Seasonal Variations In Photosynthesis Research Articles

Modeling starch dynamics from seasonal variations of photosynthesis, growth, and respiration.

Nonstructural carbohydrates (NSCs) buffer differences in plant carbon supply (photosynthesis) and demand (respiration, growth, etc.) but the regulation of their dynamics remains unresolved. Seasonal variations in NSCs are well-documented, but differences in the time-average, amplitude, phase, and other characteristics across ecosystems and functional types lack explanation; furthermore, observed dynamics do not always match expectations. The failure to match observed and expected dynamics has stimulated debate on whether carbon supply or demand drives NSC dynamics. To gain insight into how carbon supply and demand drive seasonal NSC dynamics, we derive a simple model of NSC dynamics based on carbon mass balance and linearizing the NSC demand to determine how supply-driven and demand-driven seasonal NSC dynamics differ. We find that supply-driven and demand-driven dynamics yield distinct timings of seasonal extrema, and supply overrides demand when carbon supply is low in winter (e.g., at high latitudes). Our results also suggest that NSC dynamics often lag changes carbon mass balance. We also predict differences in NSC dynamics across mass, suggesting saplings are more dynamics and respond faster to the environment than mature trees. Our findings suggest substrate-dependent regulation with environmental variation is sufficient to generate complex NSC dynamics.

Seasonal variations in leaf-level photosynthesis and water use efficiency of three isohydric to anisohydric conifers on the Tibetan Plateau

Different stomatal behaviors among species under drought conditions may affect species-specific seasonal variations in photosynthesis and water use efficiency (WUE), and could enable species to follow differing growth strategies. Here we monitored leaf gas exchange, intra-annual radial growth, leaf stoichiometry, and microclimate of three conifers with contrasting stomatal behaviors (isohydric Picea wilsonii and Pinus tabuliformis vs. anisohydric Juniperus przewalskii) over three growing seasons, and collected tree-ring records of the monitored species, on the cold and arid northeastern Tibet Plateau. We used these data to assess the species-specific seasonal variations in net photosynthetic rate (An), transpiration efficiency (TE) and intrinsic water use efficiency (WUEi), and their impacts on growth.Our observations show that the An of all three conifers was greatest in autumn. This peak was closely related to increasing stomatal conductance following alleviation of drought stress, rather than to variations in leaf nitrogen concentrations. WUEi of the three conifers did not exhibit a regular seasonal pattern, but TE was significantly lower in summer than in spring and autumn, due to the regulation by vapor pressure deficit (VPD). Increasing summer VPD reduced TE mainly by increasing transpiration rate for J. przewalskii and by decreasing An for the isohydric conifers.The increases of An from summer to autumn were much greater for the two isohydric Pinaceae plants than for J. przewalskii. This was consistent with the growth-climate relationships, in which the autumn drought in the previous year limited radial growth of the isohydric conifers more strongly than in the anisohydric conifer. Therefore, we suggest that the growth of isohydric species may profit more from the photosynthates accumulated during the previous post-growing season in the cold and arid Tibetan Plateau.

Seasonal trends in photosynthesis and leaf traits in scarlet oak.

Understanding seasonal variation in photosynthesis is important for understanding and modeling plant productivity. Here, we used shotgun sampling to examine physiological, structural and spectral leaf traits of upper canopy, sun-exposed leaves in Quercus coccinea Münchh (scarlet oak) across the growing season in order to understand seasonal trends, explore the mechanisms underpinning physiological change and investigate the impact of extrapolating measurements from a single date to the whole season. We tested the hypothesis that photosynthetic rates and capacities would peak at the summer solstice, i.e., at the time of peak photoperiod. Contrary to expectations, our results reveal a late-season peak in both photosynthetic capacity and rate before the expected sharp decrease at the start of senescence. This late-season maximum occurred after the higher summer temperatures and vapor pressure deficit and was correlated with the recovery of leaf water content and increased stomatal conductance. We modeled photosynthesis at the top of the canopy and found that the simulated results closely tracked the maximum carboxylation capacity of Rubisco. For both photosynthetic capacity and modeled top-of-canopy photosynthesis, the maximum value was therefore not observed at the summer solstice. Rather, in each case, the measurements at and around the solstice were close to the overall seasonal mean, with values later in the season leading to deviations from the mean by up to 41 and 52%, respectively. Overall, we found that the expected Gaussian pattern of photosynthesis was not observed. We conclude that an understanding of species- and environment-specific changes in photosynthesis across the season is essential for correct estimation of seasonal photosynthetic capacity.

