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Development of the mutualistic arbuscular mycorrhiza (AM) symbiosis between most land plants and fungi of the Glomeromycota is regulated by phytohormones. The role of jasmonate (JA) in AM colonization has been investigated in the dicotyledons Medicago truncatula, tomato and Nicotiana attenuata and contradicting results have been obtained with respect to a neutral, promotive or inhibitory effect of JA on AM colonization. Furthermore, it is currently unknown whether JA plays a role in AM colonization of monocotyledonous roots. Therefore we examined whether JA biosynthesis is required for AM colonization of the monocot rice. To this end we employed the rice mutant constitutive photomorphogenesis 2 (cpm2), which is deficient in JA biosynthesis. Through a time course experiment the amount and morphology of fungal colonization did not differ between wild-type and cpm2 roots. Furthermore, no significant difference in the expression of AM marker genes was detected between wild type and cpm2. However, treatment of wild-type roots with 50 μM JA lead to a decrease of AM colonization and this was correlated with induction of the defense gene PR4. These results indicate that JA is not required for AM colonization of rice but high levels of JA in the roots suppress AM development likely through the induction of defense.

Abstract An experiment was performed to investigate the effect of mycorrhizal symbiosis and foliar application of salicylic acid on quantitative and qualitative traits of maize during 2018 and 2019 in the research farm of Islamic Azad University, Chalous Branch. Split plot in a randomized complete block design with three replications was used. Experimental factors included mycorrhiza species of (G. mosseae), (G. geosporum) and (G. intraradices) at two levels (no consumption and consumption of mycorrhiza) and salicylic acid at two levels (no consumption and consumption of 1 mμ of salicylic acid). Results of interaction effects of mycorrhiza and salicylic acid on the measured traits revealed that the maximum 1000-grain weight, grain yield, biological yield, phosphorus, potassium, nitrogen percentage and yield of maize grain protein were observed in G. mosseae treatment under foliar application of salicylic acid. Foliar application of salicylic acid increases the root length and provides the necessary conditions for increasing water and nutrient uptake alongwith increase in photosynthesis and thus allocates more photosynthetic substance for development of reproductive organs. Hence, it increases maize grain weight and accordingly grain yield. In general, the results revealed that mycorrhiza and foliar application of salicylic acid increase growth indicators, yield and yield components. It also improved the quality traits of the maize plant. Based on results, the interaction effect of G. mosseae treatment and foliar application of salicylic acid yielded better results than other treatments. Mycorrhiza increases the number of grain in the ear, the number of rows in the ear, increases the plant's ability to absorb phosphorus, and the increase of mycorrhiza along with salicylic acid shows the maximum grain yield in maize. Finally, it can be concluded that the use of mycorrhiza and salicylic acid can be effective in increasing grain in the plant. Resumo Um experimento foi realizado para investigar o efeito da simbiose micorrízica e aplicação foliar de ácido salicílico em características quantitativas e qualitativas do milho durante 2018 e 2019 na fazenda de pesquisa da Universidade Islâmica Azad, Chalous Branch. Foi usada uma parcela dividida em um delineamento de blocos casualizados com três repetições. Os fatores experimentais incluíram espécies de micorrizas (G. mosseae, G. geosporum e G. intraradices) em dois níveis (sem consumo e com consumo de micorrizas) e ácido salicílico em dois níveis (sem consumo e com consumo de 1 mμ de ácido salicílico). Os resultados dos efeitos da interação de micorriza e ácido salicílico nas características medidas revelaram que peso máximo de 1.000 grãos, rendimento de grãos, rendimento biológico, fósforo, potássio, porcentagem de nitrogênio e rendimento de proteína de grão de milho foram observados no tratamento G. mosseae sob aplicação foliar de ácido salicílico. A aplicação foliar de ácido salicílico aumenta o comprimento da raiz e fornece as condições necessárias para aumentar a absorção de água e nutrientes juntamente com o aumento da fotossíntese e, assim, aloca mais substância fotossintética para o desenvolvimento dos órgãos reprodutivos. Assim, aumenta o peso do grão de milho e, consequentemente, o rendimento de grãos. Em geral, os resultados revelaram que a micorriza e a aplicação foliar de ácido salicílico aumentam os indicadores de crescimento, rendimento e componentes do rendimento. Também melhoram as características de qualidade da planta de milho. Com base nos resultados, o efeito de interação do tratamento G. mosseae e aplicação foliar de ácido salicílico produziu melhores resultados do que outros tratamentos. A micorriza aumenta o número de grãos na espiga, o número de fileiras na espiga e a capacidade da planta de absorver fósforo, e o aumento da micorriza junto com o ácido salicílico mostra o rendimento máximo de grãos no milho. Por fim, pode-se concluir que o uso de micorriza e ácido salicílico pode ser eficaz no incremento de grãos na planta.

