Search

This study aimed to evaluate effects of Funneliformis mosseae on plant growth and root exudate compositions and contents, soil and root phosphatase activity, soil glomalin concentrations, and thus soil aggregate stability and distribution in trifoliate orange under well-watered (75% of maximum water holding capacity) and drought stress (55% of maximum water holding capacity) conditions. After eight weeks of drought treatment, mycorrhizal fungal inoculation improved plant growth and exhibited altered composition of root exudates than non-inoculated treatment. Mycorrhizal fungal inoculation dramatically increased the relative abundance of phenolics (e.g., 2 H,8 H-Benzo[1,2-b:3,4-b'] dipyran-2-one,8,8-dimethyl), terpenoids (e.g., geijerene), and acids (n-hexadecanoic acid), while notably reduced the relative abundance of alkanes (e.g., tridecane, 2-methyl-), esters (e.g., hexanedioic acid and dimethyl ester), and amides (e.g.,13-docosenamide) in root exudates. Mycorrhizal fungal colonization profoundly increased easily extractable and total glomalin-related soil protein levels under two soil water regimes, which cemented soil macroaggregate (2–4 mm size) formation, thereby, improving soil aggregate stability. Mycorrhizal fungal-inoculated plants represented higher soil acid, alkaline and total phosphatase activities, irrespective of well-watered and drought treatment. The results suggest that mycorrhizal plants had improved root microenvironment to mitigate drought damage through changes in root exudate components along with glomalin, phosphatase, and soil aggregate stability in the mycorrhizosphere. [ABSTRACT FROM AUTHOR]

Soil water deficit seriously affects crop production, and soil arbuscular mycorrhizal fungi (AMF) enhance drought tolerance in crops by unclear mechanisms. Our study aimed to analyze changes in non-targeted metabolomics in roots of trifoliate orange (Poncirus trifoliata) seedlings under well-watered and soil drought after inoculation with Rhizophagus intraradices , with a focus on terpenoid profile. Root mycorrhizal fungal colonization varied from 70% under soil drought to 85% under soil well-watered, and shoot and root biomass was increased by AMF inoculation, independent of soil water regimes. A total of 643 secondary metabolites in roots were examined, and 210 and 105 differential metabolites were regulated by mycorrhizal fungi under normal water and drought stress, along with 88 and 17 metabolites being up-and down-regulated under drought conditions, respectively. KEGG annotation analysis of differential metabolites showed 38 and 36 metabolic pathways by mycorrhizal inoculation under normal water and drought stress conditions, respectively. Among them, 33 metabolic pathways for mycorrhization under drought stress included purine metabolism, pyrimidine metabolism, alanine, aspartate and glutamate metabolism, etc. We also identified 10 terpenoid substances, namely albiflorin, artemisinin (−)-camphor, capsanthin, β-caryophyllene, limonin, phytol, roseoside, sweroside, and α-terpineol. AMF colonization triggered the decline of almost all differential terpenoids, except for β-caryophyllene, which was up-regulated by mycorrhizas under drought, suggesting potential increase in volatile organic compounds to initiate plant defense responses. This study provided an overview of AMF-induced metabolites and metabolic pathways in plants under drought, focusing on the terpenoid profile. [ABSTRACT FROM AUTHOR]

Multiple functions of glomalin released by arbuscular mycorrhizal fungi are well-recognized, whereas the role of exogenous glomalins including easily extractable glomalin-related soil protein (EE-GRSP) and difficultly extractable glomalin-related soil protein (DE-GRSP) is unexplored for plant responses. Our study was carried out to assess the effects of exogenous EE-GRSP and DE-GRSP at varying strengths on plant growth and chlorophyll concentration of trifoliate orange (Poncirus trifoliata) seedlings, along with changes in root nutrient acquisition, auxin content, auxin-related enzyme and transporter protein gene expression, and element contents of purified GRSP. Sixteen weeks later, exogenous GRSP displayed differential effects on plant growth (height, stem diameter, leaf number, and biomass production): the increase by EE-GRSP and the decrease by DE-GRSP. The best positive effect on plant growth occurred at exogenous EE-GRSP at ½ strength. Similarly, the GRSP application also differently affected total chlorophyll content, root morphology (total length, surface area, and volume), and root N, P, and K content: positive effect by EE-GRSP and negative effect by DE-GRSP. Exogenous EE-GRSP accumulated more indoleacetic acid (IAA) in roots, which was associated with the upregulated expression of root auxin synthetic enzyme genes (PtTAA1, PtYUC3 , and PtYUC4) and auxin influx transporter protein genes (PtLAX1, PtLAX2 , and PtLAX3). On the other hand, exogenous DE-GRSP inhibited root IAA and indolebutyric acid (IBA) content, associated with the downregulated expression of root PtTAA1, PtLAX1 , and PtLAX3. Root IAA positively correlated with root PtTAA1, PtYUC3, PtYUC4, PtLAX1 , and PtLAX3 expression. Purified EE-GRSP and DE-GRSP showed similar element composition but varied in part element (C, O, P, Ca, Cu, Mn, Zn, Fe, and Mo) concentration. It concluded that exogenous GRSP triggered differential effects on growth response, and the effect was associated with the element content of pure GRSP and the change in auxins and root morphology. EE-GRSP displays a promise as a plant growth biostimulant in citriculture. [ABSTRACT FROM AUTHOR]

