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Photosynthetic rate of barley plants inoculated and non- inoculated with Piriformospora indica . The photosynthetic rate of barley (cv. Ingrid) was measured in P. indica colonized (black bars) and in non-colonized control plants (white bars). Values are means of three to four plants measured at three different light intensities. Error bars indicate standard deviation. Statistically significant differences between control and P. indica inoculated plants were detected only for the light intensity of 200 μ mol photons m -2 s -1 (Students t-test p <0.05) 

Photosynthetic rate of barley plants inoculated and non- inoculated with Piriformospora indica . The photosynthetic rate of barley (cv. Ingrid) was measured in P. indica colonized (black bars) and in non-colonized control plants (white bars). Values are means of three to four plants measured at three different light intensities. Error bars indicate standard deviation. Statistically significant differences between control and P. indica inoculated plants were detected only for the light intensity of 200 μ mol photons m -2 s -1 (Students t-test p <0.05) 

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Root colonization by the basidiomycete fungus Piriformospora indica induces host plant tolerance against abiotic and biotic stress, and enhances growth and yield. As P. indica has a broad host range, it has been established as a model system to study beneficial plant-microbe interactions. Moreover, its properties led to the assumption that P. indic...

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... indica enhanced the host plants`plants`photosynthetic 536 rate under low light conditions (Fig. 3) ...

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... Currently, P. indica is widely discussed as a beneficial fungus in plants such as strawberry [7], longan [10], and rice [11]. Studies have shown that the growth parameters of barley [28], king grass [29], ryegrass [30], tobacco [31], African chrysanthemum [32], and maize [33] colonized by P. indica are higher than those of non-inoculated controls. Additionally, P. indica colonization can improve the nutritional quality of black rice [34], promote the germination and growth of Arabidopsis thaliana [35], and increase the aboveground and underground biomass of sweet potato plants [3]. ...
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Numerous studies have shown that the endophytic fungus Piriformospora indica has a broad range of promoting effects on root development and plant growth in host plants. However, there are currently no reports on the application of this fungus on Cerasus humilis. This study first compared the colonization ability of P. indica on 11 C. humilis varieties and found that the colonization rate of this fungus on these varieties ranged from 90% to 100%, with the colonization rate of the varieties ‘09-01’ and ‘Nongda 7’ being as high as 100%. Subsequently, the effect of P. indica on root development and plant growth of C. humilis was investigated using cuttings of ‘09-01’ and ‘Nongda 7’ as materials. P. indica colonization was found to increase the biomass of ‘09-01’ and ‘Nongda 7’ plants; root activity, POD enzymes, and chlorophyll content were also significantly increased. In addition, indole-3-acetic acid (IAA) content in the roots of C. humilis plants increased after colonization, while jasmonic acid (JA) and 1-aminocyclopropane-1-car- boxylic acid (ACC) content decreased. In conclusion, it has been demonstrated that P. indica can promote the growth of C. humilis plants by accelerating biomass accumulation, promoting rooting, and enhancing the production of photosynthetic pigments, as well as regulating hormone synthesis.
... Serendipita indica is a potential plant growth-promoting fungus, that has been found to improve crop production under abiotic stress conditions [18,19]. This symbiotic fungus can modulate barley morphology and physiology, and change plant metabolic activities, such as CO 2 assimilation and transpiration rate [20][21][22][23][24]. It is important to understand the various factors that can influence a plant's responses to drought and the potential role of symbiotic organisms in mitigating the negative effects of water scarcity, as the knowledge of them is so far incomplete. ...
... This study hypothesizes that potassium supplementation and S. indica inoculation significantly enhances the drought resilience and productivity of spring barley genotypes adapted to local conditions, as well as those originating from arid regions. While previous studies have explored the individual effects of potassium and endophytes on plant stress responses [14,15,21,22,[28][29][30][31], their combined impact on various spring barley genotypes, particularly under field conditions, remains insufficiently understood. By investigating genotype-specific traits associated with drought resilience and evaluating the effects of these interventions on productivity and growth parameters, the study aims to offer insights into strategies for stabilizing crop production under non-standard climatic conditions. ...
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... 21,40,41 Moreover, P. indica can elevate the photosynthetic rate and growth phytohormones secretion, which helps in host's growth and development. 39,42,43 We also observed that an increase in vegetative growth was positively correlated with an increase in total yield due to an increase in grains per ear and panicle weight following P. indica inoculation (Tables 1 and 2). In addition, the heading time of plants in the SS treatment was earlier than that in the non-colonized plants (data not shown). ...
... In addition, the heading time of plants in the SS treatment was earlier than that in the non-colonized plants (data not shown). Achatz et al. (2010) reported that P. indica enhanced the grain yield of barley, which caused an increase in the number of ears per plant at early development stages. This indicates that the higher grain yield in P. indicacolonized plants not only results from the formation of more tillers but also from more rapid development, e.g., the observed earlier emergence of ears. ...