Seasonal variations in photosynthetic functions of the urban landscape tree species Gingko biloba: photoperiod is a key trait

Seasonal variations in the leaf photosynthetic traits of an urban tree, Ginkgo biloba, were almost synchronized with the photoperiod. Non-stomatal limitations were a cue for photosynthesis in Ginkgo biloba. Photosynthetic functions, which are key traits in determining the carbon uptake of urban trees, exhibit significant seasonal variations in temperate zones. It is essential to clarify the seasonal dynamics of photosynthesis to evaluate the CO2 uptake in urban areas. We investigated seasonal variations in the photosynthetic traits of Ginkgo biloba, which is a major urban deciduous tall tree often planted in Japan. Seasonal variations in the leaf photosynthetic traits, including the maximum photosynthesis rate, maximum carboxylation rate, and mesophyll and stomatal conductance, were well fitted to quadratic models, in which they peaked around the summer solstice and then declined with time. Seasonal variations in the environmental variables, such as photoperiod, temperature, and solar radiation, were compared to those of the leaf photosynthetic traits, in which the photoperiod explained well variations in the leaf photosynthetic traits. Seasonal variations in photosynthesis were largely governed by non-stomatal limitations, i.e., mesophyll and biochemical limitations. The high synchrony of the photoperiod and photosynthetic traits during leaf maturation may cause an enhancement in the daily carbon uptake of G. biloba leaves around the summer solstice, which has the longest photoperiod, and thus, will lead to an increase in the annual carbon uptake.

The impact of a simple representation of non-structural carbohydrates on the simulated response of tropical forests to drought

Abstract. Accurately representing the response of ecosystems to environmental change in land surface models (LSMs) is crucial to making accurate predictions of future climate. Many LSMs do not correctly capture plant respiration and growth fluxes, particularly in response to extreme climatic events. This is in part due to the unrealistic assumption that total plant carbon expenditure (PCE) is always equal to gross carbon accumulation by photosynthesis. We present and evaluate a simple model of labile carbon storage and utilisation (SUGAR) designed to be integrated into an LSM, which allows simulated plant respiration and growth to vary independent of photosynthesis. SUGAR buffers simulated PCE against seasonal variation in photosynthesis, producing more constant (less variable) predictions of plant growth and respiration relative to an LSM that does not represent labile carbon storage. This allows the model to more accurately capture observed carbon fluxes at a large-scale drought experiment in a tropical moist forest in the Amazon, relative to the Joint UK Land Environment Simulator LSM (JULES). SUGAR is designed to improve the representation of carbon storage in LSMs and provides a simple framework that allows new processes to be integrated as the empirical understanding of carbon storage in plants improves. The study highlights the need for future research into carbon storage and allocation in plants, particularly in response to extreme climate events such as drought.

