Diverse Optical and Photosynthetic Properties in a Neotropical Dry Forest during the Dry Season: Implications for Remote Estimation of Photosynthesis1

Using optical and photosynthetic assays from a canopy access crane, we examined the photosynthetic performance of tropical dry forest canopies during the dry season in Parque Metropolitano, Panama City, Panama. Photosynthetic gas exchange, chlorophyll fluorescence, and three indices derived from spectral reflectance (the normalized difference vegetation index, the simple ratio, and the photochemical reflectance index) were used as indicators of structural and physiological components of photosynthetic activity. Considerable interspecific variation was evident in structural and physiological behavior in this forest stand, which included varying degrees of foliage loss, altered leaf orientation, stomatal closure, and photosystem II downregulation. The normalized difference vegetation index and the simple ratio were closely related to canopy structure and absorbed radiation for most species, but failed to capture the widely divergent photosynthetic behavior among evergreen species exhibiting various degrees of downregulation. The photochemical reflectance index and chlorophyll fluorescence were related indicators of photosynthetic downregulation, which was not detectable with the normalized difference vegetation index or simple ratio. These results suggest that remote sensing methods that ignore downregulation cannot capture within‐stand variability in actual carbon flux for this diverse forest type. Instead, these findings support a sampling approach that derives photosynthetic fluxes from a consideration of both canopy light absorption (e.g., normalized difference vegetation index) and photosynthetic light‐use efficiency (e.g., photochemical reflectance index). Such sampling should improve our understanding of controls on photosynthetic carbon uptake in diverse tropical forest stands.     

Desiccation‐induced changes in photochemical processes of photosynthesis and spectral reflectance in Nostoc commune (Cyanobacteria,Nostocales) colonies from polar regions

Cyanobacterium Nostoc commune is a species highly resistant against desiccation. In this study, we investigated changes in photochemical processes of photosynthesis and spectral reflectance indices during controlled desiccation of the colonies from Antarctica. In a dehydration process, water potential (WP) reached ?3 MPa and values of potential (F _v/F _m) and effective quantum yields (ΦPSII) of photosystem II were kept to high value until 90% of water was lost from the colony, and these values decreased rapidly by further loss of water. This indicates that the colony loses water mostly from the exopolysaccharidic envelope, not from cells during the initial part of dehydration (relative water content, RWC = 100–10%). Other suggestions of inhibition of photosynthetic processes after 90% loss of water were the increase of the chlorophyll fluorescence parameter F _p/F _s. The F _m′ was higher than F _m in hydrated colonies because of state transition which change energy distribution between PS I and PS II, but decreased to same level as F _m in dehydrated colonies. The Normalized Difference Vegetation Index (NDVI) and Photochemical Reflectance Index (PRI) showed concave‐ and convex‐curvilinear relationship with RWC, respectively. The changes of NDVI values were, however, statistically insignificant. PRI values were predominantly below 0 because of phycobiliprotein involvement. These results were compared with the same species in the Arctic region. This is, according to our best knowledge, the first measurement of changes in spectral reflectance indices during desiccation of cyanobacteria.     

Diurnal patterns of photosynthesis, chlorophyll fluorescence, and PRI to evaluate water stress in the invasive species, Elaeagnus umbellata Thunb.

Photosynthesis, chlorophyll fluorescence, and hyperspectral reflectance were used to evaluate diurnal changes of Elaeagnus umbellata to quantify physiological responses of the invasive species during times of stress. Field measurements showed that E. umbellata is able to maintain higher levels of photosynthesis relative to nearby Quercus alba plants, with less water loss. Plants subjected to progressive drought were able to recover photosynthesis one day following re-watering. Laboratory and field measurements revealed decreasing ΔF/F′_m values in response to drought stress, with little corresponding decrease in photochemical reflectance index values. This research supports the view that xanthophyll cycle dissipation is not the photoprotective mechanism at work for Elaeagnus species under water stress. Elaeagnus umbellata maintains photosynthetic carbon assimilation even under drought conditions, in part, due to chemical dissipation of excess light, and in part because of morphological features that limit excess radiation while maximizing photosynthetic carbon gain. These characteristics may contribute to the invasive success of E. umbellata.     

