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Abstract Pigment dynamics in temperate evergreen forests remain poorly characterized, despite their year-round photosynthetic activity and importance for carbon cycling. Developing rapid, nondestructive methods to estimate pigment composition enables high-throughput assessment of plant acclimation states. In this study, we investigate the seasonality of eight chlorophyll and carotenoid pigments and hyperspectral reflectance data collected at both the needle (400–2400 nm) and canopy (420–850 nm) scales in Pinus palustris (longleaf pine) at the Ordway Swisher Biological Station in north-central Florida, USA. Needle spectra were obtained at three distinct times throughout the year, while tower-based spectra were collected continuously over a nine-month period. Seasonal trends in photoprotective pigments (e.g., lutein and xanthophylls) and photosynthetic pigments (e.g., chlorophylls) aligned closely with seasonal changes in photosynthetically active radiation and gross primary productivity. To track inter-tree and seasonal variability in pigment pools with hyperspectral reflectance data, we used correlation analyses and ridge regression models. Ridge regression models using the full hyperspectral range outperformed predictions using standard linear regression with specific wavelengths in a normalized difference index fashion. Ridge regression successfully predicted all pigment pools (R2 > 0.5) with comparable accuracy at both the needle and canopy scales. The models performed best for lutein, neoxanthin, antheraxanthin, and chlorophyll a and b - which had greater inter-tree and seasonal variation - and achieved moderate accuracy for violaxanthin, alpha-carotene, and beta-carotene. These results provide a foundation for scaling biochemical traits from ground-based sensors to airborne and satellite platforms, particularly in ecosystems with subtle changes in pigment dynamics. 

Abstract The seasonal timing and magnitude of photosynthesis in evergreen needleleaf forests (ENFs) has major implications for the carbon cycle and is increasingly sensitive to changing climate. Earlier spring photosynthesis can increase carbon uptake over the growing season or cause early water reserve depletion that leads to premature cessation and increased carbon loss. Determining the start and the end of the growing season in ENFs is challenging due to a lack of field measurements and difficulty in interpreting satellite data, which are impacted by snow and cloud cover, and the pervasive “greenness” of these systems. We combine continuous needle‐scale chlorophyll fluorescence measurements with tower‐based remote sensing and gross primary productivity (GPP) estimates at three ENF sites across a latitudinal gradient (Colorado, Saskatchewan, Alaska) to link physiological changes with remote sensing signals during transition seasons. We derive a theoretical framework for observations of solar‐induced chlorophyll fluorescence (SIF) and solar intensity‐normalized SIF (SIF_relative) under snow‐covered conditions, and show decreased sensitivity compared with reflectance data (~20% reduction in measured SIF vs. ~60% reduction in near‐infrared vegetation index [NIRv] under 50% snow cover). Needle‐scale fluorescence and photochemistry strongly correlated (r² = 0.74 in Colorado, 0.70 in Alaska) and showed good agreement on the timing and magnitude of seasonal transitions. We demonstrate that this can be scaled to the site level with tower‐based estimates of LUE_Pand SIF_relativewhich were well correlated across all sites (r² = 0.70 in Colorado, 0.53 in Saskatchewan, 0.49 in Alaska). These independent, temporally continuous datasets confirm an increase in physiological activity prior to snowmelt across all three evergreen forests. This suggests that data‐driven and process‐based carbon cycle models which assume negligible physiological activity prior to snowmelt are inherently flawed, and underscores the utility of SIF data for tracking phenological events. Our research probes the spectral biology of evergreen forests and highlights spectral methods that can be applied in other ecosystems. 

Abstract Evergreen needleleaf forests (ENFs) play a sizable role in the global carbon cycle, but the biological and physical controls on ENF carbon cycle feedback loops are poorly understood and difficult to measure. To address this challenge, a growing appreciation for the stress physiology of photosynthesis has inspired emerging techniques designed to detect ENF photosynthetic activity with optical signals. This Overview summarizes how fundamental plant biological and biophysical processes control the fate of photons from leaf to globe, ultimately enabling remote estimates of ENF photosynthesis. We demonstrate this using data across four ENF sites spanning a broad range of environmental conditions and link leaf- and stand-scale observations of photosynthesis (i.e., needle biochemistry and flux towers) with tower- and satellite-based remote sensing. The multidisciplinary nature of this work can serve as a model for the coordination and integration of observations made at multiple scales. 

Summary A new proliferation of optical instruments that can be attached to towers over or within ecosystems, or ‘proximal’ remote sensing, enables a comprehensive characterization of terrestrial ecosystem structure, function, and fluxes of energy, water, and carbon. Proximal remote sensing can bridge the gap between individual plants, site‐level eddy‐covariance fluxes, and airborne and spaceborne remote sensing by providing continuous data at a high‐spatiotemporal resolution. Here, we review recent advances in proximal remote sensing for improving our mechanistic understanding of plant and ecosystem processes, model development, and validation of current and upcoming satellite missions. We provide current best practices for data availability and metadata for proximal remote sensing: spectral reflectance, solar‐induced fluorescence, thermal infrared radiation, microwave backscatter, and LiDAR. Our paper outlines the steps necessary for making these data streams more widespread, accessible, interoperable, and information‐rich, enabling us to address key ecological questions unanswerable from space‐based observations alone and, ultimately, to demonstrate the feasibility of these technologies to address critical questions in local and global ecology. 

