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Introduction: The application of mineral fertilizers deteriorates soil properties and affects crop yield and nutritional properties. However, plant growthpromoting microorganisms (PGPM-Serendipita indica, phosphorus solubilizing bacteria (PSB), and vesicular arbuscular mycorrhizae (VAM)) have great potential to reduce fertilizers and improve soil fertility, crop yield, and nutrient uptake and mitigate the environmental effect of mineral fertilizers. Material and methods: Hence, a field experiment was conducted involving nine treatments to evaluate the effects of PGPM along with 50% or 100% of the recommended dose of fertilizers on plant growth, soil fertility, nutrient uptake, and onion productivity. Results and discussion: Results indicated that 100% RDF combined with S. indica or PSB led to improved plant growth, and higher nutrient concentrations in both leaves and bulbs of onions compared to RDF alone. Moreover, the application of 100% RDF with S. indica increased total dry matter yield by 11.5% and 7.6% in the 2018-2019 and 2019-2020 seasons, respectively, compared to 100% RDF alone. This treatment also resulted in the highest nutrient uptake, with N uptake increasing by 6.9%-29.9%, P by 13.7%-21.7%, K by 20.0%-23.7%, and S by 18.1%-23.4%. Additionally, the combination of 100% RDF with S. indica inoculation led to a notable increase in bulb yield, with increments of 16.2% and 13.9% observed in 2018-2019 and 2019-2020, respectively, compared to 100% RDF alone. Similarly, the application of 100% RDF along with PSB inoculation resulted in an increase in bulb yield by 7.2% and 9.4% in the respective years. However, VAM did not exhibit satisfactory performance or improvements in the onion crop. Conclusion: Overall, the study suggests that combining 100% RDF with S. indica or PSB can enhance onion productivity and nutrient use efficiency. The present study may open a new avenue of PGPM application in enhancing onion yield and improving the bulb quality as well as soil health. However, field trials across different regions and soil types are necessary to validate these findings for practical adoption by farmers. [ABSTRACT FROM AUTHOR]

Arsenic (As) is a highly toxic metal that interferes with plant growth and disrupts various biochemical and molecular processes in plants. In this study, the harmful effects of As on rice were mitigated using combined inoculation of a root endophyte Serendipita indica and an actinobacterium Zhihengliuella sp. ISTPL4. A randomized experiment was conducted, in which rice plants were grown under controlled conditions and As-stressed conditions. The control and treatment groups consisted of untreated and non-stressed plants (C1), treated and non-stressed plants (C2), stressed and untreated plants (T1), and stressed and treated plants (T2). Various phenotypic characteristics such as shoot length (SL), root length (RL), shoot fresh weight (SFW), root fresh weight (RFW), shoot dry weight (SDW), and root dry weight (RDW) and biochemical parameters such as chlorophyll content, protein content, and antioxidant enzymatic activities were evaluated. The activity of various antioxidant enzymes was increased in T2 followed by T1 plants. Furthermore, high concentrations of phytohormones such as ethylene (ET), gibberellic acid (GA), and cytokinin (CK) were found at 4.11 μmol mg-1, 2.53 μmol mg-1, and 3.62 μmol mg-1 of FW of plant, respectively. The results of AAS indicated an increased As accumulation in roots of T2 plants (131.5 mg kg-1) than in roots of T1 plants (120 mg kg-1). It showed that there was an increased As accumulation and sequestration in roots of microbial-treated plants (T2) than in uninoculated plants (T1). Our data suggest that this microbial combination can be used to reduce the toxic effects of As in plants by increasing the activity of antioxidant enzymes such as SOD, CAT, PAL, PPO and POD. Furthermore, rice plants can withstand As stress owing to the active synthesis of phytohormones in the presence of microbial combinations. [ABSTRACT FROM AUTHOR]

Programmed cell death occurring during plant development (dPCD) is a fundamental process integral for plant growth and reproduction. Here, we investigate the connection between developmentally controlled PCD and fungal accommodation in Arabidopsis thaliana roots, focusing on the root cap-specific transcription factor ANAC033/SOMBRERO (SMB) and the senescence-associated nuclease BFN1. Mutations of both dPCD regulators increase colonization by the beneficial fungus Serendipita indica, primarily in the differentiation zone. smb-3 mutants additionally exhibit hypercolonization around the meristematic zone and a delay of S. indica-induced root-growth promotion. This demonstrates that root cap dPCD and rapid post-mortem clearance of cellular corpses represent a physical defense mechanism restricting microbial invasion of the root. Additionally, reporter lines and transcriptional analysis revealed that BFN1 expression is downregulated during S. indica colonization in mature root epidermal cells, suggesting a transcriptional control mechanism that facilitates the accommodation of beneficial microbes in the roots.

Main conclusion: Upon systemic S. indica colonization in split-root system cyst and root-knot nematodes benefit from endophyte-triggered carbon allocation and altered defense responses what significantly facilitates their development in A. thaliana. Serendipita indica is an endophytic fungus that establishes mutualistic relationships with different plants including Arabidopsis thaliana. It enhances host’s growth and resistance to different abiotic and biotic stresses such as infestation by the cyst nematode Heterodera schachtii (CN). In this work, we show that S. indica also triggers similar direct reduction in development of the root-knot nematode Meloidogyne javanica (RKN) in A. thaliana. Further, to mimick the natural situation occurring frequently in soil where roots are unequally colonized by endophytes we used an in vitro split-root system with one half of A. thaliana root inoculated with S. indica and the other half infected with CN or RKN, respectively. Interestingly, in contrast to direct effects, systemic effects led to an increase in number of both nematodes. To elucidate this phenomenon, we focused on sugar metabolism and defense responses in systemic non-colonized roots of plants colonized by S. indica. We analyzed the expression of several SUSs and INVs as well as defense-related genes and measured sugar pools. The results show a significant downregulation of PDF1.2 as well as slightly increased sucrose levels in the non-colonized half of the root in three-chamber dish. Thus, we speculate that, in contrast to direct effects, both nematode species benefit from endophyte-triggered carbon allocation and altered defense responses in the systemic part of the root, which promotes their development. With this work, we highlight the complexity of this multilayered tripartite relationship and deliver new insights into sugar metabolism and plant defense responses during S. indica–nematode–plant interaction. [ABSTRACT FROM AUTHOR]

In recent years, recurrent droughts have significantly affected spring barley production, reducing the quantity and quality of grain. This study aims to identify genotype-specific traits and the drought resilience of six different Hordeum vulgare L. (spring barley) genotypes, while also examining the potential of potassium application and fungal endophyte Serendipita indica inoculation to mitigate the negative effects of dry periods during the growing season. Field experiments were conducted over a three-year period from 2020 to 2022, measuring physiological, growth, and yield parameters. To get insight into the physiological state of the plants, we measured the soluble sugars content and the ratio of stable carbon isotopes in the flag leaf tissue, which reflects conditions during its formation. The dominant factors that influenced the measured parameters were the genotypes and seasons, as well as their interaction, rather than other experimental factors. The results showed that the Spitfire and Accordine varieties were the best performing in both the 2020 and 2021 seasons, as indicated by their yield. However, in the drier 2022 season, the yield of these two varieties decreased significantly (to 55% for Spitfire and to 69% for Accordine of their yield in 2021), while for the arid-region genotypes, it remained at the same level as the previous year. This study sheds light on the potential of various genotypes to withstand periods of drought and the effectiveness of using potassium application and S. indica inoculation as mitigation approaches. [ABSTRACT FROM AUTHOR]

IntroductionThe application of mineral fertilizers deteriorates soil properties and affects crop yield and nutritional properties. However, plant growth-promoting microorganisms (PGPM- Serendipita indica, phosphorus solubilizing bacteria (PSB), and vesicular arbuscular mycorrhizae (VAM)) have great potential to reduce fertilizers and improve soil fertility, crop yield, and nutrient uptake and mitigate the environmental effect of mineral fertilizers.Material and methodsHence, a field experiment was conducted involving nine treatments to evaluate the effects of PGPM along with 50% or 100% of the recommended dose of fertilizers on plant growth, soil fertility, nutrient uptake, and onion productivity.Results and discussionResults indicated that 100% RDF combined with S. indica or PSB led to improved plant growth, and higher nutrient concentrations in both leaves and bulbs of onions compared to RDF alone. Moreover, the application of 100% RDF with S. indica increased total dry matter yield by 11.5% and 7.6% in the 2018-2019 and 2019-2020 seasons, respectively, compared to 100% RDF alone. This treatment also resulted in the highest nutrient uptake, with N uptake increasing by 6.9%-29.9%, P by 13.7%-21.7%, K by 20.0%-23.7%, and S by 18.1%-23.4%. Additionally, the combination of 100% RDF with S. indica inoculation led to a notable increase in bulb yield, with increments of 16.2% and 13.9% observed in 2018-2019 and 2019-2020, respectively, compared to 100% RDF alone. Similarly, the application of 100% RDF along with PSB inoculation resulted in an increase in bulb yield by 7.2% and 9.4% in the respective years. However, VAM did not exhibit satisfactory performance or improvements in the onion crop.ConclusionOverall, the study suggests that combining 100% RDF with S. indica or PSB can enhance onion productivity and nutrient use efficiency. The present study may open a new avenue of PGPM application in enhancing onion yield and improving the bulb quality as well as soil health. However, field trials across different regions and soil types are necessary to validate these findings for practical adoption by farmers.

