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Plants and arbuscular mycorrhizal fungi (AMF) can form complex symbiotic networks based on functional trait selection, contributing to the maintenance of ecosystem biodiversity and stability. However, the selectivity of host plants on AMF and the characteristics of plant-AMF networks remain unclear in Tibetan alpine meadows. In this study, we studied the AMF communities in 69 root samples from 23 plant species in a Tibetan alpine meadow using Illumina-MiSeq sequencing of the 18S rRNA gene. The results showed a significant positive correlation between the phylogenetic distances of plant species and the taxonomic dissimilarity of their AMF community. The plant-AMF network was characterized by high connectance, high nestedness, anti-modularity, and anti-specialization, and the phylogenetic signal from plants was stronger than that from AMF. The high connected and nested plant-AMF network potentially promoted the interdependence and stability of the plant-AMF symbioses in Tibetan alpine meadows. This study emphasizes that plant phylogeny and plant-AMF networks play an important role in the coevolution of host plants and their mycorrhizal partners and enhance our understanding of the interactions between aboveground and belowground communities. [ABSTRACT FROM AUTHOR]

Maximizing the function of indigenous arbuscular mycorrhizal (AM) fungi by choosing specific crop genotypes offers one of the few untapped opportunities to improve the sustainability of agriculture. In this study, the differences in mycorrhizal responsiveness (MR) in plant growth and shoot phosphorus (P) content among cotton (Gossypium spp. L.) genotypes from different release dates were compared and then the relationships between MR and P uptake-related traits were determined. The experimental design in a greenhouse included 24 genotypes released from 1950 to present in Xinjiang Province, inoculation with or without AM fungi, and P levels (15 and 150 mg P kg–1 added as KH2PO4). Results showed that the modern cotton genotypes exhibited a higher degree of mycorrhizal colonization, the hyphal length density (HLD), and mycorrhizae-induced changes in shoot growth than the old genotypes when inoculated with indigenous AM fungi at both the P levels. Moreover, MR was highly correlated with the HLD at low P levels and the HLD may provide useful insights for future cotton breeding aimed at delivering crop genotypes that can benefit more from AM fungi. [ABSTRACT FROM AUTHOR]

Intercropping is both a well-established and yet novel agricultural practice, depending on one's perspective. Such perspectives are principally governed by geographic location and whether monocultural practices predominate. Given the negative environmental effects of monoculture agriculture (loss of biodiversity, reliance on non-renewable inputs, soil degradation, etc.), there has been a renewed interest in cropping systems that can reduce the impact of modern agriculture while maintaining (or even increasing) yields. Intercropping is one of the most promising practices in this regard, yet faces a multitude of challenges if it is to compete with and ultimately replace the prevailing monocultural norm. These challenges include the necessity for more complex agricultural designs in space and time, bespoke machinery, and adapted crop cultivars. Plant breeding for monocultures has focused on maximizing yield in single-species stands, leading to highly productive yet specialized genotypes. However, indications suggest that these genotypes are not the best adapted to intercropping systems. Re-designing breeding programs to accommodate inter-specific interactions and compatibilities, with potentially multiple different intercropping partners, is certainly challenging, but recent technological advances offer novel solutions. We identify a number of such technology-driven directions, either ideotype-driven (i.e., "trait-based" breeding) or quantitative genetics-driven (i.e., "product-based" breeding). For ideotype breeding, plant growth modeling can help predict plant traits that affect both inter- and intraspecific interactions and their influence on crop performance. Quantitative breeding approaches, on the other hand, estimate breeding values of component crops without necessarily understanding the underlying mechanisms. We argue that a combined approach, for example, integrating plant growth modeling with genomic-assisted selection and indirect genetic effects, may offer the best chance to bridge the gap between current monoculture breeding programs and the more integrated and diverse breeding programs of the future. [ABSTRACT FROM AUTHOR]

Legumes of the Fabeae tribe form nitrogen-fixing root nodules resulting from symbiotic interaction with the soil bacteria Rhizobium leguminosarum symbiovar viciae (Rlv). These bacteria are all potential symbionts of the Fabeae hosts but display variable partner choice when co-inoculated in mixture. Because partner choice and symbiotic nitrogen fixation mostly behave as genetically independent traits, the efficiency of symbiosis is often suboptimal when Fabeae legumes are exposed to natural Rlv populations present in soil. A core collection of 32 Rlv bacteria was constituted based on the genomic comparison of a collection of 121 genome sequences, representative of known worldwide diversity of Rlv. A variable part of the nodD gene sequence was used as a DNA barcode to discriminate and quantify each of the 32 bacteria in mixture. This core collection was co-inoculated on a panel of nine genetically diverse Pisum sativum , Vicia faba , and Lens culinaris genotypes. We estimated the relative Early Partner Choice (EPC) of the bacteria with the Fabeae hosts by DNA metabarcoding on the nodulated root systems. Comparative genomic analyses within the bacterial core collection identified molecular markers associated with host-dependent symbiotic partner choice. The results revealed emergent properties of rhizobial populations. They pave the way to identify genes related to important symbiotic traits operating at this level. [ABSTRACT FROM AUTHOR]

Biological nitrogen fixation by Rhizobium -legume symbioses represents an environmentally friendly and inexpensive alternative to the use of chemical nitrogen fertilizers in legume crops. Rhizobial inoculants, applied frequently as biofertilizers, play an important role in sustainable agriculture. However, inoculants often fail to compete for nodule occupancy against native rhizobia with inferior nitrogen-fixing abilities, resulting in low yields. Strains with excellent performance under controlled conditions are typically selected as inoculants, but the rates of nodule occupancy compared to native strains are rarely investigated. Lack of persistence in the field after agricultural cycles, usually due to the transfer of symbiotic genes from the inoculant strain to naturalized populations, also limits the suitability of commercial inoculants. When rhizobial inoculants are based on native strains with a high nitrogen fixation ability, they often have superior performance in the field due to their genetic adaptations to the local environment. Therefore, knowledge from laboratory studies assessing competition and understanding how diverse strains of rhizobia behave, together with assays done under field conditions, may allow us to exploit the effectiveness of native populations selected as elite strains and to breed specific host cultivar-rhizobial strain combinations. Here, we review current knowledge at the molecular level on competition for nodulation and the advances in molecular tools for assessing competitiveness. We then describe ongoing approaches for inoculant development based on native strains and emphasize future perspectives and applications using a multidisciplinary approach to ensure optimal performance of both symbiotic partners. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) are obligate plant symbionts that have the potential to improve crop yield. These multinucleate organisms are either "homokaryotic" or "dikaryotic". In AMF dikaryons, thousands of nuclei originating from two parental strains coexist in the same cytoplasm. In other fungi, homokaryotic and dikaryotic strains show distinct life history traits (LHTs), such as variation in growth rates and fitness. However, how such traits compare between dikaryons and homokaryons of AMF is unknown. To address this, we measured 20 LHT of four dikaryons and five homokaryons of the model fungus Rhizophagus irregularis across root organ cultures of three host plants (carrot, chicory, and tobacco). Our analyses show that dikaryons have clearly distinct life history strategies (LHSs) compared to homokaryons. In particular, spores of homokaryons germinate faster and to a higher proportion than dikaryons, whereas dikaryons grow significantly faster and create a more complex hyphal network irrespective of host plant species. Our study links AMF nuclear status with key LHT with possible implications for mycorrhizal symbiotic functioning. [ABSTRACT FROM AUTHOR]

A vast majority of terrestrial plants are dependent on arbuscular mycorrhizal fungi (AMF) for their nutrient acquisition. AMF act as an extension of the root system helping phosphate uptake. In agriculture, harnessing the symbiosis can potentially increase plant growth. Application of the AMF Rhizophagus irregularis has been demonstrated to increase the yields of various crops. However, there is a paradigm that AMF colonization of roots, as well as the plant benefits afforded by inoculation with AMF, decreases with increasing phosphorus (P) supply in the soil. The paradigm suggests that when fertilized with sufficient P, inoculation of crops would not be beneficial. However, the majority of experiments demonstrating the paradigm were conducted in sterile conditions without a background AMF or soil microbial community. Interestingly, intraspecific variation in R. irregularis can greatly alter the yield of cassava even at a full application of the recommended P dose. Cassava is a globally important crop, feeding 800 million people worldwide, and a crop that is highly dependent on AMF for P uptake. In this study, field trials were conducted at three locations in Kenya and Tanzania using different AMF and cassava varieties under different P fertilization levels to test if the paradigm occurs in tropical field conditions. We found that AMF colonization and inoculation responsiveness of cassava does not always decrease with an increased P supply as expected by the paradigm. The obtained results demonstrate that maximizing the inoculation responsiveness of cassava is not necessarily only in conditions of low P availability, but that this is dependent on cassava and fungal genotypes. Thus, the modeling of plant symbiosis with AMF under different P levels in nature should be considered with caution. [ABSTRACT FROM AUTHOR]

