Induction of abiotic stress tolerance in plants by endophytic microbes

Endophytes are micro‐organisms including bacteria and fungi that survive within healthy plant tissues and promote plant growth under stress. This review focuses on the potential of endophytic microbes that induce abiotic stress tolerance in plants. How endophytes promote plant growth under stressful conditions, like drought and heat, high salinity and poor nutrient availability will be discussed. The molecular mechanisms for increasing stress tolerance in plants by endophytes include induction of plant stress genes as well as biomolecules like reactive oxygen species scavengers. This review may help in the development of biotechnological applications of endophytic microbes in plant growth promotion and crop improvement under abiotic stress conditions.

Significance and Impact of the Study

Increasing human populations demand more crop yield for food security while crop production is adversely affected by abiotic stresses like drought, salinity and high temperature. Development of stress tolerance in plants is a strategy to cope with the negative effects of adverse environmental conditions. Endophytes are well recognized for plant growth promotion and production of natural compounds. The property of endophytes to induce stress tolerance in plants can be applied to increase crop yields. With this review, we intend to promote application of endophytes in biotechnology and genetic engineering for the development of stress‐tolerant plants.     

The role of plant-associated rhizobacteria in plant growth,biocontrol and abiotic stress management

The rhizosphere is the region around the plant roots where maximum microbial activities occur. In the rhizosphere, microorganisms' beneficial and harmful activities affect plant growth and development. The mutualistic rhizospheric bacteria which improve plant growth and health are known as plant growth-promoting rhizobacteria (PGPR). They are very important due to their ability to help the plant in diverse ways. PGPR such as Pseudomonas, Bacillus, Azospirillum, Azotobacter, Arthrobacter, Achromobacter, Micrococcus, Enterobacter, Rhizobium, Agrobacterium, Pantoea and Serratia are now very well known. Rhizomicrobiome plays critical roles in nutrient acquisition and assimilation, improved soil texture, secreting and modulating extracellular molecules such as hormones, secondary metabolites, antibiotics and various signal compounds, all leading to the enhancement of plant growth and development. The microbes and compounds they secrete constitute valuable biostimulants and play pivotal roles in modulating plant stress responses. In this review, we highlight the rhizobacteria diversity and cutting-edge findings focusing on the role of a PGPR in plant growth and development. We also discussed the role of PGPR in resisting the adverse effects arising from various abiotic (drought, salinity, heat, heavy metals) stresses.     

Advances in improvement of quality and resistance in a multipurpose crop: sea buckthorn

Sea buckthorn is a berry crop with multiple uses. The berries are highly appreciated for their unique taste but are also very rich in bioactive compounds with powerful nutritional and medicinal values. In addition, the plants grow well under adverse conditions, and are often used to fight soil erosion. Utilization of sea buckthorn has therefore increased around the world but serious problems have, nevertheless, been encountered due to drought, salinity, diseases and insect pests. This review covers important aspects of sea buckthorn research, such as heritable and environmentally induced variation in biochemical compounds, causes and effects of the devastating dried-shrink disease, susceptibility to insect pests, methods for conventional breeding, and the utilization of DNA markers for taxonomical and population genetic analyses, and for investigating the inheritance of quality and resistance traits. We also present possibilities to implement innovative biotechnological breeding methods, especially metabolite profiling and MAS/GRC-based markers, for fast and efficient development of elite genotypes with specific nutritional- and health-related bioactive compounds and strong resistance to biotic and abiotic stress.     

Current status of genetic engineering in cotton (Gossypium hirsutum L): an assessment

Abstract

Cotton is considered as the foremost commercially important fiber crop and is deemed as the backbone of the textile industry. The productivity of cotton crop, worldwide, is severely hampered by the occurrence of pests, weeds, pathogens apart from various environmental factors. Several beneficial agronomic traits, viz., early maturity, improved fiber quality, heat tolerance, etc. have been successfully incorporated into cotton varieties employing conventional hybridization and mutation breeding. Crop losses, due to biotic factors, are substantial and may be reduced through certain crop protection strategies. In recent years, pioneering success has been achieved through the adoption of modern biotechnological approaches. Genetically engineered cotton varieties, expressing Bacillus thuringiensis cry genes, proved to be highly successful in controlling the bollworm complex. Various other candidate genes responsible for resistance to insect pests and pathogens, tolerance to major abiotic stress factors such as temperature, drought and salinity, have been introduced into cotton via genetic engineering methods to enhance the agronomic performance of cotton cultivars. Furthermore, genes for improving the seed oil quality and fiber characteristics have been identified and introduced into cotton cultivars. This review provides a brief overview of the various advancements made in cotton through genetic engineering approaches.     

