Novel soil-less potting mixes for the mycorrhization of Quercus pubescens Willd. seedlings with Tuber melanosporum Vittad.

Truffle cultivation has quickly grown in Europe and elsewhere as a consequence of the increase in the demand of the market. Thus the optimization of the protocols for the production of elite mycorrhized plants are also needed, keeping in consideration the economic and environmental sustainability. The suitability of two compost-based potting mixes to produce Quercus pubescens Willd. plants mycorrhized with the black Périgord truffle T. melanosporum Vittad. was tested as an alternative to the traditional potting mix used. The effects on mycorrhizal development and the morphometric assessment of the root and shoot system of the Q. pubescens seedlings were investigated eight months after the spore slurry inoculation in a glasshouse experiment. From the results obtained, the compost mix containing green organic residues from pruning and mowing (Mix 2) achieved better performance than the control and the potting mix based on composted municipal organic wastes, showing significantly higher mycorrhization percentage, root length, number of root tips, and root forks. In conclusion, a potting mix containing recycled green organic matter, which is readily available, cheap, and environmentally sustainable, can offer excellent mycorrhization performances and may be included in the mycorrhization process of downy oak seedlings with T. melanosporum under controlled conditions.

Variability in mycorrhizal status of plant species is much larger within than between plots in grassland and coastal habitats.

Community-level studies linking plant mycorrhizal status to environment usually do not account for within-plot mycorrhizal status variability; thus, patterns of plant mycorrhizal status diversity are largely unknown. Here, we assessed the relative importance of within- and between-plot variability components in mycorrhizal status and examined how plant mycorrhizal status diversity is related to soil nutrient availability. We hypothesised larger between-plot variability in mycorrhizal status and higher plant mycorrhizal status diversity in P-poor soils. To test these hypotheses, we used plant phylogenies, vegetation, soil and plant mycorrhizal status data from Czech semi-natural grasslands and Scottish coastal habitats. We divided plant mycorrhizal status diversity into divergence and evenness and tested their relations to soil P, K, Ca and Mg. Within-plot variability component of mycorrhizal status was always, on average, at least 2.2 times larger than between-plot variability in our datasets. Plant mycorrhizal status divergence was positively related to Ca (in both datasets) and Mg (only in grasslands and when accounting for phylogeny). In grasslands, the relationship between Mg and plant mycorrhizal status evenness was negative when accounting for phylogeny, while it was positive when not accounting for phylogeny. Plant mycorrhizal status diversity was not linked to P and its relation to K was inconsistent. Our results suggest that high Ca in the soil can promote coexistence of mycorrhizal, facultatively mycorrhizal and non-mycorrhizal plant species. We encourage future studies to also focus on within-plot variability in mycorrhizal status, because it appears to be highly relevant in herbaceous systems.

Cultivation of arbuscular mycorrhizal Broussonetia papyrifera seedlings by planting the mycorrhizal nurse plant downwards.

Plant mycorrhization can be achieved by transplanting new seedlings with mycorrhizal nurse plants; however, this method inevitably induces plant interactions. Transplanting nurse plants downwards may prevent light competition among new seedlings and nurse plants in the same pot. We hypothesized that seedling mycorrhization via mycorrhizal provision from plants planted downwards would be a feasible and efficient strategy. We used seedlings cultivated for 6 months after inoculation with arbuscular mycorrhizal fungi (AMF) as nurse plants, and seedlings cultivated for 1 month without AMF as recipient plants, transplanting one nurse plant and three recipient plants together in one pot. We compared two approaches for cultivating mycorrhizal Broussonetia papyrifera seedlings: planting mycorrhizal nurse plants upwards (M-NU) and downwards (M-ND). We also planted non-mycorrhizal nurse plants upwards (NM-NU) and downwards (NM-ND) as controls. We analyzed growth parameters and the mycorrhizal colonization status of recipient plants at 45, 60, and 75 days after planting (DAP). As expected, the plant growth, gas exchange, and root morphological parameters of recipient plants with mycorrhizal nurse plants were higher than those of recipient plants with non-mycorrhizal nurse plants at 60 and 75 DAP. Furthermore, the AMF colonization status and physiological growth status of M-ND recipient plants were improved compared with M-NU recipient plants. Our results demonstrate that inducing seedling mycorrhization by planting mycorrhizal nurse plants downwards is a feasible strategy for achieving AMF symbiosis while mitigating negative interactions among plants.

