Search

Arbuscular mycorrhizal fungi (AMF) in the soil have many positive effects on growth, nutrient acquisition, and stress tolerance of host plants, as well as soil fertility, soil structure, and soil ecology. Glomalin-related soil proteins (GRSP) are a mixture of humic substances and heat-stable glycoproteins, primarily of AMF origin. GRSP are as an important component of soil organic carbon (C) pools, which can stabilize and sequestrate C, thus reducing soil C emissions for slowing down global warming. Based on the CiteSpace software and the core collection of Web of Science as the database, this study made a visual analysis of GRSP's literature in the C field published from 1999 to 2022, including the number of publications, countries, institutions, co-cited literature, keywords, top cited papers, etc. The study regarding the GRSP in the C field could be divided into the initial stage (1999–2009), the steady stage (2010–2018), and the explosive stage (2019–2022). The Chinese Academy of Sciences is the organization with the most publications, and the United States, China, and India are the three leading nations in the C field of GRSP. However, there was little collaboration among the participating countries and the study's institutions. The focus of the research has shifted from the composition and content of GRSP in C to the question of whether C in GRSP affects soil properties. Future research was also prospected. [ABSTRACT FROM AUTHOR]

Background: Arbuscular mycorrhizal fungi (AMF) have a positive effect on drought tolerance of plants after establishing reciprocal resymbiosis with roots, while the underlying mechanism is not deciphered. Metabolomics can explain the mechanism of plant response to environmental stress by analyzing the changes of all small molecular weight metabolites. The purpose of this study was to use Ultra High Performance Liquid Chromatography Q Exactive Mass Spectrometer to analyze changes in root metabolites of walnut (Juglans regia) after inoculation with an arbuscular mycorrhizal fungus Diversispora spurca under well-watered (WW) and drought stress (DS). Results: Sixty days of soil drought significantly inhibited root mycorrhizal colonization rate, shoot and root biomass production, and leaf water potential in walnut, while AMF inoculation significantly increased biomass production and leaf water potential, accompanied by a higher increase magnitude under DS versus under WW. A total of 3278 metabolites were identified. Under WW, AMF inoculation up-regulated 172 metabolites and down-regulated 61 metabolites, along with no changes in 1104 metabolites. However, under DS, AMF inoculation up-regulated 49 metabolites and down-regulated 116 metabolites, coupled with no changes in 1172 metabolites. Among them, juglone (a quinone found in walnuts) as the first ranked differential metabolite was up-regulated by AMF under WW but not under DS; 2,3,5-trihydroxy-5–7-dimethoxyflavanone as the first ranked differential metabolite was increased by AMF under DS but not under WW. The KEGG annotation showed a large number of metabolic pathways triggered by AMF, accompanied by different metabolic pathways under WW and DS. Among them, oxidative phosphorylation and phenylalanine metabolism and biosynthesis were triggered by AMF in response to WW and DS, where N-acetyl-L-phenylalanine was induced by AMF to increase under DS, while decreasing under WW. Conclusion: This study provides new insights into the metabolic mechanisms of mycorrhiza-enhanced drought tolerance in walnuts. [ABSTRACT FROM AUTHOR]

Flavonoids are secondary metabolites widely found in plants with antioxidants, of which chalcone synthase (CHS) is a key enzyme required in flavonoid synthesis pathways. The objective of this study was to clone a CHS gene from trifoliate orange (Poncirus trifoliata) and analyze its biological information and partial functions. A PtCHS gene (NCBI accession: MZ350874) was cloned from the genome-wide of trifoliate orange, which has 1156 bp in length, encoding 391 amino acids, with a predicted protein relative molecular mass of 42640.19, a theoretical isoelectric point of 6.28, and a lipid coefficient of 89.82. The protein is stable, hydrophilic, and high sequence conservation (92.49% sequence homology with CHS gene of other species). PtCHS was highly expressed in stems, leaves and flowers, but very low expression in roots and seeds. Soil water deficit could up-regulate expressions of PtCHS in leaves. An arbuscular mycorrhizal fungus, Funneliformis mosseae, significantly increased plant biomass production, CHS activity, expressions of PtCHS, and total flavonoid content in leaves and roots, independent of soil water status. Total flavonoids were significantly positively correlated with PtCHS expression in leaves only and also positively with root mycorrhizal colonization. Such results provide insight into the important functions of PtCHS in trifoliate orange. [ABSTRACT FROM AUTHOR]

This study aimed to evaluate effects of Funneliformis mosseae on plant growth and root exudate compositions and contents, soil and root phosphatase activity, soil glomalin concentrations, and thus soil aggregate stability and distribution in trifoliate orange under well-watered (75% of maximum water holding capacity) and drought stress (55% of maximum water holding capacity) conditions. After eight weeks of drought treatment, mycorrhizal fungal inoculation improved plant growth and exhibited altered composition of root exudates than non-inoculated treatment. Mycorrhizal fungal inoculation dramatically increased the relative abundance of phenolics (e.g., 2 H,8 H-Benzo[1,2-b:3,4-b'] dipyran-2-one,8,8-dimethyl), terpenoids (e.g., geijerene), and acids (n-hexadecanoic acid), while notably reduced the relative abundance of alkanes (e.g., tridecane, 2-methyl-), esters (e.g., hexanedioic acid and dimethyl ester), and amides (e.g.,13-docosenamide) in root exudates. Mycorrhizal fungal colonization profoundly increased easily extractable and total glomalin-related soil protein levels under two soil water regimes, which cemented soil macroaggregate (2–4 mm size) formation, thereby, improving soil aggregate stability. Mycorrhizal fungal-inoculated plants represented higher soil acid, alkaline and total phosphatase activities, irrespective of well-watered and drought treatment. The results suggest that mycorrhizal plants had improved root microenvironment to mitigate drought damage through changes in root exudate components along with glomalin, phosphatase, and soil aggregate stability in the mycorrhizosphere. [ABSTRACT FROM AUTHOR]