Effects of drought, temperature, herbivory, and genotype on plant–insect interactions in soybean (Glycine max)

Climate change is predicted to cause continued increases in global temperatures, greater variability in precipitation and in some cases, more frequent insect pest outbreaks. Here we seek to understand how abiotic and biotic stresses associated with climate change can affect plant-herbivore interactions in a model crop species (soybean, Glycine max (L.) Merr.) by answering three questions: (1) Do the combined effects of abiotic and biotic stresses associated with climate change cause synergistic negative effects on plant biomass? (2) Can abiotic stress affect resistance of plants to insect herbivores? (3) Does genetic variation in plant traits modify a plant’s response to stress? We performed three experiments in controlled growth environments using up to 51 soybean genotypes selected to vary in numerous traits associated with drought and resistance against pests (e.g., insect herbivores, nematodes, and pathogenic fungi), and up to 3 generalist-feeding herbivorous noctuid moth species (Helicoverpa zea, Heliothis virescens, and Spodoptera exigua) that commonly feed on soybean in North America. Drought and herbivory had the largest and the most consistent negative effects on plant performance, reducing the above- and below-ground biomass by 10-45 %, whereas increased temperature had little to no effect on plants. Drought also increased susceptibility to generalist noctuid herbivores, but these results varied dramatically in magnitude and direction among plant genotypes. Our experiments show that the effects of abiotic and biotic stress on soybean biomass were largely due to the additive effects of these stresses, and there exists substantial genetic variation in the soybean germplasm pool we studied that could be used as a source of parental stock in breeding new crops that can more effectively tolerate and resist the combined negative effects of insect herbivory and drought.     

Exophiala sp. LHL08 reprograms Cucumis sativus to higher growth under abiotic stresses

Endophytic fungi are potential sources of secondary metabolites; however, they are little known for phytohormones secretion and amelioration of plant growth under abiotic stresses. We isolated a novel endophyte from the roots of Cucumis sativus and identified it as a strain of Exophiala sp. by sequencing internal transcribed spacer/large subunit rDNA and phylogenetic analysis. Prior to identification, culture filtrate (CF) of Exophiala sp. has shown significant growth promotion of Waito‐C [a gibberellins (GAs)‐deficient mutant cultivar] and Dongjin‐byeo (normal GAs biosynthesis cultivar) rice seedlings. CF analysis of Exophiala sp. showed the presence of physiologically active GAs (GA₁, GA₃, GA₄ and GA₇) and inactive GAs (GA₅, GA₈, GA₉, GA₁₂ and GA₂₀). Exophiala sp. had higher GAs in its CF than wild‐type strain of Gibberella fujikuroi except GA₃. Influence of Exophiala sp. was assessed on cucumber plant's growth and endogenous abscisic acid (ABA), salicylic acid (SA) and bioactive GAs under salinity and drought stresses. Exophiala sp.‐treated plants have shown significantly higher growth and rescued the host plants from stress promulgated water deficit, osmotic and cellular damage. The altered levels of stress‐responsive ABA showed low level of stress confined to endophyte‐applied plants than control. Elevated levels of SA and bioactive GAs (GA₃ and GA₄) in endophyte‐associated plants suggest stress‐modulating response toward salinity and drought. In conclusion, symbiotic relations between Exophiala and cucumber have reprogrammed the host plant growth under abiotic stresses, thus indicating a possible threshold role of endophytic fungi in stress alleviation. This study could be extended for improving agricultural productivity under extreme environmental conditions.     

Accumulation of terpenoid phytoalexins in maize roots is associated with drought tolerance

Maize (Zea mays) production, which is of global agro‐economic importance, is largely limited by herbivore pests, pathogens and environmental conditions, such as drought. Zealexins and kauralexins belong to two recently identified families of acidic terpenoid phytoalexins in maize that mediate defence against both pathogen and insect attacks in aboveground tissues. However, little is known about their function in belowground organs and their potential to counter abiotic stress. In this study, we show that zealexins and kauralexins accumulate in roots in response to both biotic and abiotic stress including, Diabrotica balteata herbivory, Fusarium verticillioides infection, drought and high salinity. We find that the quantity of drought‐induced phytoalexins is positively correlated with the root‐to‐shoot ratio of different maize varieties, and further demonstrate that mutant an2 plants deficient in kauralexin production are more sensitive to drought. The induction of phytoalexins in response to drought is root specific and does not influence phytoalexin levels aboveground; however, the accumulation of phytoalexins in one tissue may influence the induction capacity of other tissues.     

