Arbuscular Mycorrhiza Symbiosis Research Articles

Epigenetic control during root development and symbiosis.

The roots of plants play multiples functions that are essential for growth and development, including anchoring to the soil and water and nutrient acquisition. These underground organs exhibit the plasticity to modify their root system architecture in response to environmental cues allowing adaptation to change in water and nutrient availability. In addition, roots enter in mutualistic interactions with soil microorganisms, e.g. the root nodule symbiosis established between a limited group of plants and nitrogen fixing soil bacteria and the arbuscular mycorrhiza symbiosis involving most land plants and fungi of the Glomeromycetes phylum. In the past 20 years, genetic approaches allowed the identification and functional characterization of genes required for the specific programs of root development, root nodule and arbuscular mycorrhiza symbioses. These genetic studies provided evidence that the program of root nodule symbiosis recruited components of the arbuscular mycorrhiza symbiosis and the root developmental programs. The execution of these programs is strongly influenced by epigenetic changes -DNA methylation and histone post-translational modifications- that alter chromatin conformation modifying the expression of key genes. In this review, we summarize recent advances that highlighted how DNA methylation and histone post-translational modifications, as well as chromatin remodeling factors and long non-coding RNAs, shape the root system architecture and allow the successful establishment of both root nodule and arbuscular mycorrhiza symbioses. We anticipate that the analysis of dynamic epigenetic changes and chromatin 3D structure in specific single-cells or tissue types of root organs will illuminate our understanding of how root developmental and symbiotic programs are orchestrated, opening exciting questions and new perspectives to modulate agronomical and ecological traits linked to nutrient acquisition.

Carbon and phosphorus exchange rates in arbuscular mycorrhizas depend on environmental context and differ among co-occurring plants.

Phosphorus (P) for carbon (C) exchange is the pivotal function of arbuscular mycorrhiza (AM), but how this exchange varies with soil P availability and among co-occurring plants in complex communities is still largely unknown. We collected intact plant communities in two regions differing c. 10-fold in labile inorganic P. After a 2-month glasshouse incubation, we measured 32P transfer from AM fungi (AMF) to shoots and 13C transfer from shoots to AMF using an AMF-specific fatty acid. AMF communities were assessed using molecular methods. AMF delivered a larger proportion of total shoot P in communities from high-P soils despite similar 13C allocation to AMF in roots and soil. Within communities, 13C concentration in AMF was consistently higher in grass than in blanketflower (Gaillardia aristata Pursh) roots, that is P appeared more costly for grasses. This coincided with differences in AMF taxa composition and a trend of more vesicles (storage structures) but fewer arbuscules (exchange structures) in grass roots. Additionally, 32P-for-13C exchange ratios increased with soil P for blanketflower but not grasses. Contrary to predictions, AMF transferred proportionally more P to plants in communities from high-P soils. However, the 32P-for-13C exchange differed among co-occurring plants, suggesting differential regulation of the AM symbiosis.

Integrated transcriptomics and metabolomics reveal specific phenolic and flavonoid accumulation in licorice (Glycyrrhiza uralensis Fisch.) induced by arbuscular mycorrhiza symbiosis under drought stress

Arbuscular mycorrhizal (AM) symbiosis can strengthen plant defense against abiotic stress, such as drought, through multiple mechanisms; however, the specialized chemical defenses induced by AM symbiosis are largely unknown. In a pot experiment, licorice (Glycyrrhiza uralensis Fisch.) inoculated with and without arbuscular mycorrhizal fungus Rhizophagus irregularis Schenck & Smith were grown under well-watered or water deficit conditions. Transcriptomic and metabolomic analyses were combined to investigate licorice root specialized metabolism induced by AM symbiosis under drought stress. Results showed that mycorrhizal plants had few dead leaves, less biomass reduction, and less differentially expressed genes and metabolite features in response to drought compared with nonmycorrhizal plants. Transcriptomic and metabolomic data revealed that mycorrhizal roots generally accumulated lignin regardless of the water regime; however, the expression of genes involved in lignin biosynthesis was significantly downregulated by drought stress in mycorrhizal plants. By contrast, AM inoculation significantly decreased specialized metabolites accumulation, including phenolics and flavonoids under well-watered conditions, whereas these decreases turned to be nonsignificant under drought stress. Moreover, these specific phenolics and flavonoids showed significant drought-induced accumulation pattern in mycorrhizal roots. These results highlight that accumulation of specific root phenolics and flavonoids may support the drought tolerance of mycorrhizal plants.