Endophytes have the ability to improve plant nutrition alongside their agronomic performance, among which arbuscular mycorrhizal fungi provide the most benefits to their host. Previously, we reported for the first time that an arbuscular mycorrhizal-like fungus Piriformospora indica had the ability to colonize roots of trifoliate orange (Poncirus trifoliata) and conferred positive effects on nutrient acquisition. Present study showed the changes in fatty acids and sugars to unravel the physiological and symbiotic association of trifoliate orange with P. indica and an arbuscular mycorrhizal fungus, Funneliformis mosseae singly or in combination. All the endophytic fungi collectively increased fructose, glucose, and sucrose content in leaves and roots, along with a relatively higher increase with P. indica inoculation than with F. mosseae alone or dual inoculation. Treatment with P. indica increased the concentration of part unsaturated fatty acids such as C18:3N6, C20:2, C20:3N6, C20:4N6, C20:3N3, C20:5N3, C22:1N9, and C24:1. Additionally, P. indica induced the increase in the concentration of part saturated fatty acids such as C6:0, C8:0, C13:0, C14:0, and C24:0. F. mosseae hardly changed the content of fatty acids, except for increase in C14:0 and C20:5N3. Double inoculation only reduced the C21:0, C10:0, C12:0, C18:3N3, and C18:1 content and increased the C20:5N3 content. These endophytic fungi up-regulated the root PtFAD2, PtFAD6, PtD9, and PtD15 gene expression level, coupled with a higher expression of PtFAD2 and PtD9 by P. indica than by F. mosseae. It was concluded that P. indica exhibited a stronger response, for sugars and fatty acids, than F. mosseae on trifoliate orange. Such results also reveal the Pi (an in vitro culturable fungus) as a bio-stimulator applying to citriculture. [ABSTRACT FROM AUTHOR]

A feature of arbuscular mycorrhiza is enhanced drought tolerance of host plants, although it is unclear whether host H+-ATPase activity and gene expression are involved in the physiological process. The present study aimed to investigate the effects of an arbuscular mycorrhizal fungus (AMF), Funneliformis mosseae , on H+-ATPase activity, and gene expression of trifoliate orange (Poncirus trifoliata) seedlings subjected to well-watered (WW) and drought stress (DS), together with the changes in leaf gas exchange, root morphology, soil pH value, and ammonium content. Soil drought treatment dramatically increased H+-ATPase activity of leaf and root, and AMF inoculation further strengthened the increased effect. A plasma membrane (PM) H+-ATPase gene of trifoliate orange, PtAHA2 (MW239123), was cloned. The PtAHA2 expression was induced by mycorrhization in leaves and roots and also up-regulated by drought treatment in leaves of AMF-inoculated seedlings and in roots of AMF- and non-AMF-inoculated seedlings. And, the induced expression of PtAHA2 under mycorrhization was more prominent under DS than under WW. Mycorrhizal plants also showed greater photosynthetic rate, stomatal conductance, intercellular CO2 concentration, and transpiration rate and better root volume and diameter than non-mycorrhizal plants under DS. AMF inoculation significantly increased leaf and root ammonium content, especially under DS, whereas it dramatically reduced soil pH value. In addition, H+-ATPase activity was significantly positively correlated with ammonium contents in leaves and roots, and root H+-ATPase activity was significantly negatively correlated with soil pH value. Our results concluded that AMF stimulated H+-ATPase activity and PtAHA2 gene expression in response to DS, which resulted in great nutrient (e.g., ammonium) uptake and root growth, as well as low soil pH microenvironment. [ABSTRACT FROM AUTHOR]

