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We show that observed co-variations at sub-hourly time scales between the photochemical reflectance index (PRI) and canopy light use efficiency (LUE) over a Douglas-fir forest result directly from sub-hourly leaf reflectance changes in a 531 nm spectral window roughly 50 nm wide. We conclude then, that over a forest stand we are observing the direct effects of photosynthetic down-regulation on leaf-level reflectance at 531 nm. Key to our conclusion is our ability to simultaneously measure the LUE and reflectance of the Douglas-fir stand as a function of shadow fraction from the “hot spot” to the "dark spot" and a new finding herein, based on radiative transfer theory, that the magnitude of a normalized reflectance difference index (NDRI) such as PRI can vary with shadow fraction only in case the reflectance of the shaded and sunlit leaves differ in at least one of the NDRI bands.
Our spectrometer measurements over a nearly 6 month period show that at a forest stand scale, only two NDRIs (both containing a band near 570 nm) vary with shadow fraction and are correlated with LUE; an NDRI with a band centered at 531 nm roughly 50 nm wide, and another near 705 nm. Therefore, we are able to conclude that only these two bands' reflectance differ between the sunlit and the shaded elements of the canopy. Their reflectance changes on time scales of a few minutes or less. Our observations also show that the reflectance changes at 531 nm are more highly correlated with variations in canopy light use efficiency when only sunlit canopy elements are viewed (the hot spot), than when only shaded elements (the dark spot) are viewed. Taken together then, these results demonstrate that the observed sub-hourly changes in foliage reflectance at 531 nm and 705 nm can only result from corresponding variations in photosynthetic rates. The importance of our results are as follows: (1)We show that variations in PRI with LUE are a direct result of rapid changes in foliage reflectance at 531 nm resulting from photosynthetic down-regulation, and can be observed at forest scales. (2) Our findings also suggest a new sensor and methodology for the direct retrieval from space of changes in forest LUE by measuring PRI as a function of shadow fraction using a multi-angle spectrometer simultaneously retrieving both shadow fraction and PRI.