Quantifying root colonization by a symbiotic fungus using automated image segmentation and machine learning approaches

Arbuscular mycorrhizas (AM) are one of the most widespread symbiosis on earth. This plant-fungus interaction involves around 72% of plant species, including most crops. AM symbiosis improves plant nutrition and tolerance to biotic and abiotic stresses. The fungus, in turn, receives carbon compounds derived from the plant photosynthetic process, such as sugars and lipids. Most studies investigating AM and their applications in agriculture requires a precise quantification of the intensity of plant colonization. At present, the majority of researchers in the field base AM quantification analyses on manual visual methods, prone to operator errors and limited reproducibility. Here we propose a novel semi-automated approach to quantify AM fungal root colonization based on digital image analysis comparing three methods: (i) manual quantification (ii) image thresholding, (iii) machine learning. We recognize machine learning as a very promising tool for accelerating, simplifying and standardizing critical steps in analysing AM quantification, answering to an urgent need by the scientific community studying this symbiosis.

General and specialized metabolites in peanut roots regulate arbuscular mycorrhizal symbiosis

Arbuscular mycorrhizae (AM) fungi form symbiotic associations with plant roots, providing nutritional benefits and promoting plant growth and defenses against various stresses. Metabolic changes in the roots during AM fungal colonization are key to understanding the development and maintenance of these symbioses. Here, we investigated metabolic changes in the roots of peanut (Arachis hypogaea L.) plants during the colonization and development of AM symbiosis, and compared them to uncolonized roots. The primary changes during the initial stage of AM colonization were in the contents and compositions of phenylpropanoid and flavonoid compounds. These compounds function in signaling pathways that regulate recognition, interactions, and pre-colonization between roots and AM fungi. Flavonoid compounds decreased by 25% when the symbiosis was fully established compared to the initial colonization stage. After AM symbiosis was established, general metabolism strongly shifted toward the formation of lipids, amino acids, carboxylic acids, and carbohydrates. Lipid compounds increased by 8.5% from the pre-symbiotic stage to well-established symbiosis. Lyso-phosphatidylcholines, which are signaling compounds, were only present in AM roots, and decreased in content after the symbiosis was established. In the initial stage of AM establishment, the content of salicylic acid increased two-fold, whereas jasmonic acid and abscisic acid decreased compared to uncolonized roots. The jasmonic acid content decreased in roots after the symbiosis was well established. AM symbiosis was associated with high levels of calcium, magnesium, and D-(+)-mannose, which stimulated seedling growth. Overall, specific metabolites that favor the establishment of AM symbiosis were common in the roots, primarily during early colonization, whereas general metabolism was strongly altered when AM symbiosis was well-established. In conclusion, specialized metabolites function as signaling compounds to establish AM symbiosis. These compounds are no longer produced after the symbiosis between the roots and AM becomes fully established.

Disentangling the Belowground Web of Biotic Interactions in Temperate Coastal Grasslands: From Fundamental Knowledge to Novel Applications

Grasslands represent an essential part of terrestrial ecosystems. In particular, coastal grasslands are dominated by the influence of environmental factors resulting from sea–land interaction. Therefore, coastal grasslands are extremely heterogeneous both spatially and temporally. In this review, recent knowledge in the field of biotic interactions in coastal grassland soil is summarized. A detailed analysis of arbuscular mycorrhiza symbiosis, rhizobial symbiosis, plant–parasitic plant interactions, and plant–plant interactions is performed. The role of particular biotic interactions in the functioning of a coastal grassland ecosystem is characterized. Special emphasis is placed on future directions and development of practical applications for sustainable agriculture and environmental restoration. It is concluded that plant biotic interactions in soil are omnipresent and important constituents in different ecosystem services provided by coastal grasslands.

