The 8th Congress of the International Symbiosis Society will be held in Lisbon, Portugal in July 2015.
Plant-Microbe Symbiosis
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Beneficial associations between plants and microbes
Curated by
Jean-Michel Ané
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Emissions from transport and industry have increased faster than those from agriculture. Via Praveen Rahi 
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The knowledge of the survival of inoculated fungal and bacterial strains in field and the effects of their release on the indigenous microbial communities has been of great interest since the practical use of selected natural or genetically modified microorganisms has been developed. Soil inoculation or seed bacterization may lead to changes in the structure of the indigenous microbial communities, which is important with regard to the safety of introduction of microbes into the environment. Many reports indicate that application of microbial inoculants can influence, at least temporarily, the resident microbial communities. However, the major concern remains regarding how the impact on taxonomic groups can be related to effects on functional capabilities of the soil microbial communities. These changes could be the result of direct effects resulting from trophic competitions and antagonistic/synergic interactions with the resident microbial populations, or indirect effects mediated by enhanced root growth and exudation. Combination of inoculants will not necessarily produce an additive or synergic effect, but rather a competitive process. The extent of the inoculation impact on the subsequent crops in relation to the buffering capacity of the plant-soil-biota is still not well documented and should be the focus of future research.
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Bacterial genes associated with a single trait are often grouped in a contiguous unit of the genome known as a gene cluster. It is difficult to genetically manipulate many gene clusters because of complex, redundant, and integrated host regulation. We have developed a systematic approach to completely specify the genetics of a gene cluster by rebuilding it from the bottom up using only synthetic, well-characterized parts. This process removes all native regulation, including that which is undiscovered. First, all noncoding DNA, regulatory proteins, and nonessential genes are removed. The codons of essential genes are changed to create a DNA sequence as divergent as possible from the wild-type (WT) gene. Recoded genes are computationally scanned to eliminate internal regulation. They are organized into operons and placed under the control of synthetic parts (promoters, ribosome binding sites, and terminators) that are functionally separated by spacer parts. Finally, a controller consisting of genetic sensors and circuits regulates the conditions and dynamics of gene expression. We applied this approach to an agriculturally relevant gene cluster from Klebsiella oxytoca encoding the nitrogen fixation pathway for converting atmospheric N2 to ammonia. The native gene cluster consists of 20 genes in seven operons and is encoded in 23.5 kb of DNA. We constructed a “refactored” gene cluster that shares little DNA sequence identity with WT and for which the function of every genetic part is defined. This work demonstrates the potential for synthetic biology tools to rewrite the genetics encoding complex biological functions to facilitate access, engineering, and transferability.
Karsten Temme, Dehua Zhao, Christopher A. Voigt (2013). PNAS 109: 7085-7090.
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IvanOresnik's curator insight,
October 21, 2013 12:42 PM
Special thanks to Ben Perry (@Benji_JP) who suggested this article.
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Most biological nitrogen fixation is catalyzed by molybdenum-dependent nitrogenase, an enzyme complex comprising two component proteins that contains three different metalloclusters. Diazotrophs contain a common core of nitrogen fixation nif genes that encode the structural subunits of the enzyme and components required to synthesize the metalloclusters. However, the complement of nif genes required to enable diazotrophic growth varies significantly amongst nitrogen fixing bacteria and archaea. In this study, we identified a minimal nif gene cluster consisting of nine nif genes in the genome of Paenibacillus sp. WLY78, a gram-positive, facultative anaerobe isolated from the rhizosphere of bamboo. We demonstrate that the nif genes in this organism are organized as an operon comprising nifB,nifH, nifD, nifK, nifE, nifN, nifX, hesA and nifV and that the nif cluster is under the control of a σ70 (σA)-dependent promoter located upstream of nifB. To investigate genetic requirements for diazotrophy, we transferred the Paenibacillus nif cluster to Escherichia coli. The minimal nifgene cluster enables synthesis of catalytically active nitrogenase in this host, when expressed either from the native nifB promoter or from the T7 promoter. Deletion analysis indicates that in addition to the core nif genes, hesA plays an important role in nitrogen fixation and is responsive to the availability of molybdenum. Whereas nif transcription in Paenibacillus is regulated in response to nitrogen availability and by the external oxygen concentration, transcription from the nifB promoter is constitutive in E. coli, indicating that negative regulation of nif transcription is bypassed in the heterologous host. This study demonstrates the potential for engineering nitrogen fixation in a non-nitrogen fixing organism with a minimum set of nine nifgenes.