Mycorrhizal status is a poor predictor of the distribution of herbaceous species along the gradient of soil nutrient availability in coastal and grassland habitats.

Plant mycorrhizal status (a trait indicating the ability to form mycorrhizas) can be a useful plant trait for predicting changes in vegetation influenced by increased fertility. Mycorrhizal fungi enhance nutrient uptake and are expected to provide a competitive advantage for plants growing in nutrient-poor soils; while in nutrient-rich soils, mycorrhizal symbiosis may be disadvantageous. Some studies in natural systems have shown that mycorrhizal plants can be more frequent in P and N-poor soils (low nutrient availability) or Ca and Mg-high (high pH) soils, but empirical support is still not clear. Using vegetation and soil data from Scottish coastal habitats, and Latvian and Czech grasslands, we examined whether there is a link between plant mycorrhizal status and plant-available P, N, Ca and Mg. We performed the max test analysis (to examine the central tendency) and a combination of quantile regression and meta-analysis (to examine tendencies in different quantiles) on both community and plant species data combined with plant phylogenies. We consistently found no changes in mycorrhizal status at the community and species levels along the gradients of plant-available P, N, Ca and Mg in the central tendency and in almost all quantiles across all datasets. Thus, we found no support for the hypotheses that herbaceous species which are able to form mycorrhizas are more frequent in nutrient-poor and high pH environments. Obligatory, facultatively and non-mycorrhizal herbaceous species appear to assemble randomly along the gradients of nutrient availability in several European herbaceous habitats, suggesting that all these strategies perform similarly under non-extreme soil nutrient conditions.

Transcriptome analysis of the differential effect of the NADPH oxidase gene RbohB in Phaseolus vulgaris roots following Rhizobium tropici and Rhizophagus irregularis inoculation.

BACKGROUND: Reactive oxygen species (ROS) are generated by NADPH oxidases known as respiratory burst oxidase homologs (RBOHs) in plants. ROS regulate various cellular processes, including the mutualistic interactions between legumes and nitrogen-fixing bacteria or arbuscular mycorrhizal (AM) fungi. Rboh is a multigene family comprising nine members (RbohA-I) in common bean (Phaseolus vulgaris). The RNA interference-mediated silencing of RbohB (PvRbohB-RNAi) in this species diminished its ROS production and greatly impaired nodulation. By contrast, the PvRbohB-RNAi transgenic roots showed early hyphal root colonization with enlarged fungal hypopodia; therefore, we proposed that PvRbohB positively regulates rhizobial infection (Rhizobium tropici) and inhibits AM colonization by Rhizophagus irregularis in P. vulgaris. RESULTS: To corroborate this hypothesis, an RNA-Seq transcriptomic analysis was performed to identify the differentially expressed genes in the PvRbohB-RNAi roots inoculated with Rhizobium tropici or Rhizophagus irregularis. We found that, in the early stages, root nodule symbioses generated larger changes of the transcriptome than did AM symbioses in P. vulgaris. Genes related to ROS homeostasis and cell wall flexibility were markedly upregulated in the early stages of rhizobial colonization, but not during AM colonization. Compared with AM colonization, the rhizobia induced the expression of a greater number of genes encoding enzymes involved in the metabolism of auxins, cytokinins, and ethylene, which were typically repressed in the PvRbohB-RNAi roots. CONCLUSIONS: Our research provides substantial insights into the genetic interaction networks in the early stages of rhizobia and AM symbioses with P. vulgaris, as well as the differential roles that RbohB plays in processes related to ROS scavenging, cell wall remodeling, and phytohormone homeostasis during nodulation and mycorrhization in this legume.

