Drought stress tolerance is a complex trait regulated by multiple factors. Here, we demonstrate that the miRNA160 - Auxin Response Factor 17 - HYPONASTIC LEAVES 1 (miR160 - ARF17 - HYL1) module is crucial for apple (Malus ⅹ domestica) drought tolerance. Using stable transgenic plants, we found that drought tolerance was improved by higher levels of Mdm-miR160 or MdHYL1 and by decreased levels of MdARF17, whereas reductions in MdHYL1 or increases in MdARF17 led to greater drought sensitivity. Further study revealed that modulation of drought tolerance was achieved through regulation of drought-responsive miRNA levels by MdARF17 and MdHYL1; MdARF17 interacted with MdHYL1 and bound to the promoter of MdHYL1. Genetic analysis further suggested that MdHYL1 is a direct downstream target of MdARF17. Importantly, MdARF17 and MdHYL1 regulated the abundance of Mdm-miR160. In addition, the Mdm-miR160-MdARF17-MdHYL1 module regulated adventitious root development. We also found that Mdm-miR160 can move from the scion to the rootstock in apple and tomato (Solanum lycopersicum), thereby improving root development and drought tolerance of the rootstock. Our study revealed the mechanisms by which the positive feedback loop of Mdm-miR160-MdARF17-MdHYL1 influences apple drought tolerance.
Carbon dioxide concentration (CO2) and light intensity are known to play important roles in plant growth and carbon assimilation. Nevertheless, the underlying physiological mechanisms have not yet been fully explored. Tomato seedlings (Solanum lycopersicum Mill. cv. Jingpeng No. 1) were exposed to two levels of CO2 and three levels of light intensity and the effects on growth, leaf gas exchange and water use efficiency were investigated. Elevated CO2 and increased light intensity promoted growth, dry matter accumulation and pigment concentration and together the seedling health index. Elevated CO2 had no significant effect on leaf nitrogen content but did significantly upregulate Calvin cycle enzyme activity. Increased CO2 and light intensity promoted photosynthesis, both on a leaf‐area basis and on a chlorophyll basis. Increased CO2 also increased light‐saturated maximum photosynthetic rate, apparent quantum efficiency and carboxylation efficiency and, together with increased light intensity, it raised photosynthetic capacity. However, increased CO2 reduced transpiration and water consumption across different levels of light intensity, thus significantly increasing both leaf‐level and plant‐level water use efficiency. Among the range of treatments imposed, the combination of increased CO2 (800 µmol CO2 mol−1) and high light intensity (400 µmol m−2 s−1) resulted in optimal growth and carbon assimilation. We conclude that the combination of increased CO2 and increased light intensity worked synergistically to promote growth, photosynthetic capacity and water use efficiency by upregulation of pigment concentration, Calvin cycle enzyme activity, light energy use and CO2 fixation. Increased CO2 also lowered transpiration and hence water usage.
Summary Drought limits apple yield and fruit quality. However, the molecular mechanism of apple in response to drought is not well known. Here, we report a Cys2/His2 (C2H2)‐type zinc‐finger protein, MdZAT5, that positively regulates apple drought tolerance by regulating drought‐responsive RNAs and microRNAs (miRNAs). DNA affinity purification and sequencing and yeast‐one hybrid analysis identified the binding motifs of MdZAT5, T/ACACT/AC/A/G. Chromatin immunoprecipitation quantitative polymerase chain reaction (ChIP‐qPCR) and electrophoretic mobility shift assays (EMSAs) showed that MdZAT5 directly binds to the promoters of the drought‐responsive genes including MdRHA2a, MdLEA14, MdTPX1, and MdCAT3, and activates their expression under drought stress. MdZAT5 interacts with and directly targets HYPONASTIC LEAVES1 (MdHYL1). MdZAT5 may facilitate the interaction of MdHYL1 with pri‐miRNAs or MdDCL1 by activating MdHYL1 expression, thereby regulating the biogenesis of drought‐responsive miRNAs. Genetic dissection showed that MdHYL1 is essential for MdZAT5‐mediated drought tolerance and miRNA biogenesis. In addition, ChIP‐qPCR and EMSA revealed that MdZAT5 binds directly to the promoters of some MIR genes including Mdm‐miR171i and Mdm‐miR172c, and modulates their transcription. Taken together, our findings improve our understanding of the molecular mechanisms of drought response in apple and provide a candidate gene for the breeding of drought‐tolerant cultivars.
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