Spectral vegetation indices as the indicator of canopy photosynthetic productivity in a deciduous broadleaf forest

Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at 'Takayama' super site in central Japan.Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI × V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance.Important findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.     

Contrasting traits,contrasting environments,and considerations on population dynamics under a changing climate: an ecophysiological field study of two co‐dominant tree species

The increase in light availability resulting from canopy changes or opening is not always beneficial and can inhibit photosynthesis of tree seedlings already under other environmental stress. Tree seedlings' responses to compounded abiotic stress depend on their life‐history traits, and understanding the variations of such responses is important for understanding population dynamics under a changing climate. In this study we investigate how the photosynthesis of juveniles of two canopy tree species with different life‐history traits, Abies sachalinensis and Betula ermanii, differs in two contrasting sites at a sub‐boreal forest in northern Japan—one under a deciduous canopy (Closed site) and the other at a wide canopy opening (Open site). Seedlings at the Open site had low F_v/F_m (quantum yield of photosystem II) for a longer period than those at the Closed site. Abies sachalinensis at the Closed site showed lower F_v/F_m in spring than those at the Open site, but recovered after the canopy's new leaves flushed, indicating its acclimation to the shaded condition. Mean P_max (light‐saturated photosynthetic rate at ambient CO₂ levels) of A. sachalinensis seedlings was affected by site and air temperature, while B. ermanii seedlings were also affected by precipitation. Only B. ermanii's seedlings presented growth in the period studied, in spite of observed mid‐day drops to F_v/F_m attributed to water‐deficit‐related photoprotection. Results suggest that the climate change predicted for the Hokkaido area may increase the competitive advantage of broad‐leaved deciduous species, such as B. ermanii, in relation to evergreen conifers like A. sachalinensis.     

Predicting vegetation water content in wheat using normalized difference water indices derived from ground measurements

Vegetation water content (VWC) is an important variable for both agriculture and forest fire management. Remote sensing technology offers an instantaneous and non-destructive method for VWC assessment provided we can relate in situ measurements of VWC to spectral reflectance in a reliable way. In this paper, based on radiative transfer models, three new normalized difference water indices (NDWI) are proposed for VWC [fuel moisture content (FMC), and equivalent water thickness (EWT)] estimation, taking both leaf internal structure and dry matter content into account. Reflectance at 1,200, 1,450 and 1,940 nm were selected and normalized with reflectance at 860 nm to establish three water indices, NDWI₁₂₀₀, NDWI₁₄₅₀ and NDWI₁₉₄₀. Good correlations were observed between FMC (R ² = 0.65–0.80) and EWT (both at the leaf scale, R ² = 0.75–0.81 for EWT_L and at the canopy scale, R ² = 0.80–0.83 for EWT_C) at various stages of wheat crop development.     

Effects of photoinhibition on whole-plant carbon gain assessed with a photosynthesis model

A canopy photosynthesis model was modified to assess the effect of photoinhibition on whole‐plant carbon gain. Photoinhibitory changes in maximum quantum yield of photosystem II (F_v/F_m) could be explained solely from a parameter (Lflux) calculated from the light micro‐environment of the leaves. This relationship between F_v/F_m and the intercepted cumulative light dose, integrated and equally weighted over several hours was incorporated into the model. The effect of photoinhibition on net photosynthesis was described through relationships between photoinhibition and the shaping parameters of the photosynthetic light‐response curve (quantum use efficiency, convexity, and maximum capacity). This new aspect of the model was then validated by comparing measured field data (diurnal courses of F_v/F_m) with simulation results. Sensitivity analyses revealed that the extent of photoinhibitory reduction of whole‐plant photosynthesis was strongly dependent on the structural parameters (LAI and leaf angle). Simulations for a Mediterranean evergreen oak, Quercus coccifera, under climatic conditions which cause mild photoinhibition revealed a daily loss of 7·5–8·5% of potential carbon gain in the upper sunlit canopy layers, a 3% loss in the bottom canopy, and an overall loss of 6·1%. Thus, this canopy photoinhibition model (CANO‐PI) allows the quantitative evaluation of photoinhibition effects on primary production.     