Fungal growth promotor endophytes: a pragmatic approach towards sustainable food and agriculture

Agricultural productivity suffers a heavy loss due to plant pathogens, insect pests and various abiotic stresses. Agriculture being the world’s largest economic sector, it is the need of time to find and establish the ideal strategy for sustainable agriculture and improvement in crop growth. Endophytes are microorganisms that asymptomatically grow within the plant tissues without causing any disease to the host. Endophytic fungi live in symbiotic association with plants and play an important role in plant growth promotion, higher seed yield and plants resistant to various biotic, abiotic stresses and diseases. Many are able to produce antimicrobial compounds, plant growth hormones and various agrochemical bioactive metabolites. These mycoendophytes hold enormous potential for the development of eco-friendly and economically viable agricultural products. In this review we focused on the endophytic fungi recovered from different medicinal plants, their active principles involved in plant growth enhancement and the applications of fungal endophytes in agriculture. Moreover, we also discussed about endophytic fungi and their pragmatic approach towards sustainable food and agriculture.     

Reverse influence of riparian buffer width on herbivorous and predatory Hemiptera

Insects are major contributors to farmland biodiversity, and their economic roles are also diverse. Many herbivorous species are crop pests, while predatory insects have the potential to act as biological controls against pests. Overall insect diversity has declined as a result of intensified agricultural practices. Riparian buffers may support insect populations in intensively cultivated areas, but their actual impact on the balance between harmful pests and beneficial predators is not known. It can be postulated that this impact may vary depending on the characteristics and location of the riparian buffer itself. We investigated the possibility that the biotic and abiotic attributes of agricultural riparian buffers adjoining crop fields and watercourses can explain the species composition of hemipteran assemblages. In particular, we were interested in the abundances of species belonging to the genus Nabis (generalist predators) and recognized and potential pests of cereal crops. Riparian buffer width and the presence or absence of woody plants were not associated with hemipteran species turnover among riparian buffers. In contrast, differences in the degree of dominance by grasses, in plant species turnover, and in which crop plant was cultivated in the adjacent field, explained a significant proportion of the variance in hemipteran species turnover. The abundance of predatory Nabis species increased with increasing riparian buffer width, whereas the abundance of recognized and potential crop pests decreased. The reverse patterns in the predatory and herbivorous Heteroptera suggest that increasing riparian buffer width might enhance biological control by Nabis predators.     

An osmotin from the resurrection plant Tripogon loliiformis (TlOsm) confers tolerance to multiple abiotic stresses in transgenic rice

Osmotin is a key protein associated with abiotic and biotic stress response in plants. In this study, an osmotin from the resurrection plant Tripogon loliiformis (TlOsm) was characterized and functionally analyzed under abiotic stress conditions in T. loliiformis as well as in transgenic Nicotiana tabacum (tobacco) and Oryza sativa (rice) plants. Real‐time PCR analysis on mixed elicitor cDNA libraries from T. loliiformis showed that TlOsm was upregulated a 1000‐fold during the early stages of osmotic stresses (cold, drought, and salinity) in both shoots and roots but downregulated in shoots during heat stress. There was no change in TlOsm gene expression in roots of heat‐stressed plants and during plant development. The plasma membrane localization of TlOsm was showed in fluorescent‐tagged TlOsm tobacco plants using confocal laser scanning microscopic analysis. Transgenic rice plants expressing TlOsm were assessed for enhanced tolerance to salinity, drought and cold stresses. Constitutively expressed TlOsm in transgenic rice plants showed increased tolerance to cold, drought and salinity stress when compared with the wild‐type and vector control counterparts. This was evidenced by maintained growth, retained higher water content and membrane integrity, and improved survival rate of TlOsm‐expressing plants. The results thus indicate the involvement of TlOsm in plant response to multiple abiotic stresses, possibly through the signaling pathway, and highlight its potential applications for engineering crops with improved tolerance to cold, drought and salinity stress.     

Amaranthaceae halophytes from the French Flanders coast of the North Sea: a review of their phytochemistry and biological activities

Halophytes are plant species that tolerate high salinity levels. To adapt to these particular abiotic conditions, they develop multiple physiological, biochemical and molecular mechanisms, including the biosynthesis of osmolytes, enzymes and specialized metabolites. The French Flanders coast of the North Sea is an ideal environment for this kind of plant. Amaranthaceae is one of the most represented botanical families of halophytes present on this coast, with 15 species belonging to 7 genera, namely Atriplex, Beta, Halimione, Kali, Oxybasis, Salicornia and Suaeda. Some of these species are well known as wild edible plants, and some are used in traditional medicine. This review examines the chemistry of these species and their potential for human health.

     

Economic impact of exotic insect pests in Brazilian agriculture

Agriculture represents one of the major strengths of the economic sector in Brazil. The need to avoid economic losses because of insect pest populations is one of the greatest challenges faced by this sector. Insect pests have caused annual losses of US$12.0 billion to the Brazilian economy, of which approximately US$1.6 billion are because of exotic pest species. Furthermore, exotic insect species often show greater potential to cause harm than native species. In Brazil, since the late nineteenth century, 24 species of insect pests have been introduced into the country, and they have caused significant economic losses. Many of these species, including Bemisia tabaci, Hypothenemus hampei, Ceratitis capitata, Oryzophagus oryzae and Anthonomus grandis, are major crop pests, and they were accidentally introduced during trading of agricultural products. In this review, we present an overview of Brazilian agriculture, a brief history of the introduction of insect pests in the country and the Brazilian legislation on agricultural defence, and we estimate the economic losses caused to the Brazilian economy by the main insect pest species that have been introduced into Brazil over the last 112 years.     

