Earthworm and arbuscular mycorrhiza interactions: Strategies to motivate antioxidant responses and improve soil functionality.

Earthworms and arbuscular mycorrhizal fungi (AMF) act synergistically in the rhizosphere and may increase host plant tolerance to Cd. However, mechanisms by which earthworm-AMF-plant partnerships counteract Cd phytotoxicity are unknown. Thus, we evaluated individual and interactive effects of these soil organisms on photosynthesis, antioxidant capacity, and essential nutrient uptake by Solanum nigrum, as well as on soil quality following Cd exposure (0-120 mg kg-1). Decreases in biomass and photosynthetic activity, as well as nutrient imbalances were observed in Cd-stressed plants; however, the addition of AMF and earthworms reversed these effects. Cd exposure increased superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) activities, whereas inoculation with Rhizophagus intraradices decreased those. Soil enzymatic activity decreased by 15-60% with increasing Cd concentrations. However, Cd-mediated toxicity was partially reversed by soil organisms. Earthworms and AMF ameliorated soil quality based on soil enzyme activity. At 120 mg kg-1 Cd, the urease, catalase, and acid phosphatase activities were 1.6-, 1.4-, and 1.2-fold higher, respectively, in soils co-incubated with earthworms and AMF than in uninoculated soil. Cd inhibited shoot Fe and Ca phytoaccumulation, whereas AMF and earthworms normalized the status of essential elements in plants. Cd detoxification by earthworm-AMF-S. nigrum symbiosis was manifested by increases in plant biomass accumulation (22-117%), chlorophyll content (17-63%), antioxidant levels (SOD 10-18%, POD 9-25%, total polyphenols 17-22%, flavonoids 15-29%, and glutathione 7-61%). It also ameliorated the photosynthetic capacity, and macro- and micronutrient statuses of plants; markedly reduced the levels of malondialdehyde (20-27%), superoxide anion (29-36%), and hydrogen peroxide (19-30%); and upregulated the transcription level of FeSOD. Thus, the combined action of earthworms and AMF feasibly enhances metal tolerance of hyperaccumulating plants and improves the quality of polluted soil.

Wang G ，Wang L ，Ma F ，Yang D ，You Y ... -  《-》

Integration of earthworms and arbuscular mycorrhizal fungi into phytoremediation of cadmium-contaminated soil by Solanum nigrum L.

Arbuscular mycorrhizal fungi (AMF) and earthworms independently enhance plant growth, heavy metal (HM) tolerance, and HM uptake, thus they are potential key factors in phytoremediation. However, few studies have investigated their interactions in HM phytoextraction by hyperaccumulators. This study highlights the independent and interactive effects of earthworms and AMF on Solanum nigrum. Plants inoculated with either AMF or earthworms exhibited ameliorated growth via enhancement of productivity, metal tolerance, and phosphorus (P) acquisition. Co-inoculation with both had more pronounced effects on plant biomass and P acquisition in shoots, but not in roots, and in Cd-polluted soils it significantly promoted (P < 0.05) shoot biomass (20.7-134.6 %) and P content (20.4-112.0 %). AMF and earthworms increased Cd accumulation in plant tissues, but only AMF affected Cd partitioning between shoots and roots. Although AMF decreased root-to-shoot translocation of Cd at high Cd levels, this was counterbalanced by earthworms. Both AMF and its combination with earthworms enhanced Cd phytoavailability by altering Cd chemical fractions and decreasing pH. Co-inoculation increased Cd removal amounts up to 149.3 % in 120 mg kg-1 Cd-spiked soils. Interactions between the two organisms were synergistic in Cd phytoextraction. Thus, earthworm-AMF-plant symbiosis potentially plays an essential role in phytoremediation of HM-polluted soils.

Wang G ，Wang L ，Ma F ，You Y ，Wang Y ，Yang D ... -  《-》

Effect of arbuscular mycorrhizal fungi in roots on antioxidant enzyme activity in leaves of Robinia pseudoacacia L. seedlings under elevated CO(2) and Cd exposure.

Arbuscular mycorrhizal fungi (AMF) are easily influenced by increasing atmospheric CO2 concentration and heavy metals including cadmium (Cd), which can regulate antioxidant enzyme in host plants. Although the effect of AMF under individual conditions such as elevated CO2 (ECO2) and Cd on antioxidant enzyme in host plants has been reported widely, the effect of AMF under ECO2 + Cd receives little attention. In this study, a pot experiment was conducted to study the effect of AMF community in roots on superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD) activities in leaves of 135-d Robinia pseudoacacia L. seedlings under ECO2 + Cd. The activities of SOD and CAT increased and POD activity and the richness and diversity of AMF community decreased under ECO2 + Cd relative to Cd alone. The richness and diversity of AMF were negatively related to Cd content in roots and leaves. The richness and OTUs of AMF community positively and AMF gene abundance negatively affected POD activity under the combined treatments. Superoxide dismutase and POD activities were negatively and positively related to Archaeospora and Scutellospora, respectively, under ECO2 + Cd. Cadmium in roots and leaves was negatively and significantly related to Glomus, Scutellospora, and Claroideoglomus abundance under ECO2 + Cd. Overall, AMF diversity and Archaeospora and Scutellospora in roots significantly influenced SOD, POD, and CAT activities. The response of AM symbiosis to ECO2 might regulate antioxidant capacity in host plants upon Cd exposure. Glomus, Scutellospora, and Claroideoglomus might be applied to phytoremediation of Cd-contaminated soils.

