Chlorophyll a fluorescence (ChlF) has been used for decades to study the organization, functioning, and physiology of photosynthesis at the leaf and subcellular levels. ChlF is now measurable from remote sensing platforms. This provides a new optical means to track photosynthesis and gross primary productivity of terrestrial ecosystems. Importantly, the spatiotemporal and methodological context of the new applications is dramatically different compared with most of the available ChlF literature, which raises a number of important considerations. Although we have a good mechanistic understanding of the processes that control the ChlF signal over the short term, the seasonal link between ChlF and photosynthesis remains obscure. Additionally, while the current understanding of in vivo ChlF is based on pulse amplitude-modulated (PAM) measurements, remote sensing applications are based on the measurement of the passive solar-induced chlorophyll fluorescence (SIF), which entails important differences and new challenges that remain to be solved. In this review we introduce and revisit the physical, physiological, and methodological factors that control the leaf-level ChlF signal in the context of the new remote sensing applications. Specifically, we present the basis of photosynthetic acclimation and its optical signals, we introduce the physical and physiological basis of ChlF from the molecular to the leaf level and beyond, and we introduce and compare PAM and SIF methodology. Finally, we evaluate and identify the challenges that still remain to be answered in order to consolidate our mechanistic understanding of the remotely sensed SIF signal.

INTRODUCTION Reliable estimation of gross primary production (GPP) from landscape to global scales is pivotal to a wide range of ecological research areas, such as carbon-climate feedbacks, and agricultural applications, such as crop yield and drought monitoring. However, measuring GPP at these scales remains a major challenge. Solar-induced chlorophyll fluorescence (SIF) is a signal emitted directly from the core of photosynthetic machinery. SIF integrates complex plant physiological functions in vivo to reflect photosynthetic dynamics in real time. The advent of satellite SIF observation promises a new era in global photosynthesis research. The Orbiting Carbon Observatory-2 (OCO-2) SIF product is a serendipitous but critically complementary by-product of OCO-2’s primary mission target—atmospheric column CO 2 ( X CO 2 ). OCO-2 SIF removes some important roadblocks that prevent wide and in-depth applications of satellite SIF data sets and offers new opportunities for studying the SIF-GPP relationship and vegetation functional gradients at different spatiotemporal scales. RATIONALE Compared with earlier satellite missions with SIF capability, the OCO-2 SIF product has substantially improved spatial resolution, data acquisition, and retrieval precision. These improvements allow satellite SIF data to be validated, for the first time, directly against ground and airborne measurements and also used to investigate the SIF-GPP relationship and terrestrial ecosystem functional dynamics with considerably better spatiotemporal credibility. RESULTS Coordinated airborne measurements of SIF with the Chlorophyll Fluorescence Imaging Spectrometer (CFIS) were used to validate OCO-2 retrievals. The validation shows close agreement between OCO-2 and CFIS SIF, with a regression slope of 1.02 and R 2 of 0.71. Landscape gradients in SIF emission, corresponding to differences in vegetation types, were clearly delineated by OCO-2, a capability that was lacking in previous satellite missions. The SIF-GPP relationships at eddy covariance flux sites in the vicinity of OCO-2 orbital tracks were found to be more consistent across biomes than previously suggested. Finally, empirical orthogonal function (EOF) analyses on OCO-2 SIF and available GPP products show highly consistent spatiotemporal correspondence in their leading EOF modes across the globe, suggesting that SIF and GPP are governed by similar dynamics and controlled by similar environmental and biological conditions. CONCLUSION OCO-2 represents a major advance in satellite SIF remote sensing. Our analyses suggest that SIF is a powerful proxy for GPP at multiple spatiotemporal scales and that high-quality satellite SIF is of central importance to studying terrestrial ecosystems and the carbon cycle. Although the possibility of a universal SIF-GPP relationship across different biome types cannot be dismissed, in-depth process-based studies are needed to unravel the true nature of covariations between SIF and GPP. Of critical importance in such efforts are the potential coordinated dynamics between the light-use efficiencies of CO 2 assimilation and fluorescence emission in response to changes in climate and vegetation characteristics. Eventual synergistic uses of SIF with atmospheric CO 2 enabled by OCO-2 will lead to more reliable estimates of terrestrial carbon sources and sinks—when, where, why, and how carbon is exchanged between land and atmosphere—as well as a deeper understanding of carbon-climate feedbacks. The marked ecological gradients depicted by OCO-2’s high-resolution SIF measurements along a transect of temperate deciduous forests, crops, and urban area from Indiana to suburban Chicago, Illinois.

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The photochemical reflectance index (PRI) is regarded as a promising proxy to track the dynamics of photosynthetic light use efficiency (LUE) via remote sensing. The implementation of this approach requires the relationship between PRI and LUE to scale not only in space but also in time. The short-term relationship between PRI and LUE is well known and is based on the regulative process of non-photochemical quenching (NPQ), but at the seasonal timescale the mechanisms behind the relationship remain unclear. We examined to what extent sustained forms of NPQ, photoinhibition of reaction centres, seasonal changes in leaf pigment concentrations, or adjustments in the capacity of alternative energy sinks affect the seasonal relationship between PRI and LUE during the year in needles of boreal Scots pine. PRI and NPQ were highly correlated during most of the year but decoupled in early spring when the foliage was deeply downregulated. This phenomenon was attributed to differences in the physiological mechanisms controlling the seasonal dynamics of PRI and NPQ. Seasonal adjustments in the pool size of the xanthophyll cycle pigments, on a chlorophyll basis, controlled the dynamics of PRI, whereas the xanthophyll de-epoxidation status and other xanthophyll-independent mechanisms controlled the dynamics of NPQ at the seasonal timescale. We conclude that the PRI leads to an underestimation of NPQ, and consequently overestimation of LUE, under conditions of severe stress in overwintering Scots pine, and most likely also in species experiencing severe drought. This severe stress-induced decoupling may challenge the implementation of the PRI approach.

Partitioning of excitation energy between photochemical quenching (PQ) and non-photochemical quenching (NPQ) processes is constantly adjusted in the leaf in order to preserve the photosynthetic energy balance. Adjustments in PQ and NPQ often result from a combination of different temporal components that can be simplified into reversible and sustained components. While reversible PQ and NPQ are relatively well understood, the controls behind the sustained components of PQ and NPQ, or the interaction between sustained and reversible NPQ, remain elusive. In this study, I used a full year of high-resolution chlorophyll fluorescence (ChlF) data obtained with a Monitoring-PAM fluorometer (Walz, Effeltrich, Germany) in needles of boreal Pinus sylvestris in situ to quantitatively analyse the dynamics and interaction between temporal components of NPQ and PQ and their control by the environment. To enable the estimation of sustained and reversible components of PQ and NPQ, a number of key ChlF parameters were reviewed and adapted to the analysis of long-term monitoring data. Overall, NPQ was drastically enhanced during winter via the accumulation of sustained NPQ in a process regulated by air temperature. Reversible NPQ retained some functionality even at temperatures well below zero and was not inhibited by the presence of sustained NPQ per se but by low temperatures alone. This suggests that temporal NPQ components co-operate in an additive rather than complementary fashion, conferring additional flexibility to the photoprotective role of NPQ. Finally, the potential of the sustained photochemical quenching parameter (qL(s) ) to track photoinhibition in situ was discussed.