Jean-Michel Ané's comment,
October 21, 2013 11:29 PM
I look forward to Chris Voigt's talk on this topic at the Biochem's Synthetic Biology Seminar Series on December 9 in Madison: http://www.biochem.wisc.edu/seminars/contemporary_biochemistry/
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The interaction between legumes and rhizobia leads to the establishment of a symbiotic relationship characterized by the formation of new organs called nodules, in which bacteria have the ability to fix atmospheric nitrogen (N2) via the nitrogenase activity. Significant nitric oxide (NO) production was evidenced in the N2-fixing nodules suggesting that it may impact the symbiotic process. Indeed, NO was shown to be a potent inhibitor of nitrogenase activity and symbiotic N2 fixation. It has also been shown that NO production is increased in hypoxic nodules and this production was supposed to be linked - via a nitrate/NO respiration process - with improved capacity of the nodules to maintain their energy status under hypoxic conditions. Other data suggest that NO might be a developmental signal involved in the induction of nodule senescence. Hence, the questions were raised of the toxic effects versus signaling/metabolic functions of NO, and of the regulation of NO levels compatible with nitrogenase activity. The present review analyses the different roles of NO in functioning nodules, and discusses the role of plant and bacterial (flavo)hemoglobins in the control of NO level in nodules.
Boscari A, Meilhoc E, Castella C, Bruand C, Puppo A, Brouquisse R. (2013). Front Plant Sci. 4:384. Via IvanOresnik
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Legumes, such as soybean plants, form beneficial symbiotic relationships with Rhizobium spp., which reside in nitrogen-fixing root nodules that promote plant growth. Although it was previously thought that this interaction was limited to legumes, new research shows that non-legumes can also respond to rhizobial symbiotic signals. Similarly to their interaction with pathogens, legumes initially respond to Rhizobium spp. by mounting an immune response following the recognition of microorganism-associated molecular patterns (MAMPs), such as bacterial flagellin. However, this is quickly suppressed by Nod factors — lipochitin molecules that also activate signalling pathways which ultimately lead to nodule formation. Consistent with this, Stacey and colleagues observed that the production of reactive oxygen species (ROS) in soybean leaves on exposure to flg22 (a synthetic peptide containing the conserved 22 amino acids of most bacterial flagellins) was greatly suppressed by the addition of Nod factors or of another chitin oligomer, chitotetraose. Moreover, Nod or chitotetraose pretreatment decreased the activation of mitogen-activated protein kinase (MAPK), which occurs downstream of flg22 recognition. Surprisingly, Nod or chitotetraose pretreatment also reduced flg22-induced ROS production and MAPK activation in the leaves of the non-legume Arabidopsis thaliana, and allowed colonization of the whole plant with the pathogen Pseudomonas syringae. Similarly to legumes, which recognize Nod factors through specific receptors, A. thaliana required the LysM receptor kinase LYK3 for Nod recognition, and plants with mutated LYK3 failed to suppress flg22-mediated ROS production and MAPK activation. The presence of Nod factors or chitotetraose also suppressed the influx of Ca2+, the intracellular concentration of which is known to increase following flg22 exposure. This observation led the authors to postulate that Nod-mediated inhibition of plant immunity might occur at the level of flg22 recognition. Indeed, the levels of the flg22 receptor, FLS2, at the plasma membrane were greatly reduced after Nod factor treatment. This was mediated through proteasomal degradation of FLS2 following ubiquitylation by E3 ligases recruited by the kinase BAK1, and Nod had no effect on FLS2 levels in the presence of a proteasome inhibitor or in BAK1 mutants. As Nod factors or chitotetraose treatment also suppressed flg22-triggered ROS production in corn and tomato plants (other non-legumes), it is possible that Nod factor-mediated suppression of MAMP-induced immunity is universal in plants. Taking advantage of such potential symbiotic relationships might help to reduce the dependence on nitrogen fertilizers.