Field Performance of Saccharum × Miscanthus Intergeneric Hybrids (Miscanes) Under Cool Climatic Conditions of Northern Japan

Physiological susceptibility to early- and late-season chilling limits commercial production of sugarcane (Saccharum spp. hybrid), a major crop for lignocellulosic biomass, refined sugar, and bioethanol, to tropical and the warmest subtropical regions. Interspecific and intergeneric hybridization have been used to broaden the genetic base of sugarcane and improve its adaptation to temperate climates. Chilling tolerance can be introgressed in sugarcane through intergeneric hybridization with Miscanthus, a cold-tolerant C4 perennial grass, which is genetically homologous to sugarcane. This study evaluated intergeneric F1 hybrids of Saccharum × Miscanthus, miscanes, which included two genotypes of sugarcane × Miscanthus sinensis and sixteen genotypes of sugarcane × Miscanthus sacchariflorus, for their seasonal variation in photosynthesis and biomass production under field conditions in Hokkaido, Japan, to identify promising genotypes and traits, which can be selected to further improve sugarcane. Results showed several of the miscane genotypes had high early- and late-season photosynthesis coupled with high biomass production, which likely indicates chilling tolerance. High broad-sense heritabilities for traits, including stem diameter, tiller number, leaf width, leaf and stem dry weight, and high correlations between these traits and dry matter yield indicate selections can be made efficiently to improve sugarcane. Although none of the miscanes overwintered at the experimental location, we identified miscane “JM 14-09” as a superior genotype for introgression breeding programs and as a potential energycane cultivar for its high biomass-production capacity.

Effects of Light Component and Water Stress on Photosynthesis of Amazon Rainforests During the 2015/2016 El Niño Drought

AbstractWhether enhanced sunshine increases photosynthesis in Amazon rainforests during drought is unclear. Here we used a light component‐based two‐leaf‐photosynthesis model, driven with climate data and satellite vegetation data, to inspect the controlling mechanisms among climate factors on gross primary production (GPP) during the 2015/2016 El Niño drought event. We found that simulated GPP and Moderate Resolution Imaging Spectroradiometer enhanced vegetation index indicated an Amazonian “browning” and not a “green up” during the 2015/2016 El Niño year relative to the 2011–2014 interval. The result shows that, along with intensified sunlight, diffuse sunlight and diffuse fraction as well as canopy light use efficiency decreased, which further produced a decreased potential GPP* (determined by light components and leaf area index of shaded and sunlit leaves). The decreased GPP* and drought‐induced water stress jointly reduced canopy photosynthesis of Amazon rainforests during the 2015/2016 drought. The light component variations caused a reduction in GPP but with a magnitude inferior to the GPP reduction from water stress. These findings suggest that intensified sunlight did not enhance photosynthesis of Amazon rainforests and highlight the important role of light components in interannual and seasonal variations of photosynthesis in Amazon rainforests.

Remote Sensing Indices to Measure the Seasonal Dynamics of Photosynthesis in a Southern China Subtropical Evergreen Forest

The accurate measurement of the dynamics of photosynthesis in China's subtropical evergreen forest ecosystems is an important contribution to carbon (C) sink estimates in global terrestrial ecosystems and their responses to climate change. Eddy covariance has historically been the only direct method to assess C flux of whole ecosystems with high temporal resolution, but it suffers from limited spatial resolution. During the last decade, continuous global monitoring of plant primary productivity from spectroradiometer sensors on flux towers and satellites has extended the temporal and spatial coverage of C flux observations. In this study, we evaluated the performance of two physiological remote sensing indices, fluorescence reflectance index (FRI) and photochemical reflectance index (PRI), to measure the seasonal variations of photosynthesis in a subtropical evergreen forest ecosystem using continuous canopy spectral and flux measurements in the Dinghushan Nature Reserve in southern China. The more commonly used NDVI has been shown to be saturated and mainly affected by illumination (R2=0.88, p < 0.001), but FRI and PRI could better track the seasonal dynamics of plant photosynthetic functioning by comparison and are less affected by illumination (R2=0.13 and R2=0.51, respectively) at the seasonal scale. FRI correlated better with daily gross primary production (GPP) in the morning hours than in the afternoon hours, in contrast to PRI which correlated better with light-use efficiency (LUE) in the afternoon hours. Both FRI and PRI could show greater correlations with GPP and LUE respectively in the senescence season than in the recovery-growth season. When incident PAR was taken into account, the relationship between GPP and FRI was improved and the correlation coefficient increased from 0.22 to 0.69 (p < 0.001). The strength of the correlation increased significantly in the senescence season (R2=0.79, p < 0.001). Our results demonstrate the application of FRI and PRI as physiological indices for the accurate measurement of the seasonal dynamics of plant community photosynthesis in a subtropical evergreen forest, and suggest these indices may be applied to carbon cycle models to improve the estimation of regional carbon budgets.