Synthetic microbial communities (SynComs) are a useful tool for a more realistic understanding of the outcomes of multiple biotic interactions where microbes, plants, and the environment are players in time and space of a multidimensional and complex system. Toward a more in-depth overview of the knowledge that has been achieved using SynComs in the rhizosphere, a systematic review of the literature on SynComs was performed to identify the overall rationale, design criteria, experimental procedures, and outcomes of in vitro or in planta tests using this strategy. After an extensive bibliography search and a specific selection process, a total of 30 articles were chosen for further analysis, grouping them by their reported SynCom size. The reported SynComs were constituted with a highly variable number of members, ranging from 3 to 190 strains, with a total of 1,393 bacterial isolates, where the three most represented phyla were Proteobacteria, Actinobacteria, and Firmicutes. Only four articles did not reference experiments with SynCom on plants, as they considered only microbial in vitro studies, whereas the others chose different plant models and plant-growth systems; some of them are described and reviewed in this article. Besides, a discussion on different approaches (bottom-up and top-down) to study the microbiome role in the rhizosphere is provided, highlighting how SynComs are an effective system to connect and fill some knowledge gaps and to have a better understanding of the mechanisms governing these multiple interactions. Although the SynCom approach is already helpful and has a promising future, more systematic and standardized studies are needed to harness its full potential. [ABSTRACT FROM AUTHOR]

Orchid distribution and population dynamics are influenced by a variety of ecological factors and the formation of holobionts, which play key roles in colonization and ecological community construction. Seed germination, seedling establishment, reproduction, and survival of orchid species are strongly dependent on orchid mycorrhizal fungi (OMF), with mycorrhizal cheating increasingly observed in photosynthetic orchids. Therefore, changes in the composition and abundance of OMF can have profound effects on orchid distribution and fitness. Network analysis is an important tool for the study of interactions between plants, microbes, and the environment, because of the insights that it can provide into the interactions and coexistence patterns among species. Here, we provide a comprehensive overview, systematically describing the current research status of the effects of OMF on orchid distribution and dynamics, phylogenetic signals in orchid–OMF interactions, and OMF networks. We argue that orchid–OMF associations exhibit complementary and specific effects that are highly adapted to their environment. Such specificity of associations may affect the niche breadth of orchid species and act as a stabilizing force in plant–microbe coevolution. We postulate that network analysis is required to elucidate the functions of fungal partners beyond their effects on germination and growth. Such studies may lend insight into the microbial ecology of orchids and provide a scientific basis for the protection of orchids under natural conditions in an efficient and cost-effective manner. [ABSTRACT FROM AUTHOR]

Plant secondary metabolites (PSMs) play many roles including defense against pathogens, pests, and herbivores; response to environmental stresses, and mediating organismal interactions. Similarly, plant microbiomes participate in many of the above-mentioned processes directly or indirectly by regulating plant metabolism. Studies have shown that plants can influence their microbiome by secreting various metabolites and, in turn, the microbiome may also impact the metabolome of the host plant. However, not much is known about the communications between the interacting partners to impact their phenotypic changes. In this article, we review the patterns and potential underlying mechanisms of interactions between PSMs and plant microbiomes. We describe the recent developments in analytical approaches and methods in this field. The applications of these new methods and approaches have increased our understanding of the relationships between PSMs and plant microbiomes. Though the current studies have primarily focused on model organisms, the methods and results obtained so far should help future studies of agriculturally important plants and facilitate the development of methods to manipulate PSMs–microbiome interactions with predictive outcomes for sustainable crop productions. [ABSTRACT FROM AUTHOR]

Studies in natural ecosystems show that adaptation of arbuscular mycorrhizal (AM) fungi and other microbial plant symbionts to local environmental conditions can help ameliorate stress and optimize plant fitness. This local adaptation arises from the process of multilevel selection, which is the simultaneous selection of a hierarchy of groups. Studies of multilevel selection in natural ecosystems may inform the creation of sustainable agroecosystems through developing strategies to effectively manage crop microbiomes including AM symbioses. Field experiments show that the species composition of AM fungal communities varies across environmental gradients, and that the biomass of AM fungi and their benefits for plants generally diminish when fertilization and irrigation eliminate nutrient and water limitations. Furthermore, pathogen protection by mycorrhizas is only important in environments prone to plant damage due to pathogens. Consequently, certain agricultural practices may inadvertently select for less beneficial root symbioses because the conventional agricultural practices of fertilization, irrigation, and use of pesticides can make these symbioses superfluous for optimizing crop performance. The purpose of this paper is to examine how multilevel selection influences the flow of matter, energy, and genetic information through mycorrhizal microbiomes in natural and agricultural ecosystems, and propose testable hypotheses about how mycorrhizae may be actively managed to increase agricultural sustainability. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is an essential plant nutrient, and plants can take up N from several sources, including via mycorrhizal fungal associations. The N uptake patterns of understory plants may vary beneath different types of overstory trees, especially through the difference in their type of mycorrhizal association (arbuscular mycorrhizal, AM; or ectomycorrhizal, ECM), because soil mycorrhizal community and N availability differ beneath AM (non-ECM) and ECM overstory trees (e.g., relatively low nitrate content beneath ECM overstory trees). To test this hypothesis, we examined six co-existing AM-symbiotic understory tree species common beneath both AM-symbiotic black locust (non-ECM) and ECM-symbiotic oak trees of dryland forests in China. We measured AM fungal community composition of roots and natural abundance stable isotopic composition of N (δ15N) in plant leaves, roots, and soils. The root mycorrhizal community composition of understory trees did not significantly differ between beneath non-ECM and ECM overstory trees, although some OTUs more frequently appeared beneath non-ECM trees. Understory trees beneath non-ECM overstory trees had similar δ15N values in leaves and soil nitrate, suggesting that they took up most of their nitrogen as nitrate. Beneath ECM overstory trees, understory trees had consistently lower leaf than root δ15N, suggesting they depended on mycorrhizal fungi for N acquisition since mycorrhizal fungi transfer isotopically light N to host plants. Additionally, leaf N concentrations in the understory trees were lower beneath ECM than the non-ECM overstory trees. Our results show that, without large differences in root mycorrhizal community, the N uptake patterns of understory trees vary between beneath different overstory trees. [ABSTRACT FROM AUTHOR]

Symbiotic nitrogen fixation by legume nodules provides an abundant nitrogen source for plants, and understanding this process is key for developing green agriculture. Circular RNA (circRNA), a type of endogenous RNA produced by reverse splicing of mRNA precursors, plays important regulatory roles in plants at the transcriptional and post-transcriptional levels. However, the relationship between circRNAs and legume–rhizobium is unknown. Here, we performed comprehensive identification and expression profiling of circRNAs during nodulation in common bean (Phaseolus vulgaris) compared to uninoculated roots of corresponding ages by constructing circRNA-seq and mRNA-seq libraries. We identified 8,842 high-confident circRNAs, 3,448 of which were specifically produced during symbiosis, with the highest number at the nitrogen-fixing stage. Significantly, more circRNAs were derived from exons than from intergenic regions or introns in all samples. The lengths and GC contents of the circRNAs were similar in roots and nodules. However, circRNAs showed specific spatiotemporal expression patterns during nodule and root development. GO and other functional annotation of parental genes of differentially expressed circRNAs indicated their potential involvement in different biological processes. The expression of major circRNAs during symbiosis is independent of parental genes' expression to a certain degree, while expression of the remaining minor circRNAs showed positive correlation to parental genes. Functional annotation of the targeted mRNAs in the circRNA–miRNA–mRNA network showed that circRNAs may be involved in transmembrane transport and positive regulation of kinase activity during nodulation and nitrogen fixation as miRNA sponges. Our comprehensive analysis of the expression profile of circRNAs and their potential functions suggests that circRNAs may function as new post-transcriptional regulators in legume–rhizobium symbiosis. [ABSTRACT FROM AUTHOR]

