Nutrient and mycoremediation of a global menace 'arsenic': exploring the prospects of phosphorus and Serendipita indica-based mitigation strategies in rice and other crops.

KEY MESSAGE: Serendipita indica induced metabolic reprogramming in colonized plants complements phosphorus-management in improving their tolerance to arsenic stress on multifaceted biological fronts. Restoration of the anthropic damage done to our environment is inextricably linked to devising strategies that are not only economically sound but are self-renewing and ecologically conscious. The dilemma of heavy metal (HM) dietary ingestion, especially arsenic (As), faced by humans and animals alike, necessitates the exploitation of such technologies and the cultivation of healthy and abundant crops. The remarkable symbiotic alliance between plants and 'mycorrhizas' has evolved across eons, benefiting growth/yield aspects as well as imparting abiotic/biotic stress tolerance. The intricate interdependence of Serendipita indica (S. indica) and rice plant reportedly reduce As accumulation, accentuating the interest of microbiologists, agriculturists, and ecotoxicological scientists apropos of the remediation mechanisms of As in the soil-AMF-rice system. Nutrient management, particularly of phosphorus (P), is also praised for mitigating As phytotoxicity by deterring the uptake of As molecules due to the rhizospheric cationic competition. Taking into consideration the reasonable prospects of success in minimizing As acquisition by rice plants, this review focuses on the physiological, metabolic, and transcriptional alterations underlying S. indica symbiosis, recuperation of As stress together with nutritional management of P by gathering case studies and presenting successful paradigms. Weaving together a volume of literature, we assess the chemical forms of As and related transport pathways, discuss As-P-rice interaction and the significance of fungi in As toxicity mitigation, predominantly the role of mycorrhiza, as well as survey of the multifaceted impacts of S. indica on plants. A potential strategy for simultaneous S. indica + P administration in paddy fields is proposed, followed by future research orientation to expand theoretic comprehension and encourage field-based implementation.

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.

Endofungal bacteria and ectomycorrhizal fungi synergistically promote the absorption of organic phosphorus in Pinus massoniana.

Ectomycorrhizal fungi (ECMFs) that are involved in phosphorus mobilisation and turnover have limited ability to mineralise phytate alone. The endofungal bacteria in the ectomycorrhizal fruiting body may contribute to achieving this ecological function of ECMFs. We investigated the synergistic effect and mechanisms of endofungal bacteria and ECMF Suillus grevillea on phytate mineralisation. The results showed that soluble phosphorus content in the combined system of endofungal bacterium Cedecea lapagei and S. grevillea was 1.8 times higher than the sum of C. lapagei and S. grevillea alone treatment under the phytate mineralisation experiment. The S. grevillea could first chemotactically assist C. lapagei in adhering to the surface of S. grevillea. Then, the mineralisation of phytate was synergistically promoted by increasing the biomass of C. lapagei and the phosphatase and phytase activities of S. grevillea. The expression of genes related to chemotaxis, colonisation, and proliferation of C. lapagei and genes related to phosphatase and phytase activity of S. grevillea was also significantly upregulated. Furthermore, in the pot experiment, we verified that there might exist a ternary symbiotic system in the natural forest in which endofungal bacteria and ECMFs could synergistically promote phytate uptake in the plant Pinus massoniana via the ectomycorrhizal system.

Integrating phytoremediation and mycoremediation with biosurfactant-producing fungi for hydrocarbon removal and the potential production of secondary resources.

Treatment of petroleum-contaminated soil to a less toxic medium via physical and chemical treatment is too costly and requires posttreatment. This review focuses on the employment of phytoremediation and mycoremediation technologies in cleaning hydrocarbon-contaminated soil which is currently rare. It is considered environmentally beneficial and possibly cost-effective as it implements the synergistic interaction between plants and biosurfactant producing mycorrhiza to degrade hydrocarbon contaminants. This review also covers possible sources of hydrocarbon pollution in water and soil, toxicity effects, and current technologies for hydrocarbon removal and degradation. In addition to these problems, this review also discusses the challenges and opportunities of transforming the resultant treated sludge and treating plants into potential by-products for a higher quality of life for future generations.

Effect of foliar application of salicylic acid and mycorrhiza on quantitative and qualitative traits of maize.

