Enhanced decreases in rice evapotranspiration in response to elevated atmospheric carbon dioxide under warmer environments.

A short period of exposure to elevated CO2 is known to decrease evapotranspiration via stomatal closure. Based on theoretical evaluation of a canopy transpiration model, we hypothesized that this decrease in the evapotranspiration of rice under elevated CO2 was greater under higher temperature conditions due to an increased sensitivity of transpiration to changes in CO2 induced by the greater vapour pressure deficit. In a temperature gradient chamber-based experiment, a 200 ppm increase in CO2 concentration led to 0.4 mm (-7%) and 1.5 mm (-15%) decreases in 12 h evapotranspiration under ambient temperature and high temperature (+3.7°C) conditions, respectively. Model simulations revealed that the greater vapour pressure deficit under higher temperature conditions explained the variations in the reduction of evapotranspiration observed under elevated CO2 levels between the temperature treatments. Our study suggests the utility of a simple modelling framework for mechanistic understanding of evapotranspiration and crop energy balance system under changing environmental conditions.

Transcriptome dynamics of rice in natura: Response of above and below ground organs to microclimate.

The long-term dynamics of the transcriptome under natural field conditions remain unclear. We conducted comprehensive gene expression analyses of rice leaves and roots grown under natural field conditions for a long period, from the tillering stage to the ripening stage. In this experiment, changes in the transcriptome were captured in relation to microclimatic parameters, particularly potential evaporation (Ep), which is a multiple meteorological factor and acts as an indicator of transpirational demand. The results indicated  that many genes were regulated by changes in temperature and Ep in both leaves and roots. Furthermore, the correlation between gene expression and meteorological factors differed significantly between the vegetative and reproductive stages. Since Ep triggers transpiration, we analyzed aquaporin gene expression, which is responsible for water transport, and found that many aquaporin genes in leaves were positively correlated with Ep throughout the growth period, whereas in roots, two plasma membrane intrinsic aquaporins, PIP2;4 and PIP2;5 were strongly correlated with Ep during reproductive growth. Other genes closely related to productivity, such as those involved in nutrient absorption and photosynthesis, exhibited different responses to meteorological factors at different growth stages. The stage-dependent shift in the microclimate response provides an important perspective on crop physiology in light of climate change.

Increasing canopy photosynthesis in rice can be achieved without a large increase in water use-A model based on free-air CO2 enrichment.

Achieving higher canopy photosynthesis rates is one of the keys to increasing future crop production; however, this typically requires additional water inputs because of increased water loss through the stomata. Lowland rice canopies presently consume a large amount of water, and any further increase in water usage may significantly impact local water resources. This situation is further complicated by changing the environmental conditions such as rising atmospheric CO2 concentration ([CO2 ]). Here, we modeled and compared evapotranspiration of fully developed rice canopies of a high-yielding rice cultivar (Oryza sativa L. cv. Takanari) with a common cultivar (cv. Koshihikari) under ambient and elevated [CO2 ] (A-CO2 and E-CO2 , respectively) via leaf ecophysiological parameters derived from a free-air CO2 enrichment (FACE) experiment. Takanari had 4%-5% higher evapotranspiration than Koshihikari under both A-CO2 and E-CO2 , and E-CO2 decreased evapotranspiration of both varieties by 4%-6%. Therefore, if Takanari was cultivated under future [CO2 ] conditions, the cost for water could be maintained at the same level as for cultivating Koshihikari at current [CO2 ] with an increase in canopy photosynthesis by 36%. Sensitivity analyses determined that stomatal conductance was a significant physiological factor responsible for the greater canopy photosynthesis in Takanari over Koshihikari. Takanari had 30%-40% higher stomatal conductance than Koshihikari; however, the presence of high aerodynamic resistance in the natural field and lower canopy temperature of Takanari than Koshihikari resulted in the small difference in evapotranspiration. Despite the small difference in evapotranspiration between varieties, the model simulations showed that Takanari clearly decreased canopy and air temperatures within the planetary boundary layer compared to Koshihikari. Our results indicate that lowland rice varieties characterized by high-stomatal conductance can play a key role in enhancing productivity and moderating heat-induced damage to grain quality in the coming decades, without significantly increasing crop water use.

Prediction of future methane emission from irrigated rice paddies in central Thailand under different water management practices.

