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Showing papers in "Frontiers in Plant Science in 2018"


Journal ArticleDOI
TL;DR: The concept and role of the phytomicrobiome and the agricultural context underlying food security in the 21st century are introduced and mechanisms of plant growth promotion by PGPR are discussed, including signal exchange between plant roots and PGPR and how these relationships modulate plant abiotic stress responses via induced systemic resistance.
Abstract: Microbes of the phytomicrobiome are associated with every plant tissue and, in combination with the plant form the holobiont. Plants regulate the composition and activity of their associated bacterial community carefully. These microbes provide a wide range of services and benefits to the plant; in return, the plant provides the microbial community with reduced carbon and other metabolites. Soils are generally a moist environment, rich in reduced carbon which supports extensive soil microbial communities. The rhizomicrobiome is of great importance to agriculture owing to the rich diversity of root exudates and plant cell debris that attract diverse and unique patterns of microbial colonization. Microbes of the rhizomicrobiome play key 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 enhancement of plant growth. The microbes and compounds they secrete constitute valuable biostimulants and play pivotal roles in modulating plant stress responses. Research has demonstrated that inoculating plants with plant-growth promoting rhizobacteria (PGPR) or treating plants with microbe-to-plant signal compounds can be an effective strategy to stimulate crop growth. Furthermore, these strategies can improve crop tolerance for the abiotic stresses (e.g., drought, heat, and salinity) likely to become more frequent as climate change conditions continue to develop. This discovery has resulted in multifunctional PGPR-based formulations for commercial agriculture, to minimize the use of synthetic fertilizers and agrochemicals. This review is an update about the role of PGPR in agriculture, from their collection to commercialization as low-cost commercial agricultural inputs. First, we introduce the concept and role of the phytomicrobiome and the agricultural context underlying food security in the 21st century. Next, mechanisms of plant growth promotion by PGPR are discussed, including signal exchange between plant roots and PGPR and how these relationships modulate plant abiotic stress responses via induced systemic resistance. On the application side, strategies are discussed to improve rhizosphere colonization by PGPR inoculants. The final sections of the paper describe the applications of PGPR in 21st century agriculture and the roadmap to commercialization of a PGPR-based technology.

914 citations


Journal ArticleDOI
TL;DR: Both the efficacy and environmental and food safety of this B. oleae control method, protecting olive groves and improving olive oil quality without negative effects on the natural enemy P. concolor are revealed.
Abstract: Soil treatments with Metarhizium brunneum EAMa 01/58-Su strain conducted in both Northern and Southern Spain reduced the olive fly (Bactrocera oleae) population density emerging from the soil during spring up to 70% in treated plots compared with controls. A model to determine the influence of rainfall on the conidial wash into different soil types was developed, with most of the conidia retained at the first 5 cm, regardless of soil type, with relative percentages of conidia recovered ranging between 56 and 95%. Furthermore, the possible effect of UV-B exposure time on the pathogenicity of this strain against B. oleae adults coming from surviving preimaginals and carrying conidia from the soil at adult emergence was also evaluated. The UV-B irradiance has no significant effect on M. brunneum EAMa 01/58-Su pathogenicity with B. oleae adult mortalities of 93, 90, 79, and 77% after 0, 2, 4, and 6 of UV-B irradiance exposure, respectively. In a next step for the use of these M. brunneum EAMa 01/58-Sun soil treatments within a B. oleae IPM strategy, its possible effect of on the B. oleae cosmopolitan parasitoid Psyttalia concolor, its compatibility with the herbicide oxyfluorfen 24% commonly used in olive orchards and the possible presence of the fungus in the olive oil resulting from olives previously placed in contact with the fungus were investigated. Only the highest conidial concentration (1 × 108 conidia ml-) caused significant P. concolor adult mortality (22%) with enduing mycosis in 13% of the cadavers. There were no fungal propagules in olive oil samples resulting from olives previously contaminated by EAMa 01/58-Su conidia. Finally, the strain was demonstrated to be compatible with herbicide since the soil application of the fungus reduced the B. oleae population density up to 50% even when it was mixed with the herbicide in the same tank. The fungal inoculum reached basal levels 4 months after treatments (1.6 × 103 conidia g soil-1). These results reveal both the efficacy and environmental and food safety of this B. oleae control method, protecting olive groves and improving olive oil quality without negative effects on the natural enemy P. concolor.

460 citations


Journal ArticleDOI
TL;DR: “Rostrato Rosso” was the richest in anthocyanins whilst phenolic acids were the second class in abundance, with comparable values detected between cultivars, and was highly resistant to fungal penetration and diffusion.
Abstract: Maize is a staple food source in the world, whose ancient varieties or landraces are receiving a growing attention. In this work, two Italian maize cultivars with pigmented kernels and one inbred line were investigated for untargeted phenolic profile, in vitro antioxidant capacity and resistance to Fusarium verticillioides infection. “Rostrato Rosso” was the richest in anthocyanins whilst phenolic acids were the second class in abundance, with comparable values detected between cultivars. Tyrosol equivalents were also the highest in “Rostrato Rosso” (822.4 mg kg-1). Coherently, “Rostrato Rosso” was highly resistant to fungal penetration and diffusion. These preliminary finding might help in breeding programs, aiming to develop maize lines being more resistant to infections and with improved nutraceutical value.

