Relationship between photosynthetic radiation-use efficiency of barley canopies and the photochemical reflectance index (PRI)

Some processes of excess radiation dissipation have been associated with changes in leaf reflectance near 531 nm. We aimed to study the relations between the photochemical reflectance index (PRI) derived from this signal, and photosynthetic radiation-use efficiency (defined as net CO₂ assimilation rate/incident photon flux density) in a cereal canopy. Measurements of reflectance, fluorescence, gas exchange and xanthophyll cycle pigments were made in the morning, midday and afternoon in barley canopies with two levels of nitrogen fertilization. The photosynthetic radiation-use efficiency decreased at midday, mainly in the third leaf, in both treatments, with lower values for the nitrogen deficient leaves. The zeaxanthin content showed the inverse pattern, increasing at midday and in the nitrogen deficient treatment. The photosynthetic radiation-use efficiency was well correlated with the epoxidation state, EPS (violaxanthin + 0.5 antheraxanthin)/(violaxanthin + antheraxanthin + zeaxanthin). The PRI [here defined as (R₅₃₉ - R₅₇₀)/(R₅₃₉+ R₅₇₀)] was significantly correlated with epoxidation state and zeaxanthin and with photosynthetic radiation-use efficiency. These results validate the utility of PRI in the assessment of radiation-use efficiency at canopy level.     

Antitranspirant functions of stem periderms and their influence on corticular photosynthesis under drought stress

Transpiration and photosynthesis of current-year stems and adult leaves of different deciduous tree species were investigated to estimate their probable influence on carbon balance. Peridermal transpiration of young stems was found to be rather small as compared to the transpiration of leaves (stem/leaf like 1/5–1/20). A characteristic that was mainly attributable to the lower peridermal conductance to water and CO₂, which made up only 8–28% of stomatal conductance. Water vapour conductance was significantly lower in stems, but also non-responsive to PAR, which led to a comparatively higher water use efficiency (WUE, ratio assimilation/transpiration). Thus, although corticular photosynthesis reached only 11–37% of leaf photosynthesis, it may be a means of improving the carbon balance of stems under limited water availability. The influence of drought stress on primary photosynthetic reactions was also studied. Under simulated drought conditions the drying time needed to provoke a 50% reduction (t ₅₀) in dark- and light-adapted PSII efficiency (Fv/Fm, ΔF/Fm′) was up to ten times higher in stems than in leaves. Nevertheless, up to a relative water deficit (RWD) of around 40–50% dark-adapted PSII efficiency of leaves and stems was rather insensitive to dehydration, showing that the efficiency of open PS II reaction centres is not impaired. Thus, it may be concluded that in stems as well as in leaves the primary site of drought damage is at the level of dark enzyme reactions and not within PSII. However, enduring severe drought caused photoinhibitory damage to the photosynthetic apparatus of leaves and stems; thereby RWD₅₀ values (= RWD needed to provoke a 50% reduction in Fv/Fm ad ΔF/Fm′) were comparably lower in stems as compared to leaves, indicating a possibly higher drought sensitivity of the cortex chlorenchyma.     

Assessing photosynthetic efficiency in an experimental mangrove canopy using remote sensing and chlorophyll fluorescence

This study examined the ability of the photochemical reflectance index (PRI) to track changes in effective quantum yield (Δ F/F _m ′), non-photochemical quenching (NPQ), and the xanthophyll cycle de-epoxidation (DPS) in an experimental mangrove canopy. PRI was correlated with (Δ F/F _m ′) and NPQ over the 4-week measurement period and over the diurnal cycle. The normalised difference vegetation index (NDVI) was not correlated with any aspect of photochemical efficiency measured using chlorophyll fluorescence or xanthophyll pigments. This study demonstrated that photochemical adjustments were responsible for controlling the flow of energy through the photosynthetic apparatus in this mangrove forest canopy rather than canopy structural or chlorophyll adjustments.     

Analysis of Photosynthetic Activity at the Leaf and Canopy Levels from Reflectance Measurements: A Case Study

The photochemical reflectance index (PRI), based on reflectance signatures at 531 and 570 nm, and associated with xanthophyll pigment inter-conversion and related thylakoid energisation, was evaluated as an indicator of photosynthetic function in a Mediterranean holm oak (Quercus ilex L.) coppice. The chlorophyll fluorescence pulse-amplitude-modulation and the eddy correlation techniques were used to estimate the photosystem 2 photochemical efficiency of leaves and the CO₂ flux over the canopy, respectively. The reflectance and fluorescence techniques yielded identical estimates of the photosynthetic activity in leaves exposed to dark-light-dark cycles or to a variable irradiance in laboratory. However, there was no such correlation between photosynthetic performance and PRI when applied to a sun-exposed canopy in field conditions. Fluorescence profiles inside the canopy and especially a helpful use of multispectral reflectance imaging highlight the limitations of such method.     