SWEET transporters of Medicago lupulina in the arbuscular-mycorrhizal system in the presence of medium level of available phosphorus.

Arbuscular mycorrhiza (AM) fungi receive photosynthetic products and sugars from plants in exchange for contributing to the uptake of minerals, especially phosphorus, from the soil. The identification of genes controlling AM symbiotic efficiency may have practical application in the creation of highly productive plant-microbe systems. The aim of our work was to evaluate the expression levels of SWEET sugar transporter genes, the only family in which sugar transporters specific to AM symbiosis can be detected. We have selected a unique "host plant-AM fungus" model system with high response to mycorrhization under medium phosphorus level. This includes a plant line which is highly responsive to inoculation by AM fungi, an ecologically obligate mycotrophic line MlS-1 from black medick (Medicago lupulina) and the AM fungus Rhizophagus irregularis strain RCAM00320, which has a high efficiency in a number of plant species. Using the selected model system, differences in the expression levels of 11 genes encoding SWEET transporters in the roots of the host plant were evaluated during the development of or in the absence of symbiosis of M. lupulina with R. irregularis at various stages of the host plant development in the presence of medium level of phosphorus available for plant nutrition in the substrate. At most stages of host plant development, mycorrhizal plants had higher expression levels of MlSWEET1b, MlSWEET3c, MlSWEET12 and MlSWEET13 compared to AM-less controls. Also, increased expression relative to control during mycorrhization was observed for MlSWEET11 at 2nd and 3rd leaf development stages, for MlSWEET15c at stemming (stooling) stage, for MlSWEET1a at 2nd leaf development, stemming and lateral branching stages. The MlSWEET1b gene can be confidently considered a good marker with specific expression for effective development of AM symbiosis between M. lupulina and R. irregularis in the presence of medium level of phosphorus available to plants in the substrate.

Molecular Assessment of Metabolome Alterations in Lotus japonicus Roots Induced by Arbuscular Mycorrhiza

AbstractAn arbuscular mycorrhiza symbiosis, which is established by 80% of all vascular plants and Glomeromycota fungi, can boost the growth, stress tolerance, and yield of agricultural crop plants by increasing the mineral supply of the host plant [1]. Mineral nutrients absorbed by the fungus are exchanged for photoassimilates from the plant via highly branched fungal structures called arbuscules which are formed within root cortex cells [2]. The process of arbuscule formation and degradation is highly dynamic and accompanied by considerable cell reprogramming [3], but there is only limited knowledge on the influence of arbuscular mycorrhiza on the host metabolome.We investigated alterations of the host plant's root metabolome induced by arbuscular mycorrhizal symbiosis using the model legume Lotus japonicus. Methanol extracts of mycorrhizal (myc) and non‐mycorrhizal roots (mock) were analyzed employing ultra‐highperformance liquid chromatography–electrospray ionization–time‐of‐flight–ion mobility–mass spectrometry (UPLC–ESI–ToF–IM–MS) in an untargeted metabolomics approach. Up‐regulated marker metabolites were identified via principal components analysis (PCA) and orthogonal partial least squares–discriminant analysis (OPLS–DA) depicted as S‐plots, and their structures elucidated by co‐chromatography with authentic standards or by isolation from L. japonicus roots using preparative HPLC and one‐ and two‐dimensional nuclear magnetic resonance spectroscopy (NMR) experiments. We isolated three previously unknown mycorrhizal marker polyphenols from L. japonicus roots, one being a coumaronochromone, one pterocarp‐6a‐ene, and one aryl benzofuran carboxaldehyde. Additionally, we identified three more coumaronochromones, three flavonoids, three isoflavonoids, and one coumestan in mycorrhized L. japonicus.

The mysterious non-arbuscular mycorrhizal status of Brassicaceae species.