The circadian rhythm of plants is associated with stress responses; however, it is not clear whether increased host plant drought tolerance by arbuscular mycorrhizal fungi (AMF) is associated with changes in the circadian clock. The present study aimed to analyze the effect of Funneliformis mosseae (Nicol. & Gerd.) Schüßler & Walker on the circadian clock gene expression patterns in trifoliate orange (Poncirus trifoliata L. Raf.) along with gas exchange, abscisic acid (ABA) levels and antioxidant enzyme gene expression under well-watered (WW) and drought stress (DS) conditions. Plant growth, net photosynthetic rate, stomatal conductance and ABA levels were significantly higher in AMF- than in non-AMF-inoculated plants regardless of soil water regimes. Six circadian clock genes, including PtPRR7 , PtLHY , PtCCA1 , PtGI , PtPIF3 and PtSRR1 , were identified and showed rhythmic expression patterns over the course of the day. The AMF inoculation reduced the expression of most circadian clock genes in different time periods. However, AMF treatment significantly increased PtPRR7 and PtGI expression at 5:00 p.m. under WW and DS conditions, PtLHY expression at 1:00 a.m. and PtSRR1 expression at 9:00 p.m. At 1:00 a.m. AMF inoculation up-regulated the expression of the circadian clock genes PtPRR7 , PtCCA1 , PtLHY and PtPIF3 and the antioxidant enzyme genes PtFe-SOD , PtMn-SOD , PtCu/Zn-SOD , PtPOD and PtCAT1. Correlation analysis revealed that these changes in circadian clock gene expression were associated with antioxidant enzyme gene expression, root ABA and gas exchange. We concluded that mycorrhizal fungi have the ability to regulate the daily rhythm of the circadian clock in trifoliate orange plants in response to drought. [ABSTRACT FROM AUTHOR]

Plants are often subjected to various environmental stresses during their life cycle, among which drought stress is perhaps the most significant abiotic stress limiting plant growth and development. Arbuscular mycorrhizal (AM) fungi, a group of beneficial soil fungi, can enhance the adaptability and tolerance of their host plants to drought stress after infecting plant roots and establishing a symbiotic association with their host plant. Therefore, AM fungi represent an eco-friendly strategy in sustainable agricultural systems. There is still a need, however, to better understand the complex mechanisms underlying AM fungi-mediated enhancement of plant drought tolerance to ensure their effective use. AM fungi establish well-developed, extraradical hyphae on root surfaces, and function in water absorption and the uptake and transfer of nutrients into host cells. Thus, they participate in the physiology of host plants through the function of specific genes encoded in their genome. AM fungi also modulate morphological adaptations and various physiological processes in host plants, that help to mitigate drought-induced injury and enhance drought tolerance. Several AM-specific host genes have been identified and reported to be responsible for conferring enhanced drought tolerance. This review provides an overview of the effect of drought stress on the diversity and activity of AM fungi, the symbiotic relationship that exists between AM fungi and host plants under drought stress conditions, elucidates the morphological, physiological, and molecular mechanisms underlying AM fungi-mediated enhanced drought tolerance in plants, and provides an outlook for future research. [ABSTRACT FROM AUTHOR]

Soil water deficit seriously affects crop production, and soil arbuscular mycorrhizal fungi (AMF) enhance drought tolerance in crops by unclear mechanisms. Our study aimed to analyze changes in non-targeted metabolomics in roots of trifoliate orange (Poncirus trifoliata) seedlings under well-watered and soil drought after inoculation with Rhizophagus intraradices , with a focus on terpenoid profile. Root mycorrhizal fungal colonization varied from 70% under soil drought to 85% under soil well-watered, and shoot and root biomass was increased by AMF inoculation, independent of soil water regimes. A total of 643 secondary metabolites in roots were examined, and 210 and 105 differential metabolites were regulated by mycorrhizal fungi under normal water and drought stress, along with 88 and 17 metabolites being up-and down-regulated under drought conditions, respectively. KEGG annotation analysis of differential metabolites showed 38 and 36 metabolic pathways by mycorrhizal inoculation under normal water and drought stress conditions, respectively. Among them, 33 metabolic pathways for mycorrhization under drought stress included purine metabolism, pyrimidine metabolism, alanine, aspartate and glutamate metabolism, etc. We also identified 10 terpenoid substances, namely albiflorin, artemisinin (−)-camphor, capsanthin, β-caryophyllene, limonin, phytol, roseoside, sweroside, and α-terpineol. AMF colonization triggered the decline of almost all differential terpenoids, except for β-caryophyllene, which was up-regulated by mycorrhizas under drought, suggesting potential increase in volatile organic compounds to initiate plant defense responses. This study provided an overview of AMF-induced metabolites and metabolic pathways in plants under drought, focusing on the terpenoid profile. [ABSTRACT FROM AUTHOR]

A feature of arbuscular mycorrhiza is enhanced drought tolerance of host plants, although it is unclear whether host H+-ATPase activity and gene expression are involved in the physiological process. The present study aimed to investigate the effects of an arbuscular mycorrhizal fungus (AMF), Funneliformis mosseae , on H+-ATPase activity, and gene expression of trifoliate orange (Poncirus trifoliata) seedlings subjected to well-watered (WW) and drought stress (DS), together with the changes in leaf gas exchange, root morphology, soil pH value, and ammonium content. Soil drought treatment dramatically increased H+-ATPase activity of leaf and root, and AMF inoculation further strengthened the increased effect. A plasma membrane (PM) H+-ATPase gene of trifoliate orange, PtAHA2 (MW239123), was cloned. The PtAHA2 expression was induced by mycorrhization in leaves and roots and also up-regulated by drought treatment in leaves of AMF-inoculated seedlings and in roots of AMF- and non-AMF-inoculated seedlings. And, the induced expression of PtAHA2 under mycorrhization was more prominent under DS than under WW. Mycorrhizal plants also showed greater photosynthetic rate, stomatal conductance, intercellular CO2 concentration, and transpiration rate and better root volume and diameter than non-mycorrhizal plants under DS. AMF inoculation significantly increased leaf and root ammonium content, especially under DS, whereas it dramatically reduced soil pH value. In addition, H+-ATPase activity was significantly positively correlated with ammonium contents in leaves and roots, and root H+-ATPase activity was significantly negatively correlated with soil pH value. Our results concluded that AMF stimulated H+-ATPase activity and PtAHA2 gene expression in response to DS, which resulted in great nutrient (e.g., ammonium) uptake and root growth, as well as low soil pH microenvironment. [ABSTRACT FROM AUTHOR]