Seasonal variation in photosynthesis performance of cassava at two different growth stages under irrigated and rain-fed conditions in a tropical savanna climate

Seasonal variations in photosynthesis of cassava cv. Rayong 9 (RY9) under irrigated and rain-fed conditions were evaluated at the age of three and six months after planting (MAP). Photosynthetic light-response (PN/I) curves revealed that cassava leaves attained the highest maximum net photosynthetic rates (PNmax) in the rainy season, followed by the hot one, while the lowest PNmax was found in the cool season. Photosynthetic potential of the 3-month-old plants was mostly higher than that of the 6-month-old plants, and the seasonal variation in photosynthetic capacity was also more apparent in the younger plants. PN/I curves were used to predict daily net photosynthetic rate (PN) for each season based on daily average solar radiation data. The predicted PN were considerably lower than the PNmax values. This indicated that solar radiation is a limiting factor for photosynthesis, particularly in the rainy season. The data provided basic information for breeding cassava genotypes with enhanced photosynthesis during the period of unfavorable environment. Furthermore, the data are potentially useful in modeling photosynthesis and crop growth as affected by environmental factors.

Progress in Remote Sensing of Photosynthetic Activity over the Amazon Basin.

Although quantifying the massive exchange of carbon that takes place over the Amazon Basin remains a challenge, progress is being made as the remote sensing community moves from using traditional, reflectance-based vegetation indices, such as the Normalized Difference Vegetation Index (NDVI), to the more functional Photochemical Reflectance Index (PRI). This new index, together with satellite-derived estimates of canopy light interception and Sun-Induced Fluorescence (SIF), provide improved estimates of Gross Primary Production (GPP). This paper traces the development of these new approaches, compares the results of their analyses from multiple years of data acquired across the Amazon Basin and suggests further improvements in instrument design, data acquisition and processing. We demonstrated that our estimates of PRI are in generally good agreement with eddy-flux tower measurements of photosynthetic light use efficiency (ε) at four sites in the Amazon Basin: r2 values ranged from 0.37 to 0.51 for northern flux sites and to 0.78 for southern flux sites. This is a significant advance over previous approaches seeking to establish a link between global-scale photosynthetic activity and remotely-sensed data. When combined with measurements of Sun-Induced Fluorescence (SIF), PRI provides realistic estimates of seasonal variation in photosynthesis over the Amazon that relate well to the wet and dry seasons. We anticipate that our findings will steer the development of improved approaches to estimate photosynthetic activity over the tropics.

Seasonal variations in photosynthesis, intrinsic water-use efficiency and stable isotope composition of poplar leaves in a short-rotation plantation.

Photosynthetic carbon assimilation and transpirational water loss play an important role in the yield and the carbon sequestration potential of bioenergy-devoted cultures of fast-growing trees. For six poplar (Populus) genotypes in a short-rotation plantation, we observed significant seasonal and genotypic variation in photosynthetic parameters, intrinsic water-use efficiency (WUEi) and leaf stable isotope composition (δ13C and δ18O). The poplars maintained high photosynthetic rates (between 17.8 and 26.9 μmol m−2 s−1 depending on genotypes) until late in the season, in line with their fast-growth habit. Seasonal fluctuations were mainly explained by variations in soil water availability and by stomatal limitation upon photosynthesis. Stomatal rather than biochemical limitation was confirmed by the constant intrinsic photosynthetic capacity (Vcmax) during the growing season, closely related to leaf nitrogen (N) content. Intrinsic water-use efficiency scaled negatively with carbon isotope discrimination (Δ13Cbl) and positively with the ratio between mesophyll diffusion conductance (gm) and stomatal conductance. The WUEi – Δ13Cbl relationship was partly influenced by gm. There was a trade-off between WUEi and photosynthetic N-use efficiency, but only when soil water availability was limiting. Our results suggest that seasonal fluctuations in relation to soil water availability should be accounted for in future modelling studies assessing the carbon sequestration potential and the water-use efficiency of woody energy crops.