Altitudinal differences in UV absorbance,UV reflectance and related morphological traits of Quercus ilex and Rhododendron ferrugineum in the Mediterranean region

We studied the variations in different physiological parameters associated with UV-B radiation defense: UV-B radiation absorbance, UV-visible spectral reflectance, carotenoids concentration, leaf thickness, SLW (specific leaf weigth) and trichome density in Quercus ilex growing at 200 and 1200 m and Rhododendron ferrugineum growing at 2200 m. We examined the role of these parameters as protection mechanisms in an altitudinal gradient of increasing UV radiation in northern Catalonia and in sun and shade leaves. The concentration of UV-B radiation absorbing pigments was 15% higher in sun leaves of Q. ilex at 1200 m than in those from 200 m altitude. Sun leaves of R. ferrugineum presented concentrations three times higher than those of Q. ilex. Reflectance ranged between 5% (in the region 300–400 nm) and 12% (in the region 280–300 nm). The variation of reflectance with altitude followed an inverse trend respect to absorbance in the 280–300 nm region, with higher values the lower the altitude, but in the 300–400 nm region, reflectance of the lower site was the lowest. In both species and altitudes sun leaves presented higher concentrations of UV-B radiation absorbing pigments and UV reflectance than shade leaves. Quercus ilex trees of the higher location presented higher NDPI (Normalized Difference Pigment Reflectance Index) values, indicating higher carotenoids/chlorophyll a ratio. Actual measurements of carotenoid/chlorophyll a ratio confirmed this pattern. The photochemical reflectance index (PRI) presented higher values the higher the location indicating lower photosynthetic radiation-use efficiency. Specific leaf weight (SLW) and leaf thickness were larger in Q. ilex trees of higher location than in those of lower location. In both sites, sun leaves also presented larger SLW values than shade leaves. Adaxial leaf hair density in sun leaves was significantlly higher in the lower location. UV absorption and linked morphological traits (SLW, leaf thickness measured in Q. ilex) presented the larger differences among studied plants at different altitudes and seem to be the dominant UV protecting mechanisms.     

Functional plasticity of photosynthetic apparatus and its resistance to photoinhibition in <Emphasis Type="Italic">Plantago media</Emphasis>

Morphological and functional characteristics of Plantago media L. leaves were compared for plants growing at different light regimes on limestone outcrops in Southern Timan (62°45′N, 55°49′E). The plants grown in open areas under exposure to full sunlight had small leaves with low pigment content and high specific leaf weight; these leaves exhibited high photosynthetic capacity and elevated water use efficiency at high irradiance. The maximum photochemical activity of photosystem II (F _v/F _m) in leaves of sun plants remained at the level of about 0.8 throughout the day. The photosynthetic apparatus of sun plants was resistant to excess photosynthetically active radiation, mostly due to non-photochemical quenching of chlorophyll fluorescence (qN). This quenching was promoted by elevated deepoxiation of violaxanthin cycle pigments. Accumulation of zeaxanthin, a photoprotective pigment in sun plant leaves was observed already in the morning hours. The plant leaves grown in the shade of dense herbage were significantly larger than the sun leaves, with pigment content 1.5–2.0 times greater than in sun leaves; these leaves had low qN values and did not need extensive deepoxidation of violaxanthin cycle pigments. The data reveal the morphophysiological plasticity of plantain plants in relation to lighting regime. Environmental conditions can facilitate the formation of the ecotype with photosynthetic apparatus resistant to photoinhibition. Owing to this adjustment, hoary plantain plants are capable of surviving in ecotopes with high insolation.     

Radiation Use Efficiency,Chlorophyll Fluorescence,and Reflectance Indices Associated with Ontogenic Changes in Water-Limited Chenopodium quinoa Leaves

Net photosynthetic rate, radiation use efficiency, chlorophyll (Chl) fluorescence, photochemical reflectance index (PRI), and leaf water potential were measured during a 25-d period of progressive water deficit in quinoa plants grown in a glasshouse in order to examine effects of water stress and ontogeny. All physiological parameters except F_v/F_m were sensitive to water stress. Ontogenic variations did not exist in F_v/F_m and leaf water potential, and were moderate to high in the other parameters. The complete recovery of photosynthetic parameters after re-irrigation was related with the stability in F_v/F_m. PRI showed significant correlation with predawn leaf water potential, F_m, and midday F_v/F_m. Thus PRI and Chl fluorescence may help in assessing physiological changes in quinoa plants across different developmental stages and water status.     