Soil arbuscular mycorrhizal fungi (AMF) enhance the tolerance of plants against soil moisture deficit stress (SMDS), but the underlying mechanisms are still not fully understood. Polyamines (PAs) as low-molecular-weight, aliphatic polycations have strong roles in abiotic stress tolerance of plants. We aimed to investigate the effect of AMF (Funneliformis mosseae) inoculation on PAs, PA precursors, activities of PA synthases and degrading enzymes, and concentration of reactive oxygen species in the roots of trifoliate orange (Poncirus trifoliata) subjected to 15 days of SMDS. Leaf water potential and total chlorophyll levels were comparatively higher in AMF-inoculated than in non-AMF-treated plants exposed to SMDS. Mycorrhizal plants recorded a significantly higher concentration of precursors of PA synthesis such as L-ornithine, agmatine, and S -adenosyl methionine, besides higher putrescine and cadaverine and lower spermidine during the 15 days of SMDS. AMF colonization raised the PA synthase (arginine decarboxylase, ornithine decarboxylase, spermidine synthase, and spermine synthase) activities and PA-degrading enzymes (copper-containing diamine oxidase and FAD-containing polyamine oxidase) in response to SMDS. However, mycorrhizal plants showed a relatively lower degree of membrane lipid peroxidation, superoxide anion free radical, and hydrogen peroxide than non-mycorrhizal plants, whereas the difference between them increased linearly up to 15 days of SMDS. Our study concluded that AMF regulated PA homeostasis in roots of trifoliate orange to tolerate SMDS. [ABSTRACT FROM AUTHOR]

Walnut, an important oil fruit tree, is dependent on arbuscular mycorrhizas, while mycorrhizal roles and efficient mycorrhizal fungus in walnuts are unknown. This study was conducted to evaluate the effect of five arbuscular mycorrhizal fungi (AMF) species, including Acaulospora scrobiculata, Diversispora spurca, Glomus etunicatum, G. mosseae, and G. versiforme on plant growth, leaf gas exchange, root morphology, and root nutrient contents of walnut (Juglans regia L. Liaohe 1) seedlings. Three months of AMF inoculations later, root mycorrhizal colonisation achieved 47.0% to 76.4%. AMF treatments increased plant growth performance, dependent on AMF species. AMF- -inoculated plants with D. spurca, G. etunicatum, and G. mosseae showed higher root length, projected area, surface area, and volume than non-AMF plants. Except for G. versiforme, the other four AMF treatments almost significantly increased leaf photosynthesis rate, transpiration rate, and stomatal conductivity, while reduced intercellular CO2 concentrations and leaf temperature. AMF affected root nutrient contents, dependent on AMF and mineral nutrient species. These results, thereby, concluded that AMF had a positive role in walnuts, dependent on AMF species, and D. spurca was the best mycorrhizal fungus for walnut. Such results provide the potential possibility of a developing consortium of AMF in walnut cultivation management. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) enhance salt tolerance of host plants, while it is unclear whether aquaporins are involved in the AMF event. This study analyzed the effects of inoculation with an arbuscular mycorrhizal fungus Paraglomus occultum on plant physiological performance and relative expression of root tonoplast intrinsic proteins (TIPs, a kind of aquaporins) genes in potted trifoliate orange (Poncirus trifoliata) seedlings subjected to 0 and 100 mM NaCl concentrations. AMF inoculation significantly increased plant growth performance, leaf water status (relative water content and water potential) and gas exchange (photosynthetic rate, transpiration rate and stomatal conductance), irrespective of non-salt and salt stress. Mycorrhizal seedlings exhibited significantly higher root salicylic acid concentrations and lower leaf and root jasmonic acid concentrations than non-mycorrhizal seedlings under non-salt and salt stress. Transcript levels of PtTIP1;1, PtTIP1;2 and PtTIP1;3 were down-regulated by AMF inoculation under non-salt and salt stress. PtTIP2;1 and PtTIP2;2 expression was unchanged under non-salt stress whereas was down-regulated by AMF colonization under salt stress. AMF inoculation considerably enhanced PtTIP4;1 transcript levels under non-salt and salt stress and PtTIP5;1 expression under non-salt stress. It is concluded that mycorrhizas mitigated salt stress of trifoliate orange through improving physiological performance and regulating root PtTIPs expression. [ABSTRACT FROM AUTHOR]