Soil arbuscular mycorrhizal fungi (AMF) enhance the tolerance of plants against soil moisture deficit stress (SMDS), but the underlying mechanisms are still not fully understood. Polyamines (PAs) as low-molecular-weight, aliphatic polycations have strong roles in abiotic stress tolerance of plants. We aimed to investigate the effect of AMF (Funneliformis mosseae) inoculation on PAs, PA precursors, activities of PA synthases and degrading enzymes, and concentration of reactive oxygen species in the roots of trifoliate orange (Poncirus trifoliata) subjected to 15 days of SMDS. Leaf water potential and total chlorophyll levels were comparatively higher in AMF-inoculated than in non-AMF-treated plants exposed to SMDS. Mycorrhizal plants recorded a significantly higher concentration of precursors of PA synthesis such as L-ornithine, agmatine, and S -adenosyl methionine, besides higher putrescine and cadaverine and lower spermidine during the 15 days of SMDS. AMF colonization raised the PA synthase (arginine decarboxylase, ornithine decarboxylase, spermidine synthase, and spermine synthase) activities and PA-degrading enzymes (copper-containing diamine oxidase and FAD-containing polyamine oxidase) in response to SMDS. However, mycorrhizal plants showed a relatively lower degree of membrane lipid peroxidation, superoxide anion free radical, and hydrogen peroxide than non-mycorrhizal plants, whereas the difference between them increased linearly up to 15 days of SMDS. Our study concluded that AMF regulated PA homeostasis in roots of trifoliate orange to tolerate SMDS. [ABSTRACT FROM AUTHOR]

A pot experiment was used to evaluate the effects of an arbuscular mycorrhizal fungus (AMF) Funneliformis mosseae on plant growth performance, root-hair growth, and root hormone levels in trifoliate orange (Poncirus trifoliata) seedlings under well-watered (WW) and drought stress (DS). A 9-week mild DS treatment significantly reduced mycorrhizal colonization of 2nd- and 3rd-order lateral roots. Root mycorrhizal colonization was relatively higher in the 2nd- and 3rd-order lateral roots than in the taproot and the 1st-order lateral root under WW and DS. AMF seedlings exhibited significantly higher root-hair density, length (except for the taproot) and diameter in taproot and 1st-, 2nd-, and 3rd-order lateral roots under WW, and considerably higher root-hair density (except for 1st-order lateral root), length (except for 2nd-order lateral root) and diameter under DS. Mycorrhizal inoculation remarkably increased root abscisic acid (ABA), indole-3-acetic acid (IAA), methyl jasmonate, and brassinosteroids (BRs) concentrations under DS, in company with the decrease in root zeatin riboside and gibberellins levels and root IAA effluxes. Root-hair traits were significantly positively correlated with root colonization and root ABA and BRs levels. It is concluded that mycorrhizal plants possessed better root-hair growth to adapt mild DS, which is associated with mycorrhizal colonization and endogenous hormone changes. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizas (AMs) have the ability to enhance drought tolerance of citrus, but the underlying mechanisms have not been clearly elucidated. Considering the strong association of cell membrane fatty acid (FA) unsaturation with plant drought tolerance, the present study hypothesized that AM fungi (AMF) modulated the composition and unsaturation of FAs to enhance drought tolerance of host plants. Drought-sensitive citrus rootstocks, trifoliate orange (Poncirus trifoliata) seedlings, were inoculated with AMF (Funneliformis mosseae) for 3 months and were subsequently exposed to drought stress (DS) for 8 weeks. Mycorrhizal seedlings exhibited better plant growth performance, higher leaf water potential and lower root abscisic acid concentrations under both well-watered (WW) and DS conditions. Arbuscular mycorrhiza fungus inoculation considerably increased root methyl oleate (C18:1), methyl linoleate (C18:2) and methyl linolenate (C18:3N3) concentrations under both WW and DS conditions, and root methyl palmitoleate (C16:1) concentrations under WW, while it decreased root methyl stearate (C18:0) levels under both WW and DS. These changes in the composition of FAs of mycorrhized roots resulted in higher unsaturation index of root FAs, which later aided in reducing the oxidative damage on account of lower concentration of malondialdehyde and superoxide radicals. The changes of these FAs were a result of AMF-up-regulating root FA desaturase 2 (PtFAD2), FA desaturase 6 (PtFAD6) and Δ9 FA desaturase (PtΔ9) genes under WW and PtFAD2 , PtFAD6 and Δ15 FA desaturase (PtΔ15) genes under DS conditions. Our results confirmed that mycorrhization brought significant changes in root FA compositions, in addition to regulation of gene expression responsible for increasing the unsaturation level of FAs, a predisposing physiological event for better drought tolerance of citrus. [ABSTRACT FROM AUTHOR]