Photochemistry, remotely sensed physiological reflectance index and de-epoxidation state of the xanthophyll cycle in Quercus coccifera under intense drought

Quercus coccifera L. is a Mediterranean sclerophyllous shrub with a high capacity to resist intense drought stress. Therefore, it could be used in the study of physiological changes suffered by plants at very low water potentials. A remote sensing sensor was used to measure continuously the physiological reflectance index (PRI; defined as the changes in reflectance at 531 nm with respect to those at 570 nm; PRI = [(R531 − R570)/(R531 + R570)] at canopy level and under field conditions in an artificial carpet of seedlings of Q. coccifera during a drought cycle. Correlations between leaf level-measured chlorophyll fluorescence parameters as well as the de-epoxidation state of the xanthophyll cycle [(A + Z)/(V + A + Z)] and canopy level-measured PRI were reasonably good (R ² = 0.57–0.63, P < 0.01), and quite interesting for water stress remote sensing purposes. The instrument’s temporal resolution allowed us to follow the rapid response of PRI to changing photosynthetic active radiation, and to resolve, in response to cloud-induced changes in light intensity, a fast and a slow PRI component. We report the disappearance of the rapid one under conditions of intense drought in response to a sudden increase in light intensity. The underlying photoprotection mechanisms that Q. coccifera shows in response to intense drought stress periods seem to be related to the existence of a low intrathylakoid lumenal pH at the end of the drought cycle. Under intense drought, these mechanisms allow this species to avoid oxidative damage, which was evidenced by the maintenance of an unaltered photosynthetic pigment composition and constant photosystem II efficiency in the mornings. It is concluded that, contrary to early reports, PRI is a sensible, indirect, non-destructive water stress indicator, even in plants experiencing intense drought. Preliminary results of this work were presented at the 3rd International Workshop on Remote Sensing of Vegetation Fluorescence (February 2007, Florence, Italy).     

Spectral reflectance of Picea rubens (Pinaceae) and Abies balsamea (Pinaceae) needles along an elevational gradient, Mt. Moosilauke, New Hampshire, USA

Relationships among elevation, foliar morphology, spectral reflectance, and chlorophyll fluorescence of two co-occurring montane conifers, red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea [L.] Mill.), were investigated along two transects from 460 to 1460 m on Mt. Moosilauke in the White Mountains of New Hampshire, USA. Spectral reflectance (300-1100 nm wavelengths) and the chlorophyll fluorescence F(v)/F(m) ratio were measured on dark-adapted needles. Foliar morphology (needle size, shape, and mass) and nitrogen concentrations were measured in the laboratory. Reflectance spectra varied between species and with elevation. Two chlorophyll measures, red edge position and a chlorophyll-based difference index (Chl NDI = R750 - R705/R750 + R705), indicated more chlorophyll in fir than in spruce and decreasing chlorophyll with increasing elevation in both species. The structure-independent pigment index (SIPI = R800 - R445/R800 - R680) increased with elevation, indicating an increasing carotenoid?:?chlorophyll ratio. The photochemical reflectance index (PRI = R531 - R570/R531 + R570), a measure of photosynthetic radiation use efficiency, decreased with increasing elevation up to 1370 m. In the highest elevation site, within the stunted alpine krummholz at 1460 m, PRI was higher than at 1370 m, but still lower than at 1070 m. This same pattern was evident in the chlorophyll fluorescence F(v)/F(m) measurements. These independent indices indicate higher stress in spruce than fir, which may be related to the "spruce decline" reported in the northeastern USA. Results also indicate progressively increasing stress with increasing elevation up to 1370 m. Stress appears to be lower at 1460 m than at 1370 m, despite the harsher conditions at the very summit of Mt. Moosilauke. This may be a consequence of stress-tolerant physiology and/or prostrate architecture.     

Assessing leaf pigment content and activity with a reflectometer

This study explored reflectance indices sampled with a 'leaf reflectometer' as measures of pigment content for leaves of contrasting light history, developmental stage and functional type (herbaceous annual versus sclerophyllous evergreen). We employed three reflectance indices: a modified normalized difference vegetation index (NDVI), an index of chlorophyll content; the red/green reflectance ratio ( R _RED: R _GREEN), an index of anthocyanin content; and the change in photochemical reflectance index upon dark–light conversions (ΔPRI), an index of xanthophyll cycle pigment activity. In Helianthus annuus (sunflower), xanthophyll cycle pigment amounts were linearly related to growth light environment; leaves in full sun contained approximately twice the amount of xanthophyll cycle pigments as leaves in deep shade, and at midday a larger proportion of these pigments were in the photoprotective, de-epoxidized forms relative to shade leaves. Reflectance indices also revealed contrasting patterns of pigment development in leaves of contrasting structural types (annual versus evergreen). In H. annuus sun leaves, there was a remarkably rapid increase in amounts of both chlorophyll and xanthophyll cycle pigments along a leaf developmental sequence. This pattern contrasted with that of Quercus agrifolia (coast live oak, a sclerophyllous evergreen), which exhibited a gradual development of both chlorophyll and xanthophyll cycle pigments along with a pronounced peak of anthocyanin pigment content in newly expanding leaves. These temporal patterns of pigment development in Q. agrifolia leaves suggest that anthocyanins and xanthophyll cycle pigments serve complementary photoprotective roles during early leaf development. The results illustrate the use of reflectance indices for distinguishing divergent patterns of pigment activity in leaves of contrasting light history and functional type.