Ecosystem studies often consider the co‐benefits of biodiversity and carbon sequestration, but these carbon–biodiversity links can be complex and multifaceted. Recent findings in forest ecosystems ...emphasize the importance of looking beyond single trophic levels and the more visible, above‐ground portions to consider the full range of relationships between all ecosystem components when evaluating carbon sequestration potential. Simple engineered solutions to carbon storage based on monocultures that fail to consider all costs and benefits may be deceiving and lead to inappropriate management practices. Regenerating natural ecosystems may best enhance the co‐benefits of carbon sequestration and biodiversity.
Conceptually, plant functional types represent a classification scheme between species and broad vegetation types. Historically, these were based on physiological, structural and/or phenological ...properties, whereas recently, they have reflected plant responses to resources or environmental conditions. Often, an underlying assumption, based on an economic analogy, is that the functional role of vegetation can be identified by linked sets of morphological and physiological traits constrained by resources, based on the hypothesis of functional convergence. Using these concepts, ecologists have defined a variety of functional traits that are often context dependent, and the diversity of proposed traits demonstrates the lack of agreement on universal categories. Historically, remotely sensed data have been interpreted in ways that parallel these observations, often focused on the categorization of vegetation into discrete types, often dependent on the sampling scale. At the same time, current thinking in both ecology and remote sensing has moved towards viewing vegetation as a continuum rather than as discrete classes. The capabilities of new remote sensing instruments have led us to propose a new concept of optically distinguishable functional types ('optical types') as a unique way to address the scale dependence of this problem. This would ensure more direct relationships between ecological information and remote sensing observations.
A primary focus of this short communication is to show how the operational definition of light use efficiency (LUE) influences the results and interpretation of the LUE model. Our study was motivated ...by the observation that multiple LUE definitions are reported in the literature. The temporal behavior of three operational definitions of LUE, based on (i) incident radiation, (ii) total absorbed radiation and (iii) radiation absorbed by photosynthetically active/green vegetation was examined for two contrasting crops (soybean and maize) having different physiologies, leaf structures and canopy architectures. Over the course of a growing season, the behavior of these three contrasting LUE definitions was strikingly dissimilar, and the degree of dissimilarity varied with contrasting crops (corn and soybean). This demonstrates that LUE model behavior would vary strongly with the LUE definition used, with resulting implications both for the estimated seasonal productivity, and for the interpretation of the underlying mechanism. Based on these findings, we recommend a standard definition of the LUE model based on radiation absorbed by green vegetation. We also discuss the practical and theoretical implications of using this simple conceptual model on a dynamic biological system.
•Temporal behavior of three operational definitions of LUE in two crops was studied.•The behavior of these three contrasting LUE definitions was strikingly dissimilar.•GPP estimates and LUE model behavior varied strongly with the LUE definition used.•LUE based on radiation absorbed by green vegetation is recommended.
The photochemical reflectance index (PRI) reflects diurnal xanthophyll cycle activity and is also influenced by seasonally changing carotenoid : Chl pigment ratios. Both changing pigment pools and ...xanthophyll cycle activity contribute to photoprotection in evergreen conifers exposed to boreal winters, but they operate over different timescales, and their relative contribution to the PRI signal has often been unclear. To clarify these responses and their contribution to the PRI signal, leaf PRI, pigment composition, temperature and irradiance were monitored over 2 yr for two evergreen conifers (Pinus contorta and Pinus ponderosa) in a boreal climate. PRI was affected by three distinct processes operating over different timescales and exhibiting contrasting spectral responses. Over the 2 yr study period, the greatest change in PRI resulted from seasonally changing carotenoid : Chl pigment ratios, followed by a previously unreported shifting leaf albedo during periods of deep cold. Remarkably, the smallest change was attributable to the xanthophyll cycle. To properly distinguish these three effects, interpretation of PRI must consider temporal context, physiological responses to evolving environmental conditions, and spectral response. Consideration of the separate mechanisms affecting PRI over different timescales could greatly improve efforts to monitor changing photosynthetic activity using optical remote sensing.