Feeding on tea GH19 chitinase enhances tea defense responses induced by regurgitant derived from Ectropis grisescens

Multiple isoforms of chitinases participate in plant defense against outside invaders. However, the functions of hydrolase family 19 (GH19) chitinases on pest control remain largely unknown. Here we reported the isolation and functional analysis of a gene CsChi19, which encodes a GH19 endochitinase protein of 332 amino acid residues from tea plant (Camellia sinensis). CsChi19 expression levels were upregulated in response to mechanical wounding, infestation by two important pests: the tea geometrid Ectropis grisescens and the tea green leafhopper Empoasca (Matsumurasca) onukii, a fungal pathogen Colletotrichum fructicola, and treatment with two phytohormones: jasmonic acid (JA) and salicylic acid. CsChi19 was heterologously expressed in Escherichia coli, and its catalytic function was further elucidated. The protein could hydrolyze colloidal chitin, and the optimum temperature and pH for its activity was 40°C and pH 5.0. CsChi19 were found to be toxic to tea pests when they were fed on artificial diets containing this protein. Interestingly, the regurgitant derived from E. grisescens fed with artificial diets containing CsChi19 protein induced stronger expression of CsMPK3, more JA burst, more accumulation of defense-related secondary metabolites, and more emission of volatiles than the regurgitant derived from E. grisescens fed only with artificial diets. Our results provide first evidence that CsChi19 is involved in mediating a novel defense mechanism of tea plant through altering the composition of the regurgitant.     

The effects of drought and herbivory on plant–herbivore interactions across 16 soybean genotypes in a field experiment

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Endophytic fungal entomopathogens with activity against plant pathogens: ecology and evolution

Dual biological control, of both insect pests and plant pathogens, has been reported for the fungal entomopathogens, Beauveria bassiana (Bals.-Criv.) Vuill. (Ascomycota: Hypocreales) and Lecanicillium spp. (Ascomycota: Hypocreales). However, the primary mechanisms of plant disease suppression are different for these fungi. Beauveria spp. produce an array of bioactive metabolites, and have been reported to limit growth of fungal plant pathogens in vitro. In plant assays, B. bassiana has been reported to reduce diseases caused by soilborne plant pathogens, such as Pythium, Rhizoctonia, and Fusarium. Evidence has accumulated that B. bassiana can endophytically colonize a wide array of plant species, both monocots and dicots. B. bassiana also induced systemic resistance when endophytically colonized cotton seedlings were challenged with a bacterial plant pathogen on foliage. Species of Lecanicillium are known to reduce disease caused by powdery mildew as well as various rust fungi. Endophytic colonization has been reported for Lecanicillium spp., and it has been suggested that induced systemic resistance may be active against powdery mildew. However, mycoparasitism is the primary mechanism employed by Lecanicillium spp. against plant pathogens. Comparisons of Beauveria and Lecanicillium are made with Trichoderma, a fungus used for biological control of plant pathogens and insects. For T. harzianum Rifai (Ascomycota: Hypocreales), it has been shown that some fungal traits that are important for insect pathogenicity are also involved in biocontrol of phytopathogens.     

Root colonization by endophytic insect-pathogenic fungi

Several ascomycetous insect-pathogenic fungi, including species in the genera Beauveria and Metarhizium, are plant root symbionts/endophytes and are termed as endophytic insect-pathogenic fungi (EIPF). The endophytic capability and insect pathogenicity of Metarhizium are coupled to provide an active method of insect-derived nitrogen transfer to plant hosts via fungal mycelia. In exchange for the insect-derived nitrogen, the plant provides photosynthate to the fungus. This symbiotic interaction offers other benefits to the plant—EIPF can improve plant growth, they are antagonistic to plant pathogens and herbivores and can enhance the plant tolerance to abiotic stresses. The mechanisms and underlying biochemical and genetic features of insect pathogenesis are generally well-established. However, there is a paucity of information regarding the underlying mechanisms in this plant-symbiotic association. Here we review five aspects of EIPF interactions with host plant roots: (i) rhizosphere colonization, (ii) signalling factors from the plant and EIPF, (iii) modulation of plant defence responses, (iv) nutrient exchange and (v) tripartite interactions with insects and other micro-organisms. The elucidation of these interactions is fundamental to understanding this symbiotic association for effective application of EIPF in an agricultural setting.