Aquaporins (AQPs) involved in water and small molecule transport respond to environmental stress, while it is not clear how arbuscular mycorrhizal fungi (AMF) regulate AQP expression. Here, we investigated the change in leaf water potential and expression level of four tonoplast intrinsic proteins (TIPs), six plasma membrane intrinsic proteins (PIPs), and four nodin-26 like intrinsic proteins (NIPs) genes in trifoliate orange (Poncirus trifoliata) inoculated with Funneliformis mosseae under well-watered (WW), salt stress (SS), and waterlogging stress (WS). Root AMF colonization and soil hyphal length collectively were reduced by SS and WS. Under WW, inoculation with AMF gave diverse responses of AQPs : six AQPs up-regulated, three AQPs down-regulated, and five AQPs did not change. Such up-regulation of more AQPs under mycorrhization and WW partly accelerated water absorption, thereby, maintaining higher leaf water potential. However, under SS, all the fourteen AQPs were dramatically induced by AMF inoculation, which improved water permeability of membranes and stimulated water transport of the host. Under WS, AMF colonization almost did not induce or even down-regulated these AQPs expressions with three exceptions (PtTIP2 ;2, PtPIP1;1 , and PtNIP1;2), thus, no change in leaf water potential. As a result, mycorrhizal plants under flooding may have an escape mechanism to reduce water absorption. It is concluded that AMF had different strategies in response to environmental stresses (e.g. SS and WS) by regulating leaf AQP expression in the host (e.g. trifoliate orange). • Under ample water, AMF gave diverse responses of AQPs. • Under salinity, all the 14 AQPs were up-regulated by AMF inoculation to stimulate water uptake. • Under waterlogging, AMF almost did not induce these AQPs expressions, thus, giving an escape mechanism. • AMF have different strategies in response to environmental stress through AQPs. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) enhance plant disease resistance, while the underlying mechanisms in the molecular levels are not yet known. In this study, five-leaf-old trifoliate orange seedlings were inoculated with Funneliformis mosseae for 14 weeks and subsequently were infected by a citrus root rot pathogen Phytophthora parasitica by 7 days. The transcriptome results by Illumina HiSeq 4000 revealed that the percentage of Q30 bases reached 92.99% or above, and 29696 unigenes were annotated in a total of 63531 unigenes. 654 and 103 differentially expressed genes (DEGs) were respectively annotated in AMF-inoculated versus non-AMF-inoculated plants under non-infection and infection with P. parasitica, respectively, whilst these DEGs were related to defense mechanisms, signal transduction mechanisms and secondary metabolites biosynthesis. Forty-two genes were functionally annotated as the putative 'defense mechanism', whilst AMF inoculation induced 1 gene down-regulated and 3 genes up-regulated under P. parasitica infection. AMF inoculation stimulated more genes linked to signal transduction mechanism down-regulated than non-AMF plants. Eight genes were involved in secondary metabolites biosynthesis in AMF versus non-AMF seedlings under P. parasitica-infection conditions. Such transcriptome database provided total information in the molecular levels regarding mycorrhizal roles in tolerating Phytophthora parasitica infection. [ABSTRACT FROM AUTHOR]

Circadian clock is usually involved in many physiological processes of plants, including responses to abiotic stress, whilst pseudo-response regulator 7 (PRR7) gene is the main component of the circadian clock. In this study, the cDNA of the PRR7 gene was obtained from trifoliate orange (Poncirus trifoliata). Based on the sequence analysis, the PtPRR7 gene had an open reading frame of 2343 bp, encoded 780 amino acids, and contained proteins of the REC and CCT domains. Subcellular localization indicated that PtPRR7 was mainly localized in the nucleus and a small amount of cytoplasm. qRT-PCR analysis revealed the highest expression level of PtPRR7 in roots than in both shoots and leaves. The PtPRR7 gene during 24 hours of soil water deficit exhibited a circadian rhythmic expression pattern: the expression peak at 9:00 am in leaves and at 21:00 pm in roots. Drought treatment affected PtPRR7 gene expression. Such data provide important references for understanding the characteristics of PtPRR7 gene in citrus plants. [ABSTRACT FROM AUTHOR]