Developing and validating novel hyperspectral indices for leaf area index estimation: Effect of canopy vertical heterogeneity

Leaf area index (LAI) is one of the key biophysical parameters for understanding land surface photosynthesis, transpiration, and energy balance processes. Estimation of LAI from remote sensing data has been a premier method for a large scale in recent years. Recent studies have revealed that the within-canopy vertical variations in LAI and biochemical properties greatly affect canopy reflectance and significantly complicate the retrieval of LAI inversely from reflectance based vegetation indices, which has yet been explicitly addressed. In this study, we have used both simulated datasets (dataset I with constant vertical profiles of LAI and biochemical properties, dataset II with varied vertical profile of LAI but constant vertical biochemical properties, and dataset III with both varied vertical profiles) generated from the multiple-layer canopy radiative transfer model (MRTM) and a ground-measured dataset to identify robust spectral indices that are insensitive to such within canopy vertical variations for LAI prediction. The results clearly indicated that published indices such as normalized difference vegetation index (NDVI) had obvious discrepancies when applied to canopies with different vertical variations, while the new indices identified in this study performed much better. The best index for estimating canopy LAI under various conditions was D(920,1080), with overall RMSEs of 0.62–0.96 m²/m² and biases of 0.42–0.55 m²/m² for all three simulated datasets and an RMSE of 1.22 m²/m² with the field-measured dataset, although it was not the most conservative one among all new indices identified. This index responded mostly to the quantity of LAI but was insensitive to within-canopy variations, allowing it to aid the retrieval LAI from remote sensing data without prior information of within-canopy vertical variations of LAI and biochemical properties.     

Contribution of intercellular reflectance to photosynthesis in shade leaves

The potential contribution of intercellular light reflectance to photosynthesis was investigated by infiltrating shade leaves with mineral oil. Infiltration of leaves of Hydrophyllum canadense and Asarum canadense with mineral oil decreased adaxial leaf reflectance but increased transmittance. As a result of the large increase in transmittance, infiltration caused a decrease in absorptance of 25% and 30% at 550 and 750 nm, respectively. Thus, intercellular reflectance increased absorptance in these species by this amount. In a comparison of sun and shade leaves of Acer saccharum and Parthenocissus quinquefolia, oil infiltration decreased absorptance more in shade than in sun leaves. This difference suggests that the higher proportion of spongy mesophyll in shade leaves may increase internal light scattering and thus absorptance. The importance of the spongy mesophyll in increasing internal reflectance was also evident in comparisons of the optics of Populus leaves and in the fluorescence yield of oil-infiltrated leaves of several sun and shade species. Oil infiltration decreased the quantum yield of fluorescence (F_o) by 39–52% for shade leaves but only 21–25% for sun leaves. We conclude that the greater proportion of spongy parenchyma in shade leaves increased intercellular light scattering and thus absorptance. Direct measurements with fibre-optic light probes of the distribution of light inside leaves of Hydrophyllum canadense confirmed that oil infiltration decreased the amount of back-scattered light and that most of the light scattering for this species occurred from the middle of the palisade layer to the middle of the spongy mesophyll. We were not, however, able to assess the potential contribution of reflectance from the internal abaxial epidermis to total internal light scattering in these experiments. Using a mathematical model to compare the response of net photosynthesis (O₂, flux) to incident irradiance for control leaves of H. canadense and theoretical leaves with no intercellular reflectance, we calculated that intercellular reflectance caused a 1.97-fold increase in photosynthesis at 20 μmol m^?2s^?1 (incident photon flux density). This enhancement of absorption and photosynthesis by inter-cellular reflectance, without additional production and maintenance of photosynthetic pigments, may maintain shade leaves above the photosynthetic light compensation point between sunflecks and maintain the light induction state during protracted periods of low diffuse light.     

