Manganese accumulation in leaves of Hakea prostrata (Proteaceae) and the significance of cluster roots for micronutrient uptake as dependent on phosphorus supply

When grown at a low P supply, Hakea prostrata R.Br. (Proteaceae) develops dense clusters of determinate branch roots, termed 'proteoid' or 'cluster' roots and accumulates Mn in its leaves. The aim of this study was to vary the production of cluster roots and assess the relationship between Mn uptake and cluster-root mass. We collected native soil from a location inhabited by H. prostrata and amended this with 'high' and 'low' amounts of insoluble or soluble P. After 14 months, we measured the impact of the treatments on cluster-root development and the [P], [Mn], [Fe], [Zn] and [Cu] in young (expanding) and mature leaves. Dry mass and leaf area increased with increasing P availability in the soil, but growth decreased at the highest soluble [P], which caused symptoms of P toxicity. The [P] in young leaves (1.3-2.7 mg g(-1) DM) exceeded that in older leaves (0.28-0.85 mg g(-1) DM), except when plants were grown with soluble P (3.2-21 mg g(-1) DM). Cluster-root formation was inhibited when leaf [P] increased; [P] in young leaves, rather than that in old leaves, appeared to be the factor that determined the proportion of the root mass invested in cluster roots. Old leaves of all treatments had [Mn] from 90 to 120 mu g g(-1) DM, except for plants grown at high levels of soluble P, when [Mn] decreased below 30 mu g g(-1) DM. The [Mn] and [Zn] in old leaves and the [Cu] in young leaves were positively correlated with the fraction of roots invested in cluster roots. These findings support our hypothesis that cluster roots play a significant role in micronutrient acquisition, and also provide an explanation for Mn accumulation in leaves of H. prostrata, and presumably Proteaceae in general.