Subcellular distribution and chemical forms involved in manganese accumulation and detoxification for Xanthium strumarium L.

Xanthium strumarium L. is a candidate species for manganese (Mn)-phyto-remediation. To reveal the mechanism of this species adaptive to Mn stress, the growth, Mn subcellular distribution, chemical forms, as well as micro-structure and ultra-structure responses of the mining ecotype (ME) of X. strumarium to Mn stress were studied with the non-mining ecotype (NME) as the reference by a hydroponic experiment. The results showed the ME demonstrated a higher tolerance to Mn stress with a superior growth and a higher tolerance index (TI) when compared with the NME. The concentrations of Mn in leaves, stems, and roots of the ME were 1.1-1.8, 1.2-1.9, and 1.3-1.9 times higher than those in the corresponding organs of the NME, respectively. The micro-structure and ultra-structure showed abnormal alterations, such as shrunken ducts and sieve canals, round-shaped chloroplasts, increased starch and osmiophilic granules, as well as expanded and non-compact granum thylakoids in the NME, compared to the ME. More than 83% of Mn was localized in cell wall and soluble fraction, while the Mn concentration in all fractions had a direct linear relationship with Mn treatment in the ME. The proportions of pectates and protein integrated-Mn, phosphate-Mn, and oxalate-Mn forms were dominant in leaves and stems of the ME, whereas, in the NME the relative proportions of inorganic Mn and water-soluble Mn forms in the roots was higher than the other forms. Altogether, the combination of preferential distribution of Mn in the cell wall and soluble fraction, and storage of Mn in low toxicity forms, such as phosphate-Mn, pectates and protein-bound Mn, and oxalate-Mn, might be responsible for alleviating Mn toxicity in the ME.