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Remote sensing of solar-induced chlorophyll fluorescence (SIF) is a rapidly advancing front in terrestrial vegetation science, with emerging capability in space-based methodologies and diverse application prospects. Although remote sensing of SIF -especially from space -is seen as a contemporary new specialty for terrestrial plants, it is founded upon a multi-decadal history of research, applications, and sensor developments in active and passive sensing of chlorophyll fluorescence. Current technical capabilities allow SIF to be measured across a range of biological, spatial, and temporal scales. As an optical signal, SIF may be assessed remotely using high-resolution spectral sensors in tandem with state-of-the-art algorithms to distinguish the emission from reflected and/or scattered ambient light. Because the red to far-red SIF emission is detectable non-invasively, it may be sampled repeatedly to acquire spatio-temporally explicit information about photosynthetic light responses and steady-state behaviour in vegetation.Progress in this field is accelerating with innovative sensor developments, retrieval methods, and modelling advances. This review distills the historical and current developments spanning the last several decades. It highlights SIF heritage and complementarity within the broader field of fluorescence science, the maturation of physiological and radiative transfer modelling, SIF signal retrieval strategies, techniques for field and airborne sensing, advances in satellite-based systems, and applications of these capabilities in evaluation of photosynthesis and stress effects. Progress, challenges, and future directions are considered for this unique avenue of remote sensing.

The early light-induced proteins (ELIPs) belong to the multigenic family of light-harvesting complexes, which bind chlorophyll and absorb solar energy in green plants. ELIPs accumulate transiently in plants exposed to high light intensities. By using an Arabidopsis thaliana mutant (chaos) affected in the posttranslational targeting of light-harvesting complex-type proteins to the thylakoids, we succeeded in suppressing the rapid accumulation of ELIPs during high-light stress, resulting in leaf bleaching and extensive photooxidative damage. Constitutive expression of ELIP genes in chaos before light stress resulted in ELIP accumulation and restored the phototolerance of the plants to the wild-type level. Free chlorophyll, a generator of singlet oxygen in the light, was detected by chlorophyll fluorescence lifetime measurements in chaos leaves before the symptoms of oxidative stress appeared. Our findings indicate that ELIPs fulfill a photoprotective function that could involve either the binding of chlorophylls released during turnover of pigment-binding proteins or the stabilization of the proper assembly of those proteins during high-light stress.L ight is essential for plants through photosynthetic carbon assimilation. However, when absorbed light exceeds the photosynthetic capacities, reactive O 2 species are generated in the chloroplasts, causing oxidative damage to proteins, lipids, and photosynthetic pigments (1, 2). This effect is amplified by environmental stresses such as low temperature or drought, for example, that inhibit the photosynthetic activity, leading to strong yield reduction in crops. In green plants, solar energy is collected by chlorophyll-and carotenoid-binding lightharvesting complexes (LHCs), which are encoded by a multigene family of LHC genes. The expression of these genes is tightly regulated by light (2-4). High light intensities inhibit transcription of LHC genes and activate synthesis of the early lightinduced proteins (ELIPs), a class of proteins structurally related to the LHCs (5). The ELIPs are predicted to have three transmembrane helices, and they have sequence similarity to the LHCs in the central pair of helices (6, 7). The similarity is not only at the sequence level, however, because both LHCs and ELIPs bind chlorophyll and carotenoids (8). The ELIPs differ from the LHCs by their transient expression under high-light stress (5). Recently, a number of ELIP-type polypeptides, containing LHC motifs and inducible by high light, have been discovered in vascular plants: the one-helix high-light-induced proteins (9) and the two-helix stress-enhanced proteins (10).The physiological role of the ELIPs in vascular plants has not yet been elucidated, although there have been several suggestions (11)(12)(13)(14). The induction of ELIPs by high light intensities suggests a role in the acclimation to light stress rather than a light-harvesting function, but this has not yet been demonstrated. ELIP antisense transgenic tobacco plants did not exhibit any phenotype of sensitivity to high ...