Plants teem with microorganisms, whose tremendous diversity and role in plant–microbe interactions are being increasingly explored. Microbial communities create a functional bond with their hosts and express beneficial traits capable of enhancing plant performance. Therefore, a significant task of microbiome research has been identifying novel beneficial microbial traits that can contribute to crop productivity, particularly under adverse environmental conditions. However, although knowledge has exponentially accumulated in recent years, few novel methods regarding the process of designing inoculants for agriculture have been presented. A recently introduced approach is the use of synthetic microbial communities (SynComs), which involves applying concepts from both microbial ecology and genetics to design inoculants. Here, we discuss how to translate this rationale for delivering stable and effective inoculants for agriculture by tailoring SynComs with microorganisms possessing traits for robust colonization, prevalence throughout plant development and specific beneficial functions for plants. Computational methods, including machine learning and artificial intelligence, will leverage the approaches of screening and identifying beneficial microbes while improving the process of determining the best combination of microbes for a desired plant phenotype. We focus on recent advances that deepen our knowledge of plant–microbe interactions and critically discuss the prospect of using microbes to create SynComs capable of enhancing crop resiliency against stressful conditions. [ABSTRACT FROM AUTHOR]

Wheat is among the important crops harnessed by humans whose breeding efforts resulted in a diversity of genotypes with contrasting traits. The goal of this study was to determine whether different old and new cultivars of durum wheat (Triticum turgidum L. var. durum) recruit specific arbuscular mycorrhizal (AM) fungal communities from indigenous AM fungal populations of soil under field conditions. A historical set of five landraces and 26 durum wheat cultivars were field cultivated in a humid climate in Eastern Canada, under phosphorus-limiting conditions. To characterize the community of AMF inhabiting bulk soil, rhizosphere, and roots, MiSeq amplicon sequencing targeting the 18S rRNA gene (SSU) was performed on total DNAs using a nested PCR approach. Mycorrhizal colonization was estimated using root staining and microscope observations. A total of 317 amplicon sequence variants (ASVs) were identified as belonging to Glomeromycota. The core AM fungal community (i.e., ASVs present in > 50% of the samples) in the soil, rhizosphere, and root included 29, 30, and 29 ASVs, respectively. ASVs from the genera Funneliformis , Claroideoglomus , and Rhizophagus represented 37%, 18.6%, and 14.7% of the sequences recovered in the rarefied dataset, respectively. The two most abundant ASVs had sequence homology with the 18S sequences from well-identified herbarium cultures of Funneliformis mosseae BEG12 and Rhizophagus irregularis DAOM 197198, while the third most abundant ASV was assigned to the genus Paraglomus. Cultivars showed no significant difference of the percentage of root colonization ranging from 57.8% in Arnautka to 84.0% in AC Navigator. Cultivars were generally associated with similar soil, rhizosphere, and root communities, but the abundance of F. mosseae , R. irregularis , and Claroideoglomus sp. sequences varied in Eurostar, Golden Ball, and Wakooma. Although these results were obtained in one field trial using a non-restricted pool of durum wheat and at the time of sampling, that may have filtered the community in biotopes. The low genetic variation between durum wheat cultivars for the diversity of AM symbiosis at the species level suggests breeding resources need not be committed to leveraging plant selective influence through the use of traditional methods for genotype development. [ABSTRACT FROM AUTHOR]

The nitrogen fixing ability of common bean (Phaseolus vulgaris L.) in association with rhizobia is often characterized as poor compared to other legumes, and nitrogen fertilizers are commonly used in bean production to achieve high yields, which in general inhibits nitrogen fixation. In addition, plants cannot take up all the nitrogen applied to the soil as a fertilizer leading to runoff and groundwater contamination. The overall objective of this work is to reduce use of nitrogen fertilizer in common bean production. This would be a major advance in profitability for the common bean industry in Canada and would significantly improve the ecological footprint of the crop. In the current work, 22 bean genotypes [including recombinant inbred lines (RILs) from the Mist × Sanilac population and a non-nodulating mutant (R99)] were screened for their capacity to fix atmospheric nitrogen under four nitrogen regimes. The genotypes were evaluated in replicated field trials on N-poor soils over three years for the percent nitrogen derived from atmosphere (%Ndfa), yield, and a number of yield-related traits. Bean genotypes differed for all analyzed traits, and the level of nitrogen significantly affected most of the traits, including %Ndfa and yield in all three years. In contrast, application of rhizobia significantly affected only few traits, and the effect was inconsistent among the years. Nitrogen application reduced symbiotic nitrogen fixation (SNF) to various degrees in different bean genotypes. This variation suggests that SNF in common bean can be improved through breeding and selection for the ability of bean genotypes to fix nitrogen in the presence of reduced fertilizer levels. Moreover, genotypes like RIL_38, RIL_119, and RIL_131, being both high yielding and good nitrogen fixers, have potential for simultaneous improvement of both traits. However, breeding advancement might be slow due to an inconsistent correlation between these traits. [ABSTRACT FROM AUTHOR]

Keywords: plant beneficial microorganisms; prebiotics; microbial metabolites; formulation; microbiome management EN plant beneficial microorganisms prebiotics microbial metabolites formulation microbiome management 1 5 5 07/22/20 20200716 NES 200716 Introduction It is well known that a gram of soil contains thousands of individual microbial taxa including bacteria, fungi, protists, oomycetes and viruses. Moreover, some studies demonstrated reduced plant growth and increased microbial phytopathogenicity as a result of soil-plant systems inoculation with potentially beneficial microorganisms in conditions of nutrient saturation, changes in the microbial community, or environmental and plant genotype effects ([27]; [36]; [31]; [10]). Prebiotics and Synbiotics Prebiotics are products, which improve microbial diversity and soil microbial health by promoting the growth of soil microorganisms already present within the soil-plant system. Another, pivotal point of the new approach to understand and manage the functional and genetic role of soil microorganisms in the soil-plant systems, is the comparison between human gut microbiome and plant microbiome ([1]). [Extracted from the article]

Plant performance is strongly dependent on nitrogen (N), and thus increasing N nutrition is of great relevance for the productivity of agroecosystems. The effects of arbuscular mycorrhizal (AM) fungi on plant N acquisition are debated because contradictory results have been reported. Using 15N-labeled fertilizers as a tracer, we evaluated the effects of AM fungi on N uptake and recovery from mineral or organic sources in durum wheat. Under sufficient N availability, AM fungi had no effects on plant biomass but increased N concentrations in plant tissue, plant N uptake, and total N recovered from the fertilizer. In N-deficient soil, AM fungi led to decreased aboveground biomass, which suggests that plants and AM fungi may have competed for N. When the organic source had a low C:N ratio, AM fungi favored both plant N uptake and N recovery. In contrast, when the organic source had a high C:N ratio, a clear reduction in N recovery from the fertilizer was observed. Overall, the results indicate an active role of arbuscular mycorrhizae in favoring plant N-related traits when N is not a limiting factor and show that these fungi help in N recovery from the fertilizer. These results hold great potential for increasing the sustainability of durum wheat production. [ABSTRACT FROM AUTHOR]

A plant growing under natural conditions is always associated with a substantial, diverse, and well-orchestrated community of microbes—the phytomicrobiome. The phytomicrobiome genome is larger and more fluid than that of the plant. The microbes of the phytomicrobiome assist the plant in nutrient uptake, pathogen control, stress management, and overall growth and development. At least some of this is facilitated by the production of signal compounds, both plant-to-microbe and microbe back to the plant. This is best characterized in the legume nitrogen fixing and mycorrhizal symbioses. More recently lipo-chitooligosaccharide (LCO) and thuricin 17, two microbe-to-plant signals, have been shown to regulate stress responses in a wide range of plant species. While thuricin 17 production is constitutive, LCO signals are only produced in response to a signal from the plant. We discuss how some signal compounds will only be discovered when root-associated microbes are exposed to appropriate plant-to-microbe signals (positive regulation), and this might only happen under specific conditions, such as abiotic stress, while others may only be produced in the absence of a particular plant-to-microbe signal molecule (negative regulation). Some phytomicrobiome members only elicit effects in a specific crop species (specialists), while other phytomicrobiome members elicit effects in a wide range of crop species (generalists). We propose that some specialists could exhibit generalist activity when exposed to signals from the correct plant species. The use of microbe-to-plant signals can enhance crop stress tolerance and could result in more climate change resilient agricultural systems. [ABSTRACT FROM AUTHOR]

The specialised mutualism between Tococa guianensis and ants housed in its leaf domatia is a well-known example of myrmecophily. A pollination study on this species revealed that flowers in the bud stage exude a sugary solution that is collected by ants. Given the presence of this unexpected nectar secretion, we investigated how, where, and when floral buds of T. guianensis secret nectar and what function it serves. We studied a population of T. guianensis occurring in a swampy area in the Cerrado of Brazil by analyzing the chemical composition and secretion dynamics of the floral-bud nectar and the distribution and ultrastructure of secretory tissues. We also measured flower damage using ant-exclusion experiments. Floral bud nectar was secreted at the tip of the petals, which lack a typical glandular structure but possess distinctive mesophyll due to the presence of numerous calcium oxalate crystals. The nectar, the production of which ceased after flower opening, was composed mainly of sucrose and low amounts of glucose and fructose. Nectar was consumed by generalist ants and sporadically by stingless bees. Ant exclusion experiments resulted in significantly increased flower damage. The floral nectar of T. guianensis is produced during the bud stage. This bud-nectar has the extranuptial function of attracting generalist ants that reduce florivory. Pollen is the unique floral resource attracting pollinators during anthesis. Tococa guianensis, thus, establishes relationships with two functional groups of ant species: specialist ants acting against herbivory and generalist ants acting against florivory. [ABSTRACT FROM AUTHOR]

