Diurnal Oscillation in the Intercellular CO2 Concentration of Spring Wheat Under the Semiarid Conditions

Yields of wheat in semiarid and arid zones are limited by drought, and water condition is very important at each stage of development. Studies carried out at Loess Plateau in the northwestern part of China indicated that yield of spring wheat (Triticum aestivum L.) cv. Dingxi 81-392 was reduced by 41% when subjected to water stress. The effects of two water regimens on net photosynthetic rate (P _N), stomatal conductance (g _s), and intercellular CO₂ concentration (C _i) were investigated at the jointing, booting, anthesis, and grain filling stages. Low soil moisture in comparison to adequate one had invariably reduced P _N during the diurnal variations at the four growth stages. P _N and g _s in both soil moisture regimes was maximally reduced at midday. C _i and the stomatal limitation fluctuated remarkably during photosynthesis midday depression processes, especially at the grain filling stage. Hence atmospheric drought at midday was one of the direct causes inducing stomata closure and the g _s depression, but it was beneficial for maintaining stable intrinsic water use efficiency. Fluctuation in C _i implicated that non-stomatal limitation also plays an important role during the period of photosynthesis midday depression. Consequently stomatal and/or non-stomatal limitation are the possible cause of the midday photosynthesis decline.     

Diurnal Changes of Gas Exchange,Chlorophyll Fluorescence,and Stomatal Aperture of Hybrid Poplar Clones Subjected to Midday Light Stress

Diurnal changes in net photosynthetic rate (P _N), chlorophyll (Chl) fluorescence, and stomatal aperture of several hybrid poplar clones subjected to midday light stress were measured in July and August of 1996. Midday depression of P _N, photosystem 2 (PS2) efficiency, stomatal conductance (g _s), and stomatal aperture was observed in all clones, though at differing rates among them. Non-uniform stomatal closure occurred at noon and at other times, requiring a modification of intercellular CO₂ concentration (C ₁). A linear relationship was found between g _s and stomatal aperture. More than half of the photons absorbed by PS2 centre dissipated thermally when subjected to light stress at noon. There was a linear relationship between the rate of PS2 photochemical electron transport (PxPFD) and P _N. There was a consensus for two fluorescence indicators (1 – q_P/q_N and (F_m' – F)/F_m') in assessment of susceptibility of photoinhibition in the clones. According to P _N, Chl fluorescence, and stomatal aperture, we conclude that midday depression of photosynthesis can be attributed to both stomatal and non-stomatal limitations.     

Response of gas exchange and yield components of field-grown Triticum aestivum L. to elevated ozone in China

To assess photosynthesis and yield components’ response of field-grown wheat to increasing ozone (O₃) concentration (based on diurnal pattern of ambient O₃) in China, winter wheat (Triticum aestivum L.) cv. Jia 403 was planted in open top chambers and exposed to three different O₃ concentrations: O₃-free air (CF), ambient air (NF), and O₃-free air with additional O₃ (CF+O₃). Diurnal changes of gas exchange and net photosynthetic rate (P _N) in response to photosynthetic photon flux density (PPFD) of flag leaves were measured at the filling grain stage, and yield components were investigated at harvest. High O₃ concentration altered diurnal course of gas exchange [P _N, stomatal conductance (g _s), and intercellular CO₂ concentration (C _i)] and decreased significantly their values except for C _i. Apparent quantum yield (AQY), compensation irradiance (CI), and saturation irradiance (SI) were significantly decreased, suggesting photosynthetic capacity was also altered, characterized as reduced photon-saturated photosynthetic rate (P _Nmax). The limit of photosynthetic activity was probably dominated by non-stomatal factors in combination with stomatal closure. The significant reduction in yield was observed in CF+O₃ treatment as a result of a marked decrease in the ear length and the number of grains per ear, and a significant increase in the number of infertile florets per ear. Even though similar responses were also observed in plants exposed to ambient O₃ concentration, no statistical difference was observed at current ambient O₃ concentration in China.     

Canopy and leaf photosynthetic characteristics and water use efficiency of sweet sorghum under drought stress

The objective of this paper was to examine the relationship between Water Use Efficiency (WUE) at the canopy and leaf levels, to determine soil moisture conditions, which can optimize yield, and WUE of sweet sorghum (Sorghum bicolor (Linn.) Moench), thus providing some theoretical foundation for using marginal land effectively and developing production of sweet sorghum. Three levels of soil moisture conditions were established, and photosynthetic characteristics and yield were measured. The canopy apparent photo-synthetic rate (CAP) and leaf photosynthetic rate (P _N) were reduced gradually with increased drought stress, and the CAP was lower than the P _N under every soil moisture conditions. The P _N had a midday depression phenomenon, but the CAP did not exhibit this midday depression phenomenon under severe drought stress. The linear regression relationship of CAP and P _N was CAP = 1.5945 + 0.1496 P _N. The canopy apparent WUE_C and leaf WUE_L were the highest under moderate drought stress. The first was 5.3 and 5.8 times higher than the WUE_L in mid-July and late August, respectively. The stem fresh biomass yield was 77 tons/ha under moderate drought stress and WUE of aboveground biomass yield (WUE_B) was also the highest. Our results showed that moderate drought stress did not result in a significant reduction in biomass yield but increased WUE significantly.     