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Although many α- and some β-proteobacterial species are symbiotic with legumes, the evolutionary origin of nitrogen-fixing nodulation remains unclear. We examined α- and β-proteobacteria whose genomes were sequenced using large-scale phylogenetic profiling and revealed the evolutionary origin of two nodulation genes. These genes, nodI and nodJ(nodIJ), play key roles in the secretion of Nod factors, which are recognized by legumes during nodulation. We found that only the nodulating β-proteobacteria, including the novel strains isolated in this study, possess both nodIJ and their paralogous genes (DRA-ATPase/permease genes). Contrary to the widely accepted scenario of the α-proteobacterial origin of rhizobia, our exhaustive phylogenetic analysis showed that the entire nodIJ clade is included in the clade of Burkholderiaceae DRA-ATPase/permease genes, that is, the nodIJ genes originated from gene duplication in a lineage of the β-proteobacterial family. After duplication, the evolutionary rates of nodIJ were significantly accelerated relative to those of homologous genes, which is consistent with their novel function in nodulation. The likelihood analyses suggest that this accelerated evolution is not associated with changes in either nonsynonymous/synonymous substitution rates or transition/transversion rates, but rather, in the GC content. Although the low GC content of the nodulation genes has been assumed to reflect past horizontal transfer events from donor rhizobial genomes with low GC content, no rhizobial genome with such low GC content has yet been found. Our results encourage a reconsideration of the origin of nodulation and suggest new perspectives on the role of the GC content of bacterial genes in functional adaptation.
Mol Biol Evol (2013) doi: 10.1093/molbev/mst153First published online: September 11, 2013 Via IvanOresnik
IvanOresnik's curator insight,
October 15, 2013 11:30 AM
I would like to thank Ann Hirsch for bringing this paper to my attention.
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The default mineral nutrient acquisition strategy of land plants is the symbiosis with arbuscular mycorrhiza (AM) fungi. Research into the cell and developmental biology of AM revealed fascinating insights into the plasticity of plant cell development and of interorganismic communication. It is driven by the prospect of increased exploitation of AM benefits for sustainable agriculture. The plant cell developmental program for intracellular accommodation of AM fungi is activated by a genetically defined signaling pathway involving calcium spiking in the nucleus as second messenger. Calcium spiking is triggered by chitooligosaccharides released by AM fungi that are probably perceived via LysM domain receptor kinases. Fungal infection and calcium spiking are spatiotemporally coordinated, and only cells committed to accommodating the fungus undergo high-frequency spiking. Delivery of mineral nutrients by AM fungi occurs at tree-shaped hyphal structures, the arbuscules, in plant cortical cells. Nutrients are taken up at a plant-derived periarbuscular membrane, which surrounds fungal hyphae and carries a specific transporter composition that is of direct importance for symbiotic efficiency. An elegant study has unveiled a new and unexpected mechanism for specific protein localization to the periarbuscular membrane, which relies on the timing of gene expression to synchronize protein biosynthesis with a redirection of secretion. The control of AM development by phytohormones is currently subject to active investigation and has led to the rediscovery of strigolactones. Nearly all tested phytohormones regulate AM development, and major insights into the mechanisms of this regulation are expected in the near future.
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We studied the role of taxonomical and functional ectomycorrhizal (ECM) fungal diversity in root formation and nutrient uptake by Norway spruce (Picea abies) seedlings with fast- and slow-growing phenotypes.Seedlings were grown with an increasing ECM fungal diversity gradient from one to four species and sampled before aboveground growth differences between the two phenotypes were apparent. ECM fungal colonization patterns were determined and functional diversity was assayed via measurements of potential enzyme activities of eight exoenzymes probably involved in nutrient mobilization.Phenotypes did not vary in their receptiveness to different ECM fungal species. However, seedlings of slow-growing phenotypes had higher fine-root density and thus more condensed root systems than fast-growing seedlings, but the potential enzyme activities of ectomycorrhizas did not differ qualitatively or quantitatively. ECM species richness increased host nutrient acquisition potential by diversifying the exoenzyme palette. Needle nitrogen content correlated positively with high chitinase activity of ectomycorrhizas.Rather than fast- and slow-growing phenotypes exhibiting differing receptiveness to ECM fungi, our results suggest that distinctions in fine-root structuring and in the belowground growth strategy already apparent at early stages of seedling development may explain later growth differences between fast- and slow-growing families.