An experiment was performed to investigate the effect of mycorrhizal symbiosis and foliar application of salicylic acid on quantitative and qualitative traits of maize during 2018 and 2019 in the research farm of Islamic Azad University, Chalous Branch. Split plot in a randomized complete block design with three replications was used. Experimental factors included mycorrhiza species of (G. mosseae), (G. geosporum) and (G. intraradices) at two levels (no consumption and consumption of mycorrhiza) and salicylic acid at two levels (no consumption and consumption of 1 mµ of salicylic acid). Results of interaction effects of mycorrhiza and salicylic acid on the measured traits revealed that the maximum 1000-grain weight, grain yield, biological yield, phosphorus, potassium, nitrogen percentage and yield of maize grain protein were observed in G. mosseae treatment under foliar application of salicylic acid. Foliar application of salicylic acid increases the root length and provides the necessary conditions for increasing water and nutrient uptake alongwith increase in photosynthesis and thus allocates more photosynthetic substance for development of reproductive organs. Hence, it increases maize grain weight and accordingly grain yield. In general, the results revealed that mycorrhiza and foliar application of salicylic acid increase growth indicators, yield and yield components. It also improved the quality traits of the maize plant. Based on results, the interaction effect of G. mosseae treatment and foliar application of salicylic acid yielded better results than other treatments. Mycorrhiza increases the number of grain in the ear, the number of rows in the ear, increases the plant's ability to absorb phosphorus, and the increase of mycorrhiza along with salicylic acid shows the maximum grain yield in maize. Finally, it can be concluded that the use of mycorrhiza and salicylic acid can be effective in increasing grain in the plant.

How does phosphorus influence Cd tolerance strategy in arbuscular mycorrhizal - Phragmites australis symbiotic system?

To clarify how phosphorus (P) influences arbuscular mycorrhizal fungi (AMF) interactions with host plants, we measured the effects of variation in environmental P levels and AMF colonization on photosynthesis, element absorption, ultrastructure, antioxidant capacity, and transcription mechanisms in Phragmites australis (P. australis) under cadmium (Cd) stress. AMF maintained photosynthetic stability, element balance, subcellular integrity and enhanced antioxidant capacity by upregulating antioxidant gene expression. Specifically, AMF overcame Cd-induced stomatal limitation, and mycorrhizal dependence peaked in the high Cd-moderate P treatment (156.08%). Antioxidants and compatible solutes responded to P-level changes: the primary driving forces of removing reactive oxygen species (ROS) and maintaining osmotic balance were superoxide dismutase, catalase, and sugars at limited P levels and total polyphenol, flavonoid, peroxidase, and proline at abundant P levels, we refer to this phenomenon as "functional link." AMF and phosphorus enhanced Cd tolerance in P. australis, but the regulation of AMF was P-dependent. Phosphorus prevented increases in total glutathione content and AMF-induced GSH/GSSG ratio (reduced to oxidized glutathione ratio) by inhibiting the expression of assimilatory sulfate reduction and glutathione reductase genes. The AMF-induced flavonoid synthesis pathway was regulated by P, and AMF activated Cd-tolerance mechanisms by inducing P-dependent signaling.

Exploring the synergistic effects of biochar and arbuscular mycorrhizal fungi on phosphorus acquisition in tomato plants by using gene expression analyses.

Arbuscular mycorrhizal (AM) fungi are symbiotic organisms that contribute significantly to plant mineral nutrition, mainly phosphate. However, their benefits are constricted by the availability of phosphate in the soil, and thus they are recalcitrant as amendment in highly fertilized soils. Biochars are by-products of the pyrolysis of biomass in the absence of oxygen. They can improve soil properties and act as a source of nutrients for plants. However, depending on their origin, the final composition of biochars is extremely variable and thus, their efficiency unpredictable. In order to gain mechanistic insights into how the combined application of biochars and AM fungi contribute to plant phosphate nutrition and growth, we used gene expression analyses of key symbiotic marker genes. We compared for this analysis two biochars originated from very different feedstocks (chicken manure and wheat straw) on tomato plants with or without the AM fungus Rhizophagus irregularis. Our results show that the synergy between AM fungi and biochars as P biofertilizers is greatly governed by the origin of the biochar that determines the speed at which phosphate is released to the soil and absorbed by the plant. Thus, chicken manure biochar quickly impacted on plant growth by readily releasing P, but it turned out detrimental for symbiosis formation, decreasing colonization levels and expression of key symbiotic plant marker genes such as SlPT4 or SlFatM. In contrast, wheat straw biochar was inferior at improving plant growth but stimulated the establishment of the symbiosis, producing plants with the same concentration of phosphate as those with the chicken manure. Taken together, slow P releasing biochars from plant residues appears to be a more promising amendment for long terms experiments in which biofertilizers such as AM fungi are considered. Furthermore, our results indicate that implementing plant transcriptomic analyses might help to mechanistically dissect and better understand the effects of biochars on plant growth in different scenarios.