• Effects of symbiotic AM fungi and AM like fungi on trifoliate orange were studied. • Piriformospora indica and AMF coexisted in roots, but the colonization was inferior. • Single or dual inoculation increased plant growth and affected root nutrients. • P increase in inoculated plants was associated with soil phosphatase and PT gene. • Piriformospora indica has more applied value in citrus than AMF as a biofertilizer. Citrus plants heavily depend on soil arbuscular mycorrhizal fungi (AMF), while the application of AMF in citrus is limited because of no propagation in vitro. The present work tried to apply a culturable (in vitro) endophytic fungus Piriformospora indica into trifoliate orange (Poncirus trifoliata) and to compare the role of P. indica in plant growth and nutrient acquisition especially phosphorus (P), relative to an arbuscular mycorrhizal fungus Funneliformis mosseae. After 20 weeks, roots of trifoliate orange could be colonized by P. indica , and both P. indica and F. mosseae existed in roots together. Root fungal colonization and soil easily extractable, difficultly extractable, and total glomalin-related protein concentrations were higher under single F. mosseae inoculation than under single P. indica and dual inoculation of F. mosseae and P. indica. Single or dual microbial inoculant collectively improved plant growth including plant height, leaf number, and leaf, stem, and root biomass, compared with non-inoculation. In addition, single fungal treatment raised the content of root N, P, and Mg, and dual inoculation increased only Mg and B contents. The improvement of P by single microbial inoculant was associated with the fungi-accelerated soil phosphatase activity and the fungi-induced root phosphate transporter genes (PT3 , PT5 , and PT6) expression levels. The positive effects on growth and nutrients were more significant under single F. mosseae than under single P. indica and dual inoculations. It concluded that P. indica has the stimulated roles in plant growth and P acquisition of trifoliate orange, and thus has more applied potentiality in citriculture than AMF as a biofertilizer. [ABSTRACT FROM AUTHOR]

Arbuscular mycorrhizal fungi (AMF) produce glomalin-related soil protein (GRSP) that influences organic carbon (C) storage in soil; however, how much purified GRSP fractions contribute to soil organic carbon (SOC) is yet not known. The present study evaluated the contribution of GRSP towards changes in SOC in trifoliate orange grown in a rootbox divided into a roots + hyphae chamber (roots colonized by AMF hyphae and AMF extraradical hyphae) and a hyphae chamber (only the presence of AMF extraradical hyphae, without roots). Three AMF species (Diversispora epigaea , Paraglomus occultum and Rhizoglomus intraradices) were inoculated into the roots + hyphae chamber. Following four months of plant growth, P. occultum showed higher AMF hyphal growth in roots, soils and nylon mesh than D. epigaea or R. intraradices. Mycorrhizal inoculation improved the plant growth performance and increased easily extractable GRSP (EE-GRSP) and difficultly extractable GRSP (DE-GRSP) concentrations in both chambers, regardless of AMF species. The C content observed in total GRSP of the soil after purification was 2.71 ± 0.49 mg g−1, while purified EE-GRSP and DE-GRSP showed the C content of 1.01 ± 0.19 mg g−1 and 1.70 ± 010.34 mg g−1, respectively. The C contribution by purified EE-GRSP and DE-GRSP accounted for 8.67 ± 0.95% and 14.59 ± 2.21%, respectively, of total SOC, with a total C contribution of purified GRSPs accounting for 23.26 ± 2.67% of total SOC. A significantly higher C content of GRSP and the C contribution of GRSP to SOC were observed in DE-GRSP than in EE-GRSP, as well as the soil of the roots + hyphae chamber than the soil of the hyphae chamber. The proportionate distribution of water-stable aggregate in 2–4 and 1–2 mm sizes and their stability were higher under AMF hyphae than under non-AMF hyphae. This study thus provided a database evidence of increased contribution of GRSP towards build-up of SOC in response to mycorrhizal symbiosis. • GRSP contribution to SOC in trifoliate orange was studied. • The C content of EE-GRSP and DE-GRSP was 1.01 ± 0.19 mg g-1 and 1.70 ± 0.34 mg g. • C contribution of total GRSPs achieved 23.26 ± 2.67% of total SOC. • C content of GRSP and C contribution of GRSP to SOC were higher in DE-GRSP than in EE-GRSP. [ABSTRACT FROM AUTHOR]