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Rhizobial bacteria form symbiotic, nitrogen-fixing nodules on the roots of compatible host legume plants. One of the most well-developed model systems for studying these interactions is the plant Medicago truncatula cv. Jemalong A17 and the rhizobial bacterium Sinorhizobium meliloti 1021. Repeated imaging of plant roots and scoring of symbiotic phenotypes requires methods that are non-destructive to either plants or bacteria. The symbiotic phenotypes of some plant and bacterial mutants become apparent after relatively short periods of growth, and do not require long-term observation of the host/symbiont interaction. However, subtle differences in symbiotic efficiency and nodule senescence phenotypes that are not apparent in the early stages of the nodulation process require relatively long growth periods before they can be scored. Several methods have been developed for long-term growth and observation of this host/symbiont pair. However, many of these methods require repeated watering, which increases the possibility of contamination by other microbes. Other methods require a relatively large space for growth of large numbers of plants. The method described here, symbiotic growth of M. truncatula/S. meliloti in sterile, single-plant microcosms, has several advantages. Plants in these microcosms have sufficient moisture and nutrients to ensure that watering is not required for up to 9 weeks, preventing cross-contamination during watering. This allows phenotypes to be quantified that might be missed in short-term growth systems, such as subtle delays in nodule development and early nodule senescence. Also, the roots and nodules in the microcosm are easily viewed through the plate lid, so up-rooting of the plants for observation is not required.
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Background The new symbiotic loci of M. truncatula isolated in this study provide the foundation for further characterization of the mechanisms underpinning nodulation, in particular the later stages associated with bacterial release and nodule function.
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In this Editorial we highlight some of the innovative mycorrhizal research published in the journal and look to future challenges that lie ahead in the vraie histoire d'amour (true love story) linking New Phytologist and mycorrhizal research. In the last century, New Phytologist hosted Harley's and Mosse's seminal papers on ectomycorrhizas and arbuscular mycorrhizas (AM), and thereafter a strong association between the journal and the mycorrhizal community has developed. Via Francis Martin
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Leguminous biological nitrogen fixation (BNF) is very sensitive to environmental fluctuations. It is still contentious how BNF is regulated under stress conditions. The local or systemic control of BNF and the role played by reactive oxygen species (ROS) in such regulation have still not been elucidated completely. Cadmium, which belongs to the so-called heavy metals, is one of the most toxic substances released into the environment. The mechanisms involved in Cd toxicity are still not completely understood but the overproduction of ROS is one of its characteristic symptoms. In this work, we used a split-root system approach to study nodule BNF and the antioxidant machinery’s response to the application of a mild Cd treatment on one side of a nodulatedMedicago truncatula root system. Cd induced the majority of nodule antioxidants without generating any oxidative damage. Cd treatment also provoked BNF inhibition exclusively in nodules directly exposed to Cd, without provoking any effect on plant shoot biomass or chlorophyll content. The overall data suggest that the decline in BNF was not due to a generalized breakdown of the plant but to control exerted through leghemoglobin/oxygen availability, affecting nitrogenase function.