Fungal-bacterial combinations have a significant role in increasing and improving plant health under various stress conditions. Metabolites secreted by fungi and bacteria play an important role in this process. Our study emphasizes the significance of secondary metabolites secreted by the fungus Serendipita indica alone and by an actinobacterium Zhihengliuella sp. ISTPL4 under normal growth conditions and arsenic (As) stress condition. Here, we evaluated the arsenic tolerance ability of S. indica alone and in combination with Z. sp. ISTPL4 under in vitro conditions. The growth of S. indica and Z. sp. ISTPL4 was measured in varying concentrations of arsenic and the effect of arsenic on spore size and morphology of S. indica was determined using confocal microscopy and scanning electron microscopy. The metabolomics study indicated that S. indica alone in normal growth conditions and under As stress released pentadecanoic acid, glycerol tricaprylate, L-proline and cyclo(L-prolyl-L-valine). Similarly, d-Ribose, 2-deoxy-bis(thioheptyl)-dithioacetal were secreted by a combination of S. indica and Z. sp. ISTPL4. Confocal studies revealed that spore size of S. indica decreased by 18% at 1.9 mM and by 15% when in combination with Z. sp. ISTPL4 at a 2.4 mM concentration of As. Arsenic above this concentration resulted in spore degeneration and hyphae fragmentation. Scanning electron microscopy (SEM) results indicated an increased spore size of S. indica in the presence of Z. sp. ISTPL4 (18 ± 0.75 µm) compared to S. indica alone (14 ± 0.24 µm) under normal growth conditions. Our study concluded that the suggested combination of microbial consortium can be used to increase sustainable agriculture by combating biotic as well as abiotic stress. This is because the metabolites released by the microbial combination display antifungal and antibacterial properties. The metabolites, besides evading stress, also confer other survival strategies. Therefore, the choice of consortia and combination partners is important and can help in developing strategies for coping with As stress. [ABSTRACT FROM AUTHOR]

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Arsenic (As) is a highly toxic metal that interferes with plant growth and disrupts various biochemical and molecular processes in plants. In this study, the harmful effects of As on rice were mitigated using combined inoculation of a root endophyte Serendipita indica and an actinobacterium Zhihengliuella sp. ISTPL4. A randomized experiment was conducted, in which rice plants were grown under controlled conditions and As-stressed conditions. The control and treatment groups consisted of untreated and non-stressed plants (C1), treated and non-stressed plants (C2), stressed and untreated plants (T1), and stressed and treated plants (T2). Various phenotypic characteristics such as shoot length (SL), root length (RL), shoot fresh weight (SFW), root fresh weight (RFW), shoot dry weight (SDW), and root dry weight (RDW) and biochemical parameters such as chlorophyll content, protein content, and antioxidant enzymatic activities were evaluated. The activity of various antioxidant enzymes was increased in T2 followed by T1 plants. Furthermore, high concentrations of phytohormones such as ethylene (ET), gibberellic acid (GA), and cytokinin (CK) were found at 4.11 μmol mg−1, 2.53 μmol mg−1, and 3.62 μmol mg−1 of FW of plant, respectively. The results of AAS indicated an increased As accumulation in roots of T2 plants (131.5 mg kg−1) than in roots of T1 plants (120 mg kg−1). It showed that there was an increased As accumulation and sequestration in roots of microbial-treated plants (T2) than in uninoculated plants (T1). Our data suggest that this microbial combination can be used to reduce the toxic effects of As in plants by increasing the activity of antioxidant enzymes such as SOD, CAT, PAL, PPO and POD. Furthermore, rice plants can withstand As stress owing to the active synthesis of phytohormones in the presence of microbial combinations.

El aporte de nutrimentos al cultivo de maíz en México es principalmente con fertilizantes químicos, sin embargo, el precio de estos insumos requiere buscar otras alternativas para abatir los costos, sin disminuir la producción. El objetivo de este trabajo fue determinar el efecto individual y combinado de Azospirillum brasilense, Bacillus subtilis y Serendipita indica (ocho tratamientos) en la producción de maíz. El cultivo se estableció en Cieneguillas, Zacatecas, en condiciones de invernadero, en el año 2022. La inoculación de los microorganismos se realizó al momento de la siembra. Los ocho tratamientos con los microorganismos se evaluaron en un diseño experimental en bloques al azar con cuatro repeticiones. Con la co-inoculación de A. brasilense + S. indica se incrementó el rendimiento de toda la planta y el mayor rendimiento de grano con A. brasilense + B. subtilis, este último incremento, respecto al testigo, fue de 40%.

The main objective of this study was to investigate the influence of Serendipita indica on the growth of Tartary buckwheat plants. This study highlighted that the roots of Tartary buckwheat can be colonized by S. indica and that this fungal endophyte improved plants height, fresh weight, dry weight, and grain yield. In the meantime, the colonization of S. indica in Tartary buckwheat leaves resulted in elevated levels of photosynthesis, plant hormone content, antioxidant enzyme activity, proline content, chlorophyll content, soluble sugars, and protein content. Additionally, the introduction of S. indica to Tartary buckwheat roots led to a substantial rise in the levels of flavonoids and phenols found in the leaves and seeds of Tartary buckwheat. In addition, S. indica colonization reduced the content of malondialdehyde and hydrogen peroxide when compared to non-colonized plants. Importantly, the drought tolerance of Tartary buckwheat plants is increased, which benefits from physiology and bio-chemical changes in plants after S. indica colonized. In conclusion, we have shown that S. indica can improve systematic resistance and promote the growth of Tartary buckwheat by enhancing the photosynthetic capacity of Tartary buckwheat, inducing the production of IAA, increasing the content of secondary metabolites such as total phenols and total flavonoids, and improving the antioxidant enzyme activity of the plant. [ABSTRACT FROM AUTHOR]

In recent years, recurrent droughts have significantly affected spring barley production, reducing the quantity and quality of grain. This study aims to identify genotype-specific traits and the drought resilience of six different Hordeum vulgare L. (spring barley) genotypes, while also examining the potential of potassium application and fungal endophyte Serendipita indica inoculation to mitigate the negative effects of dry periods during the growing season. Field experiments were conducted over a three-year period from 2020 to 2022, measuring physiological, growth, and yield parameters. To get insight into the physiological state of the plants, we measured the soluble sugars content and the ratio of stable carbon isotopes in the flag leaf tissue, which reflects conditions during its formation. The dominant factors that influenced the measured parameters were the genotypes and seasons, as well as their interaction, rather than other experimental factors. The results showed that the Spitfire and Accordine varieties were the best performing in both the 2020 and 2021 seasons, as indicated by their yield. However, in the drier 2022 season, the yield of these two varieties decreased significantly (to 55% for Spitfire and to 69% for Accordine of their yield in 2021), while for the arid-region genotypes, it remained at the same level as the previous year. This study sheds light on the potential of various genotypes to withstand periods of drought and the effectiveness of using potassium application and S. indica inoculation as mitigation approaches.

Plant-endophyte symbiosis influences plant defense and growth. Serendipita indica is a root endophyte that promotes growth and induces tolerance against biotic and abiotic stress in plants. In this study, we examined the effect of S. indica colonization on herbivore (Spodoptera exigua) resistance of onion (Allium cepa L.). We found that colonization of S. indica in the roots of onion significantly reduced the feeding damage of leaves by S. exigua larvae, and also resulted in a reduction in weight gain of the larvae when fed on S. indica plants. This enhanced resistance is a result of modulation of antioxidant and defense enzymes/genes in the host by S. indica mutualism. Specifically, the activities of enzymes such as Superoxide dismutase, peroxidase, polyphenol oxidase, phenylalanine ammonia-lyase, and H2O2 content were significantly higher in the early stages of S. exigua feeding in the S. indica colonized plants compared to the non-colonized counterparts. Similarly, defense genes also showed modulation in response to this tripartite interaction of onion -S. indica mutualism and S. exigua herbivory. The hierarchical cluster analysis and principal component analysis indicated a clear difference in the onion biochemical responses, which is due to the S. indica symbiosis. Our investigation demonstrates that onion-S. indica symbiosis significantly decreases chewing injury by efficiently modulating antioxidant and defense enzyme activities and gene expression in response to S. exigua herbivory. Therefore, S. indica can be used as a potential biocontrol agent for sustainable management of this important pest of Alliums. [ABSTRACT FROM AUTHOR]

