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Background
Much of the organismal variation we observe in nature is due to differences in organ size. The observation that even closely related species can show large, stably inherited differences in organ size indicates a strong genetic component to the control of organ size. Despite recent progress in identifying factors controlling organ growth in plants, our overall understanding of this process remains limited, partly because the individual factors have not yet been connected into larger regulatory pathways or networks. To begin addressing this aim, we have studied the upstream regulation of expression of BIG BROTHER (BB), a central growth-control gene in Arabidopsis thaliana that prevents overgrowth of organs. Final organ size and BB expression levels are tightly correlated, implying the need for precise control of its expression. BB expression mirrors proliferative activity, yet the gene functions to limit proliferation, suggesting that it acts in an incoherent feedforward loop downstream of growth activators to prevent over-proliferation.
Results
To investigate the upstream regulation of BB we combined a promoter deletion analysis with a phylogenetic footprinting approach. We were able to narrow down important, highly conserved, cis-regulatory elements within the BB promoter. Promoter sequences of other Brassicaceae species were able to partially complement the A. thaliana bb-1 mutant, suggesting that at least within the Brassicaceae family the regulatory pathways are conserved.
Conclusions
This work underlines the complexity involved in precise quantitative control of gene expression and lays the foundation for identifying important upstream regulators that determine BB expression levels and thus final organ size.
Long and short-term climatic variation affect the ability of plants to simultaneously cope with increasing abiotic stress and biotic interactions. Specifically, ecotypes adapted to different climatic conditions (i.e., long-term legacy) may have to adjust their allocation to chemical defenses against enemies under acute drought (i.e., short-term response). Although several studies have addressed drought effects on chemical defense production, little is known about their intraspecific variation along resource gradients. Studying intraspecific variation is important for understanding how different environments select for defense strategies and how these may be affected directly and indirectly by changing climatic conditions. We conducted greenhouse experiments with the annual Biscutella didyma (Brassicaceae) to test the effects of long-term climatic legacy versus short-term drought stress on the concentrations of defense compounds (glucosinolates). To this aim, four ecotypes originating from a steep aridity gradient were exposed to contrasting water treatments. Concentrations of chemical defenses were measured separately in leaves of young (8 weeks) and old (14 weeks) plants, respectively. For young plants, ecotypes from the wettest climate (long-term legacy) as well as plants receiving high water treatments (short-term response) were better defended. A marginally significant interaction suggested that wetter ecotypes experienced a larger shift in defense production across water treatments. Older plants contained much lower glucosinolate concentrations and showed no differences between ecotypes and water treatments. Our results indicate that younger plants invest more resources into chemical defenses, possibly due to higher vulnerability to tissue loss compared to older plants. We propose that the strong response of wet ecotypes to water availability may be explained by a less pronounced adaptation to drought.