Abstract We examined the seasonality of photosynthesis in 46 evergreen needleleaf (evergreen needleleaf forests (ENF)) and deciduous broadleaf (deciduous broadleaf forests (DBF)) forests across North America and Eurasia. We quantified the onset and end (Start_GPPand End_GPP) of photosynthesis in spring and autumn based on the response of net ecosystem exchange of CO₂to sunlight. To test the hypothesis that snowmelt is required for photosynthesis to begin, these were compared with end of snowmelt derived from soil temperature. ENF forests achieved 10% of summer photosynthetic capacity ∼3 weeks before end of snowmelt, while DBF forests achieved that capacity ∼4 weeks afterward. DBF forests increased photosynthetic capacity in spring faster (1.95% d⁻¹) than ENF (1.10% d⁻¹), and their active season length (End_GPP–Start_GPP) was ∼50 days shorter. We hypothesized that warming has influenced timing of the photosynthesis season. We found minimal evidence for long‐term change in Start_GPP, End_GPP, or air temperature, but their interannual anomalies were significantly correlated. Warmer weather was associated with earlier Start_GPP(1.3–2.5 days °C⁻¹) or later End_GPP(1.5–1.8 days °C⁻¹, depending on forest type and month). Finally, we tested whether existing phenological models could predict Start_GPPand End_GPP. For ENF forests, air temperature‐ and daylength‐based models provided best predictions for Start_GPP, while a chilling‐degree‐day model was best for End_GPP. The root mean square errors (RMSE) between predicted and observed Start_GPPand End_GPPwere 11.7 and 11.3 days, respectively. For DBF forests, temperature‐ and daylength‐based models yielded the best results (RMSE 6.3 and 10.5 days). 

Abstract More frequent and severe droughts are driving increased forest mortality around the globe. We urgently need to describe and predict how drought affects forest carbon cycling and identify thresholds of environmental stress that trigger ecosystem collapse. Quantifying the effects of drought at an ecosystem level is complex because dynamic climate–plant relationships can cause rapid and/or prolonged shifts in carbon balance. We employ the CARbon DAta MOdel fraMework (CARDAMOM) to investigate legacy effects of drought on forest carbon pools and fluxes. Our Bayesian model‐data fusion approach uses tower observed meteorological forcing and carbon fluxes to determine the response and sensitivity of aboveground and belowground ecological processes associated with the 2012–2015 California drought. Our study area is a mid‐montane mixed conifer forest in the Southern Sierras. CARDAMOM constrained with gross primary productivity (GPP) estimates covering 2011–2017 show a ~75% reduction in GPP, compared to negligible GPP change when constrained with 2011 only. Precipitation across 2012–2015 was 45% (474 mm) lower than the historical average and drove a cascading depletion in soil moisture and carbon pools (foliar, labile, roots, and litter). Adding 157 mm during an especially stressful year (2014, annual rainfall = 293 mm) led to a smaller depletion of water and carbon pools, steering the ecosystem away from a state of GPP tipping‐point collapse to recovery. We present novel process‐driven insights that demonstrate the sensitivity of GPP collapse to ecosystem foliar carbon and soil moisture states—showing that the full extent of GPP response takes several years to arise. Thus, long‐term changes in soil moisture and carbon pools can provide a mechanistic link between drought and forest mortality. Our study provides an example for how key precipitation threshold ranges can influence forest productivity, making them useful for monitoring and predicting forest mortality events. 

Abstract Robust carbon monitoring systems are needed for land managers to assess and mitigate the changing effects of ecosystem stress on western United States forests, where most aboveground carbon is stored in mountainous areas. Atmospheric carbon uptake via gross primary productivity (GPP) is an important indicator of ecosystem function and is particularly relevant to carbon monitoring systems. However, limited ground-based observations in remote areas with complex topography represent a significant challenge for tracking regional-scale GPP. Satellite observations can help bridge these monitoring gaps, but the accuracy of remote sensing methods for inferring GPP is still limited in montane evergreen needleleaf biomes, where (a) photosynthetic activity is largely decoupled from canopy structure and chlorophyll content, and (b) strong heterogeneity in phenology and atmospheric conditions is difficult to resolve in space and time. Using monthly solar-induced chlorophyll fluorescence (SIF) sampled at ∼4 km from the TROPOspheric Monitoring Instrument (TROPOMI), we show that high-resolution satellite-observed SIF followed ecological expectations of seasonal and elevational patterns of GPP across a 3000 m elevation gradient in the Sierra Nevada mountains of California. After accounting for the effects of high reflected radiance in TROPOMI SIF due to snow cover, the seasonal and elevational patterns of SIF were well correlated with GPP estimates from a machine-learning model (FLUXCOM) and a land surface model (CLM5.0-SP), outperforming other spectral vegetation indices. Differences in the seasonality of TROPOMI SIF and GPP estimates were likely attributed to misrepresentation of moisture limitation and winter photosynthetic activity in FLUXCOM and CLM5.0 respectively, as indicated by discrepancies with GPP derived from eddy covariance observations in the southern Sierra Nevada. These results suggest that satellite-observed SIF can serve as a useful diagnostic and constraint to improve upon estimates of GPP toward multiscale carbon monitoring systems in montane, evergreen conifer biomes at regional scales. 