Earthworms and mycorrhization increase copper phytoextraction by Canavalia ensiformis in sandy soil.

Phytoremediation is an alternative for remediating soil contamination by copper, and its efficiency has been shown to increase when arbuscular mycorrhizal fungi (AMF) and earthworms are separately inoculated into the soil. This study evaluated the isolated and combined effects of inoculating earthworms and arbuscular mycorrhizal fungi into a sandy soil on copper phytoremediation by Canavalia ensiformis. The plants were grown in a greenhouse in soil contaminated with 100 mg Cu kg-1 with and without being inoculated with the arbuscular mycorrhizal fungus Rhizoglomus clarum and the earthworm Eisenia andrei. The availabilities of solid-phase Cu and other nutrients in the soil solution and plant growth were evaluated along with Cu phytotoxicity based on photochemical efficiency and oxidative stress enzyme activity. Accumulation of Cu and other nutrients in the shoots and roots; mycorrhizal colonization, nodulation, and reproduction; and Cu accumulation in the earthworm tissues were also evaluated. The copper caused photosynthetic and biochemical damage that reduced the shoot dry weight by 44% and the root dry weight by 29%. However, the arbuscular mycorrhizal fungus alleviated the Cu toxicity to the plant and increased the shoot dry weight by 81% in the contaminated soil. The earthworms increased the Cu uptake and translocation to the shoot by 31%. The combined presence of the arbuscular mycorrhizal fungus and earthworms in the contaminated soil increased the growth and Cu content of the aerial plant tissues, yielding a 200% increase in Cu accumulation (metal content × biomass) in the C. ensiformis shoots. Combined inoculation with earthworms and arbuscular mycorrhizal fungi increased copper phytoextraction by Canavalia ensiformis in a sandy soil.

Changes in the allocation of endogenous strigolactone improve plant biomass production on phosphate-poor soils.

Strigolactones (SLs) are carotenoid-derived phytohormones shaping plant architecture and inducing the symbiosis with endomycorrhizal fungi. In Petunia hybrida, SL transport within the plant and towards the rhizosphere is driven by the ABCG-class protein PDR1. PDR1 expression is regulated by phytohormones and by the soil phosphate abundance, and thus SL transport integrates plant development with nutrient conditions. We overexpressed PDR1 (PDR1 OE) to investigate whether increased endogenous SL transport is sufficient to improve plant nutrition and productivity. Phosphorus quantification and nondestructive X-ray computed tomography were applied. Morphological and gene expression changes were quantified at cellular and whole tissue levels via time-lapse microscopy and quantitative PCR. PDR1 OE significantly enhanced phosphate uptake and plant biomass production on phosphate-poor soils. PDR1 OE plants showed increased lateral root formation, extended root hair elongation, faster mycorrhization and reduced leaf senescence. PDR1 overexpression allowed considerable SL biosynthesis by releasing SL biosynthetic genes from an SL-dependent negative feedback. The increased endogenous SL transport/biosynthesis in PDR1 OE plants is a powerful tool to improve plant growth on phosphate-poor soils. We propose PDR1 as an as yet unexplored trait to be investigated for crop production. The overexpression of PDR1 is a valuable strategy to investigate SL functions and transport routes.

A role for the mevalonate pathway in early plant symbiotic signaling.