Phosphate (P) is characterized by its low availability and restricted mobility in soils, and also by a high redistribution capacity inside plants. In order to maintain P homeostasis in nutrient restricted conditions, plants have developed mechanisms which enable P acquisition from the soil solution, and an efficient reutilization of P already present in plant cells. Nitric oxide (NO) is a bioactive molecule with a plethora of functions in plants. Its endogenous synthesis depends on internal and environmental factors, and is closely tied with nitrogen (N) metabolism. Furthermore, there is evidence demonstrating that N supply affects P homeostasis and that P deficiency impacts on N assimilation. This review will provide an overview on how NO levels in planta are affected by P deficiency, the interrelationship with N metabolism, and a summary of the current understanding about the influence of this reactive N species over the processes triggered by P starvation, which could modify P use efficiency. [ABSTRACT FROM AUTHOR]

The Meloidogyne-based disease complexes (MDCs) are caused by the interaction of different root-knot nematode species and phytopathogenic fungi. These complexes are devastating several important crops worldwide including tomato and coffee. Despite their relevance, little is known about the role of the bacterial communities in the MDCs. In this study 16s rDNA gene sequencing was used to analyze the bacterial microbiome associated with healthy and infested roots, as well with females and eggs of Meloidogyne enterolobii and M. paranaensis, the causal agents of MDC in tomato and coffee, respectively. Each MDC pathosystems displayed a specific taxonomic diversity and relative abundances constituting a very complex system. The main bacterial drivers of the MDC infection process were identified for both crops at order level. While corky-root coffee samples presented an enrichment of Bacillales and Burkholderiales, the corckyroot tomato samples presented an enrichment on Saprospirales, Chthoniobacterales, Alteromonadales, and Xanthomonadales. At genus level, Nocardia was common to both systems, and it could be related to the development of tumor symptoms by altering both nematode and plant systems. Furthermore, we predicted the healthy metabolic profile of the roots microbiome and a shift that may result in an increment of activity of central metabolism and the presence of pathogenic genes in both crops. [ABSTRACT FROM AUTHOR]

Symbiosis with ectomycorrhizal (ECM) fungi is an advantageous partnership for trees in nutrient-limited environments. Ectomycorrhizal fungi colonize the roots of their hosts and improve their access to nutrients, usually nitrogen (N) and, in exchange, trees deliver a significant portion of their photosynthetic carbon (C) to the fungi. This nutrient exchange affects key soil processes and nutrient cycling, as well as plant health, and is therefore central to forest ecosystem functioning. Due to their ecological importance, there is a need to more accurately understand ECM fungal mediated C and N movement within forest ecosystems such that we can better model and predict their role in soil processes both now and under future climate scenarios. There are a number of hurdles that we must overcome, however, before this is achievable such as understanding how the evolutionary history of ECM fungi and their inter- and intra- species variability affect their function. Further, there is currently no generally accepted universal mechanism that appears to govern the flux of nutrients between fungal and plant partners. Here, we consider the current state of knowledge on N acquisition and transport by ECM fungi and how C and N exchange may be related or affected by environmental conditions such as N availability. We emphasize the role that modern genomic analysis, molecular biology techniques and more comprehensive and standardized experimental designs may have in bringing cohesion to the numerous ecological studies in this area and assist us in better understanding this important symbiosis. These approaches will help to build unified models of nutrient exchange and develop diagnostic tools to study these fungi at various scales and environments. [ABSTRACT FROM AUTHOR]

In arbuscular mycorrhizal (AM) symbiosis, key components of nutrient uptake and exchange are specialized transporters that facilitate nutrient transport across membranes. As phosphate is a nutrient and a regulator of nutrient exchanges, we investigated the effect of P availability to extraradical mycelium (ERM) on both plant and fungus transcriptomes and metabolomes in a symbiocosm system. By perturbing nutrient exchanges under the control of P, our objectives were to identify new fungal genes involved in nutrient transports, and to characterize in which extent the fungus differentially modulates its metabolism when interacting with two different plant species. We performed transportome analysis on the ERM and intraradical mycelium of the AM fungus Rhizophagus irregularis associated to Populus trichocarpa and Sorghum bicolor under high and low P availability in ERM, using quantitative RT-PCR and Illumina mRNA-sequencing. We observed that mycorrhizal symbiosis induces expression of specific phosphate and ammonium transporters in both plants. Furthermore, we identified new AM-inducible transporters and showed that a subset of phosphate transporters is regulated independently of symbiotic nutrient exchange. mRNA-Sequencing revealed that the fungal transportome was not similarly regulated in the two host plant species according to P availability. Mirroring this effect, many plant carbohydrate transporters were down-regulated in P. trichocarpa mycorrhizal root tissue. Metabolome analysis revealed further that AM root colonization led to a modification of root primary metabolism under low and high P availability and to a decrease of primary metabolite pools in general. Moreover, the down regulation of the sucrose transporters suggests that the plant limits carbohydrate long distance transport (i.e. from shoot to the mycorrhizal roots). By simultaneous uptake/reuptake of nutrients from the apoplast at the biotrophic interface, plant and fungus are both able to control reciprocal nutrient fluxes. [ABSTRACT FROM AUTHOR]

The interaction between legumes and bacteria of rhizobia type results in a beneficial symbiotic relationship characterized by the formation of new root organs, called nodules. Within these nodules the bacteria, released in plant cells, differentiate into bacteroids and fix atmospheric nitrogen through the nitrogenase activity. This mutualistic interaction has evolved sophisticated signaling networks to allow rhizobia entry, colonization, bacteroid differentiation and persistence in nodules. Nodule cysteine rich (NCR) peptides, reactive oxygen species (ROS), reactive nitrogen species (RNS), and toxin–antitoxin (TA) modules produced by the host plants or bacterial microsymbionts have a major role in the control of the symbiotic interaction. These molecules described as weapons in pathogenic interactions have evolved to participate to the intracellular bacteroid accommodation by escaping control of plant innate immunity and adapt the functioning of the nitrogen-fixation to environmental signalling cues. [ABSTRACT FROM AUTHOR]

Controlled environment studies show that arbuscular mycorrhizal fungi (AMF) may contribute to plant nitrogen (N) uptake, but the role of these near-ubiquitous symbionts in crop plant N nutrition under natural field conditions remains largely unknown. In a field trial, we tested the effects of N fertilisation and barley (Hordeum vulgare L.) cultivar identity on the contribution of AMF to barley N uptake using 15N tracers added to rhizosphere soil compartments. AMF were shown capable of significantly increasing plant 15N acquisition from root exclusion zones, and this was influenced by nitrogen addition type, N fertiliser application rate and barley cultivar identity. Our data demonstrate a previously overlooked potential route of crop plant N uptake which may be influenced substantially and rapidly in response to shifting agricultural management practices. [ABSTRACT FROM AUTHOR]

Plants may actively cultivate microorganisms in their roots and rhizosphere that enhance their nutrition. To develop cropping strategies that substitute mineral fertilizers for beneficial root symbioses, we must first understand how microbial communities associated with plant roots differ among plant taxa and how they respond to fertilization. Arbuscular mycorrhizal (AM) fungi and rhizobacteria are of particular interest because they enhance nutrient availability to plants and perform a suite of nutrient cycling functions. The purpose of this experiment is to examine the root and soil microbiome in a long-term switchgrass (Panicum virgatum) biofuel feedstock experiment and determine how AM fungi and rhizobacteria respond to plant diversity and soil fertility. We hypothesize that intra- and interspecific plant diversity, nitrogen fertilization (+N), and their interaction will influence the biomass and community composition of AM fungi and rhizobacteria. We further hypothesize that +N will reduce the abundance of nitrogenase-encoding nifH genes on the rhizoplane. Roots and soils were sampled from three switchgrass cultivars (Cave-in-Rock, Kanlow, Southlow) grown in monoculture, intraspecific mixture, and interspecific planting mixtures with either Andropogon gerardii or diverse native tallgrass prairie species. Molecular sequencing was performed on root and soil samples, fatty acid extractions were assessed to determine microbial biomass, and quantitative polymerase chain reaction (qPCR) was performed on nifH genes from the rhizoplane. Sequence data determined core AM fungal and bacterial microbiomes and indicator taxa for plant diversity and +N treatments. We found that plant diversity and +N influenced AM fungal biomass and community structure. Across all plant diversity treatments, +N reduced the biomass of AM fungi and nifH gene abundance by more than 40%. The AM fungal genus Scutellospora was an indicator for +N, with relative abundance significantly greater under +N and in monoculture treatments. Community composition of rhizobacteria was influenced by plant diversity but not by +N. Verrucomicrobia and Proteobacteria were the dominant bacterial phyla in both roots and soils. Our findings provide evidence that soil fertility and plant diversity structure the root and soil microbiome. Optimization of soil communities for switchgrass production must take into account differences among cultivars and their unique responses to shifts in soil fertility. [ABSTRACT FROM AUTHOR]