Simple Summary: Boron is an essential micronutrient for plants; however, boron toxicity has long been identified as a common nutritional problem on a global scale and is attracting increasing interest. Excessive concentrations of B significantly reduce crop yield, including that of wheat, which is a staple food throughout the world. Wheat cultivation is particularly common in arid and semi-arid regions where climate change and irrigation can aggravate the risk and severity of B toxicity. Serendipita indica is a versatile root endophytic fungus. Studies show that plants colonized by Serendipita indica exhibit a wide range of beneficial effects, encompassing enhanced growth and yield, higher tolerance to various biotic and abiotic stresses, and improved water and nutrient balance. The present study revealed that under B toxicity, the growth, membrane integrity, tissue P content, and yield of wheat can be significantly improved by this endophyte without reducing toxic B concentrations in the shoot. Considering that Serendipita indica can be axenically produced on a large scale, this endophyte can be used as a microbiological agent for promoting sustainable agriculture under adverse soil conditions like boron toxicity. Further studies are needed to evaluate the potential and specific mechanisms of Serendipita indica under field conditions. While the importance of beneficial soil microorganisms for soil health and crop performance has been receiving ever-increasing attention, Serendipita indica has been widely studied as a fungal root endophyte with significant potential for increasing the stress tolerance of host plants. Boron (B) toxicity as an adverse soil condition is particularly prevalent in arid and semi-arid regions and threatens crop production. Studies on S. indica-wheat symbiosis are limited, and effects of S. indica on crops have never been reported in the context of B toxicity. Here, two pot experiments were conducted under greenhouse conditions to investigate the effects of S. indica on the growth and yield parameters of bread (Triticum aestivum) and durum wheat (Triticum durum) grown at different levels of B toxicity in native vs. sterilized soil, and parameters related to root colonization, membrane damage, oxidative stress, chlorophyll, and mineral nutrition were measured to elucidate the physiological mechanisms of damage and benefit. Boron toxicity decreased early vegetative growth and grain yield, but it did not affect the straw dry weight of mature plants, whereas S. indica significantly enhanced the vegetative growth, straw dry weight, and the grain number of both wheat species. Membrane damage as demonstrated by increased lipid peroxidation and relative electrolyte leakage was caused by B toxicity and alleviated by S. indica. The benefits provided by S. indica could not be attributed to any significant changes in tissue concentrations of B or other minerals such as phosphorus. Soil sterilization generally improved plant performance but it did not consistently strengthen or weaken the effects of S. indica. The presented results suggest that S. indica may be used as an effective microbial inoculant to enhance wheat growth under adverse soil conditions such as B toxicity through mechanisms that are possibly unrelated to mineral homeostasis. [ABSTRACT FROM AUTHOR]

Serendipita indica, a multifunctional and useful endophyte fungus, has been intensively investigated in promoting plant growth and resistance towards biotic and abiotic stress. Multiple chitinases from microorganisms or plants have been identified to have a high antifungal activity as a biological control. However, chitinase of S. indica still needs to be characterized. We functionally characterized a chitinase (SiChi) in S. indica. The result showed that the purified SiChi protein confers high chitinase activity; importantly, SiChi inhibits the conidial germination of Magnaporthe oryzae and Fusarium moniliforme. After the successful colonization of rice roots by S. indica, both the rice blast disease and bakanae disease were significantly reduced. Interestingly, the purified SiChi could promptly induce rice disease resistance towards M. oryzae and F. moniliforme pathogens when sprayed on rice leaves. Like S. indica, SiChi could upregulate rice pathogen‐resistant proteins and defense enzymes. In conclusion, chitinase of S. indica has direct antifungal activity and indirect induced resistance activity, implying an efficient and economic strategy for rice disease control by applying S. indica and SiChi. [ABSTRACT FROM AUTHOR]

Rapid urbanization and globalization demand increasing agricultural productivity. Soil nutrient supply capacity is continuously decreasing due to soil erosion, degradation, salt deposition, undesired element, metal deposition, water scarcity, and an uneven nutrient delivery system. Rice cultivation requires a large amount of water which is becoming detrimental due to these activities. There is a need to increase its productivity. Microbial inoculants are becoming increasingly important in achieving sustainable agricultural production systems. The current study was conducted to investigate the interaction between the root endophytic fungus Serendipita indica (S. indica) and the actinobacterium Zhihengliuella sp. ISTPL4 (Z. sp. ISTPL4) and their synergistic effects on the growth of rice (Oryza sativa L). Both S. indica and Z. sp. ISTPL4 showed positive interactions. Growth of S. indica was observed at different days after Z. sp. ISTPL4 inoculation, and stimulated growth of S. indica was observed when Z. sp. ISTPL4 was inoculated at 5 dafi (days after fungal inoculation). Z. sp. ISTPL4 promoted the growth of S. indica as it increased spore germination. Furthermore, confocal and scanning electron microscopy (SEM) analyses showed a 27% increase in the spore size of S. indica in the presence of Z. sp. ISTPL4. In a liquid chromatography with tandem mass spectrometry (LC-MS/MS) analysis increased production of alanine and glutamic acid was observed in their sequential co-culture as compared with individual cultures. Sequential inoculation of S. indica and Z. sp. ISTPL4 significantly increased the biochemical and physical characteristics of rice as compared with their individual inoculum. Biochemical parameters such as chlorophyll content, total soluble sugar, and flavonoid content in the rice increased by up to 57%, 47%, and 39%, respectively, in the presence of the combined inoculum of S. indica and Z. sp. ISTPL4. This will be the first study, to the best of our knowledge, which shows the fungus and actinobacterium interaction and their synergistic roles in the growth promotion of rice. Furthermore, this novel combination can also be used to boost the growth of other crops to increase the agricultural yield. [ABSTRACT FROM AUTHOR]

Purpose Chicken eggshell as a byproduct of the egg product industry makes serious problems to the environment. The aim of this study was to evaluate the effect of eggshell as a potential option in supplementing of the organic matter and improvement of soil conditions in the pot culture and the area. Method Two pot experiments were carried out to estimate the effects of CD and CDES vermicomposts provided from cow dung and cow dung + eggshell’s powder, respectively, on symbiotic relationships of Serendipita indica with wheat and soil enzyme activities tested under sterilized and non-sterilized soil conditions, respectively. Two in vitro experiments were also conducted to test the effects of eggshell extract and humic and fulvic acids extracted from CD vermicompost on S. indica growth.Results CDES vermicompost improved soil enzyme activities of urease, phosphatase and invertase. Root colonization of wheat with multiple species of mycorrhiza and facultative symbiont S. indica was improved by CDES. S. indica growth was induced by eggshell extract. Humic and fulvic acids increased in S. indica mycelia mats dry yield. Results revealed when cow dung was amended with eggshell powder provided a better condition for earthworm growth and enhancing the colonization percentage of wheat root with indigenous mycorrhizal fungi. CDES vermicompost caused the increase in soil basal respiration and soil enzymatic activities.Conclusion Eggshell powder growth promoting effect can be attributed to the biologically active compound that exists in eggshell extract.

SNAREs (soluble N-ethylmaleimide-sensitive-factor attachment protein receptors) are engines for almost all of the membrane fusion and exocytosis events in organism cells. In this study, we identified 84 SNARE genes from banana (Musa acuminata). Gene expression analysis revealed that the expression of MaSNAREs varied a lot in different banana organs. By analyzing their expression patterns under low temperature (4 °C), high temperature (45 °C), mutualistic fungus (Serendipita indica, Si) and fungal pathogen (Fusarium oxysporum f. sp. Cubense Tropical Race 4, FocTR4) treatments, many MaSNAREs were found to be stress responsive. For example, MaBET1d was up-regulate by both low and high temperature stresses; MaNPSN11a was up-regulated by low temperature but down-regulated by high temperature; and FocTR4 treatment up-regulated the expression of MaSYP121 but down-regulated MaVAMP72a and MaSNAP33a. Notably, the upregulation or downregulation effects of FocTR4 on the expression of some MaSNAREs could be alleviated by priorly colonized Si, suggesting that they play roles in the Si-enhanced banana wilt resistance. Foc resistance assays were performed in tobacco leaves transiently overexpressing MaSYP121, MaVAMP72a and MaSNAP33a. Results showed that transient overexpression of MaSYP121 and MaSNPA33a suppressed the penetration and spread of both Foc1 (Foc Race 1) and FocTR4 in tobacco leaves, suggesting that they play positive roles in resisting Foc infection. However, the transient overexpression of MaVAMP72a facilitated Foc infection. Our study can provide a basis for understanding the roles of MaSNAREs in the banana responses to temperature stress and mutualistic and pathogenic fungal colonization. [ABSTRACT FROM AUTHOR]