There is concern about positive feedbacks between climate change and methane (CH4) emission from rice paddies. However, appropriate water management may mitigate the problem. We tested this hypothesis at six field sites in central Thailand, where the irrigated area is rapidly increasing. We used DNDC-Rice, a process-based biogeochemistry model adjusted based on rice growth data at each site to simulate CH4 emission from a rice-rice double cropping system from 2001 to 2060. Future climate change scenarios consisting of four representative concentration pathways (RCPs) and seven global climate models were generated by statistical downscaling. We then simulated CH4 emission in three water management practices: continuous flooding (CF), single aeration (SA), and multiple aeration (MA). The adjusted model reproduced the observed rice yield and CH4 emission well at each site. The simulated CH4 emissions in CF from 2051 to 2060 were 5.3 to 7.8%, 9.6 to 16.0%, 7.3 to 18.0%, and 13.6 to 19.0% higher than those from 2001 to 2010 in RCPs 2.6, 4.5, 6.0, and 8.5, respectively, at the six sites. Regionally, SA and MA mitigated CH4 emission by 21.9 to 22.9% and 53.5 to 55.2%, respectively, relative to CF among the four RCPs. These mitigation potentials by SA and MA were comparable to those from 2001 to 2010. Our results indicate that climate change in the next several decades will not attenuate the quantitative effect of water management practices on mitigating CH4 emission from irrigated rice paddies in central Thailand.

Development of a method for estimating total CH4 emission from rice paddies in Japan using the DNDC-Rice model.

Methane (CH4) is a greenhouse gas, and paddy fields are one of its main anthropogenic emission sources. To mitigate this emission based on effective management measures, CH4 emission from paddy fields must be quantified at a national scale. In Japan, country-specific emission factors have been applied since 2003 to estimate national CH4 emission from paddy fields. However, this method cannot account for the effects of weather conditions and temporal variability of nitrogen fertilizer and organic matter application rates; thus, the estimated emission is highly uncertain. To improve the accuracy of national-scale estimates, we calculated country-specific emission factors using the DeNitrification-DeComposition-Rice (DNDC-Rice) model. First, we calculated CH4 emission from 1981 to 2010 using 986 datasets that included soil properties, meteorological data, and field management data. Using the simulated site-specific emission, we calculated annual mean emission for each of Japan's seven administrative regions, two water management regimes (continuous flooding and conventional mid-season drainage), and three soil drainage rates (slow, moderate, and fast). The mean emission was positively correlated with organic carbon input to the field, and we developed linear regressions for the relationships among the regions, water management regimes, and drainage rates. The regression results were within the range of published observation values for site-specific relationships between CH4 emission and organic carbon input rates. This suggests that the regressions provide a simplified method for estimating CH4 emission from Japanese paddy fields, though some modifications can further improve the estimation accuracy.

Canopy-scale relationships between stomatal conductance and photosynthesis in irrigated rice.

Modeling stomatal behavior is critical in research on land-atmosphere interactions and climate change. The most common model uses an existing relationship between photosynthesis and stomatal conductance. However, its parameters have been determined using infrequent and leaf-scale gas-exchange measurements and may not be representative of the whole canopy in time and space. In this study, we used a top-down approach based on a double-source canopy model and eddy flux measurements throughout the growing season. Using this approach, we quantified the canopy-scale relationship between gross photosynthesis and stomatal conductance for 3 years and their relationships with leaf nitrogen content throughout each growing season above a paddy rice canopy in Japan. The canopy-averaged stomatal conductance (gsc ) increased with increasing gross photosynthesis per unit green leaf area (Ag ), as was the case with leaf-scale measurements, and 41-90% of its variation was explained by variations in Ag adjusted to account for the leaf-to-air vapor-pressure deficit and CO2 concentration using the Leuning model. The slope (m) in this model (gsc versus the adjusted Ag ) was almost constant within a 15-day period, but changed seasonally. The m values determined using an ensemble dataset for two mid-growing-season 15-day periods were 30.8 (SE = 0.5), 29.9 (SE = 0.7), and 29.9 (SE = 0.6) in 2004, 2005, and 2006, respectively; the overall mid-season value was 30.3 and did not greatly differ among the 3 years. However, m appeared to be higher during the early and late growing seasons. The ontogenic changes in leaf nitrogen content strongly affected Ag and thus gsc . In addition, we have discussed the agronomic impacts of the interactions between leaf nitrogen content and gsc . Despite limitations in the observations and modeling, our canopy-scale results emphasize the importance of continuous, season-long estimates of stomatal model parameters for crops using top-down approaches.