The nitrogen-fixing root nodule symbiosis is restricted to four plant orders: Fabales (legumes), Fagales, Cucurbitales and Rosales (Elaeagnaceae, Rhamnaceae, and Rosaceae). Interestingly all of the Rosaceae genera confirmed to contain nodulating species (i.e., Cercocarpus , Chamaebatia , Dryas , and Purshia) belong to a single subfamily, the Dryadoideae. The Dryas genus is particularly interesting from an evolutionary perspective because it contains closely related nodulating (Dryas drummondii) and non-nodulating species (Dryas octopetala). The close phylogenetic relationship between these two species makes Dryas an ideal model genus to study the genetic basis of nodulation by whole genome comparison and classical genetics. Therefore, we established methods for plant cultivation, transformation and DNA extraction for these species. We optimized seed surface sterilization and germination methods and tested growth protocols ranging from pots and Petri dishes to a hydroponic system. Transgenic hairy roots were obtained by adapting Agrobacterium rhizogenes -based transformation protocols for Dryas species. We compared several DNA extraction protocols for their suitability for subsequent molecular biological analysis. Using CTAB extraction, reproducible PCRs could be performed, but CsCl gradient purification was essential to obtain DNA in sufficient purity for high quality de novo genome sequencing of both Dryas species. Altogether, we established a basic toolkit for the culture, transient transformation and genetic analysis of Dryas sp. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi increase fitness of their host plants under Cu deficient and toxic conditions. In this study, we have characterized two Cu transporters of the CTR family (RiCTR1 and RiCTR2) and a CTR-like protein (RiCTR3A) of Rhizophagus irregularis. Functional analyses in yeast revealed that RiCTR1 encodes a plasma membrane Cu transporter, RiCTR2 a vacuolar Cu transporter and RiCTR3A a plasma membrane protein involved in Cu tolerance. RiCTR1 was more highly expressed in the extraradical mycelia (ERM) and RiCTR2 in the intraradical mycelia (IRM). In the ERM, RiCTR1 expression was up-regulated by Cu deficiency and down-regulated by Cu toxicity. RiCTR2 expression increased only in the ERM grown under severe Cu-deficient conditions. These data suggest that RiCTR1 is involved in Cu uptake by the ERM and RiCTR2 in mobilization of vacuolar Cu stores. Cu deficiency decreased mycorrhizal colonization and arbuscule frequency, but increased RiCTR1 and RiCTR2 expression in the IRM, which suggest that the IRM has a high Cu demand. The two alternatively spliced products of RiCTR3, RiCTR3A and RiCTR3B , were more highly expressed in the ERM. Up-regulation of RiCTR3A by Cu toxicity and the yeast complementation assays suggest that RiCTR3A might function as a Cu receptor involved in Cu tolerance. [ABSTRACT FROM AUTHOR]

As one of the most important limiting factors of grassland productivity, drought is predicted to increase in intensity and frequency. Greenhouse studies suggest that arbuscular mycorrhizal fungi (AMF) can improve plant drought resistance. However, whether AMF can improve plant drought resistance in field conditions and whether the effects of AMF on drought resistance differ among plants with different photosynthetic pathways remain unclear. To evaluate the effect of indigenous AMF on plant drought resistance, an in situ rainfall exclusion experiment was conducted in a temperate meadow in northeast China. The results showed that AMF significantly reduced the negative effects of drought on plant growth. On average, AMF enhanced plant biomass, photosynthetic rate (A), stomatal conductance (g s), intrinsic water use efficiency (iWUE), and superoxide dismutase (SOD) activity of the C3 species Leymus chinensis by 58, 63, 38, 15, and 45%, respectively, and reduced levels of malondialdehyde (MDA) by 32% under light and moderate drought (rainfall exclusion of 30 and 50%, respectively). However, under extreme drought (rainfall exclusion of 70%), AMF elevated only aboveground biomass and catalase (CAT) activities. Averagely, AMF increased the aboveground biomass, A , and CAT activity of Hemarthria altissima (C4) by 37, 28, and 30%, respectively, under light and moderate droughts. The contribution of AMF to plant drought resistance was higher for the C3 species than that for the C4 species under both light and moderate drought conditions. The results highlight potential photosynthetic type differences in the magnitude of AMF-associated enhancement in plant drought resistance. Therefore, AMF may determine plant community structure under future climate change scenarios by affecting the drought resistance of different plant functional groups. [ABSTRACT FROM AUTHOR]

Under fertilization levels specific to intensive farming, the impact of compensation of soil nutritional value by arbuscular mycorrhiza (AM) might be limited. Therefore, the question arises whether modern crop varieties, selected for high NPK assimilation rate, are able to gain symbiotic benefits under other challenging field conditions, such as drought. Accordingly, in this study we aimed to evaluate the contribution of Rhizophagus irregularis to the drought response of a stay-green corn hybrid in pot cultures equally fertilized until silking, compared to non-mycorrhizal (NM) counterparts. The highest tested fertilization regime not detrimental to the long-term vitality of intraradical hyphae reached the levels recommended for field cultivation of silage corn, except phosphorus application restricted to 60%. Under normal watering, mycorrhiza increased leaf nitrogen and phosphorus acquisition but only in cultures supplied with low NPK levels. At high fertilization levels, only the older leaves retained AM dependency, whereas for other leaf positions the AM-NM differences were leveled out. The similar size and nutritional status of highly fertilized AM and NM cultures, used in this study, eliminated fungal benefits before and during the 2-week drought progression. Nevertheless, mycorrhizal contribution became evident at the time of renewed watering, when AM plants showed much faster reversal of drought-induced leaf senescence symptoms: impaired photosynthesis and nitrogen management. Our results suggest that mycorrhiza can alter drought-induced senescence even in stay-green mutants. Moreover, this effect was apparently not mediated by AM-improved growth but triggered by activation of fungal transport at the time of recovery. Interestingly, the fungal protective potential was shown to be preserved at the expense of lowering AM vesicle number. It can be interpreted as engagement of hyphal nutritional resources targeted to maintain the symbiotic relationship despite the reduced vitality of the host. Finally, we compared the productivity of AM and NM cultures subjected to short-term drought at silking time and further fertilized with moderate or high NPK doses until the grain-filling stage. The yield and nutritive value of green forage showed that alleviation of drought-induced senescence by AM was not sufficient to have a significant positive effect on the final productivity compared to NM plants. [ABSTRACT FROM AUTHOR]

Soybean is an important economic and green manure crop that is widely used in intercropping and rotation systems due to its high biological nitrogen fixation (BNF) capacity and the resulting reduction in N fertilization. However, the genetic mechanisms underlying soybean BNF are largely unknown. Here, two soybean parent genotypes contrasting in BNF traits and 168 F9:11 recombinant inbred lines (RILs) were evaluated under four conditions in the field. The parent FC1 always produced more big nodules, yet fewer nodules in total than the parent FC2 in the field. Furthermore, nodulation in FC1 was more responsive to environmental changes than that in FC2. Broad-sense heritability (h2b) for all BNF traits varied from 0.48 to 0.87, which suggests that variation in the observed BNF traits was primarily determined by genotype. Moreover, two new QTLs for BNF traits, qBNF-16 and qBNF-17 , were identified in this study. The qBNF-16 locus was detected under all of the four tested conditions, where it explained 15.9–59.0% of phenotypic variation with LOD values of 6.31–32.5. Meanwhile qBNF-17 explained 12.6–18.6% of observed variation with LOD values of 4.93–7.51. Genotype group analysis indicated that the FC1 genotype of qBNF-16 primarily affected nodule size (NS), while the FC2 genotype of qBNF-16 promoted nodule number (NN). On the other hand, the FC1 genotype of qBNF-17 influenced NN and the FC2 genotype of qBNF-17 impacted NS. The results on the whole suggest that these two QTLs might be valuable markers for breeding elite soybean varieties with high BNF capacities. [ABSTRACT FROM AUTHOR]