A trade-off between space exploration and mobilization of organic phosphorus through associated microbiomes enables niche differentiation of arbuscular mycorrhizal fungi on the same root.

Ecology seeks to explain species coexistence, but experimental tests of mechanisms for coexistence are difficult to conduct. We synthesized an arbuscular mycorrhizal (AM) fungal community with three fungal species that differed in their capacity of foraging for orthophosphate (P) due to differences in soil exploration. We tested whether AM fungal species-specific hyphosphere bacterial assemblages recruited by hyphal exudates enabled differentiation among the fungi in the capacity of mobilizing soil organic P (Po). We found that the less efficient space explorer, Gigaspora margarita, obtained less 13C from the plant, whereas it had higher efficiencies in Po mobilization and alkaline phosphatase (AlPase) production per unit C than the two efficient space explorers, Rhizophagusintraradices and Funneliformis mosseae. Each AM fungus was associated with a distinct alp gene harboring bacterial assemblage, and the alp gene abundance and Po preference of the microbiome associated with the less efficient space explorer were higher than those of the two other species. We conclude that the traits of AM fungal associated bacterial consortia cause niche differentiation. The trade-off between foraging ability and the ability to recruit effective Po mobilizing microbiomes is a mechanism that allows co-existence of AM fungal species in a single plant root and surrounding soil habitat.

Hyphosphere interactions between Rhizophagus irregularis and Rahnella aquatilis promote carbon-phosphorus exchange at the peri-arbuscular space in Medicago truncatula.

Arbuscular mycorrhizal (AM) fungi form a continuum between roots and soil. One end of this continuum is comprised of the highly intimate plant-fungus interface with intracellular organelles for nutrient exchange, while on the other end the fungus interacts with bacteria to compensate for the AM fungus' inability to take up organic nutrients from soil. How both interfaces communicate in this highly complex tripartite mutualism is widely unknown. Here, the effects of phosphate-solubilizing bacteria (PSB) Rahnella aquatilis dwelling at the surface of the extraradical hyphae of Rhizophagus irregularis was analysed based on the expression of genes involved in C-P exchange at the peri-arbuscular space (PAS) in Medicago truncatula. The interaction between AM fungus and PSB resulted in an increase in uptake and transport of Pi along the extraradical hyphae and its transfer from AM fungus to plant. In return, this was remunerated by a transfer of C from plant to AM fungus, improving the C-P exchange at the PAS. These results demonstrated that a microorganism (i.e., a PSB) developing at the hyphosphere interface can affect the C-P exchange at the PAS between plant and AM fungus, suggesting a fine-tuned communication operated between three organisms via two distantly connected interfaces.

Arbuscular mycorrhizal fungi enhance plant phosphorus uptake through stimulating hyphosphere soil microbiome functional profiles for phosphorus turnover.

The extraradical hyphae of arbuscular mycorrhizal (AM) fungi are colonized by different bacteria in natural and agricultural systems, but the mechanisms by which AM fungi interact with the hyphosphere soil microbiome and influence soil organic phosphorus (P) mobilization remain unclear. We grew Medicago in two-compartment microcosms, inoculated with Rhizophagus irregularis, or not, in the root compartment and set up P treatments (without P, with P addition as KH2 PO4 or nonsoluble phytate) in the hyphal compartment. We studied the processes of soil P turnover and characterized the microbiome functional profiles for P turnover in the hyphosphere soil by metagenomic sequencing. Compared with the bulk soil, the hyphosphere soil of R. irregularis was inhabited by a specific bacterial community and their functional profiles for P turnover was stimulated. At the species level, the shift in hyphosphere soil microbiome was characterized by the recruitment of the genome bin2.39 harbouring both gcd and phoD genes and genome bin2.97 harbouring the phoD gene, which synergistically drove nonsoluble phytate mobilization in the hyphosphere soil. Our results suggest that AM fungi recruits a specific hyphosphere soil microbiome and stimulated their functional profiles for P turnover to enhance utilization of phytate.