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.     

Predicting tropical plant physiology from leaf and canopy spectroscopy

A broad regional understanding of tropical forest leaf photosynthesis has long been a goal for tropical forest ecologists, but it has remained elusive due to difficult canopy access and high species diversity. Here we develop an empirical model to predict sunlit, light-saturated, tropical leaf photosynthesis using leaf and simulated canopy spectra. To develop this model, we used partial least squares (PLS) analysis on three tropical forest datasets (159 species), two in Hawaii and one at the biosphere 2 laboratory (B2L). For each species, we measured light-saturated photosynthesis (A), light and CO₂ saturated photosynthesis (A _max), respiration (R), leaf transmittance and reflectance spectra (400–2,500 nm), leaf nitrogen, chlorophyll a and b, carotenoids, and leaf mass per area (LMA). The model best predicted A [r ²  = 0.74, root mean square error (RMSE) = 2.9 μmol m⁻² s⁻¹)] followed by R (r ²  = 0.48), and A _max (r ²  = 0.47). We combined leaf reflectance and transmittance with a canopy radiative transfer model to simulate top-of-canopy reflectance and found that canopy spectra are a better predictor of A (RMSE = 2.5 ± 0.07 μmol m⁻² s⁻¹) than are leaf spectra. The results indicate the potential for this technique to be used with high-fidelity imaging spectrometers to remotely sense tropical forest canopy photosynthesis.     

Impact of nitrogen allocation on growth and photosynthesis of Miscanthus (Miscanthus × giganteus)

Nitrogen (N) addition typically increases overall plant growth, but the nature of this response depends upon patterns of plant nitrogen allocation that vary throughout the growing season and depend upon canopy position. In this study seasonal variations in leaf traits were investigated across a canopy profile in Miscanthus (Miscanthus × giganteus) under two N treatments (0 and 224 kg ha^?1) to determine whether the growth response of Miscanthus to N fertilization was related to the response of photosynthetic capacity and nitrogen allocation. Miscanthus yielded 24.1 Mg ha^?1 in fertilized plots, a 40% increase compared to control plots. Photosynthetic properties, such as net photosynthesis (A), maximum rate of rubisco carboxylation (V_cmax), stomatal conductance (g_s) and PSII efficiency (F_v'/F_m'), all decreased significantly from the top of the canopy to the bottom, but were not affected by N fertilization. N fertilization increased specific leaf area (SLA) and leaf area index (LAI). Leaf N concentration in different canopy layers was increased by N fertilization and the distribution of N concentration within canopy followed irradiance gradients. These results show that the positive effect of N fertilization on the yield of Miscanthus was unrelated to changes in photosynthetic rates but was achieved mainly by increased canopy leaf area. Vertical measurements through the canopy demonstrated that Miscanthus adapted to the light environment by adjusting leaf morphological and biochemical properties independent of nitrogen treatments. GPP estimated using big leaf and multilayer models varied considerably, suggesting a multilayer model in which V_cmax changes both through time and canopy layer could be adopted into agricultural models to more accurately predict biomass production in biomass crop ecosystems.     

Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest

Accurate estimation of terrestrial gross primary productivity (GPP) remains a challenge despite its importance in the global carbon cycle. Chlorophyll fluorescence (ChlF) has been recently adopted to understand photosynthesis and its response to the environment, particularly with remote sensing data. However, it remains unclear how ChlF and photosynthesis are linked at different spatial scales across the growing season. We examined seasonal relationships between ChlF and photosynthesis at the leaf, canopy, and ecosystem scales and explored how leaf‐level ChlF was linked with canopy‐scale solar‐induced chlorophyll fluorescence (SIF) in a temperate deciduous forest at Harvard Forest, Massachusetts, USA. Our results show that ChlF captured the seasonal variations of photosynthesis with significant linear relationships between ChlF and photosynthesis across the growing season over different spatial scales (R² = 0.73, 0.77, and 0.86 at leaf, canopy, and satellite scales, respectively; P < 0.0001). We developed a model to estimate GPP from the tower‐based measurement of SIF and leaf‐level ChlF parameters. The estimation of GPP from this model agreed well with flux tower observations of GPP (R² = 0.68; P < 0.0001), demonstrating the potential of SIF for modeling GPP. At the leaf scale, we found that leaf F_q’/F_m’, the fraction of absorbed photons that are used for photochemistry for a light‐adapted measurement from a pulse amplitude modulation fluorometer, was the best leaf fluorescence parameter to correlate with canopy SIF yield (SIF/APAR, R² = 0.79; P < 0.0001). We also found that canopy SIF and SIF‐derived GPP (GPP_SIF) were strongly correlated to leaf‐level biochemistry and canopy structure, including chlorophyll content (R² = 0.65 for canopy GPP_SIF and chlorophyll content; P < 0.0001), leaf area index (LAI) (R² = 0.35 for canopy GPP_SIF and LAI; P < 0.0001), and normalized difference vegetation index (NDVI) (R² = 0.36 for canopy GPP_SIF and NDVI; P < 0.0001). Our results suggest that ChlF can be a powerful tool to track photosynthetic rates at leaf, canopy, and ecosystem scales.     

Non-invasive monitoring of photosynthetic activity and water content in forest lichens by spectral reflectance data and RGB colors from photographs

There is a need for non-invasive monitoring of temporal and spatial variation in hydration and photosynthetic activity of red-listed poikilohydric autotrophs. Here, we simultaneously recorded kinetics in RGB-colors (photos), reflectance spectra, water content, maximal (F_V/F_M), and effective quantum yield of PSII (Φ_PSII) during desiccation in foliose lichens differing in cortical characteristics and photobionts. The spectral absorbance peaks of chlorophyll a, phycocyanin, and phycoerythrin were clearly displayed at high hydration levels. Brightness and total RGB colors of the lichens strongly increased during desiccation. The normalized difference vegetation index (NDVI) efficiently estimated hydration level and Φ_PSII – a proxy for lichen photosynthesis – in all species, including threatened old forest lichens. Color and reflectance indices based on green wavelengths gave good estimates of water content in cephalo- and chlorolichens, but not in cyanolichens with a wider range of photosynthetic pigments. Due to species-specific characteristics, species-wise calibration is essential for non-invasive assessments of lichen functioning.     

Reproductive success of soybean (Glycine max L. Merril) cultivars and exotic lines under high daytime temperature

The objectives were to (a) quantify the effects of high daytime temperature (HDT) from gametogenesis to full bloom on photosynthesis and pod set in soybean (Glycine max L. Merril) genotypes and (b) assess the relationships among photosynthesis, cardinal temperatures for pollen germination, in vitro pollen germination percentage, canopy reflectance, and pod‐set percentage. Three field experiments were conducted, and Experiment I had HDT between gametogenesis and full bloom (36.5°C to 38.6°C) compared with Experiments II and III (29.5°C to 31.6°C; optimum temperature). HDT decreased photosynthesis (22%) and pod‐set percent (11%) compared with Experiment III. Cultivars had higher photosynthesis and pod‐set percent than plant introduction (PI) lines. The cultivars (i.e., IA3023 and KS4694) and PI lines (i.e., PI393540 and PI588026A) were HDT tolerant and susceptible, respectively. The decreased pod‐set percentage in susceptible genotypes (PI lines) was associated with pollen characteristics. Significant positive (r² ≥ 0.67) association between photosynthesis, cardinal temperatures for pollen germination (T_opt and T_max) with pod‐set percentage was observed. However, a negative (r² ≥ ?0.43) association between photosynthesis and pod set with canopy reflectance at visible spectrum was observed. In vitro pollen germination and canopy reflectance at visible spectrum can be used as a high‐throughput phenotypic tool for breeding HDT‐tolerant genotypes.     