Serendipita indica is a root endophytic fungus and can symbiose with many plant species. Effector proteins secreted by fungi show many functions during symbiosis or infection. To investigate the role of the protein PIIN 05330 secreted by S. indica, the gene PIIN_05330 was cloned and its transgenic Arabidopsis lines were constructed. The transgenic lines were evaluated for their drought tolerance. The results showed that these transgenic Arabidopsis lines were sensitive to drought stress during the germination stage. Compared with Arabidopsis wild type (WT) and the empty vector (VT) lines seedlings, the transgenic lines had higher total root surface area, root diameters, total root volumes, relative root elongation rates, total root length, and root tip numbers. In addition, the transgenic lines showed higher aboveground biomass and chlorophyll levels. Compared with WT and VT lines, activities of SOD, POD and CAT increased in the transgenic Arabidopsis lines under drought condition, especially their CAT activities. Under drought stress, levels of AsA and GSH in the transgenic lines were higher than those in control WT and VT, and concentrations of H2O2 and MDA were significantly lower than those in control WT and VT. Under drought stress, proline concentrations increased, and total concentrations of soluble sugars and proteins significantly increased in transgenic lines, compared with control WT and VT, indicating that the transgenic lines had a higher ability to reduce plant damage and protect plant cell membrane system. All these results suggest that the effector protein PIIN_05330 improved drought tolerance of the transgenic Arabidopsis lines. [ABSTRACT FROM AUTHOR]

Plant-endophyte symbiosis influences plant defense and growth. Serendipita indica is a root endophyte that promotes growth and induces tolerance against biotic and abiotic stress in plants. In this study, we examined the effect of S. indica colonization on herbivore (Spodoptera exigua) resistance of onion (Allium cepa L.). We found that colonization of S. indica in the roots of onion significantly reduced the feeding damage of leaves by S. exigua larvae, and also resulted in a reduction in weight gain of the larvae when fed on S. indica plants. This enhanced resistance is a result of modulation of antioxidant and defense enzymes/genes in the host by S. indica mutualism. Specifically, the activities of enzymes such as Superoxide dismutase, peroxidase, polyphenol oxidase, phenylalanine ammonia-lyase, and H2O2 content were significantly higher in the early stages of S. exigua feeding in the S. indica colonized plants compared to the non-colonized counterparts. Similarly, defense genes also showed modulation in response to this tripartite interaction of onion -S. indica mutualism and S. exigua herbivory. The hierarchical cluster analysis and principal component analysis indicated a clear difference in the onion biochemical responses, which is due to the S. indica symbiosis. Our investigation demonstrates that onion-S. indica symbiosis significantly decreases chewing injury by efficiently modulating antioxidant and defense enzyme activities and gene expression in response to S. exigua herbivory. Therefore, S. indica can be used as a potential biocontrol agent for sustainable management of this important pest of Alliums.

Serendipita indica is an endophytic root symbiont fungus that enhances the growth of various plants under different stress conditions, including salinity. Here, the functional characterization of two fungal Na+/H+ antiporters, SiNHA1 and SiNHX1 has been carried out to study their putative role in saline tolerance. Although their gene expression does not respond specifically to saline conditions, they could contribute, together with the previously characterized Na+ efflux systems SiENA1 and SiENA5, to relieve Na+ from the S. indica cytosol under this stressed condition. In parallel, an in-silico study has been carried out to define its complete transportome. To further investigate the repertoire of transporters expressed in free-living cells of S. indica and during plant infection under saline conditions, a comprehensive RNA-seq approach was taken. Interestingly, SiENA5 was the only gene significantly induced under free-living conditions in response to moderate salinity at all the tested time points, revealing that it is one of the main salt-responsive genes of S. indica. In addition, the symbiosis with Arabidopsis thaliana also induced SiENA5 gene expression, but significant changes were only detected after long periods of infection, indicating that the association with the plant somehow buffers and protects the fungus against the external stress. Moreover, the significant and strongest induction of the homologous gene SiENA1 occurred during symbiosis, regardless the exposure to salinity. The obtained results suggest a novel and relevant role of these two proteins during the establishment and maintenance of fungus-plant interaction.

Fungal-bacterial combinations have a significant role in increasing and improving plant health under various stress conditions. Metabolites secreted by fungi and bacteria play an important role in this process. Our study emphasizes the significance of secondary metabolites secreted by the fungus Serendipita indica alone and by an actinobacterium Zhihengliuella sp. ISTPL4 under normal growth conditions and arsenic (As) stress condition. Here, we evaluated the arsenic tolerance ability of S. indica alone and in combination with Z. sp. ISTPL4 under in vitro conditions. The growth of S. indica and Z. sp. ISTPL4 was measured in varying concentrations of arsenic and the effect of arsenic on spore size and morphology of S. indica was determined using confocal microscopy and scanning electron microscopy. The metabolomics study indicated that S. indica alone in normal growth conditions and under As stress released pentadecanoic acid, glycerol tricaprylate, L-proline and cyclo(L-prolyl-L-valine). Similarly, d-Ribose, 2-deoxy-bis(thioheptyl)-dithioacetal were secreted by a combination of S. indica and Z. sp. ISTPL4. Confocal studies revealed that spore size of S. indica decreased by 18% at 1.9 mM and by 15% when in combination with Z. sp. ISTPL4 at a 2.4 mM concentration of As. Arsenic above this concentration resulted in spore degeneration and hyphae fragmentation. Scanning electron microscopy (SEM) results indicated an increased spore size of S. indica in the presence of Z. sp. ISTPL4 (18 ± 0.75 µm) compared to S. indica alone (14 ± 0.24 µm) under normal growth conditions. Our study concluded that the suggested combination of microbial consortium can be used to increase sustainable agriculture by combating biotic as well as abiotic stress. This is because the metabolites released by the microbial combination display antifungal and antibacterial properties. The metabolites, besides evading stress, also confer other survival strategies. Therefore, the choice of consortia and combination partners is important and can help in developing strategies for coping with As stress.

Rapid urbanization and globalization demand increasing agricultural productivity. Soil nutrient supply capacity is continuously decreasing due to soil erosion, degradation, salt deposition, undesired element, metal deposition, water scarcity, and an uneven nutrient delivery system. Rice cultivation requires a large amount of water which is becoming detrimental due to these activities. There is a need to increase its productivity. Microbial inoculants are becoming increasingly important in achieving sustainable agricultural production systems. The current study was conducted to investigate the interaction between the root endophytic fungus Serendipita indica (S. indica) and the actinobacterium Zhihengliuella sp. ISTPL4 (Z. sp. ISTPL4) and their synergistic effects on the growth of rice (Oryza sativa L). Both S. indica and Z. sp. ISTPL4 showed positive interactions. Growth of S. indica was observed at different days after Z. sp. ISTPL4 inoculation, and stimulated growth of S. indica was observed when Z. sp. ISTPL4 was inoculated at 5 dafi (days after fungal inoculation). Z. sp. ISTPL4 promoted the growth of S. indica as it increased spore germination. Furthermore, confocal and scanning electron microscopy (SEM) analyses showed a 27% increase in the spore size of S. indica in the presence of Z. sp. ISTPL4. In a liquid chromatography with tandem mass spectrometry (LC-MS/MS) analysis increased production of alanine and glutamic acid was observed in their sequential co-culture as compared with individual cultures. Sequential inoculation of S. indica and Z. sp. ISTPL4 significantly increased the biochemical and physical characteristics of rice as compared with their individual inoculum. Biochemical parameters such as chlorophyll content, total soluble sugar, and flavonoid content in the rice increased by up to 57%, 47%, and 39%, respectively, in the presence of the combined inoculum of S. indica and Z. sp. ISTPL4. This will be the first study, to the best of our knowledge, which shows the fungus and actinobacterium interaction and their synergistic roles in the growth promotion of rice. Furthermore, this novel combination can also be used to boost the growth of other crops to increase the agricultural yield.