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Microbial communities play a pivotal role in the functioning of plants by influencing their physiology and development. While many members of the rhizosphere microbiome are beneficial to plant growth, also plant pathogenic microorganisms colonize the rhizosphere striving to break through the protective microbial shield and to overcome the innate plant defense mechanisms in order to cause disease. A third group of microorganisms that can be found in the rhizosphere are the true and opportunistic human pathogenic bacteria, which can be carried on or in plant tissue and may cause disease when introduced into debilitated humans. Although the importance of the rhizosphere microbiome for plant growth has been widely recognized, for the vast majority of rhizosphere microorganisms no knowledge exists. To enhance plant growth and health, it is essential to know which microorganism is present in the rhizosphere microbiome and what they are doing. Here, we review the main functions of rhizosphere microorganisms and how they impact on health and disease. We discuss the mechanisms involved in the multitrophic interactions and chemical dialogues that occur in the rhizosphere. Finally, we highlight several strategies to redirect or reshape the rhizosphere microbiome in favor of microorganisms that are beneficial to plant growth and health. Via Francis Martin, Praveen Rahi
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Nitrogen is a critical limiting element for plant growth and production. It is a major component of chlorophyll, the most important pigment needed for photosynthesis, as well as amino acids, the key building blocks of proteins. It is also found in other important biomolecules, such as ATP and nucleic acids. Even though it is one of the most abundant elements (predominately in the form of nitrogen gas (N2) in the Earth’s atmosphere), plants can only utilize reduced forms of this element. Plants acquire these forms of “combined” nitrogen by: 1) the addition of ammonia and/or nitrate fertilizer (from the Haber-Bosch process) or manure to soil, 2) the release of these compounds during organic matter decomposition, 3) the conversion of atmospheric nitrogen into the compounds by natural processes, such as lightning, and 4) biological nitrogen fixation (Vance 2001).
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Legume plants are important in agriculture because they represent an important source of protein for human and animal consumption. This high protein content results from their capacity to use atmospheric nitrogen for their nutrition as a consequence of their symbiotic interaction with rhizobia. Understanding this interaction at the molecular level is a prerequisite for its better use in agriculture and for the long term objective of its transfer to other crops. Agrobacterium-mediated transformation is a tool of choice for studying this interaction and for unraveling the function of the different genes discovered through classical genetic approaches. However, legume plants are often recalcitrant to regeneration and transformation. This paper describes the technology developments (regeneration, transformation, insertion mutagenesis) related to Agrobacterium transformations that were established in the legume plants, as well as different examples of the technology developments or gene discoveries resulting from these studies.
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In nitrogen poor soils legumes establish a symbiotic interaction with rhizobia that results in the formation of root nodules. These are unique plant organs where bacteria differentiate into bacteroids, which express the nitrogenase enzyme complex that reduces atmospheric N 2 to ammonia. Nodule metabolism requires a tight control of the concentrations of reactive oxygen and nitrogen species (RONS) so that they can perform useful signaling roles while avoiding nitro-oxidative damage. In nodules a thiol-dependent regulatory network that senses, transmits and responds to redox changes is starting to be elucidated. A combination of enzymatic, immunological, pharmacological and molecular analyses has allowed us to conclude that glutathione and its legume-specific homolog, homoglutathione, are abundant in meristematic and infected cells, that their spatio-temporally distribution is correlated with the corresponding (homo)glutathione synthetase activities, and that they are crucial for nodule development and function. Glutathione is at high concentrations in the bacteroids and at moderate amounts in the mitochondria, cytosol and nuclei. Less information is available on other components of the network. The expression of multiple isoforms of glutathione peroxidases, peroxiredoxins, thioredoxins, glutaredoxins and NADPH-thioredoxin reductases has been detected in nodule cells using antibodies and proteomics. Peroxiredoxins and thioredoxins are essential to regulate and in some cases to detoxify RONS in nodules. Further research is necessary to clarify the regulation of the expression and activity of thiol redox-active proteins in response to abiotic, biotic and developmental cues, their interactions with downstream targets by disulfide-exchange reactions, and their participation in signaling cascades. The availability of mutants and transgenic lines will be crucial to facilitate systematic investigations into the function of the various proteins in the legume-rhizobial symbiosis.
Frendo P, Matamoros MA, Alloing G, Becana M. (2013). Front Plant Sci. Sep 26;4:376.