The Brassicaceae family is unique in not fostering functional symbiosis with arbuscular mycorrhiza (AM). The family is also special in possessing glucosinolates, a class of secondary metabolites predominantly functioning for plant defence. We have reviewed what effect the glucosinolates of this non-symbiotic host have on AM or vice versa. Isothiocyanates, the toxic degradation product of the glucosinolates, particularly the indolic and benzenic glucosinolates, are known to be involved in the inhibition of AM. Interestingly, AM colonization enhances glucosinolate production in two AM-host in the Brassicales family- Moringa oleifera and Tropaeolum spp. PHOSPHATE STARVATION RESPONSE 1 (PHR1), a central transcription factor that controls phosphate starvation response also activates the glucosinolate biosynthesis in AM non-host Arabidopsis thaliana. Recently, the advances in whole-genome sequencing, enabling extensive ecological microbiome studies have helped unravel the Brassicaceae microbiome, identifying new mutualists that compensate for the loss of AM symbiosis, and reporting cues for some influence of glucosinolates on the microbiome structure. We advocate that glucosinolate is an important candidate in determining the mycorrhizal status of Brassicaceae and has played a major role in its symbiosis-defence trade-off. We also identify key open questions in this area that remain to be addressed in the future.

Long-lasting impact of chito-oligosaccharide application on strigolactone biosynthesis and fungal accommodation promotes arbuscular mycorrhiza in Medicago truncatula.

• The establishment of arbuscular mycorrhiza (AM) between plants and Glomeromycotina fungi is preceded by the exchange of chemical signals: fungal released Myc-factors, including chitoligosaccharides (CO) and lipo-chitooligosaccharides (LCO), activate plant symbiotic responses, while root exuded strigolactones stimulate hyphal branching and boost CO release. Furthermore, fungal signaling reinforcement through CO application was shown to promote AM development in Medicago truncatula, but the cellular and molecular bases of this effect remained unclear. • Here we focused on long-term M. truncatula responses to CO treatment, demonstrating its impact on the transcriptome of both mycorrhizal and non-mycorrhizal roots over several weeks and providing an insight into the mechanistic bases of the CO-dependent promotion of AM colonization. • CO treatment caused the long-lasting regulation of strigolactone biosynthesis and fungal accommodation related genes. This was mirrored by an increase in root didehydro-orobanchol content, and the promotion of accommodation responses to AM fungi in root epidermal cells. Lastly, an advanced down-regulation of AM symbiosis marker genes was observed at the latest time point in CO-treated plants, in line with an increased number of senescent arbuscules. • Overall, CO treatment triggered molecular, metabolic and cellular responses underpinning a protracted acceleration of AM development.

The utilization and molecular mechanism of arbuscular mycorrhizal symbiosis in vegetables

Vegetables are an important food and many of them can form AM (arbuscular mycorrhiza) symbiosis with AMF (AM fungi) that belongs to the sub-phylum Glomeromycota. The symbiosis with AM in vegetables enhances their tolerance to various stresses, such as low Pi (phosphate), salinity and soil-borne diseases. In the past decades, the molecular mechanism of AM symbiosis in vegetables has begun to emerge. Here, we review the key studies characterizing the molecular mechanism of AM symbiosis and highlights the huge potential of AMF in vegetable cultivation.

Arbuscular mycorrhizae reduce the response of important plant functional traits to drought and salinity. A meta-analysis study.

We aimed at exploring the plant functional traits whose responses to drought or salinity are altered by the presence of arbuscular mycorrhiza (AM). We performed a meta-analysis across 114 articles spanning 110 plant species or cultivars. We quantified the size effect of AM symbiosis on the stress response of several functional traits, using linear mixed model analysis (LMM). Correlation analysis between functional traits and total biomass responses to stresses were also performed through LMM. The literature search and further selection yielded seven functional traits, extracted from 114 laboratory studies, including 888 observations and 110 plant species/cultivars. Evidence for significant effects of predictor variables (type of stress, AM symbiosis and/or their interaction) on functional trait response were found for leaf area ratio (LAR), root mass fraction (RMF) and root-shoot (R:S) ratio. Our results provided evidence to accept the hypothesis that AM fungal inoculation may reduce the stress response of these plant functional traits by decreasing its magnitude. We also found a weak correlation between stress responses of these traits and total biomass variation. Although our literature search and data collection were intensive and our results robust, the scope of our conclusions is limited by the agronomical bias of plant species targeted by the meta-analysis. Further knowledge on non-cultivable plant species and better understanding of the mechanisms ruling resources allocation in plants would allow more generalised conclusions.