Roots are important sources of carbohydrates during flowering and fruiting in ‘Valencia’ sweet orange trees with varying fruit load

The influence of fruit loading on flowering and fruiting, CO2 assimilation, and non-structural carbohydrates (NSC) in citrus trees was evaluated in four-year-old ‘Valencia’ sweet orange grafted onto ‘Rangpur’ lime. One group of trees was completely de-fruited (DFT) on May 14 (autumn), whereas fruit were left on the remaining trees (FT). The seasonal variation in photosynthesis and stomatal conductance was unaffected by fruit loading, with the minimum diurnal-integrated CO2 assimilation (PNI) occurring in July (98.3mmolm−2day−1) and the maximum PNI occurring in November (199.5mmolm−2day−1). Fruit loading inhibited sprouting and flowering in the citrus trees, but this effect was not correlated with NSC in the leaves, branches, or roots. The DFT trees exhibited nearly four times as many reproductive structures as the FT trees, with a high remobilization of reserves. Our data showed that flowers are stronger sinks than fruit and that flowering is the most expensive phenological stage. In the DFT trees, approximately 80% of NSC were consumed prior to the end of fruit drop, primarily until flowering. NSC reserves from leaves, branches, and roots were remobilized. Between the stages of de-fruiting and the end of physiological fruit drop, the plants remobilized approximately 312g NSC, with the roots contributing more than 230g NSC per plant.

Seasonal variations of photosynthesis, gas exchange, quantum efficiency of photosystem II and biochemical responses of Jatropha curcas L. grown in semi-humid and semi-arid areas subject to water stress

This study aimed to compare the physiological adaptations of Jatropha curcas in two regions with different climates, the Agreste (a semi-arid tropical climate) and the Florest zone (a semi-humid tropical climate) subject to water stress. Measurements of gas exchange and photochemical and biochemical efficiency were performed in two seasons, the dry season (September to February) and the rainy season (April and August), between 2006 and 2007. During the dry season in the Florest zone, the J. curcas plants presented better photosynthetic performance, with smaller decreases in the rates of photosynthesis, stomatal conductance and transpiration because the conditions of water availability in the soil were slightly higher than in the semi-arid region. A greater vapor pressure deficit was observed during the dry season of the semi-arid region, showing a negative correlation with the stomatal opening of the leaves, which contributed to a reduction in the values of the net photosynthetic rate. During the dry season in the semi-arid region, dynamic photoinhibition occurred in the leaves of J. curcas around mid-day, with a rapid recovery during the afternoon. However, chronic photoinhibition may have occurred in some plants with a severe water deficit. A reduction in the effective quantum efficiency was also observed during the driest months. The increase of catalase activity and the accumulation of proline, total soluble sugars and amino acids during periods of low water availability in the soil suggest that J. curcas presents an efficient protection mechanism that allows the plant to survive under drought conditions.

Modelling leaf photosynthetic and transpiration temperature-dependent responses in Vitis vinifera cv. Semillon grapevines growing in hot, irrigated vineyard conditions.