In the current research, unique Nano-Embedded Fungus (NEF), made by the synergic association of silver nanoparticles (AgNPs) and endophytic fungus (Piriformospora indica), is studied, and the impact of NEF on black rice secondary metabolites is reported. AgNPs were synthesized by chemical reduction process using the temperature-dependent method and characterized for morphological and structural features through UV visible absorption spectroscopy, zeta potential, XRD, SEM-EDX, and FTIR spectroscopy. The NEF, prepared by optimizing the AgNPs concentration (300 ppm) in agar and broth media, showed better fungal biomass, colony diameter, spore count, and spore size than the control P. indica. Treatment with AgNPs, P. indica, and NEF resulted in growth enhancement in black rice. NEF and AgNPs stimulated the production of secondary metabolites in its leaves. The concentrations of chlorophyll, carotenoids, flavonoids, and terpenoids were increased in plants inoculated with P. indica and AgNPs. The findings of the study highlight the synergistic effect of AgNPs and the fungal symbionts in augmenting the secondary metabolites in leaves of black rice. [ABSTRACT FROM AUTHOR]

The presence of the exotic Eucalyptus tree in crop-growing soil and the accumulation of its undecomposed leaves is a significant ecological hazard. The waxy coating on the leaves and the phenolic compounds takes a long time to break down under normal conditions. It is necessary to explore various fungi that can degrade these leaves for an eco-friendly solution to this problem. In this study, spores of nine native Trichoderma strains were produced on wheat agar using a lactic acid-induced sporulation strategy (LAISS). Trichoderma biosustained spores and Serendipita indica (SI) spores were applied to a rice field with accumulated Eucalyptus leaves under continuous ponding (CP) and alternate flooding and wetting conditions (AFW). Among the strains, TI04 (Trichoderma viride) and TI15 (Trichoderma citrinoviride) showed faster (5days) and massive sporulation (1.06-1.38×1011 CFU/g) in LAISS. In vitro, TI04 and TI15 biosustained on Eucalyptus leaves and improved rice seedling growth and SI infection under greenhouse conditions. In the rice-field experiment, Trichoderma-treatment had a threefold yield (percentage) increase from control, with TI04 (CP) increasing the yield by 30.79, TI04 (AFW) by 29.45, TI15 (CP) by 32.72, and TI15 (AFW) rising by 31.91. Remarkably, unfilled grain yield significantly decreased in all the Trichoderma treatments. Under AFW conditions, TI04 and TI15 showed a higher pH increase. Furthermore, TI04 and TI15 under AFW had higher water productivity (tha-1 cm-1) of 0.0763 and 0.0791, respectively, and the highest rates (percentage) of SI colonization of 86.36 and 83.16, respectively. According to the findings, LAISS-produced Trichoderma spores can be applied to break down persistent wastes and restore agricultural ecosystems through increased mycorrhizae networking. [ABSTRACT FROM AUTHOR]

Modulating bacterial exopolysaccharides (EPS) with nanoparticles is an emerging application of nanotechnology due to the enhancement of soil fertility, encouragement of nutrient uptake, and plant growth in agriculture. This study highlights the seamless integration of nanotechnology with EPS, the fungus, Serendipita indica strain is used here to reduce selenium ions and forms selenium nanoparticles (Se NPs). Se NPs has been characterized by visual observation, UV–vis spectroscopy, FTIR, TEM, SEM-EDX, XRD, DLS and Zeta potential. Production of EPS was monitored in the presence of Se NPs. Extracted EPS was characterized by FTIR, SEM-EDX, HPLC, and NMR. Results exhibit the successful formation of spherical, crystalline Se NPs with size ranging between 20-80 nm with 92.24% selenium. Response surface methodology (RSM) was used to identify and optimise the production of exopolysaccharides (EPS) from Bacillus subtilis that was isolated from arid zone agricultural soil. The central composite rotatable design (CCRD) exhibited the highest yield of EPS 11.332 g/l. EPS was then characterized by FTIR, HPLC, NMR, and SEM which revealed the presence of carboxyl and hydroxyl groups and flakes-like structure of EPS. Hence, the present work will pave the way for the application of nanotechnology for increasing EPS production and will also offer new dimensions for further research. [Display omitted] • Mycosynthesis and characterization of Selenium Nanoparticles (Se NPs). • Impact of Se NPs on microbial exopolysaccharide production. • Characterization of exopolysaccharides extracted from arid zone microbe. [ABSTRACT FROM AUTHOR]

The main objective of this study was to investigate the influence of Serendipita indica on the growth of Tartary buckwheat plants. This study highlighted that the roots of Tartary buckwheat can be colonized by S. indica and that this fungal endophyte improved plants height, fresh weight, dry weight, and grain yield. In the meantime, the colonization of S. indica in Tartary buckwheat leaves resulted in elevated levels of photosynthesis, plant hormone content, antioxidant enzyme activity, proline content, chlorophyll content, soluble sugars, and protein content. Additionally, the introduction of S. indica to Tartary buckwheat roots led to a substantial rise in the levels of flavonoids and phenols found in the leaves and seeds of Tartary buckwheat. In addition, S. indica colonization reduced the content of malondialdehyde and hydrogen peroxide when compared to non-colonized plants. Importantly, the drought tolerance of Tartary buckwheat plants is increased, which benefits from physiology and bio-chemical changes in plants after S. indica colonized. In conclusion, we have shown that S. indica can improve systematic resistance and promote the growth of Tartary buckwheat by enhancing the photosynthetic capacity of Tartary buckwheat, inducing the production of IAA, increasing the content of secondary metabolites such as total phenols and total flavonoids, and improving the antioxidant enzyme activity of the plant.

The presence of the exotic Eucalyptus tree in crop-growing soil and the accumulation of its undecomposed leaves is a significant ecological hazard. The waxy coating on the leaves and the phenolic compounds takes a long time to break down under normal conditions. It is necessary to explore various fungi that can degrade these leaves for an eco-friendly solution to this problem. In this study, spores of nine native Trichoderma strains were produced on wheat agar using a lactic acid-induced sporulation strategy (LAISS). Trichoderma biosustained spores and Serendipita indica (SI) spores were applied to a rice field with accumulated Eucalyptus leaves under continuous ponding (CP) and alternate flooding and wetting conditions (AFW). Among the strains, TI04 (Trichoderma viride) and TI15 (Trichoderma citrinoviride) showed faster (5 days) and massive sporulation (1.06–1.38 × 1011 CFU/g) in LAISS. In vitro, TI04 and TI15 biosustained on Eucalyptus leaves and improved rice seedling growth and SI infection under greenhouse conditions. In the rice-field experiment, Trichoderma-treatment had a threefold yield (percentage) increase from control, with TI04 (CP) increasing the yield by 30.79, TI04 (AFW) by 29.45, TI15 (CP) by 32.72, and TI15 (AFW) rising by 31.91. Remarkably, unfilled grain yield significantly decreased in all the Trichoderma treatments. Under AFW conditions, TI04 and TI15 showed a higher pH increase. Furthermore, TI04 and TI15 under AFW had higher water productivity (t ha−1 cm−1) of 0.0763 and 0.0791, respectively, and the highest rates (percentage) of SI colonization of 86.36 and 83.16, respectively. According to the findings, LAISS-produced Trichoderma spores can be applied to break down persistent wastes and restore agricultural ecosystems through increased mycorrhizae networking.

• Piriformospora indica (Serendipita indica) is a novel cultivable symbiont involved in plant growth enhancement and stress resistance. • EF-hand motif containing Ca2+-binding proteins play vital role in plant growth and development. • The protein-protein interaction network analysis revealed the involvement of Pi EF-hand in important cellular processes. EF-hand domain proteins are Ca2+ binding proteins with helix-loop-helix structural pattern and classified as CaMs, CMLs, CBLs, CDPKs, Rbohs, Myosin light chains (Mlcs) and others. Mlcs are important proteins that function through binding with IQ motifs of Myosins and respond to changes in Ca2+ ion concentration. Present study reports in silico analysis of Piriformospora indica EF-hand (Pi EF-hand) protein annotated as probable myosin regulatory light chain Cdc4. The genomic analysis revealed sequence & structure attributes, orthologs identification, evolutionary perspective, protein-protein interactions and functionality. Pi EF-hand protein is 141 aa long acidic protein lacking cysteine. Structural analysis highlighted four EF-hand motifs and conservation of Myosin heavy chain IQ motif interacting sites. Further, evolutionary relationships across 79 eukaryotic organisms were drawn classifying orthologs in two groups- CaMs and Mlcs. The Pi EF-hand protein is grouped along with Mlcs underlining its possible role as myosin light chain. Separate sequence logos of each EF-hand motifs for CaMs and Mlcs revealed poor sequence conservation among Mlcs, whereas, CaMs EF-hand motifs were well conserved. The protein-protein interaction network analysis glimpsed involvement of Pi EF-hand protein in vesicular trafficking and polarized fungal hyphal growth. [ABSTRACT FROM AUTHOR]

Purpose Chicken eggshell as a byproduct of the egg product industry makes serious problems to the environment. The aim of this study was to evaluate the effect of eggshell as a potential option in supplementing of the organic matter and improvement of soil conditions in the pot culture and the area. Method Two pot experiments were carried out to estimate the effects of CD and CDES vermicomposts provided from cow dung and cow dung + eggshell's powder, respectively, on symbiotic relationships of Serendipita indica with wheat and soil enzyme activities tested under sterilized and non-sterilized soil conditions, respectively. Two in vitro experiments were also conducted to test the effects of eggshell extract and humic and fulvic acids extracted from CD vermicompost on S. indica growth. Results CDES vermicompost improved soil enzyme activities of urease, phosphatase and invertase. Root colonization of wheat with multiple species of mycorrhiza and facultative symbiont S. indica was improved by CDES. S. indica growth was induced by eggshell extract. Humic and fulvic acids increased in S. indica mycelia mats dry yield. Results revealed when cow dung was amended with eggshell powder provided a better condition for earthworm growth and enhancing the colonization percentage of wheat root with indigenous mycorrhizal fungi. CDES vermicompost caused the increase in soil basal respiration and soil enzymatic activities. Conclusion Eggshell powder growth promoting effect can be attributed to the biologically active compound that exists in eggshell extract. [ABSTRACT FROM AUTHOR]

