A theoretical analysis of the influence of heterogeneity in chlorophyll distribution on leaf reflectance.

Attempts to determine the vitality of vegetation and to detect vegetation stress from remotely sensed data have focused on chlorophyll concentration, because it influences the reflectance of vegetation and tends to correlate with vegetation health and stress. Pollution, pathogens and pests can cause localized regions of chlorosis and necrosis across a leaf surface, but the extent to which these patches influence the overall reflectance and spectral signature of the leaf and canopy has not been tested. A conifer leaf model (LIBERTY), which simulates the influence of leaf biochemical concentrations of chlorophyll, water, lignin, cellulose and protein on the reflectance of leaves from 400 to 2500 nm, was used to determine the effect of patches of chlorosis on leaf reflectance. A fraction of the leaf f is assumed to be chlorotic with a chlorophyll concentration C(1). The remainder of the leaf has chlorophyll concentration C(2) such that mean leaf chlorophyll concentration, C(mean) = fC(1) + (1 - f)C(2), is constant for a range of f and C(1) values. LIBERTY can be used to estimate the reflectance of a leaf with a particular chlorophyll concentration at a particular wavelength R(lambda,C) (assuming other leaf properties remain constant), thus we can estimate the reflectance of the chlorotic leaf as fR(lambda,C(1))+ (1 - f)R(lambda,C(2)). The model indicated that small areas of chlorosis have a disproportionately large influence on overall leaf reflectance. For example, a leaf with 25% of its area chlorotic can have the same reflectance (400-700 nm) as a homogeneous leaf with 60% less chlorophyll. Thus, determination of chlorophyll concentration from remotely sensed data is prone to underestimation when chlorophyll is nonuniformly distributed. Hence, attempts to model leaf and canopy reflectance using radiative transfer models will need to consider how to incorporate nonuniform chlorophyll distribution.