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Microbial ecology: A bacterial decoy skews plant defences Christina Tobin Kåhrström
Symbiosis: Non-legumes answer the rhizobial call Rachel David
RNA silencing suppression by plant pathogens: defence, counter-defence and counter-counter-defence Nathan Pumplin & Olivier Voinnet
On the front line: structural insights into plant–pathogen interactions Lennart Wirthmueller, Abbas Maqbool & Mark J. Banfield
Geminiviruses: masters at redirecting and reprogramming plant processes Linda Hanley-Bowdoin, Eduardo R. Bejarano, Dominique Robertson & Shahid Mansoor
Going back to the roots: the microbial ecology of the rhizosphere Laurent Philippot, Jos M. Raaijmakers, Philippe Lemanceau & Wim H. van der Putten
Filamentous plant pathogen effectors in action Martha C. Giraldo & Barbara Valent Via Nicolas Denancé
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Biotic interactions play primary roles in major theories of the distribution and abundance of species, yet the nature of these biotic interactions can depend upon the larger ecological community. Leguminous plants, for example, commonly associate with both arbuscular mycorrhizal fungi (AMF) and rhizobia bacteria and the pairwise interactions may depend upon the presence or identity of the third partner. To determine if the dynamics of plant-AMF and plant-rhizobia interactions are affected by the alternate symbiont, we manipulated the presence and identity of each symbiont, as well as levels of the nutrients supplied by each symbiont (nitrogen and phosphorus), on the growth of prairie legume Amorpha canescens. We found strong synergistic effects of AMF and rhizobia inoculation on plant biomass production that were independent of nutrient levels. AMF and rhizobia responses were influenced by the other, but not in the same direction. AMF infection increased root nodule number and weight, but rhizobia inoculation decreased AMF hyphal colonization of roots. The relative benefits of each combination of symbionts depended upon phosphorus level. The effect of nitrogen was also contingent on the biotic environment where nitrogen addition decreased nodulation but this reduction was reduced with coinfection by AMF. Our results demonstrate a strong contingency on the co-occurrence of AMF and rhizobia for the long-term fitness of A. canescens and suggest that the belowground community is critical for the success of this species in tallgrass prairies.
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Background and aims
Bradyrhizobium japonicum and Bradyrhizobium elkanii dominated soybean nodules in temperate and subtropical regions in Nepal, respectively, in our previous study. The aims of this study were to reveal the effects of temperature on the nodulation dominancy of B. japonicum and B. elkanii and to clarify the relationship between the effects of temperature and the climate-dependent distribution ofBradyrhizobium species. MethodsA laboratory competition experiment was conducted between B. japonicum and B. elkanii strains isolated from the same temperate location in Nepal. A mixture of each strain was inoculated into sterilized vermiculite with or without soybean seeds, and inoculated samples were incubated at 33/27 (day/night) and 23/17 °C. Relative populations in the non-rhizosphere, rhizosphere, and nodules were determined by competitive PCR using specific primers for each strain at 0, 1, 2, and 4 weeks after inoculation. ResultsBoth separately inoculated B. japonicum and B. elkanii strains formed nodules at both temperatures. Under competitive conditions, B. japonicum strains dominated at low temperature; however, at high temperature, both strains achieved co-nodulation in 1 week, with B. elkaniidominating after 2 weeks. The relative populations of both strains were similar in the non-rhizosphere and rhizosphere at low temperature, but B. elkanii strains dominated in these regions at high temperature. ConclusionsThe domination of B. japonicum strains in nodules at the low temperature appeared to be due to preferential infection, while the domination of B. elkanii strains at high temperature appeared to be due to the higher population of B. elkanii in the non-rhizosphere and rhizosphere, in addition to its domination in nodules after co-nodulation. The effects of temperature on the competition between B. japonicum and B. elkanii strains were remarkable and corresponded with the distribution of bradyrhizobial species in Nepal
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Auxin action is mediated by a complex signalling pathway involving transcription factors of the auxin response factor (ARF) family. In Arabidopsis, microRNA160 (miR160) negatively regulates three ARF genes (ARF10/ARF16/ARF17) and therefore controls several developmental processes, including primary and lateral root growth. Here, we analysed the role of miR160 in root development and nodulation in Medicago truncatula Gaertn. Bioinformatic analyses identified two main mtr-miR160 variants (mtr-miR160abde and mtr-miR160c) and 17 predicted ARF targets. The miR160-dependent cleavage of four predicted targets in roots was confirmed by analysis of parallel analysis of RNA ends (PARE) data and RACE-PCR experiments. Promoter-GUS analyses for mtr-miR160d and mtr-miR160c genes revealed overlapping but distinct expression profiles during root and nodule development. In addition, the early miR160 activation in roots during symbiotic interaction was not observed in mutants of the nodulation signalling or autoregulation pathways. Composite plants that overexpressed mtr-miR160a under two different promoters exhibited distinct defects in root growth and nodulation: the p35S:miR160a construct led to reduced root length associated to a severe disorganisation of the RAM, whereas pCsVMV:miR160a roots showed gravitropism defects and lower nodule numbers. Our results suggest that a regulatory loop involving miR160/ARFs governs root and nodule organogenesis in M. truncatula.