While the importance of beneficial soil microorganisms for soil health and crop performance has been receiving ever-increasing attention, Serendipita indica has been widely studied as a fungal root endophyte with significant potential for increasing the stress tolerance of host plants. Boron (B) toxicity as an adverse soil condition is particularly prevalent in arid and semi-arid regions and threatens crop production. Studies on S. indica-wheat symbiosis are limited, and effects of S. indica on crops have never been reported in the context of B toxicity. Here, two pot experiments were conducted under greenhouse conditions to investigate the effects of S. indica on the growth and yield parameters of bread (Triticum aestivum) and durum wheat (Triticum durum) grown at different levels of B toxicity in native vs. sterilized soil, and parameters related to root colonization, membrane damage, oxidative stress, chlorophyll, and mineral nutrition were measured to elucidate the physiological mechanisms of damage and benefit. Boron toxicity decreased early vegetative growth and grain yield, but it did not affect the straw dry weight of mature plants, whereas S. indica significantly enhanced the vegetative growth, straw dry weight, and the grain number of both wheat species. Membrane damage as demonstrated by increased lipid peroxidation and relative electrolyte leakage was caused by B toxicity and alleviated by S. indica. The benefits provided by S. indica could not be attributed to any significant changes in tissue concentrations of B or other minerals such as phosphorus. Soil sterilization generally improved plant performance but it did not consistently strengthen or weaken the effects of S. indica. The presented results suggest that S. indica may be used as an effective microbial inoculant to enhance wheat growth under adverse soil conditions such as B toxicity through mechanisms that are possibly unrelated to mineral homeostasis.

Low uptake efficiency, difficulty and the cost involved are major obstacles limiting micronutrients application through soil, therefore, interest exists for use of micronutrient treatment techniques such as micronutrient seed priming for biofortification of crops with micronutrients. A combination of in vitro, nursery and in field trials were conducted in 2021 to examine the influence of seed priming with different dosages (0–5%) of Zn for two soaking durations (24 and 48 h) on seedling growth and performance, grain Zn uptake, yield-related components and grain yield of two rice cultivars-Anbarbou and Shamshiri. Seed zinc concentration of two rice cultivar increased as the concentration of the priming solution increased. Natural zinc contents of two rice cultivars were significantly increased as the concentration of the priming solution increased, especially at the priming duration of 48 h. Nursery seedlings grown from primed seeds with 5% ZnSO4 at the priming duration of 48 h gave higher fresh and dry matter accumulation in shoots. The solutions used in the field trial increased the concentration of Zn in seeds from 10.5 to 29.6 µg g−1. Priming seed resulted in a significant increase in grain yield of 8.46%. Priming resulted in better yield-related parameters of panicle number plant−1 (20%), seed number panicle−1 (13.8%) and grain weight (9.1%). Further research also revealed that priming seed with zinc led the succinate dehydrogenase activity of hyphae of Serendipita indica, in symbiotic association with rice seedling roots, to increase. [ABSTRACT FROM AUTHOR]

Abstract Background Beneficial associations between plants and microbes are widespread in nature and have been studied extensively in the microbial-dominant environment of the rhizosphere. Such associations are highly advantageous for the organisms involved, benefiting soil microbes by providing them access to plant metabolites, while plant growth and development are enhanced through the promotion of nutrient uptake and/or protection against (a)biotic stresses. While the establishment and maintenance of mutualistic associations have been shown to require genetic and epigenetic reprogramming, as well as an exchange of effector molecules between microbes and plants, whether short RNAs are able to effect such changes is currently unknown. Here, we established an interaction between the model grass species Brachypodium distachyon (Bd, Pooideae) and the beneficial fungal root endophyte Serendipita indica (Si, syn. Piriformospora indica, Sebacinales) to elucidate RNA interference-based regulatory changes in gene expression and small (s)RNA profiles that occurred during establishment of a Sebacinalean symbiosis. Results Colonization of Bd roots with Si resulted in higher grain yield, confirming the mutualistic character of this interaction. Resequencing of the Si genome using the Oxford Nanopore technique, followed by de novo assembly yielded in 57 contigs and 9441 predicted genes, including putative members of several families involved in sRNA production. Transcriptome analysis at an early stage of the mutualistic interaction identified 2963 differentially expressed genes (DEG) in Si and 317 in Bd line 21-3. The fungal DEGs were largely associated with carbohydrate metabolism, cell wall degradation, and nutrient uptake, while plant DEGs indicated modulation of (a)biotic stress responses and defense pathways. Additionally, 10% of the upregulated fungal DEGs encode candidate protein effectors, including six DELD proteins typical for Sebacinales. Analysis of the global changes in the sRNA profiles of both associated organisms revealed several putative endogenous plant sRNAs expressed during colonization belonging to known micro (mi)RNA families involved in growth and developmental regulation. Among Bd- and Si-generated sRNAs with putative functions in the interacting organism, we identified transcripts for proteins involved in circadian clock and flowering regulation as well as immunity as potential targets of fungal sRNAs, reflecting the beneficial activity of Si. Conclusions We detected beneficial effects of Si colonization on Bd growth and development, and established a novel plant-mutualist interaction model between these organisms. Together, the changes in gene expression and identification of interaction-induced sRNAs in both organisms support sRNA-based regulation of defense responses and plant development in Bd, as well as nutrient acquisition and cell growth in Si. Our data suggests that a Sebacinalean symbiosis involves reciprocal sRNA targeting of genes during the interaction.

The root endophytic fungus Serendipita indica establishes beneficial symbioses with a broad spectrum of plants and enhances host resilience against biotic and abiotic stresses. However, little is known about the mechanisms underlying S. indica -mediated plant protection. Here, we report S. indica effector (SIE) 141 and its host target CDSP32, a conserved thioredoxin-like protein, and underlying mechanisms for enhancing pathogen resistance and abiotic salt tolerance in Arabidopsis thaliana. SIE141 binding interfered with canonical targeting of CDSP32 to chloroplasts, leading to its re-location into the plant nucleus. This nuclear translocation is essential for both their interaction and resistance function. Furthermore, SIE141 enhanced oxidoreductase activity of CDSP32, leading to CDSP32-mediated monomerization and activation of NON-EXPRESSOR OF PATHOGENESIS-RELATED 1 (NPR1), a key regulator of systemic resistance. Our findings provide functional insights on how S. indica transfers well-known beneficial effects to host plants and indicate CDSP32 as a genetic resource to improve plant resilience to abiotic and biotic stresses. [Display omitted] • Symbiotic Serendipita indica effector SIE141 mediates plant resistance diversity • SIE141 targets thioredoxin-like CDSP32 and enables it to depolymerize NPR1 • SIE141 binding targets plastid CDSP32 into plant nucleus • Nuclear-localized CDSP32 enhances plant resistance to biotic and abiotic stresses Zhang et al. find that effector SIE141 mediates beneficial fungus-induced plant resistance diversity by targeting the plastid protein CDSP32 into host nucleus, enhancing its oxidoreductase activity and ability to monomerize NPR1. Nuclear targeting enables CDSP32 to enhance plant resistance to biotic Phytophthora and abiotic salt stresses. [ABSTRACT FROM AUTHOR]

SNAREs (soluble N-ethylmaleimide-sensitive-factor attachment protein receptors) are engines for almost all of the membrane fusion and exocytosis events in organism cells. In this study, we identified 84 SNARE genes from banana (Musa acuminata). Gene expression analysis revealed that the expression of MaSNAREs varied a lot in different banana organs. By analyzing their expression patterns under low temperature (4 °C), high temperature (45 °C), mutualistic fungus (Serendipita indica, Si) and fungal pathogen (Fusarium oxysporum f. sp. Cubense Tropical Race 4, FocTR4) treatments, many MaSNAREs were found to be stress responsive. For example, MaBET1d was up-regulate by both low and high temperature stresses; MaNPSN11a was up-regulated by low temperature but down-regulated by high temperature; and FocTR4 treatment up-regulated the expression of MaSYP121 but down-regulated MaVAMP72a and MaSNAP33a. Notably, the upregulation or downregulation effects of FocTR4 on the expression of some MaSNAREs could be alleviated by priorly colonized Si, suggesting that they play roles in the Si-enhanced banana wilt resistance. Foc resistance assays were performed in tobacco leaves transiently overexpressing MaSYP121, MaVAMP72a and MaSNAP33a. Results showed that transient overexpression of MaSYP121 and MaSNPA33a suppressed the penetration and spread of both Foc1 (Foc Race 1) and FocTR4 in tobacco leaves, suggesting that they play positive roles in resisting Foc infection. However, the transient overexpression of MaVAMP72a facilitated Foc infection. Our study can provide a basis for understanding the roles of MaSNAREs in the banana responses to temperature stress and mutualistic and pathogenic fungal colonization.