Remote sensing has been used to detect plant biodiversity in a range of ecosystems based on the varying spectral properties of different species or functional groups. However, the most appropriate ...spatial resolution necessary to detect diversity remains unclear. At coarse resolution, differences among spectral patterns may be too weak to detect. In contrast, at fine resolution, redundant information may be introduced. To explore the effect of spatial resolution, we studied the scale dependence of spectral diversity in a prairie ecosystem experiment at Cedar Creek Ecosystem Science Reserve, Minnesota, USA. Our study involved a scaling exercise comparing synthetic pixels resampled from high-resolution images within manipulated diversity treatments. Hyperspectral data were collected using several instruments on both ground and airborne platforms. We used the coefficient of variation (CV) of spectral reflectance in space as the indicator of spectral diversity and then compared CV at different scales ranging from 1 mm² to 1 m² to conventional biodiversity metrics, including species richness, Shannon’s index, Simpson’s index, phylogenetic species variation, and phylogenetic species evenness. In this study, higher species richness plots generally had higher CV. CV showed higher correlations with Shannon’s index and Simpson’s index than did species richness alone, indicating evenness contributed to the spectral diversity. Correlations with species richness and Simpson’s index were generally higher than with phylogenetic species variation and evenness measured at comparable spatial scales, indicating weaker relationships between spectral diversity and phylogenetic diversity metrics than with species diversity metrics. High resolution imaging spectrometer data (1 mm² pixels) showed the highest sensitivity to diversity level. With decreasing spatial resolution, the difference in CV between diversity levels decreased and greatly reduced the optical detectability of biodiversity. The optimal pixel size for distinguishing α diversity in these prairie plots appeared to be around 1 mm to 10 cm, a spatial scale similar to the size of an individual herbaceous plant. These results indicate a strong scale-dependence of the spectral diversity-biodiversity relationships, with spectral diversity best able to detect a combination of species richness and evenness, and more weakly detecting phylogenetic diversity. These findings can be used to guide airborne studies of biodiversity and develop more effective large-scale biodiversity sampling methods.
In evergreen conifers, where the foliage amount changes little with season, accurate detection of the underlying “photosynthetic phenology” from satellite remote sensing has been difficult, ...presenting challenges for global models of ecosystem carbon uptake. Here, we report a close correspondence between seasonally changing foliar pigment levels, expressed as chlorophyll/carotenoid ratios, and evergreen photosynthetic activity, leading to a “chlorophyll/carotenoid index” (CCI) that tracks evergreen photosynthesis at multiple spatial scales. When calculated from NASA’s Moderate Resolution Imaging Spectroradiometer satellite sensor, the CCI closely follows the seasonal patterns of daily gross primary productivity of evergreen conifer stands measured by eddy covariance. This discovery provides a way of monitoring evergreen photosynthetic activity from optical remote sensing, and indicates an important regulatory role for carotenoid pigments in evergreen photosynthesis. Improved methods of monitoring photosynthesis from space can improve our understanding of the global carbon budget in a warming world of changing vegetation phenology.
Leaf pigment content can provide valuable insight into the physiological performance of leaves. Measurement of spectral reflectance provides a fast, nondestructive method for pigment estimation. A ...large number of spectral indices have been developed for estimation of leaf pigment content. However, in most cases these indices have been tested for only one or at most a few related species and thus it is not clear whether they can be applied across species with varying leaf structural characteristics. Our objective in this study was to develop spectral indices for prediction of leaf pigment content that are relatively insensitive to species and leaf structure variation and thus could be applied in larger scale remote-sensing studies without extensive calibration. We also quantified the degree of spectral interference between pigments when multiple pigments occur within the same leaf tissue. We found that previously published spectral indices provided relatively poor correlations with leaf chlorophyll content when applied across a wide range of species and plant functional types. Leaf surface reflectance appeared to be the most important factor in this variation. By developing a new spectral index that reduces the effect of differences in leaf surface reflectance, we were able to significantly improve the correlations with chlorophyll content. We also found that an index based on the first derivative of reflectance in the red edge region was insensitive to leaf structural variation. The presence of other pigments did not significantly affect estimation of chlorophyll from spectral reflectance. Previously published carotenoid and anthocyanin indices performed poorly across the whole data set. However, we found that the photochemical reflectance index (PRI, originally developed for estimation of xanthophyll cycle pigment changes) was related to carotenoid/chlorophyll ratios in green leaves. This result has important implications for the interpretation of PRI measured at both large and small scales. Our results demonstrate that spectral indices can be applied across species with widely varying leaf structure without the necessity for extensive calibration for each species. This opens up new possibilities for assessment of vegetation health in heterogeneous natural environments.