Influence of low air humidity and low root temperature on water uptake, growth and aquaporin expression in rice plants.

The effects of low air humidity and low root temperature (LRT) on water uptake, growth and aquaporin gene expression were investigated in rice plants. The daily transpiration of the plants grown at low humidity was 1.5- to 2-fold higher than that at high humidity. LRT at 13°C reduced transpiration, and the extent was larger at lower humidity. LRT also reduced total dry matter production and leaf area expansion, and the extent was again larger at lower humidity. These observations suggest that the suppression of plant growth by LRT is associated with water stress due to decreased water uptake ability of the root. On the other hand, the net assimilation rate was not affected by low humidity and LRT, and water use efficiency was larger for LRT. We found that low humidity induced coordinated up-regulation of many PIP and TIP aquaporin genes in both the leaves and the roots. Expression levels of two root-specific aquaporin genes, OsPIP2;4 and OsPIP2;5, were increased significantly after 6 and 13 d of LRT exposure. Taken together, we discuss the possibility that aquaporins are part of an integrated response of this crop to low air humidity and LRT.

ELPIS-JP: a dataset of local-scale daily climate change scenarios for Japan.

We developed a dataset of local-scale daily climate change scenarios for Japan (called ELPIS-JP) using the stochastic weather generators (WGs) LARS-WG and, in part, WXGEN. The ELPIS-JP dataset is based on the observed (or estimated) daily weather data for seven climatic variables (daily mean, maximum and minimum temperatures; precipitation; solar radiation; relative humidity; and wind speed) at 938 sites in Japan and climate projections from the multi-model ensemble of global climate models (GCMs) used in the coupled model intercomparison project (CMIP3) and multi-model ensemble of regional climate models form the Japanese downscaling project (called S-5-3). The capability of the WGs to reproduce the statistical features of the observed data for the period 1981-2000 is assessed using several statistical tests and quantile-quantile plots. Overall performance of the WGs was good. The ELPIS-JP dataset consists of two types of daily data: (i) the transient scenarios throughout the twenty-first century using projections from 10 CMIP3 GCMs under three emission scenarios (A1B, A2 and B1) and (ii) the time-slice scenarios for the period 2081-2100 using projections from three S-5-3 regional climate models. The ELPIS-JP dataset is designed to be used in conjunction with process-based impact models (e.g. crop models) for assessment, not only the impacts of mean climate change but also the impacts of changes in climate variability, wet/dry spells and extreme events, as well as the uncertainty of future impacts associated with climate models and emission scenarios. The ELPIS-JP offers an excellent platform for probabilistic assessment of climate change impacts and potential adaptation at a local scale in Japan.

Effect of low root temperature on hydraulic conductivity of rice plants and the possible role of aquaporins.

The role of root temperature T(R) in regulating the water-uptake capability of rice roots and the possible relationship with aquaporins were investigated. The root hydraulic conductivity Lp(r) decreased with decreasing T(R) in a measured temperature range between 10 degrees C and 35 degrees C. A single break point (T(RC) = 15 degrees C) was detected in the Arrhenius plot for steady-state Lp(r). The temperature dependency of Lp(r) represented by activation energy was low (28 kJ mol(-1)) above T(RC), but the value is slightly higher than that for the water viscosity. Addition of an aquaporin inhibitor, HgCl(2), into root medium reduced osmotic exudation by 97% at 25 degrees C, signifying that aquaporins play a major role in regulating water uptake. Below T(RC), Lp(r) declined precipitously with decreasing T(R) (E(a) = 204 kJ mol(-1)). When T(R) is higher than T(RC), the transient time for reaching the steady-state of Lp(r) after the immediate change in T(R) (from 25 degrees C) was estimated as 10 min, while it was prolonged up to 2-3 h when T(R) < T(RC). The Lp(r) was completely recovered to the initial levels when T(R) was returned back to 25 degrees C. Immunoblot analysis using specific antibodies for the major aquaporin members of PIPs and TIPs in rice roots revealed that there were no significant changes in the abundance of aquaporins during 5 h of low temperature treatment. Considering this result and the significant inhibition of water-uptake by the aquaporin inhibitor, we hypothesize that the decrease in Lp(r) when T(R) < T(RC) was regulated by the activity of aquaporins rather than their abundance.