Grapevines growing in Australia are often exposed to very high temperatures and the question of how the gas exchange processes adjust to these conditions is not well understood. The aim was to develop a model of photosynthesis and transpiration in relation to temperature to quantify the impact of the growing conditions on vine performance. Leaf gas exchange was measured along the grapevine shoots in accordance with their growth and development over several growing seasons. Using a general linear statistical modelling approach, photosynthesis and transpiration were modelled against leaf temperature separated into bands and the model parameters and coefficients applied to independent datasets to validate the model. Photosynthesis, transpiration and stomatal conductance varied along the shoot, with early emerging leaves having the highest rates, but these declined as later emerging leaves increased their gas exchange capacities in accordance with development. The general linear modelling approach applied to these data revealed that photosynthesis at each temperature was additively dependent on stomatal conductance, internal CO(2) concentration and photon flux density. The temperature-dependent coefficients for these parameters applied to other datasets gave a predicted rate of photosynthesis that was linearly related to the measured rates, with a 1 : 1 slope. Temperature-dependent transpiration was multiplicatively related to stomatal conductance and the leaf to air vapour pressure deficit and applying the coefficients also showed a highly linear relationship, with a 1 : 1 slope between measured and modelled rates, when applied to independent datasets. The models developed for the grapevines were relatively simple but accounted for much of the seasonal variation in photosynthesis and transpiration. The goodness of fit in each case demonstrated that explicitly selecting leaf temperature as a model parameter, rather than including temperature intrinsically as is usually done in more complex models, was warranted.

Seasonal variation in photosynthesis in relation to differing environmental factors of dominant plant species in three successional communities in Hangzhou Bay Wetlands, East China

We investigated the ecophysiology (physiological ecology) of plant succession within a coastal wetland community. The photosynthetic characteristics of six plants were studied; Scirpus mariqueter and Carex scabrifolia in early successional communities, Phragmites communis and Tamarix chinensis in mid successional communities, and Salix matsudana and Imperata cylindrica in later successional communities. Environmental factors were concurrently determined for each community. Relationships between photosynthetic and environmental factors were analyzed by path analysis, and the dependence of photosynthesis on particular environmental factors was shown to vary between plants in the differing successional communities. Ability to cope with irradiation appears to be one of the most important factors influencing plant succession. The increasing stratification of plant communities following colonization of new coastal land necessitates differing physiological strategies and photosynthetic dynamics in the later colonizing plants.

Diurnal and seasonal variation in photosynthesis of peach palms grown under subtropical conditions

The Amazonian peach palm (Bactris gasipaes Kunth) has been grown for heart-of-palm production under subtropical conditions. As we did not see any substantial study on its photosynthesis under Amazonian or subtropical conditions, we carried out an investigation on the diurnal and seasonal variations in photosynthesis of peach palms until the first heart-of-palm harvest, considering their relationship with key environmental factors. Spineless peach palms were grown in 80-L plastic pots, under irrigation. Gas exchange and chlorophyll fluorescence emission measurements were taken in late winter, mid spring, mid summer and early autumn, from 7:00 to 18:00 h, with an additional chlorophyll fluorescence measurement at 6:00 h. The highest net CO2 assimilation (P N), observed in mid summer, reached about 15 μmol m−2 s−1, which was about 20% higher than the maximum values found in autumn and spring, and 60% higher than that in winter The same pattern of diurnal course for P N was observed in all seasons, showing higher values from 8:00 to 9:00 h and declining gradually from 11:00 h toward late afternoon. The diurnal course of stomatal conductance (g s) followed the same pattern of P N, with the highest value of 0.6 mol m−2 s−1 being observed in February and the lowest one (0.23 mol m−2 s−1) in September. The maximal quantum yield of photosystem II (Fv/Fm) was above 0.75 in the early morning in all the months. The reversible decrease was observed around midday in September and October, suggesting the occurrence of dynamic photoinhibition. A significant negative correlation between the leaf-air vapour pressure difference (VPDleaf-air) and P N and a positive correlation between P N and g s were observed. The photosynthesis of peach palm was likely modulated mainly by the stomatal control that was quite sensible to atmospheric environmental conditions. Under subtropical conditions, air temperature (T air) and VPDleaf-air impose more significant effects over P N of peach palm than an excessive photosynthetic photon flux density (PPFD). The occurrence of dynamic photoinhibition indicates that under irrigation, peach palms appeared to be acclimated to the full-sunlight conditions under which they have been grown.

