important factor that negatively affects cane yield and reduces sugar/ethanol production from this important perennial bioenergy crop. Comparative studies have facilitated the identification and characterization of putative orthologs of key flowering time genes in sugarcane, a complex polyploid plant whose genome has yet to be sequenced completely. Using this approach we identified phosphatidylethanolamine-binding protein (PEBP) gene family members in sugarcane that are similar to the archetypical FT and TFL1 genes of Arabidopsis that play an essential role in controlling the transition from vegetative to reproductive growth. Expression analysis of ScTFL1, which falls into the TFL1-clade of floral repressors, showed transcripts in developing leaves surrounding the shoot apex but not at the apex itself. ScFT1 was detected in immature leaves and apical regions of vegetatively growing plants and, after the floral transition, expression also occurred in mature leaves. Ectopic over-expression of ScTFL1 in Arabidopsis caused delayed flowering in Arabidopsis, as might be expected for a gene related to TFL1. In addition, lines with the latest flowering phenotype exhibited aerial rosette formation. Unexpectedly, over-expression of ScFT1, which has greatest similarity to the florigen-encoding FT, also caused a delay in flowering. This preliminary analysis of divergent sugarcane FT and TFL1 gene family members from Saccharum spp. suggests that their expression patterns and roles in the floral transition has diverged from the predicted role of similar PEBP family members.