410 citations


Journal ArticleDOI
TL;DR: There is still a big room for improvement in combination of drought and chilling tolerance and future researches particularly using biotechnological and molecular approaches should be carried out to develop genetically engineered plants with enhanced tolerance against these stress factors.
Abstract: Plants face a combination of different abiotic stresses under field conditions which are lethal to plant growth and production. Simultaneous occurrence of chilling and drought stresses in plants due to the drastic and rapid global climate changes, can alter the morphological, physiological and molecular responses. Both these stresses adversely affect the plant growth and yields due to physical damages, physiological and biochemical disruptions, and molecular changes. In general, the co-occurrence of chilling and drought combination is even worse for crop production rather than an individual stress condition. Plants attain various common and different physiological and molecular protective approaches for tolerance under chilling and drought stresses. Nevertheless, plant responses to a combination of chilling and drought stresses are unique from those to individual stress. In the present review, we summarized the recent evidence on plant responses to chilling and drought stresses on shared as well as unique basis and tried to find a common thread potentially underlying these responses. We addressed the possible cross talk between plant responses to these stresses and discussed the potential management strategies for regulating the mechanisms of plant tolerance to drought and/or chilling stresses. To date, various novel approaches have been tested in minimizing the negative effects of combine stresses. Despite of the main improvements there is still a big room for improvement in combination of drought and chilling tolerance. Thus, future researches particularly using biotechnological and molecular approaches should be carried out to develop genetically engineered plants with enhanced tolerance against these stress factors.

399 citations


Journal ArticleDOI
TL;DR: Application of CRISPR/Cas9 techniques will result in the development of non-genetically modified (Non-GMO) crops with the desired trait that can contribute to increased yield potential under biotic and abiotic stress conditions.
Abstract: The availability of genome sequences for several crops and advances in genome editing approaches has opened up possibilities to breed for almost any given desirable trait. Advancements in genome editing technologies such as zinc finger nucleases (ZFNs), transcription activator-like effector nucleases (TALENs) has made it possible for molecular biologists to more precisely target any gene of interest. However, these methodologies are expensive and time-consuming as they involve complicated steps that require protein engineering. Unlike first-generation genome editing tools, CRISPR/Cas9 genome editing involves simple designing and cloning methods, with the same Cas9 being potentially available for use with different guide RNAs targeting multiple sites in the genome. After proof-of-concept demonstrations in crop plants involving the primary CRISPR-Cas9 module, several modified Cas9 cassettes have been utilized in crop plants for improving target specificity and reducing off-target cleavage (eg. Nmcas9, Sacas9, Stcas9). Further, the availability of Cas9 enzymes from additional bacterial species has made available options to enhance specificity and efficiency of gene editing methodologies. This review summarizes the options available to plant biotechnologists to bring about crop improvement using CRISPR/Cas9 based genome editing tools and also presents studies where CRISPR/Cas9 has been used for enhancing biotic and abiotic stress tolerance. Application of these techniques will result in the development of non-genetically modified (Non-GMO) crops with the desired trait that can contribute to increased yield potential under biotic and abiotic stress conditions.

385 citations


Journal ArticleDOI
TL;DR: The results suggest that predictions across breeding populations in tetraploid potato are presently unreliable, but that individual prediction models within populations can be combined in an additive fashion to obtain high quality prediction models relevant for several breeding populations.
Abstract: Genomic selection (GS) is becoming increasingly applicable to crops as the genotyping costs continue to decrease, which makes it an attractive alternative to traditional selective breeding based on observed phenotypes. With genome-wide molecular markers, selection based on predictions from genotypes can be made in the absence of direct phenotyping. The reliability of predictions depends strongly on the number of individuals used for training the predictive algorithms, particularly in a highly genetically diverse organism such as potatoes; however, the relationship between the individuals also has an enormous impact on prediction accuracy. Here we have studied genomic prediction in three different panels of potato cultivars, varying in size, design, and phenotypic profile. We have developed genomic prediction models for two important agronomic traits of potato, dry matter content and chipping quality. We used genotyping-by-sequencing to genotype 1,146 individuals and generated genomic prediction models from 167,637 markers to calculate genomic estimated breeding values with genomic best linear unbiased prediction. Cross-validated prediction correlations of 0.75-0.83 and 0.39-0.79 were obtained for dry matter content and chipping quality, respectively, when combining the three populations. These prediction accuracies were similar to those obtained when predicting performance within each panel. In contrast, but not unexpectedly, predictions across populations were generally lower, 0.37-0.71 and 0.28-0.48 for dry matter content and chipping quality, respectively. These predictions are not limited by the number of markers included, since similar prediction accuracies could be obtained when using merely 7,800 markers (<5%). Our results suggest that predictions across breeding populations in tetraploid potato are presently unreliable, but that individual prediction models within populations can be combined in an additive fashion to obtain high quality prediction models relevant for several breeding populations.

385 citations


Journal ArticleDOI
TL;DR: Recently, CRISPR/Cas9 has largely overtaken the other genome editing technologies due to the fact that it is easier to design and implement, has a higher success rate, and is more versatile and less expensive.
Abstract: Genome editing technologies have progressed rapidly and become one of the most important genetic tools in the implementation of pathogen resistance in plants. Recent years have witnessed the emergence of site directed modification methods using meganucleases, zinc finger nucleases (ZFNs), transcription activator-like effector nucleases (TALENs), and clustered regularly interspaced short palindrome repeats (CRISPR)/CRISPR-associated protein 9 (Cas9). Recently, CRISPR/Cas9 has largely overtaken the other genome editing technologies due to the fact that it is easier to design and implement, has a higher success rate, and is more versatile and less expensive. This review focuses on the recent advances in plant protection using CRISPR/Cas9 technology in model plants and crops in response to viral, fungal and bacterial diseases. As regards the achievement of viral disease resistance, the main strategies employed in model species such as Arabidopsis and Nicotiana benthamiana, which include the integration of CRISPR-encoding sequences that target and interfere with the viral genome and the induction of a CRISPR-mediated targeted mutation in the host plant genome, will be discussed. Furthermore, as regards fungal and bacterial disease resistance, the strategies based on CRISPR/Cas9 targeted modification of susceptibility genes in crop species such as rice, tomato, wheat, and citrus will be reviewed. After spending years deciphering and reading genomes, researchers are now editing and rewriting them to develop crop plants resistant to specific pests and pathogens.