Most plants in nature establish symbiotic associations with endophytic fungi in soil. Beneficial endophytic fungi induce a systemic response in the aboveground parts of the host plant, thus promoting the growth and fitness of host plants. Meanwhile, temperature elevation from climate change widely affects global plant biodiversity as well as crop quality and yield. Over the past decades, great progresses have been made in the response of plants to high ambient temperature and to symbiosis with endophytic fungi. However, little is known about their synergistic effect on host plants. The endophytic fungus Serendipita indica colonizes the roots of a wide range of plants, including Arabidopsis. Based on the Arabidopsis- S. indica symbiosis experimental system, we analyzed the synergistic effect of high ambient temperature and endophytic fungal symbiosis on host plants. By transcriptome analysis, we found that DNA replication-related genes were significantly upregulated during the systemic response of Arabidopsis aboveground parts to S. indica colonization. Plant hormones, such as jasmonic acid (JA) and ethylene (ET), play important roles in plant growth and systemic responses. We found that high ambient temperature repressed the JA and ET signaling pathways of Arabidopsis aboveground parts during the systemic response to S. indica colonization in roots. Meanwhile, PIF4 is the central hub transcription factor controlling plant thermosensory growth under high ambient temperature in Arabidopsis. PIF4 is also involving JA and/or ET signaling pathway. We found that PIF4 target genes overlapped with many differentially expressed genes (DEGs) during the systemic response, and further showed that the growth promotion efficiency of S. indica on the pif4 mutant was higher than that on the wild-type plants. In short, our data showed that high ambient temperature strengthened the growth promotion effect of S. indica fungi on the aboveground parts of the host plant Arabidopsis, and the growth promotion effect of the systemic response under high ambient temperature was regulated by PIF4. [ABSTRACT FROM AUTHOR]

Simple Summary: By using 'Zhongjiao No.3' and 'Zhongjiao No.4' banana seedlings as plant materials, we investigated the influences of Serendipita indica and bamboo fungus (Dictyophorae echinovolvata) culture substrates on the growth and wilt disease resistance of banana. S. indica colonization significantly enhanced the growth of the two banana cultivars, while banana seedlings grown in nutrient soil containing bamboo fungus culture substrates inoculated with D. echinovolvata showed suppressed growth. Moreover, both S. indica colonization and D. echinovolvata culture substrates addition can alleviate the disease symptoms caused by Fusarium oxysporum f. sp. cubense tropical race 4 (FocTR4), and their combined application showed the best disease resistance enhancement effect. The results obtained in this study can provide a basis for the application of S. indica and bamboo fungus in the prevention and control of banana Fusarium wilt disease in the future. Recently, many control methods have been tried and applied in the Fusarium wilt disease control of banana and have achieved definite progresses. In this study, by using 'Zhongjiao No.3' and 'Zhongjiao No.4' banana seedlings as materials, the effects of Serendipita indica and bamboo fungus (Dictyophorae echinovolvata) culture substrates on the growth and Fusarium wilt disease resistance of banana were investigated. Results showed that the plant height, leaf length, leaf width, root length and root thickness, aboveground part fresh weight, root fresh weight, and relative chlorophyll content and nitrogen content in leaves of banana seedlings colonized with S. indica were all greater than those of non-colonized controls, while these parameters of banana seedlings grown in nutrient soil containing D. echinovolvata culture substrates were significantly suppressed. Both S. indica non-colonized and colonized seedlings cultivated in nutrient containing 1/4 D. echinovolvata culture substrates showed much milder symptoms compared with those cultivated in normal nutrient soil, indicating that the addition of bamboo fungus substrates to the soil can enhance the Fusarium wilt resistance of banana. The results obtained in this study can provide a basis for the application of S. indica and bamboo fungus in the prevention and control of banana Fusarium wilt disease. [ABSTRACT FROM AUTHOR]

Most plants in nature establish symbiotic associations with endophytic fungi in soil. Beneficial endophytic fungi induce a systemic response in the aboveground parts of the host plant, thus promoting the growth and fitness of host plants. Meanwhile, temperature elevation from climate change widely affects global plant biodiversity as well as crop quality and yield. Over the past decades, great progresses have been made in the response of plants to high ambient temperature and to symbiosis with endophytic fungi. However, little is known about their synergistic effect on host plants. The endophytic fungus Serendipita indica colonizes the roots of a wide range of plants, including Arabidopsis. Based on the Arabidopsis-S. indica symbiosis experimental system, we analyzed the synergistic effect of high ambient temperature and endophytic fungal symbiosis on host plants. By transcriptome analysis, we found that DNA replication-related genes were significantly upregulated during the systemic response of Arabidopsis aboveground parts to S. indica colonization. Plant hormones, such as jasmonic acid (JA) and ethylene (ET), play important roles in plant growth and systemic responses. We found that high ambient temperature repressed the JA and ET signaling pathways of Arabidopsis aboveground parts during the systemic response to S. indica colonization in roots. Meanwhile, PIF4 is the central hub transcription factor controlling plant thermosensory growth under high ambient temperature in Arabidopsis. PIF4 is also involving JA and/or ET signaling pathway. We found that PIF4 target genes overlapped with many differentially expressed genes (DEGs) during the systemic response, and further showed that the growth promotion efficiency of S. indica on the pif4 mutant was higher than that on the wild-type plants. In short, our data showed that high ambient temperature strengthened the growth promotion effect of S. indica fungi on the aboveground parts of the host plant Arabidopsis, and the growth promotion effect of the systemic response under high ambient temperature was regulated by PIF4.

Juglans regia L. is a good host for Serendipita indica. Under drought condition, seedlings colonized with S. indica showed higher values in plant height, total fresh biomass, root/shoot ratio, relative growth rate, leaf relative water content and chlorophyll content, gas exchange parameters, maximal photochemical efficiency, photochemical quenching, and effective photosystem II quantum yield than the uncolonized seedlings. It suggested beneficial effects of S. indica on host plants' growth and physiological parameters in response to drought. In comparison with the uncolonized seedlings, S. indica–colonized seedlings showed lower levels in hydrogen peroxide, superoxide anion, malondialdehyde, and relative electrical conductivity under drought condition, suggesting the ability of S. indica to prevent or retard the accumulation of reactive oxygen species and to diminish the oxidative injure. Furthermore, walnut seedlings responded to drought by actively accumulating osmotic regulation substances including soluble protein, soluble sugar, and proline. Root colonization with S. indica was more conductive to the accumulation. Moreover, in response to drought stress, walnut seedlings, regardless of colonization, increased activities of superoxide dismutase, catalase, peroxidase, ascorbate peroxidase, dehydroascorbate reductase, monodehydroascorbate reductase, and glutathione reductase, levels of ascorbate and glutathione, and ratios of reduced ascorbate/dehydroascorbic acid and reduced glutathione/oxidized glutathione in leaves and roots. S. indica colonization induced much more increase in the abovementioned indicators as compared to the uncolonized seedlings. Overall, S. indica colonization alleviated the detrimental effects of drought stress by altering root system, enhancing osmotic adjustment, and repressing the accumulation of reactive oxygen species via stimulating antioxidant system including enzymatic and nonenzymatic components. Key points: • S. indica stimulated root growth of walnut seedlings under drought condition. • S. indica accelerated osmotic adjustment under drought condition. • S. indica activated antioxidant defense mechanism under drought condition. [ABSTRACT FROM AUTHOR]

Abstract Background The plant endophytic fungus Serendipita indica colonizes roots of a wide range of plant species and can enhance growth and stress resistance of these plants. Due to its ease of axenic cultivation and its broad host plant range including the model plant Arabidopsis thaliana and numerous crop plants, it is widely used as a model fungus to study beneficial fungus-root interactions. In addition, it was suggested to be utilized for commercial applications, e.g. to enhance yield in barley and other species. To produce inoculum, S. indica is mostly cultivated in a complex Hill-Käfer medium (CM medium), however, growth in this medium is slow, and yield of chlamydospores, which are often used for plant root inoculation, is relatively low. Results We tested and optimized a simple vegetable juice-based medium for an enhanced yield of fungal inoculum. The described vegetable juice (VJ) medium is based on commercially available vegetable juice and is easy to prepare. VJ medium was superior to the currently used CM medium with respect to biomass production in liquid medium and hyphal growth on agar plates. Using solid VJ medium supplemented with sucrose (VJS), a high amount of chlamydospores developed already after 8 days of cultivation, producing significantly more spores than on CM medium. Use of VJ medium is not restricted to S. indica, as it also supported growth of two pathogenic fungi often used in plant pathology experiments: the ascomycete Fusarium graminearum, the causal agent of Fusarium head blight disease on wheat and barley, and Verticillium longisporum, the causal agent of verticillium wilt. Conclusions The described VJ medium is recommended for streamlined and efficient production of inoculum for the plant endophytic fungus Serendipita indica and might prove superior for the propagation of other fungi for research purposes.