A core microbiome in the hyphosphere of arbuscular mycorrhizal fungi has functional significance in organic phosphorus mineralization.

The mycorrhizal pathway is an important phosphorus (P) uptake pathway for more than two-thirds of land plants. The arbuscular mycorrhizal (AM) fungi-associated hyphosphere microbiome has been considered as the second genome of mycorrhizal P uptake pathway and functionality in mobilizing soil organic P (Po). However, whether there is a core microbiome in the hyphosphere and how this is implicated in mining soil Po are less understood. We established on-site field trials located in humid, semiarid, and arid zones and a microcosm experiment in a glasshouse with specific AM fungi and varying soil types to answer the above questions. The hyphosphere microbiome of AM fungi enhanced soil phosphatase activity and promoted Po mineralization in all sites. Although the assemblage of hyphosphere microbiomes identified in three climate zones was mediated by environmental factors, we detected a core set in three sites and the subsequent microcosm experiment. The core members were co-enriched in the hyphosphere and dominated by Alphaproteobacteria, Actinobacteria, and Gammaproteobacteria. Moreover, these core bacterial members aggregate into stable guilds that contributed to phosphatase activity. The core hyphosphere microbiome is taxonomically conserved and provides functions, with respect to the mineralization of Po, that AM fungi lack.

Flammulina velutipes Mycorrhizae Attenuate High Fat Diet-Induced Lipid Disorder, Oxidative Stress and Inflammation in the Liver and Perirenal Adipose Tissue of Mice.

Flammulina velutipes (FV) is edible mushroom that has nutritional and medicinal values. FV mycorrhizae, the by-products of FV, are an abundant source and receive less attention. The objective of this study was to investigate the composition of FV mycorrhizae, and its effects on high fat diet (HFD)-induced lipid disorder, oxidative stress, and inflammatory cytokines, both in the liver and perirenal adipose tissue (PAT) of mice. The results showed that FV mycorrhizae contain abundant trace elements, polysaccharide, amino acids and derivatives, and organic compounds. It was found that 4% FV mycorrhizae (HFDFV) supplementation decreased HFD-induced liver weight and triglyceride (TG) in the plasma, liver and PAT, altered plasma and hepatic fatty acids profiles, promoted gene expression involved in lipid hydrolysis, fatty acid transportation and ß-oxidation in the liver and reduced lipid synthesis in the liver and PAT. HFDFV attenuated HFD-induced oxidative stress and pro-inflammatory cytokine by increasing GSH/GSSG, and decreasing levels of MDA and IL6 both in the liver and PAT, while it differentially regulated gene expression of IL1ß, IL6, and CCL2 in liver and PAT. The results indicated that FV mycorrhizae are effective to attenuate HFD-induced lipid disorder, oxidative stress and inflammation in the liver and PAT, indicating their promising constituents for functional foods and herbal medicine.

The mechanism of arbuscular mycorrhizal enhancing cadmium uptake in Phragmites australis depends on the phosphorus concentration.

Arbuscular mycorrhizal fungi (AMF) is a vital strategy to enhance the phytoremediation of cadmium (Cd) pollution. However, the function of AMF was influenced by phosphorus (P) concentration. To reveal the effect of AMF on the Cd accumulation of host plants under different P concentrations and how the AMF and P interact, this study comparatively analyzed the regulatory effects of AMF on the Cd response, extraction, and transportation processes of Phragmites australis (P. australis) under different P levels, and explored its physiological, biochemical and molecular biological mechanisms. The study showed that AMF could induce different growth allocation strategies in response to Cd stress. Moreover, AMF promoted plant Cd tolerance and detoxification by enhancing P uptake, Cd passivation, Cd retention in the cell wall, and functional group modulation. Under P starvation treatments, AMF promoted Cd uptake by inducing Cd to enter the iron pathway, increased the transport coefficient by 493.39%, and retained Cd in stems. However, these effects disappeared following the addition of P. Additionally, AMF up-regulated the expression of ZIP, ZIP, and NRAMP genes to promote cadmium uptake at low, medium, and high phosphorus levels, respectively. Thus, the Cd response mechanism of the AMF-P. australis symbiotic system was P dose-dependent.

The fine-tuning of mycorrhizal pathway in sorghum depends on both nitrogen-phosphorus availability and the identity of the fungal partner.