Root nodule symbiosis (RNS) is a feature confined to a single clade of plants, the Fabids. Among Fabids capable of RNS, legumes form root cortex-based nodules in symbioses with rhizobia, while actinorhizal species form lateral root-based nodules with actinomycetes. Cytokinin has previously been shown to be sufficient for "pseudonodule" initiation in model legumes. Here, we tested whether this response correlates with the ability to nodulate across a range of plant species. We analyzed the formation of pseudonodules in 17 nodulating and non-nodulating legume species, and 11 non-legumes, including nodulating actinorhizal species, using light and fluorescence microscopy. Cytokinin-induced pseudonodules arising from cortical cell divisions occurred in all nodulating legume species, but not in any of the other species, including non-nodulating legumes. Pseudonodule formation was dependent on the CRE1 cytokinin receptor in Medicago truncatula. Inhibition of root growth by cytokinin occurred across plant groups, indicating that pseudonodule development is the result of a specific cortical cytokinin response unique to nodulating legumes. Lack of a cortical cytokinin response from the Arabidopsis thaliana cytokinin reporter TCSn::GFP supported this hypothesis. Our results suggest that the ability to form cortical cell-derived nodules was gained in nodulating legumes, and likely lost in non-nodulating legumes, due to a specific root cortical response to cytokinin. [ABSTRACT FROM AUTHOR]

The metabolome of a biological system provides a functional readout of the cellular state, thus serving as direct signatures of biochemical events that define the dynamic equilibrium of metabolism and the correlated phenotype. Hence, to elucidate biochemical processes involved in sorghum responses to fungal infection, a liquid chromatography-mass spectrometry-based untargeted metabolomic study was designed. Metabolic alterations of three sorghum cultivars responding to Colletotrichum sublineolum , were investigated. At the 4-leaf growth stage, the plants were inoculated with fungal spore suspensions and the infection monitored over time: 0, 3, 5, 7, and 9 days post inoculation. Non-infected plants were used as negative controls. The metabolite composition of aqueous-methanol extracts were analyzed on an ultra-high performance liquid chromatography system coupled to high-definition mass spectrometry. The acquired multidimensional data were processed to create data matrices for multivariate statistical analysis and chemometric modeling. The computed chemometric models indicated time- and cultivar-related metabolic changes that reflect sorghum responses to the fungal infection. Metabolic pathway and correlation-based network analyses revealed that this multi-component defense response is characterized by a functional metabolic web, containing defense-related molecular cues to counterattack the pathogen invasion. Components of this network are metabolites from a range of interconnected metabolic pathways with the phenylpropanoid and flavonoid pathways being the central hub of the web. One of the key features of this altered metabolism was the accumulation of an array of phenolic compounds, particularly de novo biosynthesis of the antifungal 3-deoxyanthocynidin phytoalexins, apigeninidin, luteolinidin, and related conjugates. The metabolic results were complemented by qRT-PCR gene expression analyses that showed upregulation of defense-related marker genes. Unraveling key characteristics of the biochemical mechanism underlying sorghum— C. sublineolum interactions, provided valuable insights with potential applications in breeding crop plants with enhanced disease resistance. Furthermore, the study contributes to ongoing efforts toward a comprehensive understanding of the regulation and reprogramming of plant metabolism under biotic stress. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal (AM) fungi establish probably one of the oldest mutualistic relationships with the roots of most plants on earth. The wide distribution of these fungi in almost all soil ecotypes and the broad range of host plant species demonstrate their strong plasticity to cope with various environmental conditions. AM fungi elaborate fine-tuned molecular interactions with plants that determine their spread within root cortical tissues. Interactions with endomycorrhizal fungi can bring various benefits to plants, such as improved nutritional status, higher photosynthesis, protection against biotic and abiotic stresses based on regulation of many physiological processes which participate in promoting plant performances. In turn, host plants provide a specific habitat as physical support and a favorable metabolic frame, allowing uptake and assimilation of compounds required for the life cycle completion of these obligate biotrophic fungi. The search for formal and direct evidences of fungal energetic needs raised strong motivated projects since decades, but the impossibility to produce AM fungi under axenic conditions remains a deep enigma and still feeds numerous debates. Here, we review and discuss the initial favorable and non-favorable metabolic plant context that may fate the mycorrhizal behavior, with a focus on hormone interplays and their links with mitochondrial respiration, carbon partitioning and plant defense system, structured according to the action of phosphorus as a main limiting factor for mycorrhizal symbiosis. Then, we provide with models and discuss their significances to propose metabolic targets that could allow to develop innovations for the production and application of AM fungal inocula. [ABSTRACT FROM AUTHOR]

Two types of nitrogen-fixing root nodule symbioses are known, rhizobial and actinorhizal symbioses. The latter involve plants of three orders, Fagales, Rosales, and Cucurbitales. To understand the diversity of plant symbiotic adaptation, we compared the nodule transcriptomes of Datisca glomerata (Datiscaceae, Cucurbitales) and Ceanothus thyrsiflorus (Rhamnaceae, Rosales); both species are nodulated by members of the uncultured Frankia clade, cluster II. The analysis focused on various features. In both species, the expression of orthologs of legume Nod factor receptor genes was elevated in nodules compared to roots. Since arginine has been postulated as export form of fixed nitrogen from symbiotic Frankia in nodules of D. glomerata , the question was whether the nitrogen metabolism was similar in nodules of C. thyrsiflorus. Analysis of the expression levels of key genes encoding enzymes involved in arginine metabolism revealed up-regulation of arginine catabolism, but no up-regulation of arginine biosynthesis, in nodules compared to roots of D. glomerata , while arginine degradation was not upregulated in nodules of C. thyrsiflorus. This new information corroborated an arginine-based metabolic exchange between host and microsymbiont for D. glomerata , but not for C. thyrsiflorus. Oxygen protection systems for nitrogenase differ dramatically between both species. Analysis of the antioxidant system suggested that the system in the nodules of D. glomerata leads to greater oxidative stress than the one in the nodules of C. thyrsiflorus , while no differences were found for the defense against nitrosative stress. However, induction of nitrite reductase in nodules of C. thyrsiflorus indicated that here, nitrite produced from nitric oxide had to be detoxified. Additional shared features were identified: genes encoding enzymes involved in thiamine biosynthesis were found to be upregulated in the nodules of both species. Orthologous nodule-specific subtilisin-like proteases that have been linked to the infection process in actinorhizal Fagales, were also upregulated in the nodules of D. glomerata and C. thyrsiflorus. Nodule-specific defensin genes known from actinorhizal Fagales and Cucurbitales, were also found in C. thyrsiflorus. In summary, the results underline the variability of nodule metabolism in different groups of symbiotic plants while pointing at conserved features involved in the infection process. [ABSTRACT FROM AUTHOR]

Soybean (Glycine max L. Merr.) is regarded worldwide as indisputably one of the most important crops for human food and animal feed. The presence of symbiotic bacteria and fungi is essential for soybean breeding, especially in low-input agricultural systems. Research on the cooperation between different microbial symbionts is a key to understanding how the health and productivity of the plant is supported. The symbiotic effectivity of dual and tripartite symbiotic agents was investigated in two pot experiments on different soybean cultivars with special regard to compatibility. In the Selection experiment, two out of sixteen soybean cultivars (Aliz, Emese) were chosen on the basis of their drought tolerance and used in all the other investigations. In the Compatibility experiment, the compatible coupling of symbiotic partners was selected based on the efficiency of single and co-inoculation with two Bradyrhizobium japonicum strains and two commercial arbuscular mycorrhizal fungal (AMF) products. Significant differences were found in the infectivity and effectivity of the microsymbionts. The rhizobial and AMF inoculation generally improved plant production, photosynthetic efficiency and root activity, but this effect depended on the type of symbiotic assotiation. Despite the low infectivity of AMF, inocula containing fungi were more beneficial than those containing only rhizobia. In the Drought Stress (DS) experiment, co-inoculated and control plants were grown in chernozem soil originating from organic farms. Emese was more resistant to drought stress than Aliz and produced a bigger root system. Under DS, the growth parameters of both microbially inoculated cultivars were better than that of control, proving that even drought tolerant genotypes can strengthen their endurance due to inoculation with AMF and nitrogen fixing bacteria. Root electrical capacitance (CR) showed a highly significant linear correlation with root and shoot dry mass and leaf area. The same root biomass was associated with higher CR in inoculated hosts. As CR method detects the absorptive surface increasing due to inoculation, it may be used to check the efficiency of the microbial treatment. [ABSTRACT FROM AUTHOR]