A pot experiment was used to evaluate the effects of an arbuscular mycorrhizal fungus (AMF) Funneliformis mosseae on plant growth performance, root-hair growth, and root hormone levels in trifoliate orange (Poncirus trifoliata) seedlings under well-watered (WW) and drought stress (DS). A 9-week mild DS treatment significantly reduced mycorrhizal colonization of 2nd- and 3rd-order lateral roots. Root mycorrhizal colonization was relatively higher in the 2nd- and 3rd-order lateral roots than in the taproot and the 1st-order lateral root under WW and DS. AMF seedlings exhibited significantly higher root-hair density, length (except for the taproot) and diameter in taproot and 1st-, 2nd-, and 3rd-order lateral roots under WW, and considerably higher root-hair density (except for 1st-order lateral root), length (except for 2nd-order lateral root) and diameter under DS. Mycorrhizal inoculation remarkably increased root abscisic acid (ABA), indole-3-acetic acid (IAA), methyl jasmonate, and brassinosteroids (BRs) concentrations under DS, in company with the decrease in root zeatin riboside and gibberellins levels and root IAA effluxes. Root-hair traits were significantly positively correlated with root colonization and root ABA and BRs levels. It is concluded that mycorrhizal plants possessed better root-hair growth to adapt mild DS, which is associated with mycorrhizal colonization and endogenous hormone changes. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizas (AMs) have the ability to enhance drought tolerance of citrus, but the underlying mechanisms have not been clearly elucidated. Considering the strong association of cell membrane fatty acid (FA) unsaturation with plant drought tolerance, the present study hypothesized that AM fungi (AMF) modulated the composition and unsaturation of FAs to enhance drought tolerance of host plants. Drought-sensitive citrus rootstocks, trifoliate orange (Poncirus trifoliata) seedlings, were inoculated with AMF (Funneliformis mosseae) for 3 months and were subsequently exposed to drought stress (DS) for 8 weeks. Mycorrhizal seedlings exhibited better plant growth performance, higher leaf water potential and lower root abscisic acid concentrations under both well-watered (WW) and DS conditions. Arbuscular mycorrhiza fungus inoculation considerably increased root methyl oleate (C18:1), methyl linoleate (C18:2) and methyl linolenate (C18:3N3) concentrations under both WW and DS conditions, and root methyl palmitoleate (C16:1) concentrations under WW, while it decreased root methyl stearate (C18:0) levels under both WW and DS. These changes in the composition of FAs of mycorrhized roots resulted in higher unsaturation index of root FAs, which later aided in reducing the oxidative damage on account of lower concentration of malondialdehyde and superoxide radicals. The changes of these FAs were a result of AMF-up-regulating root FA desaturase 2 (PtFAD2), FA desaturase 6 (PtFAD6) and Δ9 FA desaturase (PtΔ9) genes under WW and PtFAD2 , PtFAD6 and Δ15 FA desaturase (PtΔ15) genes under DS conditions. Our results confirmed that mycorrhization brought significant changes in root FA compositions, in addition to regulation of gene expression responsible for increasing the unsaturation level of FAs, a predisposing physiological event for better drought tolerance of citrus. [ABSTRACT FROM AUTHOR]

Glomalin-related soil protein (GRSP) is a specific glycoprotein secreted into the soil by hyphae and spores of arbuscular mycorrhizal fungi that have many potential functions. It is not clear whether exogenous GRSP has an effect on plant growth and soil properties or whether the effects are related to the type of GRSP used. In this study, trifoliate orange (Poncirus trifoliata L. Raf.) seedlings were used to analyze the effects of easily extractable GRSP (EE-GRSP) and difficultly extractable GRSP (DE-GRSP) at a quarter-, half-, and full-strength concentration on shoot and root biomass as well as soil properties The results showed that, at different strengths, exogenous EE-GRSR significantly increased shoot and root biomass compared to the control, which displayed the most significant effects from the half-strength EE-GRSP. In contrast, DE-GRSP, at various strengths, significantly reduced shoot and root biomass. Furthermore, the application of exogenous EE-GRSP stimulated soil water-stable aggregate (WSA) content at 2–4 mm and 0.5–1 mm sizes, while DE-GRSP strongly reduced WSA content at the 2–4 mm, 1–2 mm, 0.5–1 mm, and 0.25–0.5 mm sizes, consequently leading to an increase or decrease in the WSA stability, according to the mean weight diameter. However, exogenous EE-GRSP decreased soil pH and DE-GRSP increased it, which was related to WSA stability. Exogenous EE-GRSP almost significantly increased soil acidic, neutral, and alkaline phosphatase activity at different strengths, while exogenous DE-GRSP, also at different strengths, significantly inhibited soil acidic phosphatase activity. The application of both exogenous EE-GRSP and DE-GRSP increased the organic carbon content of the soil. This study concluded that exogenous GRSP exerted differential effects on plant biomass and soil properties, and EE-GRSP can be considered as a soil stimulant for use in citrus plants. To our knowledge, this is the first report on the negative effects of exogenous DE-GRSP on plant biomass and soil properties. [ABSTRACT FROM AUTHOR]

The development of the mining industry and the overuse of inorganic fertilizers have led to an excess of manganese (Mn) in the soil, thereby, contaminating the soil environment and people's health. On heavy metal-contaminated soils, the combined arbuscular mycorrhizal fungi (AMF)-phytoremediation technique becomes a hotspot because of its environmentally friendly, in situ remediation. AMF inoculation often leads to a decrease in host Mn acquisition, which provides a basis for its application in phytoremediation of contaminated soils. Moreover, the utilization value of native AMF is greater than that of exotic AMF, because native AMF can adapt better to Mn-contaminated soils. In addition to the fact that AMF enhance plant Mn tolerance responses such as regionalization, organic matter chelation, limiting uptake and efflux, and so on, AMF also develop plant-independent fungal pathways such as direct biosorption of Mn by mycorrhizal hyphae, fungal Mn transporter genes, and sequestration of Mn by mycorrhizal hyphae, glomalin, and arbuscule-containing root cortical cells, which together mitigate excessive Mn toxicity to plants. Clarifying AMF-plant interactions under Mn stress will provide support for utilizing AMF as a phytoremediation in Mn-contaminated soils. The review reveals in detail how AMF develop its own mechanisms for responding to excess Mn and how AMF enhance plant Mn tolerance, accompanied by perspectives for future research. [Display omitted] • AMF inoculation diversely affects host Mn levels, dominated by reduced Mn uptake. • Mn-contaminated soils reduce AMF diversity, along with Glomus as the dominant genus and native AMF more tolerant. • AMF enhances plant Mn tolerance such as regionalization and chelation. • AMF hyphae have direct biosorption and sequestration of Mn. • Fungal Mn transporters, glomalin, and arbuscule-contained cortical cells mitigate Mn. [ABSTRACT FROM AUTHOR]