Osmotic water permeability of plasma and vacuolar membranes in protoplasts II: theoretical basis.

Water permeability of the plasma membrane (PM) and the vacuolar membrane (VM) is important for intracellular and transcellular water movement in plants, because mature plant cells have large central vacuoles. We have developed a new method for measuring the osmotic water permeability of the PM and VM (P ( f1) and P ( f2), respectively) in individual plant cells. Here, the theoretical basis and procedure of the method are discussed. Protoplasts isolated from higher plant tissues are used to measure P ( f1) and P ( f2). Because of the semi-permeability (selective permeability) of cellular membranes, protoplasts swell or shrink under hypotonic or hypertonic conditions. A theoretical three-compartment model is presented for simulating time-dependent volume changes in the vacuolar and cytoplasmic spaces in a protoplast during osmotic excursions. The model describes the theoretical relationships between P ( f1), P ( f2) and the bulk osmotic water permeability of protoplasts (P ( f(bulk))). The procedure for measuring the osmotic water permeability is: (1) P ( f(bulk)) is calculated from the time when half of the total change in protoplast volume is completed, by assuming that the protoplast has a single barrier to water movement across it (two-compartment model); (2) P ( f2) of vacuoles isolated from protoplasts is obtained in the same manner; and (3) P ( f1) is determined from P ( f(bulk)) and P ( f2) according to the three-compartment model. The theoretical relationship between P ( fl ) (m s(-1)) and L ( Pl ) (hydraulic conductivity, l=1, 2) (m s(-1) Pa(-1)) is also discussed.

Osmotic water permeability of plasma and vacuolar membranes in protoplasts I: high osmotic water permeability in radish (Raphanus sativus) root cells as measured by a new method.

Intra- and transcellular water movements in plants are regulated by the water permeability of the plasma membrane (PM) and vacuolar membrane (VM) in plant cells. In the present study, we investigated the osmotic water permeability of both PM (P ( f1)) and VM (P ( f2)), as well as the bulk osmotic water permeability of a protoplast (P ( f(bulk))) isolated from radish (Raphanus sativus) roots. The values of P ( f(bulk)) and P ( f2) were determined from the swelling/shrinking rate of protoplasts and isolated vacuoles under hypo- or hypertonic conditions. In order to minimize the effect of unstirred layer, we monitored dropping or rising protoplasts (vacuoles) in sorbitol solutions as they swelled or shrunk. P ( f1) was calculated from P ( f(bulk)) and P ( f2) by using the 'three-compartment model', which describes the theoretical relationship between P ( f1), P ( f2) and P ( f(bulk)) (Kuwagata and Murai-Hatano in J Plant Res, 2007). The time-dependent changes in the volume of protoplasts and isolated vacuoles fitted well to the theoretical curves, and solute permeation of PM and VM was able to be neglected for measuring the osmotic water permeability. High osmotic water permeability of more than 500 mum s(-1), indicating high activity of aquaporins (water channels), was observed in both PM and VM in radish root cells. This method has the advantage that P ( f1) and P ( f2) can be measured accurately in individual higher plant cells.

Differences in aquaporin levels among cell types of radish and measurement of osmotic water permeability of individual protoplasts.

We investigated tissue- and cell-specific accumulation of radish aquaporin isoforms by immunocytochemical analysis. In taproots, the plasma membrane aquaporins (RsPIP1 and RsPIP2) were accumulated at high levels in the cambium, while the tonoplast aquaporin (RsTIP) was distributed in all tissues. The three isoforms were highly accumulated in the central cylinder of seedling roots and hypocotyls, and rich in the vascular tissue of the petiole of mature plants. The results suggest that RsPIP1 and RsPIP2 are abundant in the cells surrounding the sieve tube of the radish plant. The swelling rate of protoplasts in a hypotonic solution was determined individually by a newly established method to compare the osmotic water permeability of different cell types. All cells of the cortex and endodermis in seedlings showed a high water permeability of more than 300 microm s(-1). There was no marked difference in the values between the root endodermis and cortex protoplasts, although the RsPIP level was lower in the cortex than in the endodermis. This inconsistency suggests two possibilities: (1) a low level of aquaporin is enough for high water permeability and (2) the water channel activity of aquaporin in the tissues is regulated individually. The uneven distribution of aquaporins in tissues is discussed along with their physiological roles.