A model of the seasonal pattern of carbon acquisition in two woodland herbs,Mercurialis perennis L. and Geum urbanum L.

Summary Seasonal changes in the light and temperature dependence of photosynthesis were investigated in field grown plants of Mercurialis perennis and Geum urbanum. In both species changes in photosynthetic capacity were closely related to the development of the overstorey canopy. In G. urbanum there was a marked shift in the temperature dependence of photosynthesis through the season whereas no such pattern was found in M. perennis. Model predictions of field rates of photosynthesis were made using the measurements of light and temperature dependence in the laboratory and validated against field observations. Long term continuous records of light and temperature in the field were used in conjunction with the model to make predictions of carbon acquisition in shoots of the two species through the season. These calculations indicated that G. urbanum was able to take advantage of high light levels just prior to canopy closure through a combination of high photosynthetic capacity, the ability to maintain photosynthesis at relatively low temperatures and the presence of overwintering leaves. In M. perennis leaf development was early enough to utilise the high spring light period. After canopy closure M. perennis maintained a higher average rate of CO₂ flux due to a combination of high apparent quantum efficiency and low rates of respiration.     

Temperature Sensitivity of Photosynthesis in Lolium perenne Swards: A Comparison of Two Methods for Deriving Photosynthetic Parameters from in vivo Measurements

The seasonal variation in photosynthetic rate of grass swards is partly the result of changes in the environment and partly the result of changes in the photosynthetic capacity of the sward itself. We evaluated two types of photosynthesis equations regarding their capacity to analyse seasonal and short-term temperature effects on photosynthesis of ryegrass (Lolium perenne L.). Intact cores of a field-grown ryegrass sward were taken to the laboratory 10 d after cutting for measurement of photosynthesis under controlled conditions. This was done during two four-week periods, in summer and autumn. Net photosynthetic rate (P _N) of the sward was lower in autumn than in summer. Both a simple negatively exponential photosynthesis irradiance-response curve and the Farquhar equations for photosynthesis were applied to the in vivo canopy measurements. Application of the irradiance-response curve showed that irradiance-saturated gross photosynthetic rate increased linearly with increasing temperature and was higher in summer than in autumn. The initial radiation use efficiency did not differ between the seasons but decreased with the temperature rise. This explains the observation that total canopy photosynthetic rate decreased after short-term temperature increases in both seasons. The parameters in Farquhar equations that represent the temperature sensitivity of the maximum electron transport rate and of the Michaelis-Menten constants for CO₂ and O₂ fixation could not be quantified satisfactorily. Parameterisation of the Farquhar equations was hampered by a lack of robust information on many biochemical parameters, and the use of simple empirical response-functions may be preferable in the case of in vivo canopy measurements on grass swards.     