Rhizobia and arbuscular mycorrhizal fungi produce signals that are perceived by host legume receptors at the plasma membrane and trigger sustained oscillations of the nuclear and perinuclear Ca(2+) concentration (Ca(2+) spiking), which in turn leads to gene expression and downstream symbiotic responses. The activation of Ca(2+) spiking requires the plasma membrane-localized receptor-like kinase Does not Make Infections 2 (DMI2) as well as the nuclear cation channel DMI1. A key enzyme regulating the mevalonate (MVA) pathway, 3-Hydroxy-3-Methylglutaryl CoA Reductase 1 (HMGR1), interacts with DMI2 and is required for the legume-rhizobium symbiosis. Here, we show that HMGR1 is required to initiate Ca(2+) spiking and symbiotic gene expression in Medicago truncatula roots in response to rhizobial and arbuscular mycorrhizal fungal signals. Furthermore, MVA, the direct product of HMGR1 activity, is sufficient to induce nuclear-associated Ca(2+) spiking and symbiotic gene expression in both wild-type plants and dmi2 mutants, but interestingly not in dmi1 mutants. Finally, MVA induced Ca(2+) spiking in Human Embryonic Kidney 293 cells expressing DMI1. This demonstrates that the nuclear cation channel DMI1 is sufficient to support MVA-induced Ca(2+) spiking in this heterologous system.

Ectomycorrhizae formed by three Japanese truffle species (Tuber japonicum, T. longispinosum, and T. himalayense) on indigenous oak and pine species.

Modern truffle cultivation systems started in Europe in the early 1970s and are now successfully used for several European truffles throughout the world. However, systems for indigenous novel truffle species need to be developed in several regions, especially where truffle cultivation has not been attempted so far, such as in Japan. Recently, two new and one known truffle species that are expected to be edible were reported from Japan: Tuber japonicum, T. longispinosum, and T. himalayense. Here, we conducted mycorrhization trials between these three truffle species and four native tree species in Japan (Quercus acutissima, Q. phillyraeoides, Q. serrata, and Pinus densiflora) using spore suspension and trap-plant seedling techniques under axenic pot culture conditions to understand their potential host ranges and mycorrhizal morphologies and to determine whether these inoculation methods are applicable for mycorrhization of Japanese truffles with native host plants. Of the 12 combinations, nine were successful for mycorrhization, including both oak and pine trees. The T. japonicum mycorrhiza was characterized by short, needle-shaped cystidia without septa, whereas the two black truffles, T. longispinosum and T. himalayense, were indistinguishable from each other because they shared the same morphological and anatomical characters such as brownish, long cystidia with right angle ramification. These features were similar to related black truffle species. The results of the present study indicate that the inoculation method used for European truffles can also be applied for mycorrhization between Japanese truffle species and compatible native pine and/or oak hosts in Japan.

Transcriptional induction of two phosphate transporter 1 genes and enhanced root branching in grape plants inoculated with Funneliformis mosseae.

We investigated the effects of the arbuscular mycorrhizal fungus (AMF) Funneliformis mosseae on the growth and root architecture of plantlets of the grape rootstock 41B MGt under hydroponic conditions, and analyzed the concomitant expression of putative mycorrhizal-specific phosphate transporter 1 (PHT1) genes. In vitro propagated plantlets were acclimatized to ex vitro culture before AMF inoculation and grown under low phosphate (Pi) nutrition conditions during 6 weeks. Grape roots could be efficiently colonized by F. mosseae in this culture system, as shown by high mycorrhization frequency and intensity. The presence of many arbuscules in the cortex was coupled with high-level expression of two PHT1 genes in grape roots. These two very similar genes, named VvPht1-1 and VvPht1-2, present P1BS and MYCS regulatory motifs in their promoter, consistent with a specific role in the mycorrhizal pathway of Pi uptake. Although AMF inoculation significantly increased shoot growth, no effect on root biomass was observed. However, inoculated grapes exhibited an enhanced branched root system compared with non-inoculated controls, with a twofold higher number of tips and a higher proportion of fine roots usually involved in nutrient uptake from the soil. Taken together, these results suggest that root colonization by F. mosseae improved grape growth by favoring the uptake of Pi from the substrate via VvPht1-1 and VvPht1-2 high-level expression.