Seasonal variation in photosynthesis and diel carbon balance under natural conditions in two Peperomia species that differ with respect to leaf anatomy1

Abstract The occurrence of crassulacean acid metabolism (CAM) and C3 photosynthetic activities in Peperomia species has been linked to the differential distribution of C3 and C4 photosynthetic enzymes within the distinct leaf tissue layers. Our objectives were to test if relative thickness of the spongy mesophyll layer is an indication of CAM and if the distribution of water in the leaf tissue changes in response to drought. Under field conditions we studied photosynthesis and water relations of Peperomia obtusifolia and P. macrostachya, species that differ in leaf anatomy. Gas exchange and titratable acidity were measured over a complete 24 hour photoperiod. Plant water status was determined at dawn while chlorophyll a fluorescence data were collected during the day (12 noon). Thickness of the distinct leaf layers was determined with the aid of a calibrated light microscope. Gas exchange and titratable acidity showed that when well-hydrated, P. obtusifolia has typical C3 activity and P. macrostachya has ...

Seasonal variation in photosynthesis in six woody species with different leaf phenology in a valley savanna in southwestern China

During 2003–2005, we examined the effect of seasonal drought on water status, gas exchange, δ13C, chlorophyll fluorescence and spectral reflectance in six woody species in a valley savanna near the Yuanjiang River (the upper Red River) in southwestern China. Three different phenological types of these woody species were compared, i.e., an evergreen species, Cyclobalanopsis helferiana, two winter-deciduous (WD) species, Buchanania latifolia and Symplocos racemosa, and three drought-deciduous (DD) species, Terminthia paniculata, Wendlandia tinctoria and Woodfordia fruticosa. We aimed to test the following three hypotheses: (1) the evergreen and WD species employ a drought avoidance strategy, whereas DD species employ a drought tolerance strategy; (2) the evergreen and WD species have a more economical water use strategy than the DD species and (3) the evergreen and WD species have a stronger photoprotection capacity through thermal dissipation than the DD species. At the end of a prolonged drought, the predawn leaf water potential (Ψpd) in C. helferiana and S. racemosa dropped to ca. −0.8 MPa, whereas the Ψpd in B. latifolia remained close to zero and DD species were leafless. In the rainy seasons, maximal photosynthetic rates of the evergreen (18.4 μmol m−2 s−1) and W. fruticosa (18.0 μmol m−2 s−1) were higher than those of the other four species (12.2−13.8 μmol m−2 s−1). The evergreen and WD species responded to drought by closing stomata and thus maintained a constant relative water content (RWC), which is a typical drought avoidance strategy; however, it is at the expense of carbon gain. DD species maintained a high photosynthetic capacity with a decrease in both stomatal conductance and RWC until the driest period, and then shifted from the drought tolerance strategy to the avoidance mechanism by shoot dieback. There was no significant difference in the means of δ13C across the phenological groups. The evergreen and WD species had stronger heat dissipation than the DD species in dry seasons. All species increased leaf spectral reflectance, probably because of degradation of chlorophyll as indicated by the leaf reflectance index, which should reduce light harvesting. All species showed a strong increase in the ratio of red to green spectral reflectance of leaves during dry seasons, indicating the accumulation of anthocyanin, which may contribute to screening sunlight and scavenging reactive oxygen species. Different responses to drought of savanna woody species with different leaf phenologies may facilitate the partitioning of resource use and hence their co-existence.