Root nodule symbiosis (RNS) is a symbiotic interaction established between angiosperm hosts and nitrogen-fixing soil bacteria in specialized organs called root nodules. The host plants provide photosynthate and the microsymbionts supply fixed nitrogen. The origin of RNS represents a major evolutionary event in the angiosperms, and understanding the genetic underpinnings of this event is of major economic and agricultural importance. Plants that engage in RNS are restricted to a single angiosperm clade known as the nitrogen-fixing clade (NFC), yet occur in multiple lineages scattered within the NFC. It has been postulated that RNS evolved in two steps: a gain-of-predisposition event occurring at the base of the NFC, followed by a gain-of-function event in each host plant lineage. Here, we first explore the premise that RNS has evolved from a single common background, and then we explore whether a two-step process better explains the evolutionary origin of RNS than either a single-step process, or multiple origins. We assembled the transcriptomes of root and nodule of two actinorhizal plants, Ceanothus thyrsiflorus and Datisca glomerata. Together with the corresponding published transcriptomes of the model legume Medicago truncatula , the gene expression patterns in roots and nodules were compared across the three lineages. We found that orthologs of many genes essential for RNS in the model legumes are expressed in all three lineages, and that the overall nodule gene expression patterns were more similar to each other than expected by random chance, a finding that supports a common evolutionary background for RNS shared by the three lineages. Moreover, phylogenetic analyses suggested that a substantial portion of the genes experiencing selection pressure changes at the base of the NFC also experienced additional changes at the base of each host plant lineage. Our results (1) support the occurrence of an event that led to RNS at the base of the NFC, and (2) suggest a subsequent change in each lineage, most consistent with a two-step origin of RNS. Among several conserved functions identified, strigolactone-related genes were down-regulated in nodules of all three species, suggesting a shared function similar to that shown for arbuscular mycorrhizal symbioses. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The rhizosphere microbiome is known to play a crucial role in promoting plant growth, partly by countering soil-borne phytoparasites and by improving nutrient uptake. The abundance and composition of the rhizosphere and root-associated microbiota are influenced by several factors, including plant species and genotype. We hypothesize that crop domestication might influence the composition and diversity of plant-associated microbiomes. We tested the contribution of domestication to the bacterial and archaeal root and soil composition associated with six genotypes of domesticated Setaria italica and four genotypes of its wild ancestor, S. viridis. The bacterial microbiome in the rhizoplane and root endophyte compartments, and the archaea in the endophyte compartment, showed major composition differences. For instance, members of the Betaproteobacteria and Firmicutes were overrepresented in S. italica root samples compared to S. viridis. Metagenomic analysis of samples that contained both root surface-bound (rhizoplane) and inside-root (endophytic) bacteria defined two unique microbial communities only associated with S. italica roots and one only associated with S. viridis roots. Root endophytic bacteria were found in six discernible communities, of which four were primarily on S. italica and two primarily on S. viridis. Among archaea, Methanobacteria, and Methanomicrobia exhibited species-associated differences in the rhizosphere and root compartments, but most detected archaea were not classified more specifically than at the level of phylum. These results indicate a host genetic contribution to the microbial composition in Setaria , and suggest that domestication has selected for specific associations in the root and in the rhizosphere. [ABSTRACT FROM AUTHOR]

Direct and indirect negative interactions between ant guards and pollinators on ant-plants are expected for two reasons. First, aggressive ants may deter pollinators directly. Second, pollinators benefit from plant investment in reproduction whilst ants benefit from plant investment in indirect defense, and resource allocation trade-offs between these functions could lead to indirect conflict. We explored the potential for ant-pollinator conflict in a Mexican myrmecophile, Turnera velutina, which rewards ants with extrafloral nectar and pollinators with floral nectar. We characterized the daily timing of ant and pollinator activity on the plant and used experiments to test for direct and indirect conflict between these two groups of mutualists. We tested for direct conflict by quantifying pollinator responses to flowers containing dead specimens of aggressive ant species, relative to unoccupied control flowers. We assessed indirect conflict by testing for the existence of a trade-off in sugar allocation between ant and pollinator rewards, evidenced by an increase in floral nectar secretion when extrafloral nectar secretion was prevented. Secretion of floral and extrafloral nectar, activity of ants and pollinators, and pollen deposition all overlapped in daily time and peaked within the first 2 h after flowers opened. We found evidence of direct conflict, in that presence of ants inside the flowers altered pollinator behavior and reduced visit duration, although visit frequency was unchanged. We found no evidence for indirect conflict, with no significant difference in the volume or sugar content of floral nectar between control plants and those in which extrafloral nectar secretion was prevented. The presence of ants in flowers alters pollinator behavior in ways that are likely to affect pollination dynamics, though there is no apparent tradeoff between plant investment in nectar rewards for pollinators and ant guards. Further studies are required to quantify the effect of the natural abundance of ants in flowers on pollinator behavior, and any associated impacts on plant reproductive success. [ABSTRACT FROM AUTHOR]

Plants in more than 100 families secrete extrafloral nectar (EFN) to establish food-for-protection mutualisms with ants. Facultative ant-plants secrete EFN as a jasmonic acid (JA)-dependent response to attract generalist ants. In contrast, obligate ant-plants like the Central American “Swollen-Thorn Acacias” are colonized by specialized ants, although an individual host can carry ant colonies from different species that differ in the degree of protection they provide. We hypothesized that hosts that associate simultaneously with various partners should produce rewards in a modular manner to preferentially reward high quality partners. To test this hypothesis, we applied JA to distinct leaves and quantified cell wall invertase activity (CWIN; a regulator of nectar secretion) and EFN secretion by these “local” (i.e., treated) and the “systemic” (i.e., non-treated) leaves of the same branch. Both CWIN activity and EFN secretion increased in local and systemic leaves of the facultative ant-plant Acacia cochliacantha, but only in the local leaves of the obligate ant-plant, A. cornigera. The systemic EFN secretion in A. cochliacantha was associated with an enhanced emission of volatile organic compounds (VOCs). Such VOCs function as “external signals” that control systemic defense responses in diverse plant species. Indeed, the headspace of JA-treated branches of A. cochliacantha induced EFN secretion in both plant species, whereas the headspace of A. cornigera caused no detectable induction effect. Analyses of the headspace using GC-MS identified six VOCs in the headspace of A. cochliacantha that were not emitted by A. cornigera. Among these VOCs, β-caryophyllene and (cis)-hexenyl isovalerate have already been reported in other plant species to induce defense traits, including EFN secretion. Our observations underline the importance of VOCs as systemic within-plant signals and show that the modular rewarding in A. cornigera is likely to result from a reduced emission of the systemic signal, rather than from a reduced responsiveness to the signal. We suggest that modular rewarding allows hosts to restrict the metabolic investment to specific partners and to efficiently sanction potential exploiters. [ABSTRACT FROM AUTHOR]

Nitrate leaching is a pressing environmental problem in intensive agriculture. Especially after the crop harvest, leaching risk is greatest due to decomposing plant residues, and low plant nutrient uptake and evapotranspiration. The specific crop also matters: grain legumes and canola commonly result in more leftover N than the following winter crop can take up before spring. Addition of a high carbon amendment (HCA) could potentially immobilize N after harvest. We set up a 2-year mesocosm experiment to test the effects of N fertilization (40 or 160 kg N/ha), HCA addition (no HCA, wheat straw, or sawdust), and precrop plant functional group identity on winter barley yield and soil C/N ratio. Four spring precrops were sown before winter barley (white lupine, faba bean, spring canola, spring barley), which were selected based on a functional group approach (colonization by arbuscular mycorrhizal fungi [AMF] and/or N2-fixing bacteria). We also measured a subset of faba bean and spring barley for leaching over winter after harvest. As expected, N fertilization had the largest effect on winter barley yield, but precrop functional identity also significantly affected the outcome. The non-AMF precrops white lupine and canola had on average a positive effect on yield compared to the AMF precrops spring barley and faba bean under high N (23% increase). Under low N, we found only a small precrop effect. Sawdust significantly reduced the yield compared to the control or wheat straw under either N level. HCAs reduced nitrate leaching over winter, but only when faba bean was sown as a precrop. In our setup, short-term immobilization of N by HCA addition after harvest seems difficult to achieve. However, other effects such as an increase in SOM or nutrient retention could play a positive role in the long term. Contrary to the commonly found positive effect of AMF colonization, winter barley showed a greater yield when it followed a non-AMF precrop under high fertilization. This could be due to shifts of the agricultural AMF community toward parasitism. [ABSTRACT FROM AUTHOR]