371 citations


Journal ArticleDOI
TL;DR: A synthesis of observed trends in recent studies is provided and possible directions for future research are discussed that are hoped will provide for more knowledgeable predictions about community responses to future drought events.
Abstract: Root-associated bacterial communities play a vital role in maintaining health of the plant host. These communities exist in complex relationships, where composition and abundance of community members is dependent on a number of factors such as local soil chemistry, plant genotype and phenotype, and perturbations in the surrounding abiotic environment. One common perturbation, drought, has been shown to have drastic effects on bacterial communities, yet little is understood about the underlying causes behind observed shifts in microbial abundance. As drought may affect root bacterial communities both directly by modulating moisture availability, as well as indirectly by altering soil chemistry and plant phenotypes, we provide a synthesis of observed trends in recent studies and discuss possible directions for future research that we hope will provide for more knowledgeable predictions about community responses to future drought events.

353 citations


Journal ArticleDOI
TL;DR: This review aimed at presenting an overview of defensive systems and the regulatory network involving upstream signaling molecules including stress hormones, reactive oxygen species, gasotransmitters, polyamines, phytochromes, and calcium, as well as downstream gene regulation factors, particularly transcription factors.
Abstract: Abiotic stresses, such as low or high temperature, deficient or excessive water, high salinity, heavy metals, and ultraviolet radiation, are hostile to plant growth and development, leading to great crop yield penalty worldwide. It is getting imperative to equip crops with multistress tolerance to relieve the pressure of environmental changes and to meet the demand of population growth, as different abiotic stresses usually arise together in the field. The feasibility is raised as land plants actually have established more generalized defenses against abiotic stresses, including the cuticle outside plants, together with unsaturated fatty acids, reactive species scavengers, molecular chaperones, and compatible solutes inside cells. In stress response, they are orchestrated by a complex regulatory network involving upstream signaling molecules including stress hormones, reactive oxygen species, gasotransmitters, polyamines, phytochromes, and calcium, as well as downstream gene regulation factors, particularly transcription factors. In this review, we aimed at presenting an overview of these defensive systems and the regulatory network, with an eye to their practical potential via genetic engineering and/or exogenous application.

314 citations


Journal ArticleDOI
TL;DR: New developments in the industry for AM-related products for agriculture, horticulture, and landscaping are discussed, and future potential and limits toward the use of AM fungi for plant production are highlighted.
Abstract: Arbuscular mycorrhiza (AM) is the most common symbiotic association of plants with microbes AM fungi occur in the majority of natural habitats and they provide a range of important ecological services, in particular by improving plant nutrition, stress resistance and tolerance, soil structure and fertility AM fungi also interact with most crop plants including cereals, vegetables, and fruit trees, therefore, they receive increasing attention for their potential use in sustainable agriculture Basic research of the past decade has revealed the existence of a dedicated recognition and signaling pathway that is required for AM Furthermore, recent evidence provided new insight into the exchange of nutritional benefits between the symbiotic partners The great potential for application of AM has given rise to a thriving industry for AM-related products for agriculture, horticulture, and landscaping Here, we discuss new developments in these fields, and we highlight future potential and limits toward the use of AM fungi for plant production

299 citations


Journal ArticleDOI
TL;DR: Findings in various food crops are highlighted, showing how their seed composition is drastically impacted at various cellular levels due to drought and heat stresses, applied separately, or in combination.
Abstract: Drought (water deficits) and heat (high temperatures) stress are the prime abiotic constraints, under the current and climate change scenario in future. Any further increase in the occurrence, and extremity of these stresses, either individually or in combination, would severely reduce the crop productivity and food security, globally. Although, they obstruct productivity at all crop growth stages, the extent of damage at reproductive phase of crop growth, mainly the seed filling phase, is critical and causes considerable yield losses. Drought and heat stress substantially affect the seed yields by reducing seed size and number, eventually affecting the commercial trait ‘100 seed weight’ and seed quality. Seed filling is influenced by various metabolic processes occurring in the leaves, especially production and translocation of photoassimilates, importing precursors for biosynthesis of seed reserves, minerals and other functional constituents. These processes are highly sensitive to drought and heat, due to involvement of array of diverse enzymes and transporters, located in the leaves and seeds. We highlight here the findings in various food crops showing how their seed composition is drastically impacted at various cellular levels due to drought and heat stresses, applied separately, or in combination. The combined stresses are extremely detrimental for seed yield and its quality, and thus need more attention. Understanding the precise target sites regulating seed filling events in leaves and seeds, and how they are affected by abiotic stresses, is imperative to enhance the seed quality. It is vital to know the physiological, biochemical and genetic mechanisms, which govern the various seed filling events under stress environments, to devise strategies to improve stress tolerance. Converging modern advances in physiology, biochemistry and biotechnology, especially the “omics” technologies might provide a strong impetus to research on this aspect. Such application, along with effective agronomic management system would pave the way in developing crop genotypes/varieties with improved productivity under drought and/or heat stresses.