Heavy metal accumulation in plants is a severe environmental problem, rising at an expeditious rate. Heavy metals such as cadmium, arsenic, mercury and lead are known environmental pollutants that exert noxious effects on the morpho-physiological and biological attributes of a plant. Due to their mobile nature, they have become an extended part of the food chain and affect human health. Arbuscular mycorrhizal fungi ameliorate metal toxicity as they intensify the plant's ability to tolerate metal stress. Mycorrhizal fungi have vesicles, which are analogous to fungal vacuoles and accumulate massive amount of heavy metals in them. With the help of a pervasive hyphal network, arbuscular mycorrhizal fungi help in the uptake of water and nutrients, thereby abating the use of chemical fertilizers on the plants. They also promote resistance parameters in the plants, secrete a glycoprotein named glomalin that reduces the metal uptake in plants by forming glycoprotein–metal complexes, and improve the quality of the soil. They also assist plants in phytoremediation by increasing the absorptive area, increase the antioxidant response, chelate heavy metals and stimulate genes for protein synthesis that reduce the damage caused by free radicals. The current manuscript focuses on the uptake of heavy metals, accumulation, and arbuscular mycorrhizal impact in ameliorating heavy metal stress in plants. [ABSTRACT FROM AUTHOR]

A potted experiment was carried out to evaluate the effect of an arbuscular mycorrhizal fungus (AMF), Acaulospora scrobiculata, on peach seedlings grown in non-replant (NR) and replant (R) soils, to establish whether AMF inoculation alleviated soil replant disease through changes in physiological levels and relevant gene expression. After 15 weeks of mycorrhization, root mycorrhizal colonization was heavily inhibited by R treatment versus NR treatment. AMF plants under NR and R soil conditions displayed significantly higher total plant biomass than non-AMF plants. AMF inoculation significantly increased root sucrose and fructose concentrations and root catalase, peroxidase, polyphenol oxidase, and phenylalanine ammonialyase activities under R conditions. Likewise, salicylic acid, jasmonic acid, chitinase, total soluble phenol, and lignin concentrations in roots were significantly higher in AMF than in non-AMF seedlings grown in R soil. Over-expression of PpCHI, PpLOX1, PpLOX5, PpAOC3, PpAOC4, and PpOPR2 in roots was observed in AMF-inoculated seedlings, as compared to that of non-AMF-inoculated seedlings grown in R soils. Thus, mycorrhizal fungal inoculation conferred a greater tolerance to peach plants in R soil by stimulating antioxidant enzyme activities, disease-resistance substance levels, and the expression of relevant genes. [ABSTRACT FROM AUTHOR]

Earthworms and arbuscular mycorrhizal fungi (AMF) are beneficial organisms in the soil and play an important role in improving soil fertility and plant growth. The objective of this study was to analyze the response of single versus dual inoculation of earthworms (Pheretima guillelmi) and AMF (Funneliformis mosseae) on mycorrhizal growth, root morphology, and rhizosphere properties of white clover (Trifolium repens). Addition of earthworms significantly increased the degree of root AMF colonization, while decreased hyphal length and spore number in the soil. Single or dual inoculation of F. mosseae and P. guillelmi significantly increased root traits (total length, area, volume, and average diameter), soil phosphatase activities (acid, neutral, and alkaline), easily extractable and total glomalin-related soil protein content, the percentage of soil water-stable aggregates at the size of 2–4 mm, 1–2 mm and 0.5–1 mm, aggregate stability, Bray-P, and soil organic carbon (SOC) contents. Among them, the dual inoculation further amplified the effect on glomalin production, phosphatase activity, and aggregate stability, but did not show a superposition effect on root improvement, Bray-P, and SOC. Our study, hence, suggested that AMF and earthworms have synergistic roles in elevating rhizosphere properties vis-à-vis agronomic responses of white clover. • Effects of earthworm-AMF (singly and in combination) on rhizosphere environment and crop response were studied. • Earthworms increased root AMF colonization, but reduced hyphae and spores in soils. • AMF or earthworms singly improved root traits, Bray-P, and soil organic carbon contents, but dual application of AMF and earthworms did not show a major effect than single application. • Single AMF or earthworms increased aggregation, phosphatase, GRSP, and dual addition amplified these responses. [ABSTRACT FROM AUTHOR]

Medicinal plants are well known to have the advantages of high concentration of medicinal ingredients having clinical importance, curative value, small toxic and side effects. Important compounds viz., paclitaxel, camptothecin, and vincristine have been developed from medicinal plants as first-line of clinical drugs, leading to their consistently increasing demand globally. However, the destruction of natural environment due to excessive mining threatened such resources jeopardizing the successful growing of medicinal plants. A group of beneficial arbuscular mycorrhizal (AM) fungi is known to exist in the rhizosphere of medicinal plants, which can establish a reciprocal symbiosis with their roots, namely arbuscular mycorrhizas. These AM fungi are pivotal in the habitat adaptation of medicinal plants. Studies have demonstrated that AM fungi aided in growth promotion and nutrient absorption of medicinal plants, thereby, accelerating the accumulation of medicinal ingredients and aiding resistance against abiotic stresses such as drought, low temperature, and salinity. An AMlike fungus Piriformospora indica is known to be cultured in vitro without roots, later showed analogous effects of AM fungi on medicinal plants. These fungi provide new mechanistic pathways towards the artificial cultivation of medicinal plants loaded with ingredients in huge demand in international market. This review provides an overview of the diversity of AM fungi inhabiting the rhizosphere of medicinal plants, and analyzes the functioning of AM fungi and P. indica, coupled with future lines of research. [ABSTRACT FROM AUTHOR]

