Exogenous Caffeine (1,3,7-Trimethylxanthine) Application Diminishes Cadmium Toxicity by Modulating Physio-Biochemical Attributes and Improving the Growth of Spinach ( Spinacia oleracea L.)

Leafy vegetables usually absorb and retain heavy metals more readily than most of the other crop plants, and thus contribute ≥70% of the total cadmium (Cd) intake of humans. Caffeine mediates plant growth and has proved to be beneficial against pathogens and insects. Therefore, it was hypothesized that foliar applications of caffeine could alter metabolism and reduce Cd toxicity in spinach ( Spinacia oleracea L.). Seven-day old spinach seedlings were provided with Cd (0, 50, and 100 µM) stress. Caffeine (0, 5, or 10 mM) foliar spray was given twice at after 20 days of seeds germination with an interval of one week. In results, Cd stress reduced photosynthetic pigments biosynthesis, increased oxidative stress, imbalanced nutrient retention, and inhibited plant growth. On the other hand, the caffeine-treated spinach plants showed better growth owing to the enhanced biosynthesis of chlorophylls, better oxidative defense systems, and lower accumulation and transport of Cd within the plant tissues. Furthermore, caffeine application enhanced the accumulation of the proline and ascorbic acid, but reduced MDA and H 2 O 2 contents and Cd in plant leaves, and ultimately improved mineral nutrition of spinach plants exposed to different Cd regimes. In conclusion, exogenous application of caffeine significantly diminishes Cd stress by modulating physiological, biochemical, and growth attributes of spinach ( Spinacia oleracea L.)