Journal ArticleDOI
TL;DR: The interactions within the rhizosphere and subsequent above-ground ‘signalomics’ are reviewed, and the contributions that mass spectrometric-based metabolomic approaches can bring to the study of plant-beneficial – and priming events are emphasized.
Abstract: Plant roots communicate with microbes in a sophisticated manner through chemical communication within the rhizosphere, thereby leading to biofilm formation of beneficial microbes and, in the case of plant growth-promoting rhizomicrobes / -bacteria (PGPR), resulting in priming of defense or induced resistance in the plant host. The knowledge of plant-plant and plant-microbe interactions have been greatly extended over recent years; however, the chemical communication leading to priming is far from being well understood. Furthermore, linkage between below- and above-ground plant physiological processes adds to the complexity. In metabolomics studies, the main aim is to profile and annotate all exo- and endo-metabolites in a biological system that drive and participate in physiological processes. Recent advances in this field has enabled researchers to analyze hundreds of compounds in one sample over a short time period. Here, from a metabolomics viewpoint, we review the interactions within the rhizosphere and subsequent above-ground ‘signalomics’, and emphasize the contributions that mass spectrometric-based metabolomic approaches can bring to the study of plant-beneficial – and priming events.

Journal ArticleDOI
TL;DR: This review takes into account the current state of knowledge of the harmful effects of heavyMetal stress, the signaling responses to metal stress, and the role of plant-associated microbes in metal stress tolerance, and highlights the challenges and opportunities.
Abstract: Several anthropogenic activities including mining, modern agricultural practices, and industrialization have long-term detrimental effect on our environment. All these factors lead to increase in heavy metal concentration in soil, water, and air. Soil contamination with heavy metals cause several environmental problems and imparts toxic effect on plant as well as animals. In response to these adverse conditions, plants evolve complex molecular and physiological mechanisms for better adaptability, tolerance, and survival. Nowadays conventional breeding and transgenic technology are being used for development of metal stress resistant varieties which, however, are time consuming and labor intensive. Interestingly the use of microbes as an alternate technology for improving metal tolerance of plants is gaining momentum recently. The use of these beneficial microorganisms is considered as one of the most promising methods for safe crop-management practices. Interaction of plants with soil microorganisms can play a vital role in acclimatizing plants to metalliferous environments, and can thus be explored to improve microbe-assisted metal tolerance. Plant-associated microbes decrease metal accumulation in plant tissues and also help to reduce metal bioavailability in soil through various mechanisms. Nowadays, a novel phytobacterial strategy, i.e., genetically transformed bacteria has been used to increase remediation of heavy metals and stress tolerance in plants. This review takes into account our current state of knowledge of the harmful effects of heavy metal stress, the signaling responses to metal stress, and the role of plant-associated microbes in metal stress tolerance. The review also highlights the challenges and opportunities in this continued area of research on plant-microbe-metal interaction.

Journal ArticleDOI
TL;DR: This review focused on three basic HM phytoextraction strategies that differ in the type of plant species being employed: natural hyperaccumulators, fast-growing plant species with high-biomass production and, potentially, plants genetically engineered toward a phenotype that favors efficient HM uptake and boosted HM tolerance.
Abstract: Pollution by heavy metals (HM) represents a serious threat for both the environment and human health. Due to their elemental character, HM cannot be chemically degraded, and their detoxification in the environment mostly resides either in stabilization in situ or in their removal from the matrix, e.g., soil. For this purpose, phytoremediation, i.e., the application of plants for the restoration of a polluted environment, has been proposed as a promising green alternative to traditional physical and chemical methods. Among the phytoremediation techniques, phytoextraction refers to the removal of HM from the matrix through their uptake by a plant. It possesses considerable advantages over traditional techniques, especially due to its cost effectiveness, potential treatment of multiple HM simultaneously, no need for the excavation of contaminated soil, good acceptance by the public, the possibility of follow-up processing of the biomass produced, etc. In this review, we focused on three basic HM phytoextraction strategies that differ in the type of plant species being employed: natural hyperaccumulators, fast-growing plant species with high-biomass production and, potentially, plants genetically engineered toward a phenotype that favors efficient HM uptake and boosted HM tolerance. Considerable knowledge on the applicability of plants for HM phytoextraction has been gathered to date from both lab-scale studies performed under controlled model conditions and field trials using real environmental conditions. Based on this knowledge, many specific applications of plants for the remediation of HM-polluted soils have been proposed. Such studies often also include suggestions for the further processing of HM-contaminated biomass, therefore providing an added economical value. Based on the examples presented here, we recommend that intensive research be performed on the selection of appropriate plant taxa for various sets of conditions, environmental risk assessment, the fate of HM-enriched biomass, economical aspects of the process, etc.

Journal ArticleDOI
TL;DR: The current knowledge on the epidemiology of FW of banana is summarized, highlighting knowledge gaps in pathogen survival and dispersal, factors driving disease intensity, soil and plant microbiome and the dynamics of the disease.
Abstract: Banana production is seriously threatened by Fusarium wilt (FW), a disease caused by the soil-borne fungus Fusarium oxysporum f. sp. cubense (Foc). In the mid-twentieth century FW, also known as "Panama disease", wiped out the Gros Michel banana industry in Central America. The devastation caused by Foc race 1 was mitigated by a shift to resistant Cavendish cultivars, which are currently the source of 99% of banana exports. However, a new strain of Foc, the tropical race 4 (TR4), attacks Cavendish clones and a diverse range of other banana varieties. Foc TR4 has been restricted to East and parts of Southeast Asia for more than 20 years, but since 2010 the disease has spread westward into five additional countries in Southeast and South Asia (Vietnam, Laos, Myanmar, India, and Pakistan) and at the transcontinental level into the Middle East (Oman, Jordan, Lebanon, and Israel) and Africa (Mozambique). The spread of Foc TR4 is of great concern due to the limited knowledge about key aspects of disease epidemiology and the lack of effective management models, including resistant varieties and soil management approaches. In this review we summarize the current knowledge on the epidemiology of FW of banana, highlighting knowledge gaps in pathogen survival and dispersal, factors driving disease intensity, soil and plant microbiome and the dynamics of the disease. Comparisons with FW in other crops were also made to indicate possible differences and commonalities. Our current understanding of the role of main biotic and abiotic factors on disease intensity is reviewed, highlighting research needs and futures directions. Finally, a set of practices and their impact on disease intensity are discussed and proposed as an integrative management approach that could eventually be used by a range of users, including plant protection organizations, researchers, extension workers and growers.