Soybean is one of the leading oilseed crop in the world and is showing a remarkable surge in its utilization in formulating animal feeds and supplements. Its dietary consumption, however, is incongruent with its existing industrial demand due to the presence of anti-nutritional factors in sufficiently large amounts. Phytic acid in particular raises concern as it causes a concomitant loss of indigestible complexed minerals and charged proteins in the waste and results in reduced mineral bioavailability in both livestock and humans. Reducing the seed phytate level thus seems indispensable to overcome the nutritional menace associated with soy grain consumption. In order to conceive our objective we designed and expressed a inositol polyphosphate 6-/3-/5-kinase gene-specific RNAi construct in the seeds of Pusa-16 soybean cultivar. We subsequently conducted a genotypic, phenotypic and biochemical analysis of the developed putative transgenic populations and found very low phytic acid levels, moderate accumulation of inorganic phosphate and elevated mineral content in some lines. These low phytic acid lines did not show any reduction in seedling emergence and displayed an overall good agronomic performance. [ABSTRACT FROM AUTHOR]

Plants face numerous challenges from both aboveground and belowground stressors, and defend themselves against harmful insects and microorganisms in many ways. Because plant responses to biotic stresses are not only local but also systemic, belowground interactions can influence aboveground interactions in both natural and agricultural ecosystems. Arbuscular mycorrhizal fungi (AMF) are soilborne organisms that form symbiotic associations with many plant roots and are thought to play a central role in plant nutrition, growth, and fitness. In the present study, we focused on the influence of AMF on rice defense against pests. We inoculated rice plants with AMF in several field and greenhouse experiments to test whether the interaction of AMF with rice roots changes the resistance of rice against two chewing insects, the rice water weevil (Lissorhoptrus oryzophilus Kuschel, RWW) and the fall armyworm (Spodoptera frugiperda, FAW), and against infection by sheath blight (Rhizoctonia solani, ShB). Both in field and greenhouse experiments, the performance of insects and the pathogen on rice was enhanced when plants were inoculated with AMF. In the field, inoculating rice plants with AMF resulted in higher numbers of RWW larvae on rice roots. In the greenhouse, more RWW first instars emerged from AMF-colonized rice plants than from non-colonized control plants. Weight gains of FAW larvae were higher on rice plants treated with AMF inoculum. Lesion lengths and susceptibility to ShB infection were higher in rice plants colonized by AMF. Although AMF inoculation enhanced the growth of rice plants, the nutritional analyses of root and shoot tissues indicated no major increases in the concentrations of nutrients in rice plants colonized by AMF. The large effects on rice susceptibility to pests in the absence of large effects on plant nutrition suggest that AMF colonization influences other mechanisms of susceptibility (e.g., defense signaling processes). This study represents the first study conducted in the U.S. in rice showing AMF-induced plant susceptibility to several antagonists that specialize on different plant tissues. Given the widespread occurrence of AMF, our findings will help to provide a different perspective into the causal basis of rice systemic resistance/susceptibility to insects and pathogens. [ABSTRACT FROM AUTHOR]

The main objective of this study was to investigate Artemisia annua plant property variations in terms of plant biomass, glandular trichome numbers, artemisinin production and Guaiacol peroxidase (GPOX) activity when plants are in mutualism with AMF. According to the results, A. annua mutualism with AMF significantly increased the most important and pharmaceutically relevant factors of fresh and dry plant biomass. This increase, especially in the biomass of plant herba (leaves), was 30% higher during the vegetation period and remained high (29% higher than for control) when plants were harvested at the end of the vegetation period. Similar differences in dry biomass were also detected. Glandular trichomas numbers increased by 40%, and the artemisinin content by 17% under AMF colonization. No effects due to AMF on chlorophyll variations were detected, while GPOX enzyme concentrations increased significantly under AMF colonization. Altogether the Artemisia plant properties with high pharmaceutically importance (fresh and dry biomass of leaves and artemisinin, number of trichomes and the artemisinin content) were significantly improved by AMF, the application in Artemisia cultivation can be an effective and cheap method. The high GPOX activity under AMF colonization indicate an enhanced oxidative stress alleviation, therefore a higher resistance to water deficiency, mechanisms important under climate conditions with low water supply where Artemisia is usually cultivated. [ABSTRACT FROM AUTHOR]

Common mycorrhizal networks (CMNs) formed by arbuscular mycorrhizal fungi (AMF) interconnect plants of the same and/or different species, redistributing nutrients and draining carbon (C) from the different plant partners at different rates. Here, we conducted a plant co-existence (intercropping) experiment testing the role of AMF in resource sharing and exploitation by simplified plant communities composed of two congeneric grass species (Panicum spp.) with different photosynthetic metabolism types (C3 or C4). The grasses had spatially separated rooting zones, conjoined through a root-free (but AMF-accessible) zone added with 15N-labeled plant (clover) residues. The plants were grown under two different temperature regimes: high temperature (36/32 °C day/night) or ambient temperature (25/21 °C day/night) applied over 49 days after an initial period of 26 days at ambient temperature. We made use of the distinct C-isotopic composition of the two plant species sharing the same CMN (composed of a synthetic AMF community of five fungal genera) to estimate if the CMN was or was not fed preferentially under the specific environmental conditions by one or the other plant species. Using the C-isotopic composition of AMF-specific fatty acid (C16:1ω5) in roots and in the potting substrate harboring the extraradical AMF hyphae, we found that the C3-Panicum continued feeding the CMN at both temperatures with a significant and invariable share of C resources. This was surprising because the growth of the C3 plants was more susceptible to high temperature than that of the C4 plants and the C3-Panicum alone suppressed abundance of the AMF (particularly Funneliformis sp.) in its roots due to the elevated temperature. Moreover, elevated temperature induced a shift in competition for nitrogen between the two plant species in favor of the C4-Panicum, as demonstrated by significantly lower 15N yields of the C3-Panicum but higher 15N yields of the C4-Panicum at elevated as compared to ambient temperature. Although the development of CMN (particularly of the dominant Rhizophagus and Funneliformis spp.) was somewhat reduced under high temperature, plant P uptake benefits due to AMF inoculation remained well visible under both temperature regimes, though without imminent impact on plant biomass production that actually decreased due to inoculation with AMF. [ABSTRACT FROM AUTHOR]

Parasponia represents five fast-growing tropical tree species in the Cannabaceae and is the only plant lineage besides legumes that can establish nitrogen-fixing nodules with rhizobium. Comparative analyses between legumes and Parasponia allows identification of conserved genetic networks controlling this symbiosis. However, such studies are hampered due to the absence of powerful reverse genetic tools for Parasponia. Here, we present a fast and efficient protocol for Agrobacterium tumefaciens-mediated transformation and CRISPR/Cas9 mutagenesis of Parasponia andersonii. Using this protocol, knockout mutants are obtained within 3 months. Due to efficient micro-propagation, bi-allelic mutants can be studied in the T0 generation, allowing phenotypic evaluation within 6 months after transformation. We mutated four genes - PanHK4, PanEIN2, PanNSP1, and PanNSP2 - that control cytokinin, ethylene, or strigolactone hormonal networks and that in legumes commit essential symbiotic functions. Knockout mutants in Panhk4 and Panein2 displayed developmental phenotypes, namely reduced procambium activity in Panhk4 and disturbed sex differentiation in Panein2 mutants. The symbiotic phenotypes of Panhk4 and Panein2 mutant lines differ from those in legumes. In contrast, PanNSP1 and PanNSP2 are essential for nodule formation, a phenotype similar as reported for legumes. This indicates a conserved role for these GRAS-type transcriptional regulators in rhizobium symbiosis, illustrating the value of Parasponia trees as a research model for reverse genetic studies. [ABSTRACT FROM AUTHOR]

In legume nodules, symbiosomes containing endosymbiotic rhizobial bacteria act as temporary plant organelles that are responsible for nitrogen fixation, these bacteria develop mutual metabolic dependence with the host legume. In most legumes, the rhizobia infect post-mitotic cells that have lost their ability to divide, although in some nodules cells do maintain their mitotic capacity after infection. Here, we review what is currently known about legume symbiosomes from an evolutionary and developmental perspective, and in the context of the different interactions between diazotroph bacteria and eukaryotes. As a result, it can be concluded that the symbiosome possesses organelle-like characteristics due to its metabolic behavior, the composite origin and differentiation of its membrane, the retargeting of host cell proteins, the control of microsymbiont proliferation and differentiation by the host legume, and the cytoskeletal dynamics and symbiosome segregation during the division of rhizobia-infected cells. Different degrees of symbiosome evolution can be defined, specifically in relation to rhizobial infection and to the different types of nodule. Thus, our current understanding of the symbiosome suggests that it might be considered a nitrogen-fixing link in organelle evolution and that the distinct types of legume symbiosomes could represent different evolutionary stages toward the generation of a nitrogen-fixing organelle. [ABSTRACT FROM AUTHOR]