Abstract:The photochemical reflectance index (PRI) is a proxy for light use efficiency (LUE), and is used in remote sensing to measure plant stress and photosynthetic downregulation in plant canopies. It is known to depend on local light conditions within a canopy indicating non-photosynthetic quenching of incident radiation. Additionally, when measured from a distance, canopy PRI depends on shadow fraction-the fraction of shaded foliage in the instantaneous field of view of the sensor-due to observation geometry. Our aim is to quantify the extent to which sunlit fraction alone can describe variations in PRI so that it would be possible to correct for its variation and identify other possible factors affecting the PRI-sunlit fraction relationship. We used a high spatial and spectral resolution Aisa Eagle airborne imaging spectrometer above a boreal Scots pine site in Finland (Hyytiälä forest research station, 61 • 50 N, 24 • 17 E), with the sensor looking in nadir and tilted (off-nadir) directions. The spectral resolution of the data was 4.6 nm, and the spatial resolution was 0.6 m. We compared the PRI for three different scatter angles (β = 19 • , 55 • and 76 • , defined as the angle between sensor and solar directions) at the forest stand level, and observed a small (0.006) but statistically significant (p < 0.01) difference in stand PRI. We found that stand mean PRI was not a direct function of sunlit fraction. However, for each scatter angle separately, we found a clear non-linear relationship between PRI and sunlit fraction. The relationship was systematic and had a similar shape for all of the scatter angles. As the PRI-sunlit fraction curves for the different scatter angles were shifted with respect to each other, no universal curve could be found causing the observed independence of canopy PRI from the average sunlit fraction of each view direction. We found the shifts of the curves to be related to a leaf structural effect on canopy scattering: the ratio of needle spectral reflectance to transmittance. We demonstrate that modeling PRI-sunlit fraction relationships using high spatial resolution imaging spectroscopy data is suitable and needed in order to quantify PRI variations over forest canopies.

BackgroundThe Photochemical Reflectance Index (PRI) calculated from narrow-band spectral reflectance data is a vegetation index which is increasingly used as an indicator of photosynthetic activity. The leaf-level link between the status of photosynthetic apparatus and PRI has been robustly established under controlled light conditions. However, when a whole canopy is measured instantaneously, the PRI signal is heavily modified by vegetation structure and local variations in incident light conditions. To apply PRI for monitoring the photosynthesis of whole canopies under natural conditions, these large-scale measurements need to be validated against simultaneous leaf PRI. Unfortunately, PRI changes dynamically with incident light and has a large natural variation. No generally accepted procedure exists today for determining the PRI of canopy elements in situ.ResultsWe present a successful procedure for in situ measurements of needle PRI. We describe, characterize and test an optical measurement protocol and demonstrate its applicability in field conditions. The measurement apparatus consisted of a light source, needle clip, spectroradiometer and a controlling computer. The light level inside the clip was approximately two-thirds of that on sunlit needle surfaces at midday. During each measurement the needle was inserted into the clip for approximately 5 s. We found no near-instantaneous changes (sub-second scale jumps) in PRI during the measurements. The time constants for PRI variation in light to full shade acclimations were approximately 10 s. The procedure was successfully applied to monitor the greening-up of Scots pine trees. We detected both facultative (diurnal) PRI changes of 0.02 (unitless) and constitutive (seasonal) variations of 0.1. In order to reliably detect the facultative PRI change of 0.02, 20 needles need to be sampled from both sunlit and shaded locations.ConclusionsWe established a robust procedure for irradiance-dependent leaf (needle) PRI measurements, facilitating empirical scaling of PRI from leaf (needle) to full canopy level and the application of PRI to monitoring the changes in highly structured vegetation. The measured time constants, and facultative and constitutive PRI variations support the use of an artificial light for in situ PRI measurements at leaf (needle) level.

Abstract:The photochemical reflectance index (PRI) is calculated from vegetation narrowband reflectance in two bands in the visible part of the spectrum. Variations in PRI are associated with changes in the xanthophyll cycle pigments which regulate the light use efficiency of vegetation. Correlations have been found between remotely-sensed PRI and various photosynthetic productivity parameters at the scales from leaves to landscapes. Environmental satellites can provide only an instantaneous value of this index at the time of overpass. The diurnal course of needle (leaf) PRI needs to be known in order to link the instantaneous values robustly with photosynthetic parameters at time scales exceeding one day. This information is not currently available in the scientific literature. Here we present the daily cycle of Scots pine needle and canopy PRI in a southern boreal forest zone in the presence of direct solar radiation during the peak growing season of two consecutive years. We found the PRI of the needles which are exposed to direct radiation to have a distinct diurnal cycle with constant or slightly increasing values before noon and a daily minimum in the afternoon. The cycle in needle PRI was not correlated with that in the incident photosynthetic photon flux density (PPFD). However, when PPFD was above 1000 µmol m −2 s −1 , approximately between 8 a.m. and 5 p.m., needle PRI was correlated with the light use efficiency (LUE), measured with shoot chambers. The timing of the minimum needle PRI coincided with the minimum canopy value, as measured by an independent sensor above the canopy, but the correlation between the two variables was not significant. Our field results corroborate the applicability of needle PRI in monitoring the daily variation in LUE. However, to apply this to remote sensing of seasonal photosynthetic productivity, the daily cycle of leaf PRI needs to be known for the specific vegetation type.

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The spectral properties of plant leaves relate to the state of their photosynthetic apparatus and the surrounding environment. An example is the well known photosynthetic downregulation, active on the time scale from minutes to hours, caused by reversible changes in the xanthophyll cycle pigments. These changes affect leaf spectral absorption and are frequently quantified using the photochemical reflectance index (PRI). This index can be used to remotely monitor the photosynthetic status of vegetation, and allows for a global satellite-based measurement of photosynthesis. Such earth observation satellites in near-polar orbits usually cover the same geographical location at the same local solar time at regular intervals. To facilitate the interpretation of these instantaneous remote PRI measurements and upscale them temporally, we measured the daily course of leaf PRI in two evergreen biomes-a European boreal forest and an Amazon rainforest. The daily course of PRI was different for the two locations: At the Amazonian forest, the PRI of Manilkara elata leaves was correlated with the average photosynthetic photon flux density (PPFD) (R 2 = 0.59, p < 0.01) of the 40 minutes preceding the leaf measurement. In the boreal location, the variations in Pinus sylvestris needle PRI were only weakly (R 2 = 0.27, p < 0.05) correlated with mean PPFD of the preceding two hours; for Betula pendula, the correlation was insignificant (p > 0.5) regardless of the averaging period. The measured daily PRI curves were specific to species and/or environmental conditions. Hence, for a proper

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