Journal ArticleDOI
TL;DR: The role of Glutathione S-transferases (GSTs) in abiotic and biotic stresses with an emphasis on As uptake, metabolism, and detoxification in plants is discussed in this paper.
Abstract: Arsenic (As), naturally occurring metalloid and a potential hazardous material, is found in low concentrations in the environment and emerges from natural sources and anthropogenic activities. The presence of As in ground water, which is used for irrigation, is a matter of great concern since it affects crop productivity and contaminates food chain. In plants, As alters various metabolic pathways in cells including the interaction of substrates/enzymes with the sulfhydryl groups of proteins and the replacement of phosphate in ATP for energy. In addition, As stimulates the generation of free radicals and reactive oxygen species (ROS), resulting in oxidative stress. Glutathione S-transferases (GSTs) quench reactive molecules with the addition of glutathione (GSH) and protect the cell from oxidative damage. GSTs are a multigene family of isozymes, known to catalyze the conjugation of GSH to miscellany of electrophilic and hydrophobic substrates. GSTs have been reported to be associated with plant developmental processes and are responsive to multitude of stressors. In past, several studies suggested involvement of plant GST gene family in As response due to the requirement of sulfur and GSH in the detoxification of this toxic metalloid. This review provides updated information about the role of GSTs in abiotic and biotic stresses with an emphasis on As uptake, metabolism, and detoxification in plants. Further, the genetic manipulations that helped in enhancing the understanding of the function of GSTs in abiotic stress response and heavy metal detoxification has been reviewed.

Journal ArticleDOI
TL;DR: The review identifies several perspectives for future research between the scientific community and private industry to design and develop a second generation of PBs products (biostimulant 2.0) with specific biostimulatory action to render agriculture more sustainable and resilient.
Abstract: Over the past ten years, interest in plant biostimulants (PBs) has been on the rise compelled by the growing interest of scientists, extension specialists, private industry and growers in integrating these products in the array of environmentally-friendly tools that secure improved crop performance and yield stability. Based on the new EU regulation PBs are defined through claimed agronomic effects, such as improvement of nutrient use efficiency, tolerance to abiotic stressors and crop quality. This definition entails diverse organic and inorganic substances and/or microorganisms such as humic acids, protein hydrolysates, seaweed extracts, mycorrhizal fungi and N-fixing bacteria. The current mini-review provides an overview of the direct (stimulatory on C and N metabolism) and indirect (enhancing nutrient uptake and modulating root morphology) mechanisms by which microbial and non-microbial PBs improve nutrient efficiency, plant performance and physiological status, resilience to environmental stressors and stimulate plant microbiomes. The scientific advances underlying synergistic and additive effects of microbial and non-microbial PBs are compiled and discussed for the first time. The review identifies several perspectives for future research between the scientific community and private industry to design and develop a second generation of PBs products (biostimulant 2.0) with specific biostimulatory action to render agriculture more sustainable and resilient.

Journal ArticleDOI
TL;DR: Functional studies revealed that overexpression or silencing of specific GSTs can markedly modify disease symptoms and also pathogen multiplication rates, but very limited information is available about the exact metabolic functions of disease-induced GST isoenzymes and about their endogenous substrates.
Abstract: Plant glutathione S-transferases (GSTs) are ubiquitous and multifunctional enzymes encoded by large gene families. A characteristic feature of GST genes is their high inducibility by a wide range of stress conditions including biotic stress. Early studies on the role of GSTs in plant biotic stress showed that certain GST genes are specifically up-regulated by microbial infections. Later numerous transcriptome-wide investigations proved that distinct groups of GSTs are markedly induced in the early phase of bacterial, fungal and viral infections. Proteomic investigations also confirmed the accumulation of multiple GST proteins in infected plants. Furthermore, functional studies revealed that overexpression or silencing of specific GSTs can markedly modify disease symptoms and also pathogen multiplication rates. However, very limited information is available about the exact metabolic functions of disease-induced GST isoenzymes and about their endogenous substrates. The already recognized roles of GSTs are the detoxification of toxic substances by their conjugation with glutathione, the attenuation of oxidative stress and the participation in hormone transport. Some GSTs display glutathione peroxidase activity and these GSTs can detoxify toxic lipid hydroperoxides that accumulate during infections. GSTs can also possess ligandin functions and participate in the intracellular transport of auxins. Notably, the expression of multiple GSTs is massively activated by salicylic acid and some GST enzymes were demonstrated to be receptor proteins of salicylic acid. Furthermore, induction of GST genes or elevated GST activities have often been observed in plants treated with beneficial microbes (bacteria and fungi) that induce a systemic resistance response (ISR) to subsequent pathogen infections. Further research is needed to reveal the exact metabolic functions of GST isoenzymes in infected plants and to understand their contribution to disease resistance.