Sorghum is an important worldwide source of food, feed and fibres. Like most plants, it forms mutualistic symbioses with arbuscular mycorrhizal fungi (AMF), but the nutritional basis of mycorrhiza-responsiveness is largely unknown. Here, we investigated the transcriptional and physiological responses of sorghum to two different AMF species, Rhizophagus irregularis and Funneliformis mosseae, under 16 different conditions of nitrogen (N) and phosphorus (P) supply. Our experiment reveals fine-scale differences between two AMF species in the nutritional interactions with sorghum plants. Physiological and gene expression patterns (ammonium transporters: AMT; phosphate transporters: PHT) indicate the existence of generalist or specialist mycorrhizal pathway. While R. irregularis switched on the mycorrhizal pathway independently of the plant nutritional status, F. mosseae influenced the mycorrhizal pathway depending on the N-to-P plant ratio and soil supply. The differences between both AMF species suggest some AMT and PHT as ideal candidates to develop markers for improving efficiency of nutrient acquisition in sorghum under P and N limitation, and for the selection of plant genotypes.

Temporal complementarity between roots and mycorrhizal fungi drives wheat nitrogen use efficiency.

Improving nitrogen (N) use efficiency (NUE) to reduce the application of N fertilisers in a way that benefits the environment and reduces farmers' costs is an ongoing objective for sustainable wheat production. However, whether and how arbuscular mycorrhizal fungi (AMF) affect NUE in wheat is still not well explored. Three independent but complementary experiments were conducted to decipher the contribution of roots and AMF to the N uptake and utilisation efficiency in wheat. We show a temporal complementarity pattern between roots and AMF in shaping NUE of wheat. Pre-anthesis N uptake efficiency mainly depends on root functional traits, but the efficiency to utilise the N taken up during pre-anthesis for producing grains (EN,g ) is strongly affected by AMF, which might increase the uptake of phosphorus and thereby improve photosynthetic carbon assimilation. Root association with AMF reduced the N remobilisation efficiency in varieties with high EN,g ; whilst the overall grain N concentration increased, due to a large improvement in post-anthesis N uptake supported by AMF and/or other microbes. The findings provide evidence for the importance of managing AMF in agroecosystems, and an opportunity to tackle the contradiction between maximising grain yield and protein concentration in wheat breeding.

Combination of biochar and AMF promotes phosphorus utilization by stimulating rhizosphere microbial co-occurrence networks and lipid metabolites of Phragmites.

Agricultural biochar and arbuscular mycorrhizal fungi were used to promote the growth of Phragmites in the structural damaged and nutritional imbalanced littoral zone soils. Wheat straw biochar played a significant role in improving soil porosity and supplementing available phosphorus to 79.20 ± 3.20 mg/kg, compared with CK at 17.50 ± 0.88 mg/kg. The addition of Diversispora versiformis improved the plant net photosynthetic rate reaching up to 25.66 ± 0.65 µmol·m-2·s-1, which was 36.60 % higher than CK. The combination of biochar and fungi contributed to the whole plant dry weight biomass of 32.30 % and 234.00 % higher than the single biochar or AMF amendment groups, respectively. Meanwhile, the analysis of microbial co-occurrence networks showed the most relevant networks node species were mainly Talaromyces, Chaetomiacea and Gemmatimonadetes etc. Root lipid metabolite of Glycerophospholines further proved that phosphorus utilization was also enhanced endogenously in the rhizosphere soil. These results indicate that the combination of biochar and arbuscular mycorrhizal fungi play synergic role in enhancing phosphorus utilization endogenously and exogenously.

Laser Light Treatment Improves the Mineral Composition, Essential Oil Production and Antimicrobial Activity of Mycorrhizal Treated Pelargoniumgraveolens.

Pelargonium graveolens, rose-scented geranium, is commonly used in the perfume industry. P. graveolens is enriched with essential oils, phenolics, flavonoids, which account for its tremendous biological activities. Laser light treatment and arbuscular mycorrhizal fungi (AMF) inoculation can further enhance the phytochemical content in a significant manner. In this study, we aimed to explore the synergistic impact of these two factors on P. graveolens. For this, we used four groups of surface-sterilized seeds: (1) control group1 (non-irradiated; non-colonized group); (2) control group2 (mycorrhizal colonized group); (3) helium-neon (He-Ne) laser-irradiated group; (4) mycorrhizal colonization coupled with He-Ne laser-irradiation group. Treated seeds were growing in artificial soil inculcated with Rhizophagus irregularis MUCL 41833, in a climate-controlled chamber. After 6 weeks, P. graveolens plants were checked for their phytochemical content and antibacterial potential. Laser light application improved the mycorrhizal colonization in P. graveolens plants which subsequently increased biomass accumulation, minerals uptake, and biological value of P. graveolens. The increase in the biological value was evident by the increase in the essential oils production. The concomitant application of laser light and mycorrhizal colonization also boosted the antimicrobial activity of P. graveolens. These results suggest that AMF co-treatment with laser light could be used as a promising approach to enhance the metabolic content and yield of P. graveolens for industrial and pharmaceutical use.