[Nutrient transfer and growth of Pinus greggii Engelm. inoculated with edible ectomycorrhizal mushrooms in two substrates]. / Transferencia de nutrientes y crecimiento de Pinus greggii Engelm. inoculado con hongos comestibles ectomicorrícicos en dos sustratos.

An ectomycorrhiza is a mutualistic symbiosis of paramount importance in forestry and tree production. One of the selection criteria of ectomycorrhizal fungi that has currently gained importance is their edibility due to the economic, ecological and cultural relevance of edible ectomycorrhizal mushrooms as a non-timber forest product. The effect of the inoculation with three edible ectomycorrhizal mushrooms: Laccaria laccata, Laccaria bicolor y Hebeloma leucosarx, which are widely sold in Mexico, on the growth and nutrient contents of Pinus greggii grown in an experimental substrate and a commercial substrate enriched with a slow-release fertilizer, was evaluated. Two years after sowing, differences in terms of shoot and root biomass and macro and micronutrient contents between inoculated and non-inoculated plants, were recorded independently of the fungal species and the substrate. Despite the fact that plants grown in the commercial substrate had higher growth and nutrient contents, their ectomycorrhizal colonization percentages were smaller than those of the plants grown in the experimental substrate. The differences in the nutrient transfer to the inoculated plant shoots among the evaluated fungal species were recorded. Ca mobilization by L. laccata, Na by L. bicolor and Mn by H. leucosarx were observed in the plants growing in the experimental substrate. It has been demonstrated that the selection of substrates constitutes an important factor in the production of ectomycorrhizal plants and that the three evaluated species of edible ectomycorrhizal mushrooms have an enormous potential in the controlled mycorrhization of P. greggii.

Ectomycorrhizae of Tuber huidongense and T. liyuanum with Castanea mollissima and Pinus armandii.

Tuber huidongense and T. liyuanum are common commercial white truffles in China that belong to the Rufum and Puberulum groups of the genus Tuber, respectively. Their mycorrhizae were successfully synthesized with two native trees--Castanea mollissima and Pinus armandii--under greenhouse conditions. The identities of the mycorrhizae were confirmed through internal transcribed spacer (ITS) sequence analyses, and their morphological characteristics were described. All of the obtained mycorrhizae have an interlocking pseudoparenchymatous mantle, which is a typical feature of truffle mycorrhizae. The mycorrhizae of T. huidongense on the two trees have hyaline branched emanating hyphae, similar to the documented mycorrhizae of the Rufum group. The unramified, spiky, and hyaline cystidia on the mycorrhizae of T. liyuanum with both C. mollissima and P. armandii further confirmed that this characteristic is constant for the mycorrhizae of the Puberulum group. The successful mycorrhizal syntheses on the two nut-producing trees will be of economic importance in the cultivation of the two truffles.

Phyto-Extraction of Nickel by Linum usitatissimum in Association with Glomus intraradices.

Plants show enhanced phytoremediation of heavy metal contaminated soils particularly in response to fungal inoculation. Present study was conducted to find out the influence of Nickel (Ni) toxicity on plant biomass, growth, chlorophyll content, proline production and metal accumulation by L. usitatissimum (flax) in the presence of Glomus intraradices. Flax seedlings of both inoculated with G. intraradices and non-inoculated were exposed to different concentrations i.e., 250, 350 and 500 ppm of Ni at different time intervals. Analysis of physiological parameters revealed that Ni depressed the growth and photosynthetic activity of plants. However, the inoculation of plants with arbuscular mycorrhizae (G. intraradices) partially helped in the alleviation of Ni toxicity as indicated by improved plant growth under Ni stress. Ni uptake of non- mycorrhizal flax plants was increased by 98% as compared to control conditions whereas inoculated plants showed 19% more uptake when compared with the non-inoculated plants. Mycorrhizal plants exhibited increasing capacity to remediate contaminated soils along with improved growth. Thus, AM assisted phytoremediation helps in the accumulation of Ni in plants to reclaim Ni toxic soils. Based on our findings, it can be concluded that the role of flax plants and mycorrhizal fungi is extremely important in phytoremediation.