Arbuscular mycorrhizal (AM) fungi facilitate plant uptake of mineral nutrients and draw organic nutrients fromthe plant. Organic nutrients are thought to be supplied primarily in the formof sugars. Here we show that the AM fungus Rhizophagus irregularis is a fatty acid auxotroph and that fatty acids synthesized in the host plants are transferred to the fungus to sustain mycorrhizal colonization. The transfer is dependent on RAM2 (REQUIRED FOR ARBUSCULAR MYCORRHIZATION 2) and the ATP binding cassette transporter–mediated plant lipid export pathway. We further show that plant fatty acids can be transferred to the pathogenic fungus Golovinomyces cichoracerum and are required for colonization by pathogens. We suggest that themutualistic mycorrhizal and pathogenic fungi similarly recruit the fatty acid biosynthesis program to facilitate host invasion.

Key message This article describes the composition of root exudates, how these metabolites are released to the rhizosphere and their importance in the recruitment of benefcial microbiota that alleviate plant stress. Abstract Metabolites secreted to the rhizosphere by roots are involved in several processes. By modulating the composition of the root exudates, plants can modify soil properties to adapt and ensure their survival under adverse conditions. They use several strategies such as (1) changing soil pH to solubilize nutrients into assimilable forms, (2) chelating toxic compounds, (3) attracting benefcial microbiota, or (4) releasing toxic substances for pathogens, etc. In this work, the composition of root exudates as well as the diferent mechanisms of root exudation have been reviewed. Existing methodologies to collect root exudates, indicating their advantages and disadvantages, are also described. Factors afecting root exudation have been exposed, including physical, chemical, and biological agents which can produce qualitative and quantitative changes in exudate composition. Finally, since root exudates play an important role in the recruitment of mycorrhizal fungi and plant growth-promoting rhizobacteria (PGPR), the mechanisms of interaction between plants and the benefcial microbiota have been highlighted.

