A physiological signal derived from sun-induced chlorophyll fluorescence quantifies crop physiological response to environmental stresses in the U.S. Corn Belt
Hyungsuk KimmKaiyu GuanChongya JiangGuofang MiaoGenghong WuAndrew E. SuykerElizabeth A. AinsworthCarl J. BernacchiChristopher M. MontesJoseph A. BerryXi YangChristian FrankenbergMin ChenPhilipp Köhler
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Abstract Sun-induced chlorophyll fluorescence (SIF) measurements have shown unique potential for quantifying plant physiological stress. However, recent investigations found canopy structure and radiation largely control SIF, and physiological relevance of SIF remains yet to be fully understood. This study aims to evaluate whether the SIF-derived physiological signal improves quantification of crop responses to environmental stresses, by analyzing data at three different spatial scales within the U.S. Corn Belt, i.e. experiment plot, field, and regional scales, where ground-based portable, stationary and space-borne hyperspectral sensing systems are used, respectively. We found that, when controlling for variations in incoming radiation and canopy structure, crop SIF signals can be decomposed into non-physiological (i.e. canopy structure and radiation, 60% ∼ 82%) and physiological information (i.e. physiological SIF yield, Φ F , 17% ∼ 31%), which confirms the contribution of physiological variation to SIF. We further evaluated whether Φ F indicated plant responses under high-temperature and high vapor pressure deficit (VPD) stresses. The plot-scale data showed that Φ F responded to the proxy for physiological stress (partial correlation coefficient, r p = 0.40, p < 0.001) while non-physiological signals of SIF did not respond ( p > 0.1). The field-scale Φ F data showed water deficit stress from the comparison between irrigated and rainfed fields, and Φ F was positively correlated with canopy-scale stomatal conductance, a reliable indicator of plant physiological condition (correlation coefficient r = 0.60 and 0.56 for an irrigated and rainfed sites, respectively). The regional-scale data showed Φ F was more strongly correlated spatially with air temperature and VPD ( r = 0.23 and 0.39) than SIF ( r = 0.11 and 0.34) for the U.S. Corn Belt. The lines of evidence suggested that Φ F reflects crop physiological responses to environmental stresses with greater sensitivity to stress factors than SIF, and the stress quantification capability of Φ F is spatially scalable. Utilizing Φ F for physiological investigations will contribute to improve our understanding of vegetation responses to high-temperature and high-VPD stresses.Keywords:
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Under natural conditions, the use of vapor pressure deficit between mesophyll cell surface and ambient air ( VPD s ) instead of atmospheric humidity factors in some stomatal models may markedly promote the applicability of stomatal models. It has been pointed out from theoretical analysis that the expression of the responses of stomatal conductance to VPD s is equivalent to the expression of responses of stomatal conductance to water loss of transpiration in stomatal models.
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We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.
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Stomatal behavior under global climate change is a central topic of plant ecophysiological research. Vapor pressure deficit (VPD) and phytohormones can affect stomata of leaves which can affect gas exchange characteristics of plant. The role of VPD in regulating leaf gas exchange of three tree species was investigated in Jinan, China. Experiments were performed in June, August, and October. Levels of three phytohormones (GA3, IAA, ABA) in the leaves of the three trees were determined by high-performance liquid chromatography in three seasons. The responses of stomatal conductance (gs) to an increasing VPD in the leaves of the three trees had peak curves under different seasons, which differed from the prevailing response pattern of gs to VPD in most literature. The peak curve could be fitted with a Log-Normal Model (R2 = 0.838-0.995). The VPD/RH values of the corresponding maximum of gs (gs-max-VPD/RH) could be calculated by fitted models. The gs-max-RH could be affected by environmental conditions, because of positive correlation between gs-max-RH and the mean monthly temperature in 2010 (R2 > 0.81). Two typical stomatal models (the Leuning model and the optimal stomatal behavior model) were used to estimate gs values, but they poorly predicted gs in the three trees. The concentration of ABA was positively correlated to sensitivity in response of stomatal conductance to VPD in the leaves of the tree species during the different seasons.