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... The outcomes of endophytic relationships cover the whole range from negative to beneficial effects on plant performance (Schulz and Boyle, 2005;Deshmukh et al., 2006). Beneficial plant-fungus interactions hold potential for an application in sustainable agriculture to achieve higher yields (Achatz et al., 2010a(Achatz et al., , 2010b and increased resistance and tolerance to pathogens, parasites, and abiotic stresses (Harman, 2011;Lugtenberg et al., 2016). ...
... The increase in the dry matter content means that lower amounts of water were present in the tissues, possibly due to a higher transpiration rate not covered by increased water uptake. Water uptake is balanced with the photosynthetic rate, and it has previously been shown in barley and has also been proposed for in tomato that plants inoculated with S. indica showed enhanced photosynthetic activity even at low light intensities (Achatz et al., 2010b;Fakhro et al., 2010a). ...
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... S. indica, in addition to increasing crop production in plants such as rice, barley, black cabbage, and melon [31,37,38], can increase secondary metabolites and phytochemicals in Inoculation of Arabidopsis, Chinese cabbage, rice, and corn roots with S. indica causes root proliferation and biomass changes, according to Tsai et al. [27], and it may be an increase in root biomass due to increased indole-3 acetic acid (IAA) production by S. indica [28]. Komis et al. found that S. indica colonized maize roots after 15 days, which led to their further growth due to stimulating genes involved in microtubule-based processes [29]. ...
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... The effects of P. indica colonization on increasing biomass have been proven in several plant species such as rice, Chinese cabbage and barley (Lee et al. 2011;Hilbert et al. 2012;Tsai et al. 2020). The capability of P. indica to accelerate plant growth and to enhance crop yield could result from promoting root development and nutrient uptake (Achatz et al. 2010;Ansari et al. 2013). P. indica colonization not only benefits plant growth but also improves environmental stress tolerance (Sherameti et al. 2008;Jiang et al. 2020). ...
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... It is an axenically cultivable root colonizing endosymbiotic fungus with many biotechnological applications by acting as a biofertilizer, a bioprotector against biotic and abiotic stresses, as a bioregulator for plant growth and secondary metabolite production . Various studies have reported enhanced biomass and secondary metabolism after successful root colonization with this fungus (Baldi et al., 2008;Achatz et al., 2010;Satheesan et al., 2012;Su et al., 2017;Khalvandi et al., 2019). There are also reports on the stimulatory effects of P.indica derived elicitors on growth and secondary metabolite production in host plants (Kumar et al., 2012;Tashackori et al., 2018). ...
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Andrographis paniculata [(Burm. f.) Wall. ex Nees, family Acanthaceae], is an important medicinal plant used in Indian and Chinese traditional systems of medicines. The therapeutic activity of A. paniculata is due to the presence of andrographolide, a diterpenoid lactone with potent pharmacological activities. Andrographolide and its various semi synthetic derivatives are of high therapeutic potential and therefore of high demand. Due to the complex structure of andrographolide, chemical synthesis has not been effective till date and thus the pharmaceutical industry depends on A. paniculata for extraction of pure andrographolide. The content of andrographolide in A. paniculata is low (14.28 ± 1.62 mg/g DW) and thus biotechnological interventions are essential for enhanced production of andrographolide. The potential mutualistic role of endosymbiosis in enhancing the yield of andrographolide in in vitro cultures of A. paniculata was studied and the salient findings obtained are reported. It was consistently observed that colonization of roots of A. paniculata seedlings by the mutualistic endosymbiotic fungus Piriformospora indica resulted in significant enhancement of andrographolide (58.2 ± 6.5 mg/g DW) in comparison to control plants (19.5 ± 2.3 mg/g DW), along with a correlated increase in plant biomass. Fractions derived from P. indica such as cell wall and culture exudate were also employed to study their effect on andrographolide and biomass yield. It was observed that these fractions successfully improved the growth rate of the plant along with elevated andrographolide production, with maximum andrographolide content (55.8 ± 3.2 mg/g DW) observed in cell wall fraction treated seedlings. Under all the treatment conditions the transcriptional regulation of andrographolide biosynthetic pathway genes were analysed using RT-qPCR and correlated with andrographolide content. Andographolide biosynthesis pathway genesGGPS, HMGS and DXR showed a significantly high transcript level expression and correlation with andrographolide accumulation over a period of 35 days in case of all the treatments studied. The potential protective role of P. indica colonization under host abiotic stress was demonstrated in A. paniculata seedlings exposed to abiotic stress conditions induced by poly ethylene glycol. Piriformospora indica colonized plants under drought stress exhibited normal morphology, high antioxidant enzyme content, low malondialdehyde accumulation and high proline content when compared with the control plants. The observations prove that P. indica and P. indica derived fractions can be used as an ideal bio stimulator to enhance the biomass (2.344 fold) and andrographolide content (3.91 fold) in A. paniculata. The findings also showed that as A. paniculata is significantly affected by drought stress, P. indica can also be used as a helpful eco-friendly tool to confer drought stress tolerance, thereby improving biomass and andrographolide yield. Moreover, the biostimulants present in the fungal cell wall fractions and culture exudates can be purified and characterized for exploring their further functional roles in order to develop products of commercial value.