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Nitrogen, particularly nitrate is an important yield determinant for crops. However, current agricultural practice with excessive fertilizer usage has detrimental effects on the environment. Therefore, legumes have been suggested as a sustainable alternative for replenishing soil nitrogen. Legumes can uniquely form nitrogen-fixing nodules through symbiotic interaction with specialized soil bacteria. Legumes possess a highly plastic root system which modulates its architecture according to the nitrogen availability in the soil. Understanding how legumes regulate root development in response to nitrogen availability is an important step to improving root architecture. The nitrogen-mediated root development pathway starts with sensing soil nitrogen level followed by subsequent signal transduction pathways involving phytohormones, microRNAs and regulatory peptides that collectively modulate the growth and shape of the root system. This review focuses on the current understanding of nitrogen-mediated legume root architecture including local and systemic regulations by different N-sources and the modulations by phytohormones and small regulatory molecules.
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Small polypeptides are widely used as signaling molecules in cell-to-cell communication in animals and plants. The CLAVATA3/EMBRYO SURROUNDING REGION-RELATED (CLE) gene family is composed of numerous genes that contain conserved CLE domains in various plant species and plant-parasitic nematodes. Here, we review recent progress in our understanding of CLE signaling during stem cell maintenance in Arabidopsis and grasses. We also summarize the roles of CLE signaling in the legume-Rhizobium symbiosis and infection by plant-parasitic nematodes. CLE signaling is important for diverse aspects of cell-to-cell signaling and long-distance communication, which are critical for survival, and the basic components of the CLE signaling pathway are evolutionarily conserved in both plants and animals.
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Arbuscular mycorrhizal symbiosis (AMS) and root nodule symbiosis (RNS) are mutualistic plant–microbe interactions that confer nutritional benefits to both partners. Leguminous plants possess a common genetic system for intracellular symbiosis with AM fungi and with rhizobia. Here we show that CERBERUS and NSP1, which respectively encode an E3 ubiquitin ligase and a GRAS transcriptional regulator and which have previously only been implicated in RNS, are involved in AM fungal infection in Lotus japonicus. Hyphal elongation along the longitudinal axis of the root was reduced in the cerberus mutant, giving rise to a lower colonization level. Knockout ofNSP1 decreased the frequency of plants colonized by AM fungi or rhizobia. CERBERUS and NSP1 showed different patterns of expression in response to infection with symbiotic microbes. A low constitutive level ofCERBERUS expression was observed in the root and an increased level ofNSP1 expression was detected in arbuscule-containing cells. Induction of AM marker gene was triggered in both cerberus and nsp1 mutants by infection with symbiotic microbes; however, the mutants showed a weaker induction of marker gene expression than the wild type, mirroring their lower level of colonization. The common symbiosis genes are believed to act in an early signaling pathway for recognition of symbionts and for triggering early symbiotic responses. Our quantitative analysis of symbiotic phenotypes revealed developmental defects of the novel common symbiosis mutants in both symbioses, which demonstrates that common symbiosis mechanisms also contribute to a range of functions at later or different stages of symbiont infection.
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