Orbiliomycetes is one of the earliest diverging branches of the filamentous ascomycetes. The class contains nematode-trapping fungi that form unique infection structures, called traps, to capture and kill free-living nematodes. The traps have evolved differently along several lineages and include adhesive traps (knobs, nets or branches) and constricting rings. We show, by genome sequencing of the knob-forming species Monacrosporium haptotylum and comparison with the net-forming species Arthrobotrys oligospora, that two genomic mechanisms are likely to have been important for the adaptation to parasitism in these fungi. Firstly, the expansion of protein domain families and the large number of species-specific genes indicated that gene duplication followed by functional diversification had a major role in the evolution of the nematode-trapping fungi. Gene expression indicated that many of these genes are important for pathogenicity. Secondly, gene expression of orthologs between the two fungi during infection indicated that differential regulation was an important mechanism for the evolution of parasitism in nematode-trapping fungi. Many of the highly expressed and highly upregulated M. haptotylum transcripts during the early stages of nematode infection were species-specific and encoded small secreted proteins (SSPs) that were affected by repeat-induced point mutations (RIP). An active RIP mechanism was revealed by lack of repeats, dinucleotide bias in repeats and genes, low proportion of recent gene duplicates, and reduction of recent gene family expansions. The high expression and rapid divergence of SSPs indicate a striking similarity in the infection mechanisms of nematode-trapping fungi and plant and insect pathogens from the crown groups of the filamentous ascomycetes (Pezizomycotina). The patterns of gene family expansions in the nematode-trapping fungi were more similar to plant pathogens than to insect and animal pathogens. The observation of RIP activity in the Orbiliomycetes suggested that this mechanism was present early in the evolution of the filamentous ascomycetes.
Hydroponic experiments were conducted to investigate the role of exogenous silicon (Si) addition in increasing hexavalent chromium (Cr VI) tolerance in rice seedlings. Rice seedlings were grown under 100 lM Cr(VI) stress without or with 10 lM Si. Chromium treatment decreased growth, photosynthetic pigments and protein, which was accompanied by a significant increase in Cr accumulation and lipid peroxidation (as malondialdehyde; MDA). However, Si addition alleviated Cr toxicity and promoted growth of rice by decreasing Cr accumulation, root-to-shoot Cr transport and MDA level. Contents of macro (Mg, Ca and K) as well as micronutrients (Zn and Fe) were decreased by Cr except Mn while Si addition prevented decrease in these nutrients induced by Cr. Antioxidant capacity and total phenolic contents were decreased by Cr while these indices improved by Si addition. Treatment of Cr decreased the length of leaf epidermal cells and stomatal frequency, and adversely affected chloroplasts containing mesophyll cells and integrity of xylem and phloem, and Si addition minimized these abnormalities. However, frequency of root hairs was increased by Cr treatment. Results showed that exogenous Si addition enhanced Cr(VI) tolerance in rice seedlings by decreasing Cr accumulation, root-to-shoot Cr transport and MDA level, and by increasing content of some mineral elements (K, Fe and Zn) and antioxidant capacity compared to the Cr treatment alone.
For better understanding of silicon (Si)-mediated increase in metal tolerance in plants, the alterations in growth, mineral elements and anatomical features were investigated in rice seedlings exposed to Si (10 μM) and aluminium (Al; 50 μM). Al decreased seedling growth which was accompanied by an increase in Al accumulation. In contrast, addition of Si alleviated toxic effects of Al and led to the decrease in Al accumulation. Magnesium (Mg) and zinc (Zn) contents were decreased by Al; however, Si addition prevented decrease in Mg and Zn contents. Manganese (Mn) content was increased by Al while Si addition did not have a significant effect on Mn content under Al treatment. Al exposure decreased frequency of stomata and root hairs, length of root hairs and leaf epidermal cells, and distorted the structure and integrity of mesophyll cells and phloem; however, addition of Si reduced these abnormalities. Results showed that Si addition protected rice seedlings against Al toxicity by decreasing Al accumulation and by maintaining level of some mineral elements, and the key structures of leaf and root.
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