... In Arabidopsis and tobacco roots, P. indica stimulated the expression of enzymes involved in nitrate metabolism including nitrate reductase, 41 and a positive effect of P. indica on phosphate uptake was shown in maize. 42 Achatz et al. (2010) reported that P. indica enhanced grain yield of barley, 43 and this was caused by growth promotion at early development stages. In our study, the panicle and filled grain number of P. indica-colonized rice plants were significantly higher compared to noninoculated control plants, whereas the ten-grain weight was not changed (Table 1). ...
... In Arabidopsis and tobacco roots, P. indica stimulated the expression of enzymes involved in nitrate metabolism including nitrate reductase, 41 and a positive effect of P. indica on phosphate uptake was shown in maize. 42 Achatz et al. (2010) reported that P. indica enhanced grain yield of barley, 43 and this was caused by growth promotion at early development stages. In our study, the panicle and filled grain number of P. indica-colonized rice plants were significantly higher compared to noninoculated control plants, whereas the ten-grain weight was not changed (Table 1). ...
... In barley, P. indica colonization increased the number of ears per plant. 43 Filling efficiency of rice spikelet is dependent on photosynthesis of the mature leaves during grain filling and the mobilization of carbohydrate in the leaf sheath and culm of the upper leaves. 44,45 Stimulation of the carbohydrate metabolism and expression of sugar transporter genes was described for several plant species associated with mycorrhizal fungi. ...
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Global water shortage seriously threatens rice growth especially in irrigated production areas. Association of plants with beneficial soil microbes is one strategy for plant adaption to environmental stresses. In this study, rice (Oryza sativa L.) plants were colonized by the beneficial root-colonizing endophytic fungus Piriformospora indica (P. indica). We demonstrate that grain yield were higher in P. indica-colonized rice plants compared to the uncolonized plants grown in soil. Moreover, P. indica effect on improving water stress tolerance in rice and its physiological mechanism were investigated in a hydroponic culture system. Polyethylene glycol (PEG) was applied to the culture solution to conduct the water stress condition. Water stress-induced leaf wilting and impairments in photosynthetic efficiency were diminished in P. indica-colonized plants. Furthermore, P. indica colonization promotes stomata closure and increases the leaf surface temperature under water stress. The malondialdehyde level (as an indicator for oxidative stress) was lower and the reduced to oxidized glutathione ratio was higher in P. indica-colonized and PEG-exposed rice plants compared to the uncolonized plants. Furthermore, the activities of the antioxidant enzymes catalase and glutathione reductase were up-regulated in inoculated rice seedlings under water stress. In conclusion, P. indica promotes rice performance under water stress by stomata closure and lower oxidative stress.
... When inspecting the beneficial effect of the co-cultivation of plants with S. indica it becomes obvious that there is a discrepancy between the observed effects and the classical growth-defense tradeoff concept. Although S. indica infection triggers defense responses, plant biomass production and productivity in terms of seed yield are increased Achatz et al., 2010). It therefore has to be concluded that, along with the basic initial plant defense response, further mechanisms are triggered by the fungus, allowing the host plant to grow despite all of the metabolic restrictions. ...
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... In some cases nutrient status has been reported to play a role in the interaction of the model sebacinoid fungus Serendipita indica with some plant species Sherameti et al., 2005;Nautiyal et al., 2010;Yadav et al., 2010;Kumar et al., 2012). However, S. indica colonization of Nicotiana attenuata and barley had no effect on host phosphorus (P) and nitrogen (N) content (Barazani et al., 2005;Achatz et al., 2010), suggesting that nutrient exchange is not central to the beneficial effects conferred by sebacinoid fungi. S. indica colonizes the rhizodermis and outer root cortex (Deshmukh et al., 2006;Jacobs et al., 2011;Weiss et al., 2016). ...
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Loss-of-function alleles of MLO (Mildew Resistance Locus O) confer broad-spectrum resistance to foliar infections by powdery mildew pathogens. Like pathogens, microbes that establish mutually beneficial relationships with their plant hosts, trigger the induction of some defense responses. Initially, barley colonization by the root endophyte Serendipita indica (syn. Piriformospora indica) is associated with enhanced defense gene expression and the formation of papillae at sites of hyphal penetration attempts. This phenotype is reminiscent of mlo-conditioned immunity in barley leaf tissue and raises the question whether MLO plays a regulatory role in the establishment of beneficial interactions. Here we show that S. indica colonization was significantly reduced in plants carrying mlo mutations compared to wild type controls. The reduction in fungal biomass was associated with the enhanced formation of papillae. Moreover, epidermal cells of S. indica-treated mlo plants displayed an early accumulation of iron in the epidermal layer suggesting increased basal defense activation in the barley mutant background. Correspondingly, the induction of host cell death during later colonization stages was impaired in mlo colonized plants, highlighting the importance of the early biotrophic growth phase for S. indica root colonization. In contrast, the arbuscular mycorrhizal fungus Funneliformis mosseae displayed a similar colonization morphology on mutant and wild type plants. However, the frequency of mycorrhization and number of arbuscules was higher in mlo-5 mutants. These findings suggest that MLO differentially regulates root colonization by endophytic and AM fungi.