Plant MAPK cascade performs a critical role in the regulation of plant immunity and disease resistance. Although the function of MAPK cascade in immunity regulation is partially conserved between different species, the mechanism varies in different host and pathogen combinations. To date, the MAPK cascade function of woody plants in the regulation of disease resistance has seldom been reported. Here, we present evidence to show that apple MdMAPKKK1 performed an important role in the regulation of apple resistance to Botryosphaeria dothidea, the causal agent of apple ring rot. B. dothidea infection leads to enhanced MdMAPKKK1 expression and MAPK cascade activation, indicating that the MAPK cascade is involved in the defense against B. dothidea. MdMAPKKK1 overexpression-induced pathogen-independent cell death. MdMAPKKK1 silencing decreases the resistance of apple calli and fruits to B. dothidea. Further analysis indicates that MdMAPKKK1 can bind MdBSK1 and is likely phosphorylated by it. The MdBSK1-mediated phosphorylation of MdMAPKKK1 is important for resistance to B. dothidea. These results collectively indicate that apple resistance to B. dothidea is regulated by the interaction between MAPKKK1 and MdBSK1.
Glomerella leaf spot (GLS), caused by the fungus Colletotrichum fructicola, is one of the most devastating apple diseases. Our previous study reported that the GLS resistance locus was defined on the chromosome 15 region. Here, we further found a single-nucleotide polymorphism (SNP) site (SNP7309212) in the GLS resistance that was able to distinguish resistant cultivars (lines) from susceptible ones. On the basis of the SNP site, we cloned a TNL gene from the GLS resistant locus and named it MdTNL1 (NCBI Accession Number: ON402514). This gene contains a toll/interleukin-1 receptor transmembrane domain (TIR), nucleotide-binding sites (NBS), and leucine-rich repeat (LRR) domain. Subcellular location indicated that MdTNL1 was expressed in the nucleus and cell membrane. Ectopic overexpression of MdTNL1 in Nicotiana benthamiana caused cell death. We further demonstrated allelic polymorphisms in MdTNL1. It is noteworthy that NBS and LRR domains of the MdTNL1 protein serve as the repository for generating allelic diversity. Quantitative real-time PCR (qRT-PCR) assay revealed that MdTNL1 was highly expressed in resistant apple cultivar ‘Fuji’ after inoculation with C. fructicola, whereas susceptible cultivar ‘Golden Delicious’ exhibited low expression after inoculation. Over-expression of MdTNL1-1 in susceptible apple fruits and leaves improved disease resistance, while in ‘Orin’ calli, silencing the MdTNL1-1 gene conversely decreased GLS resistance. In conclusion, we identified a GLS associated with SNP7309212 and demonstrated that a TIR-NBS-LRR gene MdTNL1-1 positively regulates GLS resistance in apple.
Ring rot is a destructive apple disease caused by Botryosphaeria dothidea. The resistance mechanism of apple plants to B. dothidea remains unclear. Here, we show that apple vacuolar processing enzyme 4 (MdVPE4) is involved in fruit resistance to B. dothidea. MdVPE4 silencing reduced fruit resistance, whereas its overexpression improved fruit resistance. Gene expression analysis revealed that MdVPE4 influenced the expression of fruit resistance-related genes, such as apple polygalacturonase 1 (MdPG1), apple polygalacturonase inhibitor protein 1 (MdPGIP1), apple endochitinase 1 (MdCHI1), and apple thaumatin-like protein 1 (MdTHA1). The expression of the four genes responding to B. dothidea infection decreased in MdVPE4-silenced fruits. Further analysis demonstrated that B. dothidea infection induced MdVPE4 expression and enzyme activation in apple fruits. Moreover, MdVPE4 activity was modulated by apple cysteine proteinase inhibitor 1 (MdCPI1), which also contributed to fruit resistance to B. dothidea as revealed by gene overexpression and silencing analysis. MdCPI1 interacted with MdVPE4 and inhibited its activity. However, MdCPI1 expression was decreased by B. dothidea infection. Taken together, our findings indicate that the interaction of MdVPE4 and MdCPI1 plays an important role in modulating fruit resistance to B. dothidea.
Titanium alloy has been widely used in Marine pipeline system because of its excellent corrosion resistance. However, there are differences in microstructure and electrochemical properties because of the heterogeneous structure of the welded joint, the corrosion behavior is often different. In this paper, the corrosion behavior of TA2 titanium alloy welded joint in seawater at different temperatures was studied by traditional macro electrochemical test analysis combined with microelectrode array test and surface morphology analysis. Conventional macroscopic electrochemical analysis results show that the corrosion resistance of heat-affected zone is always the best, followed by the base metal and the weld. And the higher the temperature, the easier the formation of passivation film. The results of microelectrode array test show that the heat-affected zone is always the cathode region of the whole welded joint, and part of the cathode near the base metal region has the largest current density, which acts as the main cathode to slow down corrosion. At slightly higher temperatures, the polarity deflection will occur in the base metal zone and weld zone due to the different formation speeds of passivation film in early corrosion stage. With the prolongation of corrosion time, the base metal eventually becomes the cathode zone and the weld zone eventually becomes the anode zone.
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