Citrate, oxalate, polyamines and other ligand forming metabolites can chelate heavy metals reducing their effective concentration. Ascorbate and polyphenols (like vanillin and other lignin derived compounds) are good at protecting against oxidative stress that can be caused by the heavy metals.
One primary metabolite that has been shown to be effective in mitigating heavy metal stress in plants is proline. Proline is an amino acid that is synthesized in response to various abiotic stresses, including heavy metal stress. Proline helps to maintain cell membrane stability, regulate the osmotic balance, and scavenge free radicals generated by heavy metal stress.
When plants are exposed to heavy metals, proline accumulates in the cytoplasm and helps to protect the plant cells from the toxic effects of heavy metals. Proline can also chelate heavy metals, reducing their toxicity and facilitating their transport out of the plant cells.
Studies have shown that exogenous application of proline can enhance the tolerance of plants to heavy metal stress. For example, when wheat plants were treated with cadmium, the addition of exogenous proline reduced the accumulation of cadmium in the roots and leaves, and improved plant growth and photosynthetic activity.
In addition to proline, other primary metabolites, such as organic acids and amino acids, can also play a role in mitigating heavy metal stress in plants. For example, citric acid and malic acid have been shown to chelate heavy metals and facilitate their transport out of plant cells. Amino acids such as cysteine and methionine can also chelate heavy metals and prevent their accumulation in plant tissues.