In the current research, unique Nano-Embedded Fungus (NEF), made by the synergic association of silver nanoparticles (AgNPs) and endophytic fungus (Piriformospora indica), is studied, and the impact of NEF on black rice secondary metabolites is reported. AgNPs were synthesized by chemical reduction process using the temperature-dependent method and characterized for morphological and structural features through UV visible absorption spectroscopy, zeta potential, XRD, SEM-EDX, and FTIR spectroscopy. The NEF, prepared by optimizing the AgNPs concentration (300 ppm) in agar and broth media, showed better fungal biomass, colony diameter, spore count, and spore size than the control P. indica. Treatment with AgNPs, P. indica, and NEF resulted in growth enhancement in black rice. NEF and AgNPs stimulated the production of secondary metabolites in its leaves. The concentrations of chlorophyll, carotenoids, flavonoids, and terpenoids were increased in plants inoculated with P. indica and AgNPs. The findings of the study highlight the synergistic effect of AgNPs and the fungal symbionts in augmenting the secondary metabolites in leaves of black rice.

There is mounting evidence that the root-colonizing endosymbiotic fungus Serendipita indica improves plant growth. The beneficial effects have been observed when plants are growing in optimal conditions or under nutritionally deficient soils (e.g., phosphate poor soil) or exposed to stressful environmental conditions such as drought or salinity. However, until now its role in the nutrition of other plant essential macronutrient, such as K+, has not been fully clarified. Here, we study the role of the fungus in the K+ nutrition of Arabidopsis thaliana plants, during growth under K+ limiting conditions. As a first step, we studied the high-affinity K+ uptake of the plant and fungus when growing separately and in symbiosis. In the search for putative fungal actors involved in K+ nutrition, we also have cloned and functionally characterized the K+ transporters of S. indica SiHAK1, SiTRK1, SiTRK2, and SiTOK1, among which it has been shown that SiHAK1 is the main transporter involved in the K+ uptake in the high affinity range of concentrations. In addition, a gene expression study of these transporters and other candidates that could participate in the K+ homeostasis of the fungus has been carried out. The results indicated that, contrary to what happens with P nutrition, S. indica seems not to improve neither the growth nor the plant K+ reserves during K+ starvation. Instead, this nutritionally restrictive condition favored fungal colonization, suggesting that the fungus obtains the greatest benefit in K+ supply during symbiosis.

Background: Beneficial associations between plants and microbes are widespread in nature and have been studied extensively in the microbial-dominant environment of the rhizosphere. Such associations are highly advantageous for the organisms involved, benefiting soil microbes by providing them access to plant metabolites, while plant growth and development are enhanced through the promotion of nutrient uptake and/or protection against (a)biotic stresses. While the establishment and maintenance of mutualistic associations have been shown to require genetic and epigenetic reprogramming, as well as an exchange of effector molecules between microbes and plants, whether short RNAs are able to effect such changes is currently unknown. Here, we established an interaction between the model grass species Brachypodium distachyon (Bd, Pooideae) and the beneficial fungal root endophyte Serendipita indica (Si, syn. Piriformospora indica, Sebacinales) to elucidate RNA interference-based regulatory changes in gene expression and small (s)RNA profiles that occurred during establishment of a Sebacinalean symbiosis. Results: Colonization of Bd roots with Si resulted in higher grain yield, confirming the mutualistic character of this interaction. Resequencing of the Si genome using the Oxford Nanopore technique, followed by de novo assembly yielded in 57 contigs and 9441 predicted genes, including putative members of several families involved in sRNA production. Transcriptome analysis at an early stage of the mutualistic interaction identified 2963 differentially expressed genes (DEG) in Si and 317 in Bd line 21-3. The fungal DEGs were largely associated with carbohydrate metabolism, cell wall degradation, and nutrient uptake, while plant DEGs indicated modulation of (a)biotic stress responses and defense pathways. Additionally, 10% of the upregulated fungal DEGs encode candidate protein effectors, including six DELD proteins typical for Sebacinales. Analysis of the global changes in the sRNA profiles of both associated organisms revealed several putative endogenous plant sRNAs expressed during colonization belonging to known micro (mi)RNA families involved in growth and developmental regulation. Among Bd- and Si-generated sRNAs with putative functions in the interacting organism, we identified transcripts for proteins involved in circadian clock and flowering regulation as well as immunity as potential targets of fungal sRNAs, reflecting the beneficial activity of Si. Conclusions: We detected beneficial effects of Si colonization on Bd growth and development, and established a novel plant-mutualist interaction model between these organisms. Together, the changes in gene expression and identification of interaction-induced sRNAs in both organisms support sRNA-based regulation of defense responses and plant development in Bd, as well as nutrient acquisition and cell growth in Si. Our data suggests that a Sebacinalean symbiosis involves reciprocal sRNA targeting of genes during the interaction. [ABSTRACT FROM AUTHOR]

Abstract In this study, a pot experiment was carried out in greenhouse to investigate the potentials of Xerocomus badius and Serendipita indica to penetrate and colonize roots of ryegrass (Lolium multiflorum Lam.) and to induce beneficial effects on seed germination and seedling growth. The results showed that X. badius and S. indica successfully colonized in the root system of L. multiflorum seedlings and the root colonization rate was 72.65% and 88.42%, respectively. By microscopy, the hyphae, chlamydospores and spores produced by S. indica were observed in roots cortex of L. multiflorum seedlings. In comparison with the non-inoculated seedlings, seedlings inoculated with X. badius and S. indica showed significant increase in growth parameters with plant height, basal diameter, biomass accumulation, relative growth rate, leaf relative water content and chlorophyll content. Also, we found that seedlings inoculated with S. indica exhibited a greater growth-promotion as compared with X. badius-inoculated seedlings. No significant influence of the two fungus application has been observed with respect to seed germination. It suggested that well establishments of mutualistic symbiosis between L. multiflorum and X. badius or S. indica were not so essential to seed germination but contributed highly to the survival and growth of the seedlings.

Recently, many control methods have been tried and applied in the Fusarium wilt disease control of banana and have achieved definite progresses. In this study, by using ‘Zhongjiao No.3’ and ‘Zhongjiao No.4’ banana seedlings as materials, the effects of Serendipita indica and bamboo fungus (Dictyophorae echinovolvata) culture substrates on the growth and Fusarium wilt disease resistance of banana were investigated. Results showed that the plant height, leaf length, leaf width, root length and root thickness, aboveground part fresh weight, root fresh weight, and relative chlorophyll content and nitrogen content in leaves of banana seedlings colonized with S. indica were all greater than those of non-colonized controls, while these parameters of banana seedlings grown in nutrient soil containing D. echinovolvata culture substrates were significantly suppressed. Both S. indica non-colonized and colonized seedlings cultivated in nutrient containing 1/4 D. echinovolvata culture substrates showed much milder symptoms compared with those cultivated in normal nutrient soil, indicating that the addition of bamboo fungus substrates to the soil can enhance the Fusarium wilt resistance of banana. The results obtained in this study can provide a basis for the application of S. indica and bamboo fungus in the prevention and control of banana Fusarium wilt disease.

Serendipita (=Piriformospora) indica is a fungal endophytic symbiont with the capabilities to enhance plant growth and confer resistance to different stresses. However, the application of this fungus in the field has led to inconsistent results, perhaps due to antagonism with other microbes. Here, we studied the impact of individual bacterial isolates from the endophytic bacterial community on the in vitro growth of S. indica. We further analyzed how combinations of bacteria and S. indica influence plant growth and protection against the phytopathogens Fusarium oxysporum and Rhizoctonia solani. Bacterial strains of the genera Bacillus, Enterobacter and Burkholderia negatively affected S. indica growth on plates, whereas Mycolicibacterium, Rhizobium, Paenibacillus strains and several other bacteria from different taxa stimulated fungal growth. To further explore the potential of bacteria positively interacting with S. indica, four of the most promising strains belonging to the genus Mycolicibacterium were selected for further experiments. Some dual inoculations of S. indica and Mycolicibacterium strains boosted the beneficial effects triggered by S. indica, further enhancing the growth of tomato plants, and alleviating the symptoms caused by the phytopathogens F. oxysporum and R. solani. However, some combinations of S. indica and bacteria were less effective than individual inoculations. By analyzing the genomes of the Mycolicibacterium strains, we revealed that these bacteria encode several genes predicted to be involved in the stimulation of S. indica growth, plant development and tolerance to abiotic and biotic stresses. Particularly, a high number of genes related to vitamin and nitrogen metabolism were detected. Taking into consideration multiple interactions on and inside plants, we showed in this study that some bacterial strains may induce beneficial effects on S. indica and could have an outstanding influence on the plant-fungus symbiosis.