Selenium uptake and accumulation in winter wheat as affected by level of phosphate application and arbuscular mycorrhizal fungi.

Selenium (Se) is an advantageous element to crops. However, the influence of arbuscular mycorrhizal fungi (AMF), phosphate (P) and selenite in soil on Se uptake by winter wheat remain elusive. Pot trials were carried out including seven levels of P (0, 12.5, 25, 50, 100, 200 or 400 mg kg-1) and non-mycorrhizal inoculation (NM), inoculation of Funneliformis mosseae (F.m) or Glomus versiforme (G.v). The present results found that grain phosphorus concentration increased with increase of P level from 0 to 100 mg kg-1 and then tended to plateau, while grain Se concentration decreased with the level of P from 0 to 400 mg kg-1. Based on mathematical modeling, inoculation of F.m or G.v dramatically improved grain Se concentration by 16.90% or 12.53% under the lower level of P (48.76 mg kg-1). Furthermore, partial least squares path modeling (PLS-PM) identified that both up-regulated of the expression of AMF-inducible phosphate transporter and improved Se bioavailability in rhizosphere soil contributed to enhancing plant Se concentration under P levels ≤ 100 mg kg-1. The present study demonstrated that AMF combined with 48.76 mg kg-1 P applied in soil can not only achieve high grain yield, but also fully exploit the biological potential of Se uptake in wheat.

Phosphorus Acquisition and Utilization in Plants.

Tremendous progress has been made on molecular aspects of plant phosphorus (P) nutrition, often without heeding information provided by soil scientists, ecophysiologists, and crop physiologists. This review suggests ways to integrate information from different disciplines. When soil P availability is very low, P-mobilizing strategies are more effective than mycorrhizal strategies. Soil parameters largely determine how much P roots can acquire from P-impoverished soil, and kinetic properties of P transporters are less important. Changes in the expression of P transporters avoid P toxicity. Plants vary widely in photosynthetic P-use efficiency, photosynthesis per unit leaf P. The challenge is to discover what the trade-offs are of different patterns of investment in P fractions. Less investment may save P, but are costs incurred? Are these costs acceptable for crops? These questions can be resolved only by the concerted action of scientists working at both molecular and physiological levels, rather than pursuing these problems independently.

Two-component system in Rahnella aquatilis is impacted by the hyphosphere of the arbuscular mycorrhizal fungus Rhizophagus irregularis.

Two-component systems (TCS) are ubiquitous among bacteria, playing key roles in signalling events. However, to what extent the TCS of Rahnella aquatilis (a Phosphate solubilizing bacteria) is influenced by the hyphosphere of the arbuscular mycorrhizal (AM) fungus Rhizophagus irregularis is totally unknown. Here, the expression of 16 genes encoding the TCS of R. aquatilis (i.e. involved in carbon-sensing and nutrient-sensing) and of eight genes regulated by the PhoR TCS (i.e. involved in inorganic and organic phosphorus mobilization) were analysed at regular intervals in presence of hyphae of R. irregularis. The study was conducted under in vitro culture conditions with phytate as the unique source of phosphorus. In presence of the AM fungus, the expression of TCS genes involved in carbon-sensing and nutrient-sensing were stimulated. Only, BaeS at 30 and 120 min, and BaeR at 60 min were inhibited. In addition, the PhoR TCS stimulated the expression of genes encoding phosphatase but inhibited the expression of genes involved in gluconic acid production. As the mechanism of coupling environmental changes with cellular physiological changes, TCS plays a pivotal role in regulating specific gene expression in R. aquatilis, recognizing environmental signals. More importantly, TCS genes may regulate bacteria response to hyphal carbon to mobilize phosphorus efficiently in the hyphosphere.