Abstract Located at northern latitudes and subject to large seasonal temperature fluctuations, boreal forests are sensitive to the changing climate, with evidence for both increasing and decreasing productivity, depending upon conditions. Optical remote sensing of vegetation indices based on spectral reflectance offers a means of monitoring vegetation photosynthetic activity and provides a powerful tool for observing how boreal forests respond to changing environmental conditions. Reflectance‐based remotely sensed optical signals at northern latitude or high‐altitude regions are readily confounded by snow coverage, hampering applications of satellite‐based vegetation indices in tracking vegetation productivity at large scales. Unraveling the effects of snow can be challenging from satellite data, particularly when validation data are lacking. In this study, we established an experimental system in Alberta, Canada including six boreal tree species, both evergreen and deciduous, to evaluate the confounding effects of snow on three vegetation indices: the normalized difference vegetation index (NDVI), the photochemical reflectance index (PRI), and the chlorophyll/carotenoid index (CCI), all used in tracking vegetation productivity for boreal forests. Our results revealed substantial impacts of snow on canopy reflectance and vegetation indices, expressed as increased albedo, decreased NDVI values and increased PRI and CCI values. These effects varied among species and functional groups (evergreen and deciduous) and different vegetation indices were affected differently, indicating contradictory, confounding effects of snow on these indices. In addition to snow effects, we evaluated the contribution of deciduous trees to vegetation indices in mixed stands of evergreen and deciduous species, which contribute to the observed relationship between greenness‐based indices and ecosystem productivity of many evergreen‐dominated forests that contain a deciduous component. Our results demonstrate confounding and interacting effects of snow and vegetation type on vegetation indices and illustrate the importance of explicitly considering snow effects in any global‐scale photosynthesis monitoring efforts using remotely sensed vegetation indices. 

Abstract Although our observing capabilities of solar‐induced chlorophyll fluorescence (SIF) have been growing rapidly, the quality and consistency of SIF datasets are still in an active stage of research and development. As a result, there are considerable inconsistencies among diverse SIF datasets at all scales and the widespread applications of them have led to contradictory findings. The present review is the second of the two companion reviews, and data oriented. It aims to (1) synthesize the variety, scale, and uncertainty of existing SIF datasets, (2) synthesize the diverse applications in the sector of ecology, agriculture, hydrology, climate, and socioeconomics, and (3) clarify how such data inconsistency superimposed with the theoretical complexities laid out in (Sun et al., 2023) may impact process interpretation of various applications and contribute to inconsistent findings. We emphasize that accurate interpretation of the functional relationships between SIF and other ecological indicators is contingent upon complete understanding of SIF data quality and uncertainty. Biases and uncertainties in SIF observations can significantly confound interpretation of their relationships and how such relationships respond to environmental variations. Built upon our syntheses, we summarize existing gaps and uncertainties in current SIF observations. Further, we offer our perspectives on innovations needed to help improve informing ecosystem structure, function, and service under climate change, including enhancing in‐situ SIF observing capability especially in “data desert” regions, improving cross‐instrument data standardization and network coordination, and advancing applications by fully harnessing theory and data. 

Abstract Solar‐induced chlorophyll fluorescence (SIF) is a remotely sensed optical signal emitted during the light reactions of photosynthesis. The past two decades have witnessed an explosion in availability of SIF data at increasingly higher spatial and temporal resolutions, sparking applications in diverse research sectors (e.g., ecology, agriculture, hydrology, climate, and socioeconomics). These applications must deal with complexities caused by tremendous variations in scale and the impacts of interacting and superimposing plant physiology and three‐dimensional vegetation structure on the emission and scattering of SIF. At present, these complexities have not been overcome. To advance future research, the two companion reviews aim to (1) develop an analytical framework for inferring terrestrial vegetation structures and function that are tied to SIF emission, (2) synthesize progress and identify challenges in SIF research via the lens of multi‐sector applications, and (3) map out actionable solutions to tackle these challenges and offer our vision for research priorities over the next 5–10 years based on the proposed analytical framework. This paper is the first of the two companion reviews, and theory oriented. It introduces a theoretically rigorous yet practically applicable analytical framework. Guided by this framework, we offer theoretical perspectives on three overarching questions: (1)The forward (mechanism) question—How are the dynamics of SIF affected by terrestrial ecosystem structure and function? (2)The inference question: What aspects of terrestrial ecosystem structure, function, and service can be reliably inferred from remotely sensed SIF and how? (3)The innovation question: What innovations are needed to realize the full potential of SIF remote sensing for real‐world applications under climate change? The analytical framework elucidates that process complexity must be appreciated in inferring ecosystem structure and function from the observed SIF; this framework can serve as a diagnosis and inference tool for versatile applications across diverse spatial and temporal scales.