Seasonal and annual respiration of a ponderosa pine ecosystem

The net ecosystem exchange of CO₂ between forests and the atmosphere, measured by eddy covariance, is the small difference between two large fluxes of photosynthesis and respiration. Chamber measurements of soil surface CO₂ efflux (F_s), wood respiration (F_w) and foliage respiration (F_f) help identify the contributions of these individual components to net ecosystem exchange. Models developed from the chamber data also provide independent estimates of respiration costs. We measured CO₂ efflux with chambers periodically in 1996–97 in a ponderosa pine forest in Oregon, scaled these measurements to the ecosystem, and computed annual totals for respiration by component. We also compared estimated half-hourly ecosystem respiration at night (F_nc) with eddy covariance measurements. Mean foliage respiration normalized to 10 °C was 0.20 μmol m^–2 (hemi-leaf surface area) s^–1, and reached a maximum of 0.24 μmol m^–2 HSA s^–1 between days 162 and 208. Mean wood respiration normalized to 10 °C was 5.9 μmol m^–3 sapwood s^–1, with slightly higher rates in mid-summer, when growth occurs. There was no significant difference (P > 0.10) between wood respiration of young (45 years) and old trees (250 years). Soil surface respiration normalized to 10 °C ranged from 0.7 to 3.0 μmol m^–2 (ground) s^–1 from days 23 to 329, with the lowest rates in winter and highest rates in late spring. Annual CO₂ flux from soil surface, foliage and wood was 683, 157, and 54 g C m^–2 y^–1, with soil fluxes responsible for 76% of ecosystem respiration. The ratio of net primary production to gross primary production was 0.45, consistent with values for conifer sites in Oregon and Australia, but higher than values reported for boreal coniferous forests. Below-ground carbon allocation (root turnover and respiration, estimated as F_s– litterfall carbon) consumed 61% of GPP; high ratios such as this are typical of sites with more water and nutrient constraints. The chamber estimates were moderately correlated with change in CO₂ storage in the canopy (F_stor) on calm nights (friction velocity u* < 0.25 m s^–1; R² = 0.60); F_stor was not significantly different from summed chamber estimates. On windy nights (u* > 0.25 m s^–1), the sum of turbulent flux measured above the canopy by eddy covariance and F_stor was only weakly correlated with summed chamber estimates (R² = 0.14); the eddy covariance estimates were lower than chamber estimates by 50%.     

Partitioning net ecosystem carbon exchange with isotopic fluxes of CO2

Because biological and physical processes alter the stable isotopic composition of atmospheric CO₂, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of ¹³CO₂ above terrestrial ecosystems can potentially be used to separate net ecosystem CO₂ exchange (NEE) into its component fluxes, net photosynthetic assimilation (F_A) and ecosystem respiration (F_R). In this paper theory is developed to partition measured NEE into F_A and F_R, using measurements of fluxes of CO₂ and ¹³CO₂, and isotopic composition of respired CO₂ and forest air. The theory is then applied to fluxes measured (or estimated, for ¹³CO₂) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in ¹³CO₂ fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in F_A and F_R were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night‐time NEE and temperature (a standard technique which provides alternate estimates of F_R and F_A). The light response curve for photosynthesis (F_A vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation‐weighted discrimination against ¹³CO₂ for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole‐canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of ¹³CO₂ flux are needed to assess the validity of the assumptions used and provide defensible isotope‐based estimates of the component fluxes of net ecosystem exchange.     

Spectral reflectance of Thalassia testudinum (Hydrocharitaceae) seagrass: low salinity effects

Massive anthropogenic changes in estuarine salinities, from manipulations of freshwater flows, are again occurring through governmental projects “correcting” past freshwater alterations. The downstream effects of increased freshwater on seagrass meadows, a major fisheries and ecosystem habitat, are not clear. Spectral responses to low salinities were quantitatively delimited for the important habitat seagrass Thalassia testudinum utilizing spectral reflectance measurements for the first time (non-invasive sampling). Over a range of salinities (32–16 parts per thousand sea salts [ppt] for 24 h) and spectra (308–1138 nm), Thalassia specimens showed statistically significant differences in spectral values (P < 0.05) between treatments at normal (32 ppt) and 50% reduced (16 ppt) seawater. Mature blades yellowed at low salinities. Reflectance changes at 525 nm and 650–680 nm at low salinities suggested changes in xanthophylls and chlorophylls. Four indices were also used to characterize the reflectance spectra to delineate the effect of the salinity changes: (1) The normalized difference vegetation index (NDVI) for mature blades reduced at 16 ppt from that at 32 ppt. (2) The chlorophyll normalized difference index (Chl NDI) suggested chlorophyll content decreases in response to reduced salinity. (3) The structure independent pigment index (SIPI), higher in mature blades at 16 ppt than new blades, indicates a higher carotenoid : chlorophyll ratio in mature blades. (4) The photochemical reflectance index (PRI) suggested a lower photochemical efficiency at lower salinities. The main low-salinity effect on Thalassia physiology delineated herein is likely through changes in pigmentation (decreases in chlorophyll and changes in xanthophyll cycle epoxidation).