In evergreens, the seasonal down‐regulation and reactivation of photosynthesis is largely invisible and difficult to assess with remote sensing. This invisible phenology may be changing as a result ...of climate change. To better understand the mechanism and timing of these hidden physiological transitions, we explored several assays and optical indicators of spring photosynthetic activation in conifers exposed to a boreal climate. The photochemical reflectance index (PRI), chlorophyll fluorescence, and leaf pigments for evergreen conifer seedlings were monitored over 1 yr of a boreal climate with the addition of gas exchange during the spring. PRI, electron transport rate, pigment levels, light‐use efficiency and photosynthesis all exhibited striking seasonal changes, with varying kinetics and strengths of correlation, which were used to evaluate the mechanisms and timing of spring activation. PRI and pigment pools were closely timed with photosynthetic reactivation measured by gas exchange. The PRI provided a clear optical indicator of spring photosynthetic activation that was detectable at leaf and stand scales in conifers. We propose that PRI might provide a useful metric of effective growing season length amenable to remote sensing and could improve remote‐sensing‐driven models of carbon uptake in evergreen ecosystems.
The goal of this study was to explore the relationships between the photochemical reflectance index (PRI) at the leaf level and pigment pools, focusing on the constitutive role of pigments in ...influencing PRI over seasonal or ontogenetic time frames. The purpose was to re-evaluate the role of PRI as an indicator of seasonally shifting pigment (chlorophyll, carotenoid and anthocyanin) contents, and hence photosynthetic activity, across a range of tree and crop species. We studied natural vegetation - three tree species (maple, chestnut and beech) and two managed irrigated and rainfed crop species (maize and soybean), contrasting in photosynthetic pathway and leaf structure, and having wide variation of pigment content and composition. In anthocyanin-free leaves, PRI related to both total chlorophyll (Chl) and carotenoid (Car) contents, however, much closer relationships were found between PRI and Car to Chl ratio (Car/Chl). The sensitivity of PRI to Car/Chl varied widely in tree species with the degree of secondary carotenoids. In crop leaves where the Car vs. Chl relationship was very close, the slopes of PRI vs. Car/Chl relationships for maize and soybean were almost identical. PRI vs. Car/Chl relationships for leaves of different tree species formed a significant, uniform relationship with PRI. Two crops also formed a significant, uniform PRI vs. Car/Chl relationship with a slope half the value found for trees. In anthocyanin-containing leaves, PRI did not clearly relate to any pigment content because reflectance values at both PRI wavebands are affected by anthocyanin content. The findings of a strong link between leaf level PRI and Car/Chl over seasonal and ontogenetic time spans supports recent findings calling for a more careful evaluation of the relationship between PRI and either LUE or photosynthetic activity. In particular, studies that contrast short-term (e.g. diurnal) vs. long-term (e.g. seasonal) pigment, PRI, and photosynthetic responses in contrasting vegetation types are needed.
•Seasonal change of PRI relates significantly to contents of both Chl and Car.•Car/Chl drives PRI seasonally, during ontogenesis, and under stress.•PRI vs. Car/Chl relationships for trees converge forming unified relationship.•PRI vs. Car/Chl relationships for crops converge forming unified relationship.•Presence of anthocyanins invalidates the tight relationship of PRI vs. Car/Chl.
Tree species in the boreal forest cycle between periods of active growth and dormancy alter their photosynthetic processes in response to changing environmental conditions. For deciduous species, ...these changes are readily visible, while evergreen species have subtler foliar changes during seasonal transitions. In this study, we used remotely sensed optical indices to observe seasonal changes in photosynthetic activity, or photosynthetic phenology, of six boreal tree species. We evaluated the normalized difference vegetation index (NDVI), the photochemical reflectance index (PRI), the chlorophyll/carotenoid index (CCI), and steady-state chlorophyll fluorescence (FS) as a measure of solar-induced fluorescence (SIF), and compared these optical metrics to gas exchange to determine their efficacy in detecting seasonal changes in plant photosynthetic activity. The NDVI and PRI exhibited complementary responses. The NDVI paralleled photosynthetic phenology in deciduous species, but not in evergreens. The PRI closely paralleled photosynthetic activity in evergreens, but less so in deciduous species. The CCI and FS tracked photosynthetic phenology in both deciduous and evergreen species. The seasonal patterns of optical metrics and photosynthetic activity revealed subtle differences across and within functional groups. With the CCI and fluorescence becoming available from satellite sensors, they offer new opportunities for assessing photosynthetic phenology, particularly for evergreen species, which have been difficult to assess with previous methods.
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