Abstract
Leaf anatomy varies with abiotic factors and is an important trait for understanding plant adaptive responses to environmental conditions. Leaf mass per area (LMA) is a key morphological trait and is related to leaf performance, such as light-saturated photosynthetic rate per leaf mass, leaf mechanical strength, and leaf lifespan. LMA is the multiplicative product of leaf thickness (LT) and leaf density (LD), both of which vary with leaf anatomy. Nevertheless, how LMA, LT, and LD covary with leaf anatomy is largely unexplored along natural environmental gradients. Slope aspect is a topographic factor that underlies variations in solar irradiation, air temperature, humidity, and soil fertility. In the present study, we examined (1) how leaf anatomy varies with different slope aspects and (2) how leaf anatomy is related to LMA, LD, and LT. Leaf anatomy was measured for 30 herbaceous species across three slope aspects (south-, west-, and north-facing slopes; hereafter, SFS, WFS, and NFS, respectively) in an eastern Tibetan subalpine meadow. For 18 of the 30 species, LMA data were available from previous studies. LD was calculated as LMA divided by LT. Among the slope aspects, the dominant species on the SFS exhibited the highest LTs with the thickest spongy mesophyll layers. The thicker spongy mesophyll layer was related to a lower LD via larger intercellular airspaces. In contrast, LD was the highest on NFS among the slope aspects. LMA was not significantly different among the slope aspects because higher LTs on SFS were effectively offset by lower LDs. These results suggest that the relationships between leaf anatomy and LMA were different among the slope aspects. Mechanisms underlying the variations in leaf anatomy may include different solar radiation, air temperatures, soil water, and nutrient availabilities among the slope aspects.