Journal ArticleDOI
TL;DR: The findings indicated that CRISPR/Cas9 is a site-specific genome editing technology that allows highly efficient target mutagenesis in multiple genes of interest, and lycopene content in tomato fruit subjected to genome editing was successfully increased to about 5.1-fold.
Abstract: Numerous studies have been focusing on breeding tomato plants with enhanced lycopene accumulation, considering its positive effects of fruits on the visual and functional properties. In this study, we used a bidirectional strategy: promoting the biosynthesis of lycopene, while inhibiting the conversion from lycopene to β- and α-carotene. The accumulation of lycopene was promoted by knocking down some genes associated with the carotenoid metabolic pathway. Finally, five genes were selected to be edited in genome by CRISPR/Cas9 system using Agrobacterium tumefaciens-mediated transformation. Our findings indicated that CRISPR/Cas9 is a site-specific genome editing technology that allows highly efficient target mutagenesis in multiple genes of interest. Surprisingly, the lycopene content in tomato fruit subjected to genome editing was successfully increased to about 5.1-fold. The homozygous mutations were stably transmitted to subsequent generations. Taken together, our results suggest that CRISPR/Cas9 system can be used for significantly improving lycopene content in tomato fruit with advantages such as high efficiency, rare off-target mutations, and stable heredity.

Journal ArticleDOI
TL;DR: The main aim of the present review is to discuss the major factors determining stress-related protein accumulation and their final biological functions to provide an overview of themajor factors which have to be considered when interpreting data from proteomic studies on stress-treated plants.
Abstract: HIGHLIGHTS: Major environmental and genetic factors determining stress-related protein abundance are discussed.Major aspects of protein biological function including protein isoforms and PTMs, cellular localization and protein interactions are discussed.Functional diversity of protein isoforms and PTMs is discussed. Abiotic stresses reveal profound impacts on plant proteomes including alterations in protein relative abundance, cellular localization, post-transcriptional and post-translational modifications (PTMs), protein interactions with other protein partners, and, finally, protein biological functions. The main aim of the present review is to discuss the major factors determining stress-related protein accumulation and their final biological functions. A dynamics of stress response including stress acclimation to altered ambient conditions and recovery after the stress treatment is discussed. The results of proteomic studies aimed at a comparison of stress response in plant genotypes differing in stress adaptability reveal constitutively enhanced levels of several stress-related proteins (protective proteins, chaperones, ROS scavenging- and detoxification-related enzymes) in the tolerant genotypes with respect to the susceptible ones. Tolerant genotypes can efficiently adjust energy metabolism to enhanced needs during stress acclimation. Stress tolerance vs. stress susceptibility are relative terms which can reflect different stress-coping strategies depending on the given stress treatment. The role of differential protein isoforms and PTMs with respect to their biological functions in different physiological constraints (cellular compartments and interacting partners) is discussed. The importance of protein functional studies following high-throughput proteome analyses is presented in a broader context of plant biology. In summary, the manuscript tries to provide an overview of the major factors which have to be considered when interpreting data from proteomic studies on stress-treated plants.

Journal ArticleDOI
TL;DR: It is suggested that building SOC enhances agronomic productivity at only 0–100 kg N ha-1, to reduce the off-farm N losses depending on the environmental zones, land use and the production system.
Abstract: SOC (soil organic carbon) has a vital role to enhance agricultural productivity and for mitigation of climate change. To quantify SOC effects on productivity, process models serve as a robust tool to keep track of multiple plant and soil factors and their interactions affecting SOC dynamics. We used soil-plant-atmospheric model viz. DAISY, to assess effects of SOC on nitrogen (N) supply and plant available water (PAW) under varying N fertilizer rates in winter wheat(Triticum aestivum) in Denmark. The study objective was assessment of SOC effects on winter wheat grain and aboveground biomass accumulation at three SOC levels (low:0.7% SOC ; reference: 1.3% SOC and high: 2% SOC) with five nitrogen rates (0-200 kg N ha-1) and PAW at low, reference and high SOC levels. The three SOC levels had significant effects on grain yields and aboveground biomass accumulation at only 0-100 kg N ha-1 and the SOC effects decreased with increasing N rates until no effects at 150-200 kg N ha-1. PAW had significant positive correlation with SOC content, with high SOC retaining higher PAW compared to low and reference SOC. The mean PAW and SOC correlation was given by PAW% = 1.0073 ×SOC% + 15.641. For the 0.7-2% SOC range, the PAW increase was small with no significant effects on grain yields and aboveground biomass accumulation. The higher winter wheat grain and aboveground biomass was attributed to higher N supply in N deficient wheat production system. Our study suggested that building SOC enhances agronomic productivity at only 0-100 kg N ha-1. Maintenance of SOC stock will require regular replenishment of SOC, to compensate for the mineralization process degrading SOC overtime. Hence, management can maximize realisation of SOC benefits by building up SOC and maintaining N rates in the range 0-100 kg N ha-1, to reduce the off-farm N losses depending on the environmental zones, land use and the production system.

Journal ArticleDOI
TL;DR: This review demonstrates the key role of MAPKs in synchronously controlling the other signaling components and regulators in metal stress and attempts have been made to focus on metal transporters and chelators that are regulated by MAPK signaling.
Abstract: Plants confront multifarious environmental stresses widely divided into abiotic and biotic stresses, of which heavy metal stress represents one of the most damaging abiotic stresses Heavy metals cause toxicity by targeting crucial molecules and vital processes in the plant cell One of the approaches by which heavy metals act in plants is by over production of reactive oxygen species (ROS) either directly or indirectly Plants act against such overdose of metal in the environment by boosting the defense responses like metal chelation, sequestration into vacuole, regulation of metal intake by transporters, and intensification of antioxidative mechanisms This response shown by plants is the result of intricate signaling networks functioning in the cell in order to transmit the extracellular stimuli into an intracellular response The crucial signaling components involved are calcium signaling, hormone signaling, and mitogen activated protein kinase (MAPK) signaling that are discussed in this review Apart from signaling components other regulators like microRNAs and transcription factors also have a major contribution in regulating heavy metal stress This review demonstrates the key role of MAPKs in synchronously controlling the other signaling components and regulators in metal stress Further, attempts have been made to focus on metal transporters and chelators that are regulated by MAPK signaling

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TL;DR: Application of this system will provide new opportunities to deliver improved genotypes and agronomic interventions via more efficient and reliable phenotyping of these important traits in large experiments through rapid multi-temporal and non-destructive estimation of canopy height, ground cover and above-ground biomass.
Abstract: The authors acknowledge the financial support of the Australian Government National Collaborative Research Infrastructure Strategy (Australian Plant Phenomics Facility) and the Grains Research and Development Corporation (GRDC).