Arbuscular Mycorrhiza Symbiosis as a Factor of Asteraceae Species Invasion

Invasive weeds of the Asteraceae family are widespread in the world. Arbuscular mycorrhiza (AM) is one of the main factors contributing to the successful distribution of these species that is most clearly manifested in the subfamily Asteroideae. The benefits of plant-AMF symbiosis are most significant under unfavorable biotic and abiotic conditions. The specificity of the relationship between arbuscular mycorrhizal fungi (AMF) communities and plants and is determined at the presymbiotic stage. The AMF colonization level is higher in invasive species than in native ones, but AMF communities associated with Asteraceae invasive species are less diverse. AMF communities of Asteraceae invaders often include fewer common species (e.g., species belonging to Diversisporales). Invaders also reduce native AMF species richness in new areas. Arbuscular mycorrhizal fungi can form mycorrhizal networks that allow the redistribution of nutrients in plant communities. The most significant influence of AMF associated with invasive Asteraceae plants is seen in the formation of soil and rhizosphere microbiota, including the suppression of beneficial soil bacteria and fungi. This review could be useful in the development of practical recommendations for the use of AMF-based fertilizers.

Arbuscular mycorrhizal colonization leads to a change of hormone profile in micropropagated plantlet Satureja khuzistanica Jam

Phytohormones are supposed to contribute to the establishment of mutualistic Arbuscular mycorrhiza (AM) symbioses. However, their role in the acclimation of micropropagated plantlet inoculated with AM is still unknown. To address this question, we performed a hormone profiling during the acclimation of Satureja khuzistanica plantlets inoculated with Rhizoglomus fasciculatum. The levels of indoleacetic acid (IAA), methyl indole acetic acid, cis-zeatin, cis zeatin ribose, jasmonate, jasmonoyl isoleucine, salicylic acid, abscisic acid (ABA) were analyzed. Further, the relative gene expression of AOS (Allene oxide synthase) as a key enzyme of jasmonate biosynthesis, in either inoculated or non-inoculated micropropagated plantlets was evaluated during acclimation period. The concentrations of IAA and cis-zeatin increased in the plantlets inoculated by AM whereas the concentration of ABA decreased upon 60 days acclimation in the whole shoot of plantlets of S. khuzistanica. The relative expression of AOS gene resulted in an increase of isoleucine jasmonate, the bioactive form of jasmonate. Based on our results, IAA and cis-zeatin probably contribute to maintaining growth, and AM reduces transition stress by modifying ABA and jasmonate concentrations.

Identification of microRNAs responsive to arbuscular mycorrhizal fungi in Panicum virgatum (switchgrass)

