Intra-Plant Variation in Leaf Dry Mass per Area (LMA): Effects of Leaf–Shoot Orientation and Vertical Position on Dry Mass and Area Scaling

The intra-plant plasticity of leaves plays a vital role in enabling plants to adapt to changing climatic conditions. However, limited research has investigated the extent of intra-plant leaf trait variation and leaf biomass allocation strategies in herbaceous plants. To address this gap, we collected a total of 1746 leaves from 217 Lamium barbatum Siebold and Zucc. plants and measured their leaf dry mass (M) and leaf area (A). Leaves were categorized by vertical position (upper vs. lower canopy layer) and leaf–shoot orientation (east, south, west, north). ANOVA with Tukey’s HSD test was used to compare differences in M, A, and leaf dry mass per unit area (LMA). Reduced major axis regression was employed to evaluate the scaling relationship between M and A, and the bootstrap percentile method was used to determine differences in scaling exponents. The data indicated that: (i) M, A, LMA, and the scaling exponents of M versus A did not differ significantly among leaf–shoot orientations, and (ii) lower layer leaves exhibited significantly greater M, A, and LMA than upper layer leaves, but their scaling exponents were significantly smaller. These findings highlight that plant vertical growth brings significant intra-plant plasticity in leaf traits and their scaling relationships in herbaceous plants. This plasticity differs from that observed in trees, but is also critical for balancing weight load and optimizing light-use efficiency, potentially enhancing stress resilience in herbaceous plants.