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Canopy stomatal conductance is a key physiological factor controlling transpiration from plant canopies, but it is extremely difficult to determine in field environments. The objective of this study was to develop a radiometric method for calculating canopy stomatal conductance for two plant species—wheat and soybean from direct measurements of bulk surface conductance to water vapor and the canopy aerodynamic conductance in controlled-environment chambers. The chamber provides constant net radiation, temperature, humidity, and ventilation rate to the plant canopy. In this method, stepwise changes in chamber CO2 alter canopy temperature, latent heat, and sensible heat fluxes simultaneously. Sensible heat and the radiometric canopy-to-air temperature difference are computed from direct measurements of net radiation, canopy transpiration, photosynthesis, radiometric temperature, and air temperature. The canopy aerodynamic conductance to the transfer of water vapor is then determined from a plot of sensible heat versus radiometric canopy-to-air temperature difference. Finally, canopy stomatal conductance is calculated from canopy surface and aerodynamic conductances. The canopy aerodynamic conductance was 5.5 mol m−2 s−1 in wheat and 2.5 mol m−2 s−1 in soybean canopies. At 400 umol mol−1 of CO2 and 86 kPa atmospheric pressure, canopy stomatal conductances were 2.1 mol m−2 s−1 for wheat and 1.1 mol m−2 s−1 for soybean, comparable to canopy stomatal conductances reported in field studies. This method measures canopy aerodynamic conductance in controlled-environment chambers where the log-wind profile approximation does not apply and provides an improved technique for measuring canopy-level responses of canopy stomatal conductance and the decoupling coefficient. The method was used to determine the response of canopy stomatal conductance to increased CO2 concentration and to determine the sensitivity of canopy transpiration to changes in canopy stomatal conductance. These responses are useful for improving the prediction of ecosystem-level water fluxes in response to climatic variables.
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Based on the diurnal change of stomatal conductance (Gs) of ginseng under forest, the qualities of the environmental factors to Gs were confirmed with correlation analysis. The interaction among factors was analysed with the method of multiple regression analysis. A conclusion was the main factors impacting on stomatal conductance (Gs) of ginseng under forest were photosynthetically active radiation (PAR), air temperature (Ta) and vapor pressure deficit (VPD). The relationship between stomatal conductance (Gs) of the leaf and environmental factors could be expressed as:Gs=4.712PAR×10-4+1.559Ta×10-2-0.002VPD-3.657,(R=0.971**,F=59.074).
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Yellow-cedar (Chamaecyparis nootkatensis (D. Don) Spach) gas exchange processes were measured in response to the following primary environmental variables: photosynthetically active radiation, vapour pressure deficit, root temperature, and soil moisture. Under nonlimiting edaphic conditions, maximum stomatal conductance and maximum CO 2 assimilation increased rapidly as photosynthetically active radiation increased from 0 to 200 μmol∙m −2 ∙s −1 and from 0 to 500 μmol∙m −2 ∙s −1 , respectively. Thereafter, greater photosynthetically active radiation levels only resulted in minor increases in stomatal conductance and CO 2 assimilation. Maximum stomatal conductance and maximum CO 2 assimilation declined in a concave manner as vapour pressure deficit increased from 1 to 5 kPa. Response surface model for stomatal conductance showed vapour pressure deficit was the primary influence after light had caused initial stomatal opening. Response surface modeling approach showed CO 2 assimilation increased as photosynthetically active radiation increased, but increased vapour pressure deficit resulted in a suppression of CO 2 assimilation. Response surface model showed internal CO 2 concentration declined sharply as photosynthetically active radiation increased from 0 to 500 μmol∙m −2 ∙s −1 , but it remained constant with increasing vapour pressure deficit. Decreasing root temperature resulted in a continual decline in CO 2 assimilation and stomatal conductance from 22 to 1 °C, while internal CO 2 concentration declined from 22 to 13 °C with little change between 13 and 1 °C. As predawn shoot water potential decreased from −0.5 to −2.0 MPa, CO 2 assimilation declined in a linear manner, while stomatal conductance and internal CO 2 concentration declined in a concave manner. Key words: Chamaecyparis nootkatensis, CO 2 assimilation, stomatal conductance, internal CO 2 concentration, photosynthetically active radiation, vapour pressure deficit, root temperature, predawn shoot water potential.
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