BackgroundMicroRNAs (miRNAs) are important post-transcriptional regulators involved in the control of a range of processes, including symbiotic interactions in plants. MiRNA involvement in arbuscular mycorrhizae (AM) symbiosis has been mainly studied in model species, and our study is the first to analyze global miRNA expression in the roots of AM colonized switchgrass (Panicum virgatum), an emerging biofuel feedstock. AM symbiosis helps plants gain mineral nutrition from the soil and may enhance switchgrass biomass production on marginal lands. Our goals were to identify miRNAs and their corresponding target genes that are controlling AM symbiosis in switchgrass.ResultsThrough genome-wide analysis of next-generation miRNA sequencing reads generated from switchgrass roots, we identified 122 mature miRNAs, including 28 novel miRNAs. By comparing miRNA expression profiles of AM-inoculated and control switchgrass roots, we identified 15 AM-responsive miRNAs across lowland accession “Alamo”, upland accession “Dacotah”, and two upland/lowland F1 hybrids. We used degradome sequencing to identify target genes of the AM-responsive miRNAs revealing targets of miRNAs residing on both K and N subgenomes. Notably, genes involved in copper ion binding were targeted by downregulated miRNAs, while upregulated miRNAs mainly targeted GRAS family transcription factors.ConclusionThrough miRNA analysis and degradome sequencing, we revealed that both upland and lowland switchgrass genotypes as well as upland-lowland hybrids respond to AM by altering miRNA expression. We demonstrated complex GRAS transcription factor regulation by the miR171 family, with some miR171 family members being AM responsive while others remained static. Copper miRNA downregulation was common amongst the genotypes tested and we identified superoxide dismutases and laccases as targets, suggesting that these Cu-miRNAs are likely involved in ROS detoxification and lignin deposition, respectively. Other prominent targets of the Cu miRNAs were blue copper proteins. Overall, the potential effect of AM colonization on lignin deposition pathways in this biofuel crop highlights the importance of considering AM and miRNA in future biofuel crop development strategies.

Characterization of lncRNAs in mycorrhizal tomato and elucidation of the role of lncRNA69908 in disease resistance

Long noncoding RNAs (lncRNAs) have attracted widespread attention because of their meaningful roles in various plant biological processes. However, the potential functions of lncRNAs in the plant-beneficial microorganism interactions have not been fully explored. Arbuscular mycorrhiza (AM) symbiosis is accompanied by the systemic induction of defense responses in the host leaves. In the present study, we globally profiled lncRNA expression and explored their potential regulatory roles in AM fungi-inoculated tomato leaves. Among 851 differentially expressed lncRNAs, a novel lncRNA (lncRNA69908) that was significantly downregulated in the leaves of AM fungi inoculated tomato, affected tomato resistance after pathogen infection. One of the competing endogenous RNA networks, lncRNA69908-sly-miR319c, was verified by using a coexpression system. Silencing of lncRNA69908 or overexpression of sly-miR319c enhanced tomato resistance to Phytophthora infestans, whereas overexpression of lncRNA69908 decreased the reactive oxygen species scavenging. As above, we speculated that lncRNA69908 may be involved in mycorrhiza-induced defense responses. Our findings can broaden the knowledge on the potential regulatory roles of ncRNAs in AM symbiosis.

The Role of Medicago lupulina Interaction with Rhizophagus irregularis in the Determination of Root Metabolome at Early Stages of AM Symbiosis.

The nature of plant–fungi interaction at early stages of arbuscular mycorrhiza (AM) development is still a puzzling problem. To investigate the processes behind this interaction, we used the Medicago lupulina MlS-1 line that forms high-efficient AM symbiosis with Rhizophagus irregularis. AM fungus actively colonizes the root system of the host plant and contributes to the formation of effective AM as characterized by a high mycorrhizal growth response (MGR) in the host plant. The present study is aimed at distinguishing the alterations in the M. lupulina root metabolic profile as an indicative marker of effective symbiosis. We examined the root metabolome at the 14th and 24th day after sowing and inoculation (DAS) with low substrate phosphorus levels. A GS-MS analysis detected 316 metabolites. Results indicated that profiles of M. lupulina root metabolites differed from those in leaves previously detected. The roots contained fewer sugars and organic acids. Hence, compounds supporting the growth of mycorrhizal fungus (especially amino acids, specific lipids, and carbohydrates) accumulated, and their presence coincided with intensive development of AM structures. Mycorrhization determined the root metabolite profile to a greater extent than host plant development. The obtained data highlight the importance of active plant–fungi metabolic interaction at early stages of host plant development for the determination of symbiotic efficiency.

SlSPX1-SlPHR complexes mediate the suppression of arbuscular mycorrhizal symbiosis by phosphate repletion in tomato.

