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Arbuscular mycorrhizal fungi (AMF) and the antioxidant germanium (Ge) are promising tools for boosting bioactive compound synthesis and producing healthier foods. However, their combined effect remains unexplored. This study demonstrates the synergistic impact of AMF and Ge on the growth, metabolite accumulation, biological activities, and nutritional qualities of Spinacia oleracea L. (spinach), a globally significant leafy vegetable. Individually, Ge and AMF increased biomass by 68.1% and 22.7%, respectively, while their combined effect led to an 86.3% increase. AMF and Ge also improved proximate composition, with AMF-Ge interaction enhancing crude fiber and mineral content ( p < 0.05). Interestingly, AMF enhanced photosynthesis-related parameters (e.g., total chlorophyll) in Ge treated plants, which in turn increased carbohydrate accumulation. This accumulation could provide a route for the biosynthesis of amino acids, organic acids, and fatty acids, as evidenced by increased essential amino acid and organic acid levels. Consistently, the activity of key enzymes involved in amino acids biosynthesis (e.g., glutamine synthase (GS), methionine biosynthase (MS), lysine biosynthase (LS)) showed significant increments. Furthermore, AMF improved fatty acid levels, particularly unsaturated fatty acids in Ge-treated plants compared to the control. In addition, increased phenylalanine provided a precursor for the production of antioxidants (e.g., phenols and flavonoids), through the action of the enzyme phenylalanine ammonia-lyase (PAL), resulting in improved antioxidant activity gains as indicated by FRAP, ABTS, and DPPH assays. This study is the first to show that Ge enhances the beneficial effect of AMF on spinach, improving growth and nutritional quality, with promising implications for agricultural practices.

Heavy metal pollution is one of the major agronomic challenges. Tungsten (W) exposure leads to its accumulation in plants, which in turn reduces plant growth, inhibits photosynthesis and induces oxidative damage. In addition, the predicted increase in CO 2 could boost plant growth under both optimal and heavy metal stress conditions. The aim of the present study was to investigate the effect of W on growth, photosynthetic parameters, oxidative stress and redox status in rye plants under ambient and elevated (eCO 2 ) levels. To this end, rye plants were grown under the following conditions: ambient CO 2 (aCO 2 , 420 ppm), elevated CO 2 (eCO 2 , 720 ppm), W stress (350 mg kg -1 soil) and W+eCO 2 . W stress induced significant ( p < 0.05) decreases in growth and photosynthesis, increases in oxidative damages (lipid peroxidation) and the antioxidant defense system, i.e., ascorbate (ASC), reduced glutathione (GSH), GSH reductase (GR), peroxidase (POX), catalase (CAT), superoxide dismutase (SOD), ASC peroxide (APX) and dehydroascorbate reductase (DHAR). On the other hand, eCO 2 decreased W uptake and improved photosynthesis, which sequentially improved plant growth. The obtained results showed that eCO 2 can decrease the phytotoxicity risks of W in rye plants. This positive impact of eCO 2 on reducing the negative effects of soil W was related to their ability to enhance plant photosynthesis, which in turn provided energy and a carbon source for scavenging the reactive oxygen species (ROS) accumulation caused by soil W stress.

Increases in atmospheric CO 2 is known to promote plant growth under heavy metals stress conditions. However, vanadium (V) stress mitigating the impact of eCO 2 as well as the physiological and biochemical bases of this stress mitigation have not been well studied. To this end, this study investigated the growth, photosynthetic parameters, oxidative damages antioxidants, and antioxidants enzymes in wheat plants grown under ambient (420 PPM) and high eCO 2 (720 ppm) levels. Exposing wheat plants to higher V increased its accumulation in plants which consequentially inhibited plant growth and induced oxidative damage. An increase in antioxidant and detoxification defense systems was observed but it was not enough to reduce V stress toxicity. On the other hand, wheat growth was improved as a result of reduced V uptake and toxicity on photosynthesis under eCO 2 . To reduce V uptake, wheat accumulated citric acid, and oxalic acid in soil preferentially under both treatments but to more extend under V and eCO 2 . Additionally, improved photosynthesis induced high carbon availability that was directed to produce chelating proteins (metallothioneins, phytochelatin) and antioxidants (phenolics, flavonoids, total antioxidant capacity). This study advances our knowledge of the processes behind the variations in the physiological and biochemical responses of the wheat crop under V and eCO 2 conditions.