We have studied the time-resolved fluorescence properties of the light-harvesting complexes (Lhc) of photosystem II (Lhcb) in order to obtain information on the mechanism of energy dissipation (non-photochemical quenching) which is correlated to the conversion of violaxanthin to zeaxanthin in excess light conditions. The chlorophyll fluorescence decay of Lhcb proteins LHCII, CP29, CP26, and CP24 in detergent solution is mostly determined by two lifetime components of 1.2-1.5 and 3.6-4 ns while the contribution of the faster component is higher in CP29, CP26, and CP24 with respect to LHCII. The xanthophyll composition of Lhc proteins affects the ratio of the lifetime components: when zeaxanthin is bound into the site L2 of LHCII, the relative amplitude of the faster component is increased and, consequently, the chlorophyll fluorescence quenching is enhanced. Analysis of quenching in mutants of Arabidopsis thaliana, which incorporate either violaxanthin or zeaxanthin in their Lhc proteins, shows that the extent of quenching is enhanced in the presence of zeaxanthin. The origin of the two fluorescence lifetimes was analyzed by their temperature dependence: since lifetime heterogeneity was not affected by cooling to 77 K, it is concluded that each lifetime component corresponds to a distinct conformation of the Lhc proteins. Upon incorporation of Lhc proteins into liposomes, a quenching of chlorophyll fluorescence was observed due to shortening of all their lifetime components: this indicates that the equilibrium between the two conformations of Lhcb proteins is displaced toward the quenched conformation in lipid membranes or thylakoids with respect to detergent solution. By increasing the protein density in the liposomes, and therefore the probability of protein-protein interactions, a further decrease of fluorescence lifetimes takes place down to values typical of quenched leaves. We conclude that at least two major factors determine the quenching of chlorophyll fluorescence in Lhcb proteins, i.e., intrasubunit conformational change and intersubunit interactions within the lipid membranes, and that these processes are both important in the photoprotection mechanism of nonphotochemical quenching in vivo.The photosynthetic apparatus of higher plants is composed of two photosystems (PS), 1 PSI and PSII, each consisting of a core complex moiety, binding chlorophyll (Chl) a and -carotene, and a peripheral antenna system, binding Chl a, Chl b, and xanthophylls (Cars). The major component of the PSII antenna system is LHCII, a heterotrimer of the Lhcb1, -2, and -3 gene products (also called LHCIIb) binding half of the thylakoid pigments. Minor components are CP29 (LHCIIa), CP26 (LHCIIc), and CP24 (LHCIId), respectively the products of the Lhcb4, -5, and -6 genes, which are monomeric and connect the PSII core complex to the peripheral LHCII trimers.The large absorption cross-section of the antenna systems allows plants to grow in dim light. However, the light intensity experienced by a plant varies largely over ...

Water stress experiments were performed with grapevines (Vitis vinifera L.) and other C3 plants in the field, in potted plants in the laboratory, and with detached leaves. It was found that, in all cases, the ratio of steady state chlorophyll fluorescence (Fs) normalized to dark-adapted intrinsic fluorescence (Fo) inversely correlated with non-photochemical quenching (NPQ). Also, at high irradiance, the ratio Fs/Fo was positively correlated with CO2 assimilation in air, with electron transport rate calculated from fluorescence, and with stomatal conductance, but no clear correlation was observed with qP. The significance of these relationships is discussed. The ratio Fs/Fo, measured with a portable instrument (PAM-2000) or with a remote sensing FIPAM system, provides a good method for the early detection of water stress, and may become a useful guide to irrigation requirements.

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Remote sensing of far-red sun-induced chlorophyll fluorescence (SIF) has emerged as an important tool for studying gross primary productivity (GPP) at the global scale. However, the relationship between SIF and GPP at the canopy scale lacks a clear mechanistic explanation. This is largely due to the poorly characterized role of the relative contributions from canopy structure and leaf physiology to the variability of the top-of-canopy, observed SIF signal. In particular, the effect of the canopy structure beyond light absorption is that only a fraction (f esc) of the SIF emitted from all leaves in the canopy can escape from the canopy due to the strong scattering of near-infrared radiation. We combined rice, wheat and corn canopy-level in-situ datasets to study how the physiological and structural components of SIF individually relate to measures of photosynthesis. At seasonal time scales, we found a considerably strong positive correlation (R 2 =0.4-0.6) of f esc to the seasonal dynamics of the photosynthetic light use efficiency (LUE P), while the estimated physiological SIF yield was almost entirely uncorrelated to LUE P both at seasonal and diurnal time scales, with the partial exception of wheat. Consistent with these findings, the canopy structure and radiation component of SIF, defined as the product of APAR and f esc , explained the relationship of observed SIF to GPP and even outperformed GPP estimation based on observed SIF at two of the three sites investigated. These results held for both half-hourly and daily mean values. In contrast, the total emitted SIF, obtained by normalizing observed SIF for f esc , improved only the relationship to APAR but considerably decreased the correlation to GPP for all three crops. Our findings demonstrate the dominant role of canopy structure in the SIF-GPP relationship and establish a strong, mechanistic link between the near-infrared reflectance of vegetation (NIR V) and the relevant canopy structure information contained in the SIF signal. These insights are expected to be useful in improving remote sensing based GPP estimates.

In this study a method was designed to assess non-destructively the type of UV-screening compounds present in the leaf epidermis. The method is based on the recording and calculation of the ratio of UV-excitation spectra of chlorophyll fluorescence (FER) from the adaxial and abaxial sides of bifacial leaves, or from older and younger segments of monocotyledonous leaves. The logarithm of this ratio (logFER) matched the absorption spectrum of the UVabsorbers present in the leaf, as confirmed by its overlap with the absorption spectrum of the methanolic extract of the leaf or of the isolated epidermis. By using the logFER approach, it was possible to demonstrate that the concentration but not the classes of compounds present in the epidermis that are responsible for UV-screening is affected by the side and the age of the leaves. In contrast, measurements from the leaves of seven dicots and one monocot indicated large difference in the classes of these compounds between species. Finally, it was shown that the logFER in the UV is independent of the emission wavelength, and that the method can be used for quantitative measurements. This method expands to the spectral domain the use of ChlF for the estimation of the leaf epidermal transmittance.