Foliar-applied manganese and phosphorus in deficient barley: Linking absorption pathways and leaf nutrient status.

Foliar fertilization delivers essential nutrients directly to plant tissues, reducing excessive soil fertilizer applications that can lead to eutrophication following nutrient leaching. Foliar nutrient absorption is a dynamic process affected by leaf surface structure and composition, plant nutrient status, and ion physicochemical properties. We applied multiple methods to study the foliar absorption behaviors of manganese (Mn) and phosphorus (P) in nutrient-deficient spring barley (Hordeum vulgare) at two growth stages. Nutrient-specific chlorophyll a fluorescence assays were used to visualize leaf nutrient status, while laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS) was used to visualize foliar absorption pathways for P and Mn ions. Rapid Mn absorption was facilitated by a relatively thin cuticle with a low abundance of waxes and a higher stomatal density in Mn-deficient plants. Following absorption, Mn accumulated in epidermal cells and in the photosynthetically active mesophyll, enabling a fast (6 h) restoration of Mn-dependent photosynthetic processes. Conversely, P-deficient plants developed thicker cuticles and epidermal cell walls, which reduced the penetration of P across the leaf surface. Foliar-applied P accumulated in trichomes and fiber cells above leaf veins without reaching the mesophyll and, as a consequence, no restoration of P-dependent photosynthetic processes was observed. This study reveals new links between leaf surface morphology, foliar-applied ion absorption pathways, and the restoration of affected physiological processes in nutrient-deficient leaves. Understanding that ions may have different absorption pathways across the leaf surface is critical for the future development of efficient fertilization strategies for crops in nutrient-limited soils.