• This study analyzed how PAs were involved in AMF-enhanced drought tolerance of plants. • AM trifoliate orange had higher Put and Cad and lower Spd and Spm under drought. • Mycorrhizal plants had higher activity of PA catabolic enzymes and Put synthetase. • AM plants showed overexpression of PtSPMS , PtPAO , PtSOD , and PtCAT1 under DS. • AM plants represented down-regulated expression of PtCuAO under DS. Arbuscular mycorrhizas enhance the drought tolerance of host plants through several underlying mechanisms. Polyamines (PAs) are known to protect plant cells from the damages of drought by enhancing the antioxidant defense system, restricting ethylene synthesis, maintaining cell pH and ion homeostasis, preventing chlorophyll loss, and so on; concomitantly, the mechanism through which arbuscular mycorrhizal (AM) fungi modulate PA metabolism to enhance drought tolerance of plants remains unclear. A pot experiment was conducted to assess how an AM fungus, Funneliformis mosseae , affects root PA homeostasis, activities and gene expressions of PA-related synthesizing and degrading enzymes, hydrogen peroxide (H 2 O 2 , a PA metabolite) production, and antioxidant enzyme gene expressions in trifoliate orange (Poncirus trifoliata) exposed to drought stress (DS). AM seedlings showed higher growth traits, leaf water potential, two plasma membrane intrinsic protein aquaporin gene expressions, and chlorophyll concentrations than non-AM seedlings under well-watered (WW) and DS conditions. Mycorrhizal treatment induced higher putrescine and cadaverine but lower spermidine and spermine concentrations, with higher activity of PA catabolic enzymes (copper-containing diamine oxidase, CuAO; polyamine oxidase, PAO) and putrescine synthases (ornithine decarboxylaseby; arginine decarboxylase, ADC). Mycorrhizas up-regulated the expression of the spermine synthase gene, PtSPMS , under DS, and down-regulated the transcript levels of PA catabolic enzyme genes (PtCuAO1 , PtCuAO2 , PtCuAO6 , and PtCuAO8) and PA synthase genes (PtADC1 and PtADC2) under DS. PtPAO1 , PtPAO2 , and PtPAO3 had higher expression levels in AMF-inoculated seedlings, as compared to non-AMF-inoculated seedlings, under DS, triggering reactive oxygen species-related signalling for stress responsiveness through low H 2 O 2 levels by up-regulating the expression of PtMn-SOD , PtCu/Zn-SOD, and PtCAT1. This study demonstrated that mycorrhizas have the capacity to modulate PA metabolism to enhance the drought tolerance of plants. [ABSTRACT FROM AUTHOR]

• Responses of antioxidant enzyme defense system to AMF under DS were studied. • Leaf Fe-SOD, Mn-SOD, Cu/Zn-SOD and CAT activity was increased by AMF under DS. • Leaf PtMn-SOD , PtCAT1 and PtPOD expression was up-regulated by AMF under DS. • AM plants had lower leaf MDA, O 2 − and H 2 O 2 levels under DS. • AM plants had greater capacity on antioxidant enzyme defense systems under DS. A pot experiment was carried out to investigate the responses of leaf antioxidant enzyme defense systems to an arbuscular mycorrhizal fungus (AMF), Funneliformis mosseae , in trifoliate orange (Poncirus trifoliata) seedlings exposed to drought stress (DS). An eight-week soil drought treatment notably decreased root mycorrhizal colonization, root surface area and volume, total plant biomass, leaf relative water content, and leaf chlorophyll a , b , and a + b concentrations, while these variables were significantly greater in mycorrhizal seedlings than in non-mycorrhizal seedlings. There was significantly higher leaf catalase (CAT) activity in AMF-inoculated seedlings versus non-AMF-inoculated seedlings under well-watered (WW) conditions. Under DS, leaf superoxide dismutase (Fe-SOD, Mn-SOD, and Cu/Zn-SOD) and CAT activities were dramatically increased with AMF inoculation. Leaf PtMn-SOD and peroxidase (PtPOD) gene expressions were significantly increased by AMF treatment under WW. Under DS, the relative expression of leaf PtMn-SOD , PtCAT1 , and PtPOD was increased by AMF, while leaf PtFe-SOD and PtCu/Zn-SOD expressions were down-regulated. Such higher antioxidant enzyme activities and corresponding gene expressions in mycorrhizal plants under DS indicated their greater capacity to mitigate the oxidative damage of drought, as evidenced by the lower concentrations of malondialdehyde, superoxide radical, and hydrogen peroxide in AMF-inoculated plants. [ABSTRACT FROM AUTHOR]