Aquaporins (AQPs) involved in water and small molecule transport respond to environmental stress, while it is not clear how arbuscular mycorrhizal fungi (AMF) regulate AQP expression. Here, we investigated the change in leaf water potential and expression level of four tonoplast intrinsic proteins (TIPs), six plasma membrane intrinsic proteins (PIPs), and four nodin-26 like intrinsic proteins (NIPs) genes in trifoliate orange (Poncirus trifoliata) inoculated with Funneliformis mosseae under well-watered (WW), salt stress (SS), and waterlogging stress (WS). Root AMF colonization and soil hyphal length collectively were reduced by SS and WS. Under WW, inoculation with AMF gave diverse responses of AQPs : six AQPs up-regulated, three AQPs down-regulated, and five AQPs did not change. Such up-regulation of more AQPs under mycorrhization and WW partly accelerated water absorption, thereby, maintaining higher leaf water potential. However, under SS, all the fourteen AQPs were dramatically induced by AMF inoculation, which improved water permeability of membranes and stimulated water transport of the host. Under WS, AMF colonization almost did not induce or even down-regulated these AQPs expressions with three exceptions (PtTIP2 ;2, PtPIP1;1 , and PtNIP1;2), thus, no change in leaf water potential. As a result, mycorrhizal plants under flooding may have an escape mechanism to reduce water absorption. It is concluded that AMF had different strategies in response to environmental stresses (e.g. SS and WS) by regulating leaf AQP expression in the host (e.g. trifoliate orange). • Under ample water, AMF gave diverse responses of AQPs. • Under salinity, all the 14 AQPs were up-regulated by AMF inoculation to stimulate water uptake. • Under waterlogging, AMF almost did not induce these AQPs expressions, thus, giving an escape mechanism. • AMF have different strategies in response to environmental stress through AQPs. [ABSTRACT FROM AUTHOR]

The aim of the present study was to analyze the effects of two arbuscular mycorrhizal fungi (AMF), Funneliformis mosseae and Paraglomus occultum, on leaf water status, root morphology, root sugar accumulation, root abscisic acid (ABA) levels, root malondialdehyde (MDA) content, and root antioxidant enzyme activities in white clover ( Trifolium repens L.) exposed to well-watered (WW) and drought stress (DS) conditions. The results showed that root colonization by F. mosseae and P. occultum was significantly decreased by 7-week soil drought treatment. Under drought stress conditions, mycorrhizal fungal treatment considerably stimulated root total length, surface area and volume, as compared with non-mycorrhizal controls. In addition, inoculation with arbuscular mycorrhizal fungi also increased leaf relative water content and accelerated the accumulation of root glucose and fructose under drought stress. Mycorrhizal plants under drought stress registered higher activities of superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD) and ABA levels in roots, while lower MDA contents, relative to non-mycorrhizal plants. As a result, mycorrhiza-inoculated plants represented better physiological activities (e.g. antioxidant defense systems, root morphology, and sugar accumulation) than non-inoculated plants in response to soil drought, whilst P. occultum had superior effects than F. mosseae. [ABSTRACT FROM AUTHOR]

The present study aimed to evaluate the effect of an arbuscular mycorrhizal fungus (AMF), Clariodeoglomus etunicatum, on leaf food quality and relevant gene expression levels of tea (Camellia sinensis cv. 'Fuding Dabaicha') seedlings exposed to 0.5 µM P (P0.5) and 50 µM P (P50) levels. Twenty-four weeks later, the seedlings recorded higher root mycorrhizal fungal colonization in P50 than in P0.5. AMF-inoculated tea plants represented significantly higher leaf fructose and glucose contents and lower sucrose content than noninoculated plants, irrespective of substate P levels. AMF treatment also increased total amino acids content in P0.5 and P50, accompanied with higher expression of glutamate dehydrogenase (CsGDH) and lower expression of glutamine synthetase (CsGS) and glutamine oxoglutarate aminotransferase (CsGOGAT). The total flavonoid content was higher in mycorrhizal versus non-mycorrhizal plants under P0.5 and P50, together with induced expression of phenylalanine ammonia-lyase (CsPAL) and cinnamic acid 4-hydroxylase (CsC4H). Mycorrhizal fungal inoculation improved catechins content, which is due to the up-regulated expression of flavanone 3-hydroxylase ( CsF3H), flavonoid 3'-hydroxylase ( CsF3'H), dihydroflavonol 4-reductase ( CsDFR), leucoanthocyanidin reductase ( CsLAR), anthocyanidin reductase ( CsANR), and chalcone isomerase ( CsCHI) under P0.5. However, under P50, the gene involved in catechins synthesis was not affected or down-regulated by mycorrhization, implying a complex mechanism (e.g. nutrient improvement). AMF also inhibited the tea caffeine synthase 1 (CsTCS1) expression regardless of P levels. Therefore, the results of this study concluded that inoculation with C. etunicatum improves leaf food quality of tea exposed to P stress, but the improved mechanisms were different between P0.5 and P50. [ABSTRACT FROM AUTHOR]