In vitro Scleroderma laeve and Eucalyptus grandis mycorrhization and analysis of atp6, 17S rDNA, and ras gene expression during ectomycorrhizal formation.

The interaction between fungi and plants that form ectomycorrhizae (ECM) promotes alterations in the gene expression profiles of both organisms. Fungal genes expression related to metabolism were evaluated at the pre-symbiotic stage and during the ECM development between Scleroderma laeve and Eucalyptus grandis. Partial sequences of ATP synthase (atp6), translation elongation factor (ef1α), the RAS protein (ras), and the 17S rDNA genes were isolated. The expression of the atp6 and 17S rDNA genes during the pre-symbiotic stage showed an approximately threefold increase compared to the control. During ECM development, the expression of the 17S rDNA gene showed a 4.4-fold increase after 3 days of contact, while the expression of the atp6 gene increased 7.23-fold by the 15th day, suggesting that protein synthesis and respiratory chain activities are increased during the formation of the mantle and the Hartig net. The ras gene transcripts were only detected by RT-PCR 30 days after fungus-plant contact, suggesting that RAS-mediated signal transduction pathways are functional during the establishment of symbiosis. The present study demonstrates that alterations in gene expression occur in response to stimuli released by the plant during ECM association and increases the understanding of the association between S. laeve and E. grandis.

An Overview of Mycorrhiza in Pines: Research, Species, and Applications.

In the latest literature, climate models show that the conditions for pines, spruces, larches, and birches will deteriorate significantly. In Poland, as well as in other European countries, there are already signs of the decline of these species. This review article deals with the symbiotic relationships between fungi and plants, which can hardly be overestimated, using the example of pine trees. These are the oldest known symbiotic relationships, which are of great benefit to both components and can help plants, in particular, survive periods of severe drought and the attack of pathogens on the roots. This article describes symbioses and their causal conditions, as well as the mycorrhizal components of pine trees and their properties; characterizes ectomycorrhizal fungi and their mushroom-forming properties; and provides examples of the cultivation of pure fungal cultures, with particular attention to the specificity of the mycorrhizal structure and its effects on the growth and development of Pinus species. Finally, the role of mycorrhiza in plant protection and pathogen control is described.

A tripartite bacterial-fungal-plant symbiosis in the mycorrhiza-shaped microbiome drives plant growth and mycorrhization.

BACKGROUND: Plant microbiomes play crucial roles in nutrient cycling and plant growth, and are shaped by a complex interplay between plants, microbes, and the environment. The role of bacteria as mediators of the 400-million-year-old partnership between the majority of land plants and, arbuscular mycorrhizal (AM) fungi is still poorly understood. Here, we test whether AM hyphae-associated bacteria influence the success of the AM symbiosis. RESULTS: Using partitioned microcosms containing field soil, we discovered that AM hyphae and roots selectively assemble their own microbiome from the surrounding soil. In two independent experiments, we identified several bacterial genera, including Devosia, that are consistently enriched on AM hyphae. Subsequently, we isolated 144 pure bacterial isolates from a mycorrhiza-rich sample of extraradical hyphae and isolated Devosia sp. ZB163 as root and hyphal colonizer. We show that this AM-associated bacterium synergistically acts with mycorrhiza on the plant root to strongly promote plant growth, nitrogen uptake, and mycorrhization. CONCLUSIONS: Our results highlight that AM fungi do not function in  isolation and that the plant-mycorrhiza symbiont can recruit beneficial bacteria that support the symbiosis. Video Abstract.

Enhancing tomato plants' tolerance to combined heat and salt stress - The role of arbuscular mycorrhizae and biochar.

