Phytohormones play vital roles in the growth and development of plants as well as in interactions of plants with microbes such as endophytic fungi. The endophytic root-colonizing fungus Piriformospora indica promotes plant growth and performance, increases resistance of colonized plants to pathogens, insects and abiotic stress. Here, we discuss the roles of the phytohormones (auxins, cytokinin, gibberellins, abscisic acid, ethylene, salicylic acid, jasmonates, and brassinosteroids) in the interaction of P. indica with higher plant species, and compare available data with those from other (beneficial) microorganisms interacting with roots. Crosstalks between different hormones in balancing the plant responses to microbial signals is an emerging topic in current research. Furthermore, phytohormones play crucial roles in systemic signal propagation as well as interplant communication. P. indica interferes with plant hormone synthesis and signaling to stimulate growth, flowering time, differentiation and local and systemic immune responses. Plants adjust their hormone levels in the roots in response to the microbes to control colonization and fungal propagation. The available information on the roles of phytohormones in beneficial root–microbe interactions opens new questions of how P. indica manipulates the plant hormone metabolism to promote the benefits for both partners in the symbiosis.

Drought stress is one of the most severe environmental constraints to plant growth and crop productivity. Plant growth is greatly affected by drought stress, and plants, to survive, adapt to this stress by invoking different pathways. Piriformospora indica, a root-colonizing endophytic fungus of Sebacinales, promotes plant growth and confers resistance to biotic and abiotic stresses, including drought stress, by affecting the physiological properties of the host plant. The fungus strongly colonizes the roots of maize (Zea mays L.) and promotes shoot and root growth under both normal growth conditions and drought stress. We used polyethylene glycol (PEG-6000) to mimic drought stress and found that root fresh and dry weight, leaf area, SPAD value, and leaf number were increased in P. indica-colonized plants. The antioxidative activities of catalases and superoxide dismutases were upregulated within 24h in the leaves of P. indica-colonized plants. Drought-related genes DREB2A, CBL1, ANAC072, and RD29A were upregulated in drought-stressed leaves of P. indica-colonized plants. Furthermore, after drought treatment, proline content increased, whereas accumulation of malondialdehyde (MDA), an indicator of membrane damage, decreased in P. indica-colonized maize. We conclude that P. indica-mediated plant protection against the detrimental effects of drought may result from enhanced antioxidant enzyme activity, proline accumulation, and expression of drought-related genes and lower membrane damage in maize plants

As an endophytic fungus of Sebacinales, Piriformospora indica promotes plant growth and resistance to abiotic stress, including drought. Colonization of maize roots promoted the leaf size, root length and number of tap roots. Under drought stress, the maize seedlings profited from the presence of the fungus and performed visibly better than the uncolonized controls. To identify genes and biological processes involved in growth promotion and drought tolerance conferred by P. indica, the root transcriptome of colonized and uncolonized seedlings was analyzed 0, 6 and 12 h after drought stress (20% polyethylene glycol 6000). The number of P. indica-responsive genes increased from 464 (no stress at 0 h) to 1337 (6 h drought) and 2037 (12 h drought). Gene Ontology analyses showed that the carbon and sulfur metabolisms are major targets of the fungus. Furthermore, the growth promoting effect of P. indica is reflected by higher transcript levels for microtubule associated processes. Under drought stress, the fungus improved the oxidative potential of the roots, and stimulated genes for hormone functions, including those which respond to abscisic acid, auxin, salicylic acid and cytokinins. The comparative analyses of our study provides systematic insight into the molecular mechanism how P. indica promotes plant performance under drought stress, and presents a collection of genes which are specifically targeted by the fungus under drought stress in maize roots.

The dwarfing gene sdw1 has been widely used throughout the world to develop commercial barley varieties. There are at least four different alleles at the sdw1 locus. Mutations in the gibberellin 20-oxidase gene (HvGA20ox2) resulted in multiple alleles at the sdw1 locus. The sdw1.d allele from Diamant is due to a 7-bp deletion in exon 1, while the sdw1.c allele from Abed Denso has 1-bp deletion and a 4-bp insertion in the 5’ untranslated region. The sdw1.a allele from Jotun resulted from a total deletion of the HvGA20ox2 gene. The structural changes result in lower gene expression in sdw1.d and lack of expression in sdw1.a. There are three HvGA20ox genes in the barley genome. The partial or total loss of function of the HvGA20ox2 gene could be compensated by enhanced expression of its homolog HvGA20ox1and HvGA20ox3. A diagnostic molecular marker was developed to differentiate between the wild-type, sdw1.d and sdw1.a alleles and another molecular marker for differentiation of sdw1.c and sdw1.a. The markers were further tested in 197 barley varieties, out of which 28 had the sdw1.d allele and two varieties the sdw1.a allele. To date, the sdw1.d and sdw1.a alleles have only been detected in the modern barley varieties and lines. The results provided further proof that the gibberellin 20-oxidase gene (HvGA20ox2) is the functional gene of the barley sdw1 mutants. Different deletions resulted in different functional alleles for different breeding purposes. Truncated protein could maintain partial function. Partial or total loss of function of the HvGA20ox2 gene could be compensated by enhanced expression of its homolog HvGA20ox1 and HvGA20ox3.