The demand for ornamental plants is increasing due to urban greening and rural construction, while the growing environment of plants, especially the soil environment, is deteriorating. Hence, sustainable methods of ornamental plant cultivation need to be developed quickly. The application of arbuscular mycorrhizal fungi (AMF) to ornamental plants can be one of the eco-friendly ways to achieve the objective. Soil AMF establish mycorrhizal symbiosis with roots of ornamental plants, which can develop a marvelous mycorrhizal mycelium network in the rhizosphere to stimulate nutrient and water acquisition of host plants. Numerous researches have proven that AMF improved the quality of ornamental plants, like fruit yield, height, biomass, seed quality, the size and number of flowers, leaf, and root. In addition, mycorrhizal fungi also improve nutrient uptake and endogenous hormone balance of host plants. Another important function of AMF is to regulate the physiological, biochemical, and molecular responses of host plants to adversity, including drought stress, temperature stress, heavy-metal stress, and insect and disease stress. From the perspective of the ecological garden landscape, AMF richness would maintain plant abundance, nutrient and energy balance, and higher productivity in normal and soil environment stress, thus, establishing a friendly-environmental ecosystem. This review also provides the basis to exploit and improve the commercial application of AMF in ornamental plants in the future. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) enhance plant disease resistance, while the underlying mechanisms in the molecular levels are not yet known. In this study, five-leaf-old trifoliate orange seedlings were inoculated with Funneliformis mosseae for 14 weeks and subsequently were infected by a citrus root rot pathogen Phytophthora parasitica by 7 days. The transcriptome results by Illumina HiSeq 4000 revealed that the percentage of Q30 bases reached 92.99% or above, and 29696 unigenes were annotated in a total of 63531 unigenes. 654 and 103 differentially expressed genes (DEGs) were respectively annotated in AMF-inoculated versus non-AMF-inoculated plants under non-infection and infection with P. parasitica, respectively, whilst these DEGs were related to defense mechanisms, signal transduction mechanisms and secondary metabolites biosynthesis. Forty-two genes were functionally annotated as the putative 'defense mechanism', whilst AMF inoculation induced 1 gene down-regulated and 3 genes up-regulated under P. parasitica infection. AMF inoculation stimulated more genes linked to signal transduction mechanism down-regulated than non-AMF plants. Eight genes were involved in secondary metabolites biosynthesis in AMF versus non-AMF seedlings under P. parasitica-infection conditions. Such transcriptome database provided total information in the molecular levels regarding mycorrhizal roles in tolerating Phytophthora parasitica infection. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizas absorb water from soil to host plants, while the relationship between mycorrhizas and aquaporins (AQPs, membrane water channel proteins, which function in water transport) in mycorrhizal plants is unclear. In this study, Funneliformis mosseae -colonized trifoliate orange (Poncirus trifoliata) seedlings were grown in pots fitted with 37-μm nylon meshes at the bottom of each pot to allow mycorrhizal hyphae absorb water from an outer beaker. The expression of seven plasma membrane intrinsic proteins (PIPs) genes, six tonoplast intrinsic proteins (TIPs) genes, and four nodulin-26 like intrinsic proteins (NIPs) genes were analyzed in roots of both well-watered (WW) and drought stressed (DS) plants. The six-week DS plants dramatically increased hyphal water absorption rate by 1.4 times, as compared with WW plants. Mycorrhizal plants exhibited greater plant growth performance, leaf water status (water potential and relative water content), and gas exchange under both WW and DS conditions. Mycorrhizal inoculation induced diverse expression patterns in these AQPs under WW: up-regulation of PtNIP1 ; 1 , PtPIP2 ; 1 , and PtPIP2 ; 5 , down-regulation of PtNIP1;2 , PtNIP6 ; 1 , PtPIP1;2 , PtPIP1 ; 5 , PtPIP2 ; 8 , PtTIP1 ; 1 , PtTIP1 ;2, PtTIP1 ; 3 , and PtTIP5;1 , and no changes in other AQPs. However, the expression of PtPIPs and PtNIPs was down-regulated by mycorrhizal inoculation under DS, and PtTIPs was not induced by mycorrhizal colonization under DS. The expression pattern of AQPs in response to mycorrhizas under DS is a way of mycorrhizal plants to minimize water loss. • Mycorrhizas and AQPs are involved in water absorption of AM plants. • AM trifoliate orange had higher hyphal water absorption rate under DS than under WW. • AQP gene expression was down-regulated or not changed by AMF under DS and hyphae. [ABSTRACT FROM AUTHOR]

Beneficial fungi - including difficult - to - culture arbuscular mycorrhizal fungi (AMF) and the culturable endophytic fungus Piriformospora indica - play positive roles in hosts under controlled conditions, whereas information regarding the application of these fungi in field citrus (a mycorrhizal - dependent plant) is scarce. The present study aims to analyze the effects of two AMF species (Diversispora spurca and D. versiformis) and an endophytic fungus (P. indica) on the fruit quality and soil properties of Newhall navel oranges (Citrus sinensis Osbeck cv. Newhall) grafted on trifoliate oranges in the field. The results showed the improvement of root fungal colonization following fungal inoculation. The fungal inoculations increased the soil's acid, neutral, alkaline, and total phosphatase activity, as well as soil Bray-P content; increased the distribution of water-stable aggregates, mainly in the size range of 2–4 mm; and stimulated the production of easily extractable, difficultly extractable, and total glomalin-related soil proteins in a fungal-species-dependent manner, which was positively correlated with soil aggregate stability. Fungal inoculations, to some extent, improved external fruit quality, depending on the fungal species. Among the three fungi, only D. versiformis significantly increased the fruits' soluble solid content, but all three together promoted fruit glucose, fructose, and sucrose content. The two AMF species dramatically increased fruit phosphorus (P) content, while P. indica distinctly reduced it. D. versiformis also significantly elevated fruit potassium (K), magnesium (Mg), and manganese (Mn) content, D. spurca improved fruit copper (Cu), zinc (Zn), K, and Mg content, and P. indica increased fruit iron (Fe) and Zn content, compared with non-inoculated controls. Our results indicate that the field inoculation of citrus with AMF and P. indica partially improved fruit quality and soil properties, providing a pathway for P. indica to partially replace AMF in field citrus applications. • Effects of field inoculation with two AMF and an AM-like fungus on Newhall navel orange were studied. • Root fungal colonization was improved by the three fungal inoculations. • These fungi increased soil phosphatases, Olsen-P, EE-GRSP, T-GRSP, and aggregate stability. • Fungal inoculations partly improved fruit quality and mineral element contents, depending on fungi species. • A cultural mycorrhiza-like fungus Piriformospora indica partially replace AMF in applications of field citrus. [ABSTRACT FROM AUTHOR]