The Mediterranean basin is highly susceptible to climate change, with soil salinization and the increase in average temperatures being two of the main factors affecting crop productivity in this region. Following our previous studies on describing the detrimental effects of heat and salt stress co-exposure on tomato plants, this study aimed to understand if substrate supplementation with a combination of arbuscular mycorrhizal fungi (AMF) and biochar could mitigate the negative consequences of these stresses. Upon 21 days of exposure, stressed tomato plants grown under supplemented substrates showed increased tolerance to heat (42 °C for 4 h/day), salt (100 mM NaCl), and their combination, presenting increased biomass and flowering rate. The beneficial effects of AMF and biochar were associated with a better ionic balance (i.e. lower sodium accumulation and higher uptake of calcium and magnesium) and increased photosynthetic efficiency. Indeed, these plants presented higher chlorophyll content and improved CO2 assimilation rates. Biochemical data further supported that tomato plants grown with AMF and biochar were capable of efficiently modulating their defence pathways, evidenced by the accumulation of proline, ascorbate, and glutathione, coupled with a lower dependency on energy-costly enzymatic antioxidant players. In summary, the obtained data strongly point towards a beneficial role of combined AMF and biochar as sustainable tools to improve plant growth and development under a climate change scenario, where soil salinization and heat peaks often occur together.

Effect of Bacteria Inoculation on Colonization of Roots by Tuber melanosporum and Growth of Quercus ilex Seedlings.

Tuber melanosporum is an ascomycete that forms ectomycorrhizal (ECM) symbioses with a wide range of host plants, producing edible fruiting bodies with high economic value. The quality of seedlings in the early symbiotic stage is important for successful truffle cultivation. Numerous bacterial species have been reported to take part in the truffle biological cycle and influence the establishment of roots symbiosis in plant hosts and the development of the carpophore. In this work, three different bacteria formulations were co-inoculated in Quercus ilex L. seedlings two months after T. melanosporum inoculation. At four months of bacterial application, the T. melanosporum ECM root tip rate of colonization and bacterial presence were assessed using both morphological and molecular techniques. A 2.5-fold increase in ECM colonization rate was found in the presence of Pseudomonas sp. compared to the seedlings inoculated only with T. melanosporum. The same treatment caused reduced plant growth either for the aerial and root part. Meanwhile, the ECM colonization combined with Bradyrhizobium sp. and Pseudomonas sp. + Bradyrhizobium sp. reduced the relative density of fibrous roots (nutrient absorption). Our work suggests that the role of bacteria in the early symbiotic stages of ECM colonization involves both the mycorrhizal symbiosis rate and plant root development processes, both essential for improve the quality of truffle-inoculated seedlings produced in commercial nurseries.

Mycorrhization in trees: ecology, physiology, emerging technologies and beyond.

Mycorrhization has been an integral part of plants since colonization by the early land plants. Over decades, substantial research has highlighted its potential role in improving nutritional efficiency and growth, development and survival of crop plants. However, the focus of this review is trees. Evidence have been provided to explain ecological and physiological significance of mycorrhization in trees. Advances in recent technologies (e.g., metagenomics, artificial intelligence, machine learning, agricultural drones) may open new windows to apply this knowledge in promoting tree growth in forest ecosystems. Dual mycorrhization relationships in trees and even triple relationships among trees, mycorrhizal fungi and bacteria offer an interesting physiological system to understand how plants interact with other organisms for better survival. Besides, studies indicate additional roles of mycorrhization in learning, memorizing and communication between host trees through a common mycorrhizal network (CMN). Recent observations in trees suggest that mycorrhization may even promote tolerance to multiple abiotic (e.g., drought, salt, heavy metal stress) and biotic (e.g. fungi) stresses. Due to the extent of physiological reliance, local adaptation of trees is heavily impacted by the mycorrhizal community. This knowledge opens the possibility of a non-GMO avenue to promote tree growth and development. Indeed, mycorrhization could impact growth of trees in nurserys and subsequent survival of the inoculated trees in field conditions. Future studies might integrate hyperspectral imaging and drone technologies to identify tree communities that are deficient in nitrogen and spray mycorrhizal spore formulations on them.