Forming mutualistic symbioses with arbuscular mycorrhizae (AMs) improves the acquisition of mineral nutrients for most terrestrial plants. However, the formation of AM symbiosis usually occurs under phosphate (Pi)-deficient conditions. Here, we identify SlSPX1 (SYG1 (suppressor of yeast GPA1)/Pho81(phosphate 81)/XPR1 (xenotropic and polytropic retrovirus receptor 1) as the major repressor of the AM symbiosis in tomato (Solanum lycopersicum) under phosphate-replete conditions. Loss of SlSPX1 function promotes direct Pi uptake and enhances AM colonization under phosphate-replete conditions. We determine that SlSPX1 integrates Pi signaling and AM symbiosis by directly interacting with a set of arbuscule-induced SlPHR proteins (SlPHR1, SlPHR4, SlPHR10, SlPHR11, and SlPHR12). The association with SlSPX1 represses the ability of SlPHR proteins to activate AM marker genes required for the arbuscular mycorrhizal symbiosis. SlPHR proteins exhibit functional redundancy, and no defective AM symbiosis was detected in the single mutant of SlPHR proteins. However, silencing SlPHR4 in the Slphr1 mutant background led to reduced AM colonization. Therefore, our results support the conclusion that SlSPX1-SlPHRs form a Pi-sensing module to coordinate the AM symbiosis under different Pi-availability conditions.

Multifarious and Interactive Roles of GRAS Transcription Factors During Arbuscular Mycorrhiza Development

Arbuscular mycorrhiza (AM) is a mutualistic symbiotic interaction between plant roots and AM fungi (AMF). This interaction is highly beneficial for plant growth, development and fitness, which has made AM symbiosis the focus of basic and applied research aimed at increasing plant productivity through sustainable agricultural practices. The creation of AM requires host root cells to undergo significant structural and functional modifications. Numerous studies of mycorrhizal plants have shown that extensive transcriptional changes are induced in the host during all stages of colonization. Advances have recently been made in identifying several plant transcription factors (TFs) that play a pivotal role in the transcriptional regulation of AM development, particularly those belonging to the GRAS TF family. There is now sufficient experimental evidence to suggest that GRAS TFs are capable to establish intra and interspecific interactions, forming a transcriptional regulatory complex that controls essential processes in the AM symbiosis. In this minireview, we discuss the integrative role of GRAS TFs in the regulation of the complex genetic re-programming determining AM symbiotic interactions. Particularly, research being done shows the relevance of GRAS TFs in the morphological and developmental changes required for the formation and turnover of arbuscules, the fungal structures where the bidirectional nutrient translocation occurs.

Karakteristik Mikoriza Arbuskula Tanaman Serai Wangi (Cymbopogon nardus L.) di Lapangan Ternaungi dan Tidak Ternaungi

Arbuscular mycorrhiza (AM) characteristics of citronella grass in the field have not been reported. This research aimed to study the AM characteristics of citronella grass grown in unshaded and shaded fields. The roots of citronella grass were collected from citronella grass plantations in Cianjur, West Java. The root samples were analyzed for AM structures, namely entry points, intercellular hyphae, arbuscules, and vesicles. The results showed that the citronella grass form AM colonization. The quality of root colonization differed between the two cultivation systems. The unshaded citronella grass had higher root colonization compared to shaded citronella grass. In the unshaded citronella grass, the number of arbuscules was 7 per cm of root length, whereas in the shaded citronella grass was 4 per cm of root length. The types of arbuscules observed were arum and intermediate. There were no differences in the number of entry points in the two cultivated systems, which was 3,5 entry points per cm of root length. The numbers of vesicles and internal hyphae in unshaded citronella grass were lower than that of in the shaded citronella grass. In the unshaded citronella grass, the number of vesicles and intracellular hyphae were 1,5 and 8,5 per cm root length, whereas in the shaded citronella grass were 3,5 and 11 per cm root length, respectively. Shading plants grown in the field were bamboo, banana, coffee, tea, and sugar palm. All the shading plants formed AM symbiosis with a colonization value of 7 to 30%. This research indicates that arbuscular mycorrhiza is an important component in the citronella grass cultivation in unshaded and shaded fields.
 
 Keywords: Arbuscule, entry point, intercellular hyphae, root colonization, vesicle