Wang L ，Jia X ，Zhao Y ，Zhang C ，Zhao J ... -  《-》

Arbuscular mycorrhizal fungi and nitric oxide alleviate cadmium phytotoxicity by improving internal detoxification mechanisms of corn plants.

Plants develop several external and internal mechanisms to increase their tolerance to heavy metals (HMs) toxicity including cadmium (Cd). Symbiosis with arbuscular mycorrhizae fungi (AMF) is one of the plants' strategies to tolerate HMs toxicity. Nitric oxide (NO), as a signaling molecule, is also involved in physiological responses of plants to various stresses. The present work was conducted as a factorial completely randomized design with three replications to study the effects of Funneliformis mosseae fungi and Sodium nitroprusside (SNP, 100 mM) as a donor of NO alone, in combination (AMF + SNP) on corn plant growth, and internal detoxification mechanisms of Cd toxicity in a Cd-contaminated calcareous soil (0, 25, 50, and 100 mg Cd kg-1). The results showed that under Cd stress, AMF inoculation and/or foliar application of SNP significantly increased plant growth (32% to 103% for shoot and 44% to 84% for root) by decreasing Cd concentration in corn plant tissues (23% to 46% for shoot and 19% to 40% for root). Cd-induced oxidative stress was mitigated by AMF and/or SNP by enhancing the activities of antioxidant enzymes, including superoxide dismutase (SOD) and catalase (CAT), and concentration of non-enzymatic antioxidants such as glutathione (GSH) and phytochelatin (PC). Increasing the tolerance index (TI) and decreasing the transfer factor (TF) in the corn plants treated with AMF and/or SNP, confirm the efficient role of SNP and AMF in stimulating the detoxification mechanisms of Cd within the plant cells, which was more pronounced at the lowest Cd level (25 mg Cd kg-1). In conclusion, symbiotic associations of corn plants with AMF alone or in combination with SNP mitigated the detrimental effect of Cd toxicity in corn grown in Cd-contaminated calcareous soil. The corn's internal detoxification mechanisms lowered the Cd concentration in plant tissue which resulted in the improvement of the corn's growth parameters.

Zare L ，Ronaghi A ，Ghasemi-Fasaei R ，Zarei M ，Sepehri M ... -  《-》

Cd induced generation of free radical species in Brassica juncea is regulated by supplementation of earthworms in the drilosphere.

The antioxidant defense system of Brassica juncea under Cd stress was examined on supplementation of earthworms in the rhizosphere at different concentrations of Cd (0.50, 0.75, 1.00 and 1.25 mM i.e. 56, 84, 112 and 140 mg kg-1 respectively). Seedlings were raised in small pots containing soil spiked with Cd and earthworms under controlled conditions for 15 days. Improved Cd accumulation, as well as enhanced plant dry weight and metal tolerance were observed following the addition of earthworms. Earthworm supplementation reduced reactive oxygen species (ROS) generation by 7.3% for hydrogen peroxide (H2O2), 7.1% for superoxide anion (O2-), and 8.4% for malondialdehyde (MDA) in plants treated with 1.25 mM (140 mg kg-1) Cd. Confocal microscopy revealed improved cell viability and reduced H2O2 content due to enhanced antioxidative activity. Activity and expression levels of genes coding for antioxidative enzymes (superoxide dismutase; SOD, catalase; CAT, guaicol peroxidase; POD, glutathione reductase; GR, and glutathione-S-transferase; GST) were higher in plants raised in soils inoculated with earthworms, with expression of SOD increasing by 58.8%, CAT by 75%, POD by 183%, GR by 106.6%, and GST by 11.8%. Moreover, plant pigment (chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids) concentrations increased by 8%, 9.1%, 9.1%, and 7.7% respectively, in plants grown in soils supplemented with earthworms. The results of our study suggest that the addition of earthworms to soil increases antioxidative enzyme activities, gene expression in plants, and ROS inhibition, which enhances tolerance to Cd during the phytoextraction process.

Kaur P ，Bali S ，Sharma A ，Kohli SK ，Vig AP ，Bhardwaj R ，Thukral AK ，Abd Allah EF ，Wijaya L ，Alyemeni MN ，Ahmad P ... -  《-》