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TL;DR: The mechanisms of crosstalk between MAPK cascades and plant hormone signaling pathways are highlighted and recent findings on MAPK regulation and function in various cellular processes are summarized.
Abstract: Mitogen-activated protein kinase (MAPK) modules play key roles in the transduction of environmental and developmental signals through phosphorylation of downstream signaling targets, including other kinases, enzymes, cytoskeletal proteins or transcription factors, in all eukaryotic cells. A typical MAPK cascade consists of at least three sequentially acting serine/threonine kinases, a MAP kinase kinase kinase (MAPKKK), a MAP kinase kinase (MAPKK) and finally, the MAP kinase (MAPK) itself, with each phosphorylating, and hence activating, the next kinase in the cascade. Recent advances in our understanding of hormone signaling pathways have led to the discovery of new regulatory systems. In particular, this research has revealed the emerging role of crosstalk between the protein components of various signaling pathways and the involvement of this crosstalk in multiple cellular processes. Here we provide an overview of current models and mechanisms of hormone signaling with a special emphasis on the role of MAPKs in cell signaling networks. One-sentence summary: In this review we highlight the mechanisms of crosstalk between MAPK cascades and plant hormone signaling pathways and summarize recent findings on MAPK regulation and function in various cellular processes.

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TL;DR: The canonical lignin biosynthesis pathway, the recently discovered tyrosine shortcut pathway, and the biosynthesis of unconventional C-lignin are reviewed and the existence of a transcriptional co-regulation mechanism coordinating energy allowance among growth, defense and lignIn biosynthesis is proposed.
Abstract: Plant growth-defense tradeoffs are fundamental for optimizing plant performance and fitness in a changing biotic/abiotic environment This process is thought to involve readjusting resource allocation to different pathways It has been frequently observed that among secondary cell wall components, alteration in lignin biosynthesis results in changes in both growth and defense How this process is regulated, leading to growth or defense, remains largely elusive In this article, we review the canonical lignin biosynthesis pathway, the recently discovered tyrosine shortcut pathway, and the biosynthesis of unconventional C-lignin We summarize the current model of the hierarchical transcriptional regulation of lignin biosynthesis Moreover, the interface between recently identified transcription factors and the hierarchical model are also discussed We propose the existence of a transcriptional co-regulation mechanism coordinating energy allowance among growth, defense and lignin biosynthesis

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TL;DR: A meta-analysis to quantify benefits of biofertilizers in terms of yield increase, nitrogen and phosphorus use efficiency, based on 171 peer reviewed publications that met eligibility criteria provides a basis and guidance for proper choice and application of bio fertilizer.
Abstract: The application of microbial inoculants (biofertilizers) is a promising technology for future sustainable farming systems in view of rapidly decreasing phosphorus stocks and the need to more efficiently use available nitrogen (N). Various microbial taxa are currently used as biofertilizers, based on their capacity to access nutrients from fertilizers and soil stocks, to fix atmospheric nitrogen, to improve water uptake or to act as biocontrol agents. Despite the existence of a considerable knowledge on effects of specific taxa of biofertilizers, a comprehensive quantitative assessment of the performance of biofertilizers with different traits such as phosphorus solubilization and N fixation applied to various crops at a global scale is missing. We conducted a meta-analysis to quantify benefits of biofertilizers in terms of yield increase, nitrogen and phosphorus use efficiency, based on 171 peer reviewed publications that met eligibility criteria. Major findings are: (i) the superiority of biofertilizer performance in dry climates over other climatic regions (yield response: dry climate +20.0 ± 1.7%, tropical climate +14.9 ± 1.2%, oceanic climate +10.0 ± 3.7%, continental climate +8.5 ± 2.4%); (ii) meta-regression analyses revealed that yield response due to biofertilizer application was generally small at low soil P levels; efficacy increased along higher soil P levels in the order arbuscular mycorrhizal fungi (AMF), P solubilizers, and N fixers; (iii) meta-regressions showed that the success of inoculation with AMF was greater at low organic matter content and at neutral pH. Our comprehensive analysis provides a basis and guidance for proper choice and application of biofertilizers.

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TL;DR: This review is aimed to focus on the current knowledge of protein persulfidation and addresses the regulation mechanisms that are disclosed based on the knowledge from other cysteine modifications.
Abstract: Hydrogen sulfide (H2S) has been largely referred as a toxic gas and environmental hazard, but recent years, it has emerged as an important gas-signaling molecule with effects on multiple physiological processes in both animal and plant systems. The regulatory functions of H2S in plants are involved in important processes such as the modulation of defense responses, plant growth and development, and the regulation of senescence and maturation. The main signaling pathway involving sulfide has been proven to be through protein persulfidation (alternatively called S-sulfhydration), in which the thiol group of cysteine (-SH) in proteins is modified into a persulfide group (-SSH). This modification may cause functional changes in protein activities, structures, and subcellular localizations of the target proteins. New shotgun proteomic approaches and bioinformatic analyses have revealed that persulfidated cysteines regulate important biological processes, highlighting their importance in cell signaling, since about one in 20 proteins in Arabidopsis is persulfidated. During oxidative stress, an increased persulfidation has been reported and speculated that persulfidation is the protective mechanism for protein oxidative damage. Nevertheless, cysteine residues are also oxidized to different post-translational modifications such S-nitrosylation or S-sulfenylation, which seems to be interconvertible. Thus, it must imply a tight cysteine redox regulation essential for cell survival. This review is aimed to focus on the current knowledge of protein persulfidation and addresses the regulation mechanisms that are disclosed based on the knowledge from other cysteine modifications.