• Effects of symbiotic AM fungi and AM like fungi on trifoliate orange were studied. • Piriformospora indica and AMF coexisted in roots, but the colonization was inferior. • Single or dual inoculation increased plant growth and affected root nutrients. • P increase in inoculated plants was associated with soil phosphatase and PT gene. • Piriformospora indica has more applied value in citrus than AMF as a biofertilizer. Citrus plants heavily depend on soil arbuscular mycorrhizal fungi (AMF), while the application of AMF in citrus is limited because of no propagation in vitro. The present work tried to apply a culturable (in vitro) endophytic fungus Piriformospora indica into trifoliate orange (Poncirus trifoliata) and to compare the role of P. indica in plant growth and nutrient acquisition especially phosphorus (P), relative to an arbuscular mycorrhizal fungus Funneliformis mosseae. After 20 weeks, roots of trifoliate orange could be colonized by P. indica , and both P. indica and F. mosseae existed in roots together. Root fungal colonization and soil easily extractable, difficultly extractable, and total glomalin-related protein concentrations were higher under single F. mosseae inoculation than under single P. indica and dual inoculation of F. mosseae and P. indica. Single or dual microbial inoculant collectively improved plant growth including plant height, leaf number, and leaf, stem, and root biomass, compared with non-inoculation. In addition, single fungal treatment raised the content of root N, P, and Mg, and dual inoculation increased only Mg and B contents. The improvement of P by single microbial inoculant was associated with the fungi-accelerated soil phosphatase activity and the fungi-induced root phosphate transporter genes (PT3 , PT5 , and PT6) expression levels. The positive effects on growth and nutrients were more significant under single F. mosseae than under single P. indica and dual inoculations. It concluded that P. indica has the stimulated roles in plant growth and P acquisition of trifoliate orange, and thus has more applied potentiality in citriculture than AMF as a biofertilizer. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) produce glomalin-related soil protein (GRSP) that influences organic carbon (C) storage in soil; however, how much purified GRSP fractions contribute to soil organic carbon (SOC) is yet not known. The present study evaluated the contribution of GRSP towards changes in SOC in trifoliate orange grown in a rootbox divided into a roots + hyphae chamber (roots colonized by AMF hyphae and AMF extraradical hyphae) and a hyphae chamber (only the presence of AMF extraradical hyphae, without roots). Three AMF species (Diversispora epigaea , Paraglomus occultum and Rhizoglomus intraradices) were inoculated into the roots + hyphae chamber. Following four months of plant growth, P. occultum showed higher AMF hyphal growth in roots, soils and nylon mesh than D. epigaea or R. intraradices. Mycorrhizal inoculation improved the plant growth performance and increased easily extractable GRSP (EE-GRSP) and difficultly extractable GRSP (DE-GRSP) concentrations in both chambers, regardless of AMF species. The C content observed in total GRSP of the soil after purification was 2.71 ± 0.49 mg g−1, while purified EE-GRSP and DE-GRSP showed the C content of 1.01 ± 0.19 mg g−1 and 1.70 ± 010.34 mg g−1, respectively. The C contribution by purified EE-GRSP and DE-GRSP accounted for 8.67 ± 0.95% and 14.59 ± 2.21%, respectively, of total SOC, with a total C contribution of purified GRSPs accounting for 23.26 ± 2.67% of total SOC. A significantly higher C content of GRSP and the C contribution of GRSP to SOC were observed in DE-GRSP than in EE-GRSP, as well as the soil of the roots + hyphae chamber than the soil of the hyphae chamber. The proportionate distribution of water-stable aggregate in 2–4 and 1–2 mm sizes and their stability were higher under AMF hyphae than under non-AMF hyphae. This study thus provided a database evidence of increased contribution of GRSP towards build-up of SOC in response to mycorrhizal symbiosis. • GRSP contribution to SOC in trifoliate orange was studied. • The C content of EE-GRSP and DE-GRSP was 1.01 ± 0.19 mg g-1 and 1.70 ± 0.34 mg g. • C contribution of total GRSPs achieved 23.26 ± 2.67% of total SOC. • C content of GRSP and C contribution of GRSP to SOC were higher in DE-GRSP than in EE-GRSP. [ABSTRACT FROM AUTHOR]

• This study analyzed how PAs were involved in AMF-enhanced drought tolerance of plants. • AM trifoliate orange had higher Put and Cad and lower Spd and Spm under drought. • Mycorrhizal plants had higher activity of PA catabolic enzymes and Put synthetase. • AM plants showed overexpression of PtSPMS , PtPAO , PtSOD , and PtCAT1 under DS. • AM plants represented down-regulated expression of PtCuAO under DS. Arbuscular mycorrhizas enhance the drought tolerance of host plants through several underlying mechanisms. Polyamines (PAs) are known to protect plant cells from the damages of drought by enhancing the antioxidant defense system, restricting ethylene synthesis, maintaining cell pH and ion homeostasis, preventing chlorophyll loss, and so on; concomitantly, the mechanism through which arbuscular mycorrhizal (AM) fungi modulate PA metabolism to enhance drought tolerance of plants remains unclear. A pot experiment was conducted to assess how an AM fungus, Funneliformis mosseae , affects root PA homeostasis, activities and gene expressions of PA-related synthesizing and degrading enzymes, hydrogen peroxide (H 2 O 2 , a PA metabolite) production, and antioxidant enzyme gene expressions in trifoliate orange (Poncirus trifoliata) exposed to drought stress (DS). AM seedlings showed higher growth traits, leaf water potential, two plasma membrane intrinsic protein aquaporin gene expressions, and chlorophyll concentrations than non-AM seedlings under well-watered (WW) and DS conditions. Mycorrhizal treatment induced higher putrescine and cadaverine but lower spermidine and spermine concentrations, with higher activity of PA catabolic enzymes (copper-containing diamine oxidase, CuAO; polyamine oxidase, PAO) and putrescine synthases (ornithine decarboxylaseby; arginine decarboxylase, ADC). Mycorrhizas up-regulated the expression of the spermine synthase gene, PtSPMS , under DS, and down-regulated the transcript levels of PA catabolic enzyme genes (PtCuAO1 , PtCuAO2 , PtCuAO6 , and PtCuAO8) and PA synthase genes (PtADC1 and PtADC2) under DS. PtPAO1 , PtPAO2 , and PtPAO3 had higher expression levels in AMF-inoculated seedlings, as compared to non-AMF-inoculated seedlings, under DS, triggering reactive oxygen species-related signalling for stress responsiveness through low H 2 O 2 levels by up-regulating the expression of PtMn-SOD , PtCu/Zn-SOD, and PtCAT1. This study demonstrated that mycorrhizas have the capacity to modulate PA metabolism to enhance the drought tolerance of plants. [ABSTRACT FROM AUTHOR]