Components of ecosystem respiration and an estimate of net primary productivity of an intermediate-aged Douglas-fir stand

Continuous half-hourly chamber-measured belowground ( R s) and eddy covariance (EC)-measured total ecosystem ( R e) respiration in a 56-year-old Pacific west coast Douglas-fir stand from 2003 to 2005 were analyzed to study their seasonal and interannual variability. Seasonal variation in both R s and R e was better predicted by soil temperature at the 5 cm depth than at any other depth and air temperature. R e was more responsive than R s to changes in temperature and soil water content. While nighttime ( R sn) and daytime ( R sd) soil respiration showed no difference in their response to soil temperature, daytime ecosystem respiration ( R ed) was less responsive than its nighttime counterpart ( R en) to changes in soil temperature. Half-hourly R ed was almost always smaller than R en possibly due to photoinhibition while daily total R ed was higher than daily total R en during summer and spring months but smaller in winter and autumn months with the latter due to fewer daylight hours. A distinct seasonal pattern in the R s to R e ratio was observed. The 3-year mean seasonal R s/ R e was at its minimum of 0.52 in spring followed by 0.63 in summer, 0.81 in autumn and a maximum of 0.86 in winter. Both daily total and half-hourly R sd/ R ed were larger than R sn/ R en. It appears that the different responses of R s and R e to environmental variables arise as a result of seasonal variations in photosynthesis, mobilization and use of stored carbohydrates, and differences in the phenology of aboveground and belowground plant tissues. On an annual basis, R s accounted for 62% of R e with the latter accounting for 86% of the carbon (C) assimilated in annual photosynthesis or gross primary productivity (GPP), leaving the net C sequestration efficiency (1 − R e/GPP) at 14%. In the relatively dry year of 2003, both R s and R e as well as GPP were the lowest but with the highest net ecosystem productivity (NEP) of the 3 years. The relatively wetter growing season soil moisture regime in 2005 resulted in the highest R e and GPP. Partitioning of R s into its autotrophic and heterotrophic components indicated that 54% of GPP was respired back to the atmosphere as autotrophic respiration and an additional 32% was lost in the decomposition of litterfall and soil organic matter. The mean annual estimate of net primary productivity (NPP) at 843 g C m −2 accounted for 47% of the mean annual GPP of 1815 g C m −2.

Hourly and seasonal variation in photosynthesis and stomatal conductance of soybean grown at future CO2 and ozone concentrations for 3 years under fully open‐air field conditions

It is anticipated that enrichment of the atmosphere with CO(2) will increase photosynthetic carbon assimilation in C3 plants. Analysis of controlled environment studies conducted to date indicates that plant growth at concentrations of carbon dioxide ([CO(2)]) anticipated for 2050 ( approximately 550 micromol mol(-1)) will stimulate leaf photosynthetic carbon assimilation (A) by 20 to 40%. Simultaneously, concentrations of tropospheric ozone ([O(3)]) are expected to increase by 2050, and growth in controlled environments at elevated [O(3)] significantly reduces A. However, the simultaneous effects of both increases on a major crop under open-air conditions have never been tested. Over three consecutive growing seasons > 4700 individual measurements of A, photosynthetic electron transport (J(PSII)) and stomatal conductance (g(s)) were measured on Glycine max (L.) Merr. (soybean). Experimental treatments used free-air gas concentration enrichment (FACE) technology in a fully replicated, factorial complete block design. The mean A in the control plots was 14.5 micromol m(-2) s(-1). At elevated [CO(2)], mean A was 24% higher and the treatment effect was statistically significant on 80% of days. There was a strong positive correlation between daytime maximum temperatures and mean daily integrated A at elevated [CO(2)], which accounted for much of the variation in CO(2) effect among days. The effect of elevated [CO(2)] on photosynthesis also tended to be greater under water stress conditions. The elevated [O(3)] treatment had no statistically significant effect on mean A, g(s) or J(PSII) on newly expanded leaves. Combined elevation of [CO(2)] and [O(3)] resulted in a slightly smaller increase in average A than when [CO(2)] alone was elevated, and was significantly greater than the control on 67% of days. Thus, the change in atmospheric composition predicted for the middle of this century will, based on the results of a 3 year open-air field experiment, have smaller effects on photosynthesis, g(s) and whole chain electron transport through photosystem II than predicted by the substantial literature on relevant controlled environment studies on soybean and likely most other C3 plants.