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TL;DR: This article provides a review of the efforts to date on the exploration of CWRs as a source of tolerance to multiple biotic and abiotic stresses in four global crops of importance; maize, rice, cotton, and soybean.
Abstract: Global food demand is expected to nearly double by 2050 due to an increase in the world's population. The Green Revolution has played a key role in the past century by increasing agricultural productivity worldwide, however, limited availability and continued depletion of natural resources such as arable land and water will continue to pose a serious challenge for global food security in the coming decades. High yielding varieties with proven tolerance to biotic and abiotic stresses, superior nutritional profiles, and the ability to adapt to the changing environment are needed for continued agricultural sustainability. The narrow genetic base of modern cultivars is becoming a major bottleneck for crop improvement efforts and, therefore, the use of crop wild relatives (CWRs) is a promising approach to enhance genetic diversity of cultivated crops. This article provides a review of the efforts to date on the exploration of CWRs as a source of tolerance to multiple biotic and abiotic stresses in four global crops of importance; maize, rice, cotton, and soybean. In addition to the overview of the repertoire and geographical spread of CWRs in each of the respective crops, we have provided a comprehensive discussion on the morphological and/or genetic basis of the traits along with some examples, when available, of the research in the transfer of traits from CWRs to cultivated varieties. The emergence of modern molecular and genomic technologies has not only accelerated the pace of dissecting the genetics underlying the traits found in CWRs, but also enabled rapid and efficient trait transfer and genome manipulation. The potential and promise of these technologies has also been highlighted in this review.

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TL;DR: Overall, inoculations with Trichoderma may be considered as a viable strategy to manage the nutrient content of leafy horticulture crops cultivated in low fertility soils, and assist vegetable growers in reducing the use of synthetic fertilizers, developing sustainable management practices to optimize N use efficiency.
Abstract: Microbial inoculants such as Trichoderma-based products are receiving great interest among researchers and agricultural producers for their potential to improve crop productivity, nutritional quality as well as resistance to plant pathogens/pests and numerous environmental stresses. Two greenhouse experiments were conducted to assess the effects of Trichoderma-based biostimulants under suboptimal, optimal and supraoptimal levels of nitrogen (N) fertilization in two leafy vegetables: Iceberg lettuce (Lactuca sativa L.) and rocket (Eruca sativa Mill.). The yield, nutritional characteristics, N uptake and mineral composition were analyzed for each vegetable crop after inoculation with Trichoderma strains T. virens (GV41) or T. harzianum (T22), and results were compared to non-inoculated plants. In addition, the effect of the Trichoderma-based biostimulants on microbes associated with the rhizosphere in terms of prokaryotic and eurkaryotic composition and concentration using DGGE was also evaluated. Trichoderma-based biostimulants, in particular GV41, positively increased lettuce and rocket yield in the unfertilized plots. The highest marketable lettuce fresh yield was recorded with either of the biostimulant inoculations when plants were supplied with optimal levels of N. The inoculation of rocket with GV41, and to a lesser degree with T22, elicited an increase in total ascorbic acid under both optimal and high N conditions. T. virens GV41 increased N-use efficiency of lettuce, and favoured the uptake of native N present in the soil of both lettuce and rocket. The positive effect of biostimulants on nutrient uptake and crop growth was species-dependent, being more marked with lettuce. The best biostimulation effects from the Trichoderma treatments were observed in both crops when grown under low N availability. The Trichoderma inoculation strongly influenced the composition of eukaryotic populations in the rhizosphere, in particularly exerting different effects with low N levels in comparison to the N fertilized plots. Overall, inoculations with Trichoderma may be considered as a viable strategy to manage the nutrient content of leafy horticulture crops cultivated in low fertility soils, and assist vegetable growers in reducing the use of synthetic fertilizers, developing sustainable management practices to optimize N use efficiency.

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TL;DR: The literature on how GA and ABA control different stages of plant development is reviewed, with focus on seed germination and selected abiotic stresses.
Abstract: Overall growth and development of a plant is regulated by complex interactions among various hormones, which is critical at different developmental stages. Some of the key aspects of plant growth include seed development, germination and plant survival under unfavorable conditions. Two of the key phytohormones regulating the associated physiological processes are gibberellins (GA) and abscisic acid (ABA). GAs participate in numerous developmental processes, including, seed development and seed germination, seedling growth, root proliferation, determination of leaf size and shape, flower induction and development, pollination and fruit expansion. Despite the association with abiotic stresses, ABA is essential for normal plant growth and development. It plays a critical role in different abiotic stresses by regulating various downstream ABA-dependent stress responses. Plants maintain a balance between GA and ABA levels constantly throughout the developmental processes at different tissues and organs, including under unfavorable environmental or physiological conditions. Here, we will review the literature on how GA and ABA control different stages of plant development, with focus on seed germination and selected abiotic stresses. The possible crosstalk of ABA and GA in specific events of the above processes will also be discussed, with emphasis on downstream stress signaling components, kinases and transcription factors (TFs). The importance of several key ABA and GA signaling intermediates will be illustrated. The knowledge gained from such studies will also help to establish a solid foundation to develop future crop improvement strategies.