Arbuscular mycorrhiza (AM) symbioses contribute to global carbon cycles as plant hosts divert up to 20% of photosynthate to the obligate biotrophic fungi. Previous studies suggested carbohydrates as the only form of carbon transferred to the fungi. However, de novo fatty acid (FA) synthesis has not been observed in AM fungi in absence of the plant. In a forward genetic approach, we identified two Lotus japonicus mutants defective in AM-specific paralogs of lipid biosynthesis genes (KASI and GPAT6). These mutants perturb fungal development and accumulation of emblematic fungal 16:1ω5 FAs. Using isotopolog profiling we demonstrate that 13C patterns of fungal FAs recapitulate those of wild-type hosts, indicating cross-kingdom lipid transfer from plants to fungi. This transfer of labelled FAs was not observed for the AM-specific lipid biosynthesis mutants. Thus, growth and development of beneficial AM fungi is not only fueled by sugars but depends on lipid transfer from plant hosts. Most land plants are able to form partnerships with certain fungi – known as arbuscular mycorrhiza fungi – that live in the soil. These fungi supply the plant with mineral nutrients, especially phosphate and nitrogen, in return for receiving carbon-based food from the plant. To exchange nutrients, the fungi grow into the roots of the plant and form highly branched structures known as arbuscules inside plant cells. Due to the difficulties of studying this partnership, it has long been believed that plants only provide sugars to the fungus. However, it has recently been discovered that these fungi lack important genes required to make molecules known as fatty acids. Fatty acids are needed to make larger fat molecules that, among other things, store energy for the organism and form the membranes that surround each of its cells. Therefore, these results raise the possibility that the plant may provide the fungus with some of the fatty acids the fungus needs to grow. Keymer, Pimprikar et al. studied how arbuscules form in a plant known as Lotus japonicus, a close relative of peas and beans. The experiments identified a set of mutant L. japonicus plants that had problems forming arbuscules. These plants had mutations in several genes involved in fat production that are only active in plant cells containing arbuscules. Further experiments revealed that certain fat molecules that are found in fungi, but not plants, were present at much lower levels in samples from mutant plants colonized with the fungus, compared to samples from normal plants colonized with the fungus. This suggests that the fungi colonizing the mutant plants may be starved of fat molecules. Using a technique called stable isotope labelling it was possible to show that fatty acids made in normal plants can move into the colonizing fungus. The findings of Keymer, Pimprikar et al. provide evidence that the plant feeds the fungus not only with sugars but also with fat molecules. The next challenge will be to find out exactly how the fat molecules are transferred from the plant cell to the fungus. Many crop plants are able to form partnerships with arbuscular mycorrhizal fungi. Therefore, a better understanding of the role of fat molecules in these relationships may help to breed crop plants that, by providing more support to their fungal partner, may grow better in the field.

Soil microbes comprise a large portion of the genetic diversity on Earth and influence a large number of important ecosystem processes. Increasing atmospheric nitrogen (N) deposition represents a major global change driver; however, it is still debated whether the impacts of N deposition on soil microbial biomass and respiration are ecosystem-type dependent. Moreover, the extent of N deposition impacts on microbial composition remains unclear. Here we conduct a global meta-analysis using 1408 paired observations from 151 studies to evaluate the responses of soil microbial biomass, composition, and function to N addition. We show that nitrogen addition reduced total microbial biomass, bacterial biomass, fungal biomass, biomass carbon, and microbial respiration. Importantly, these negative effects increased with N application rate and experimental duration. Nitrogen addition reduced the fungi to bacteria ratio and the relative abundances of arbuscular mycorrhizal fungi and gram-negative bacteria and increased gram-positive bacteria. Our structural equation modeling showed that the negative effects of N application on soil microbial abundance and composition led to reduced microbial respiration. The effects of N addition were consistent across global terrestrial ecosystems. Our results suggest that atmospheric N deposition negatively affects soil microbial growth, composition, and function across all terrestrial ecosystems, with more pronounced effects with increasing N deposition rate and duration.

Mycorrhizal fungi form mutualistic associations with the roots of most land plants and provide them with mineral nutrients from the soil in exchange for fixed carbon derived from photosynthesis. The common symbiosis pathway (CSP) is a conserved molecular signaling pathway in all plants capable of associating with arbuscular mycorrhizal fungi. It is required not only for arbuscular mycorrhizal symbiosis but also for rhizobia-legume and actinorhizal symbioses. Given its role in such diverse symbiotic associations, we hypothesized that the CSP also plays a role in ectomycorrhizal associations. We showed that the ectomycorrhizal fungus Laccaria bicolor produces an array of lipochitooligosaccharides (LCOs) that can trigger both root hair branching in legumes and, most importantly, calcium spiking in the host plant Populus in a CASTOR/POLLUX-dependent manner. Nonsulfated LCOs enhanced lateral root development in Populus in a calcium/calmodulin-dependent protein kinase (CCaMK)-dependent manner, and sulfated LCOs enhanced the colonization of Populus by L. bicolor. Compared with the wildtype Populus, the colonization of CASTOR/POLLUX and CCaMK RNA interference lines by L. bicolor was reduced. Our work demonstrates that similar to other root symbioses, L. bicolor uses the CSP for the full establishment of its mutualistic association with Populus.

The association between plants and arbuscular mycorrhizal fungi (AMF) affects plant performance and ecosystem functioning. Recent studies have identified AMF-associated bacteria as cooperative partners that participate in AMF–plant symbiosis: specific endobacteria live inside AMF, and hyphospheric bacteria colonize the soil that surrounds the extraradical hyphae. In this Review, we describe the concept of a plant–AMF–bacterium continuum, summarize current advances and provide perspectives on soil microbiology. First, we review the top-down carbon flow and the bottom-up mineral flow (especially phosphorus and nitrogen) in this continuum, as well as how AMF–bacteria interactions influence the biogeochemical cycling of nutrients (for example, carbon, phosphorus and nitrogen). Second, we discuss how AMF interact with hyphospheric bacteria or endobacteria to regulate nutrient exchange between plants and AMF, and the possible molecular mechanisms that underpin this continuum. Finally, we explore future prospects for studies on the hyphosphere to facilitate the utilization of AMF and hyphospheric bacteria in sustainable agriculture.Most plants form symbioses with arbuscular mycorrhizal fungi, which themselves harbour endobacteria and hyphospheric bacteria. In this Review, Duan et al. explore how nutrients are transferred between the partners in this plant–fungus–bacterium continuum.

During arbuscular mycorrhizal symbiosis (AMS), considerable amounts of lipids are generated, modified and moved within the cell to accommodate the fungus in the root, and it has also been suggested that lipids are delivered to the fungus. To determine the mechanisms by which root cells redirect lipid biosynthesis during AMS we analyzed the roles of two lipid biosynthetic enzymes (FatM and RAM2) and an ABC transporter (STR) that are required for symbiosis and conserved uniquely in plants that engage in AMS. Complementation analyses indicated that the biochemical function of FatM overlaps with that of other Fat thioesterases, in particular FatB. The essential role of FatM in AMS was a consequence of timing and magnitude of its expression. Lipid profiles of fatm and ram2 suggested that FatM increases the outflow of 16:0 fatty acids from the plastid, for subsequent use by RAM2 to produce 16:0 β-monoacylglycerol. Thus, during AMS, high-level, specific expression of key lipid biosynthetic enzymes located in the plastid and the endoplasmic reticulum enables the root cell to fine-tune lipid biosynthesis to increase the production of β-monoacylglycerols. We propose a model in which β-monoacylglycerols, or a derivative thereof, are exported out of the root cell across the periarbuscular membrane for ultimate use by the fungus.

Arbuscular mycorrhizal fungi (AMF) establish symbiotic interaction with 80% of known land plants. It has a pronounced impact on plant growth, water absorption, mineral nutrition, and protection from abiotic stresses. Plants are very dynamic systems having great adaptability under continuously changing drying conditions. In this regard, the function of AMF as a biological tool for improving plant drought stress tolerance and phenotypic plasticity, in terms of establishing mutualistic associations, seems an innovative approach towards sustainable agriculture. However, a better understanding of these complex interconnected signaling pathways and AMF-mediated mechanisms that regulate the drought tolerance in plants will enhance its potential application as an innovative approach in environmentally friendly agriculture. This paper reviews the underlying mechanisms that are confidently linked with plant–AMF interaction in alleviating drought stress, constructing emphasis on phytohormones and signaling molecules and their interaction with biochemical, and physiological processes to maintain the homeostasis of nutrient and water cycling and plant growth performance. Likewise, the paper will analyze how the AMF symbiosis helps the plant to overcome the deleterious effects of stress is also evaluated. Finally, we review how interactions between various signaling mechanisms governed by AMF symbiosis modulate different physiological responses to improve drought tolerance. Understanding the AMF-mediated mechanisms that are important for regulating the establishment of the mycorrhizal association and the plant protective responses towards unfavorable conditions will open new approaches to exploit AMF as a bioprotective tool against drought.