Photosynthetic Responses of Four Hosta Cultivars to Shade Treatments

The effects of shade on the gas exchange of four Hosta cultivars were determined under differing irradiances (5, 30, 50, and 100 % of full irradiance) within pots. Irradiance saturation ranged between 400–800 mol m^–2 s^–1 among the four cultivars, of which H. sieboldiana cv. Elegans and H. plantagenea cv. Aphrodite exerted the lowest saturation and compensation irradiances. The maximal photosynthetic rate (P _max) was significantly higher in shade than in full irradiance in Elegans and Aphrodite, and was at maximum in seedlings grown in 30 % of full irradiance. The best shade treatment for cvs. Antioch and Golden Edger was 50 % of full irradiance. The diurnal gas exchange patterns in four cultivars were greatly influenced by the irradiance. Single-peak patterns of net photosynthetic rate (P _N) and stomatal conductance (g _s) were observed under 5 and 30 % full irradiance for all the cultivars while Elegans and Aphrodite suffered from midday depression in 50 % of full irradiance. Under open sky, all four cultivars showed two-peak patters in their diurnal gas exchange, but the midday depression was less in Antioch and Golden Edger than in Elegans and Aphrodite. According to their photosynthetic responses to shade, the shade tolerance of the four cultivars was in the order: Elegans>Aphrodite>Antioch>Golden Edger.     

Photosynthetic responses of wheat (Triticum aestivum L.) to combined effects of drought and exogenous methyl jasmonate

Drought stress limits wheat growth and productivity. The response of wheat (Triticum aestivum L.) to different water supply conditions (well-watered and drought-stressed) and exogenous methyl jasmonate (MeJA; 0 and 0.25 μM) was studied. The application of MeJA enhanced wheat adaptability to drought stress by physiological and metabolic adjustments. Drought stress reduced net photosynthetic rate (P _N), stomatal conductance (g _s), transpiration rate (E), and water-use efficiency (WUE) in wheat. The application of exogenous MeJA decreased also g _s and E, but stimulated WUE. Meanwhile, MeJA mitigated the decline of P _N, g _s, and WUE induced by drought stress and midday depression by 6–183%. Both drought stress and exogenous MeJA induced stomatal closure, which improved water status and delayed plant senescence. MeJA enhanced the activities of superoxide dismutase, peroxidase, catalase, and reduced malondialdehyde content. P _N-PAR response curves showed that MeJA mitigated the decline of maximum P _N, apparent quantum yield, and saturation irradiance, and the increase of compensation irradiance. Drought stress and exogenous MeJA increased dark respiration rate and showed an additive effect. These results indicated that 0.25 μM MeJA enhanced the photosynthesis under drought stress mainly by improving the water status and antioxidant capacity of wheat.     

Diurnal Gas Exchange and Superior Resources Use Efficiency of Typical C4 Species in Hunshandak Sandland,China

Net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), leaf water potential (ψ_leaf), leaf nitrogen content, and photosynthetic nitrogen use efficiency (PNUE) were compared between a typical C₄ plant, Agriophyllum squarrosum and a C₃ plant, Leymus chinensis, in Hunshandak Sandland, China. The plant species showed different diurnal gas exchange patterns on June 12–14 when photosynthetic photon flux density (PPFD), air temperature (T _air), and water potential were moderate. P _N, E, and g _s of A. squarrosum showed distinct single peak while those of L. chinensis were depressed at noon and had two peaks in their diurnal courses. Gas exchange traits of both species showed midday depression under higher photosynthetic photon flux density (PPFD) and T _air when Ψ_leaf was significantly low down on August 6–8. However, those of A. squarrosum were depressed less seriously. Moreover, A. squarrosum had higher P _N, Ψ_leaf, water use efficiency (WUE), and PNUE than L. chinensis. Thus A. squarrosum was much more tolerant to heat and high irradiance and could utilise the resources on sand area more efficiently than L. chinensis. Hence species like A. squarrosum may be introduced and protected to reconstruct the degraded sand dunes because of their higher tolerance to stress and higher resource use efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

Discrimination Against 13CO2 in Leaves,Pod Walls,and Seeds of Water-stressed Chickpea

The rate of photosynthesis (P _N) in leaves and pods as well as carbon isotope content in leaves, pod walls, and seeds was measured in well-watered (WW) and water-stressed (WS) chickpea plants. The P _N, on an area basis, was negligible in pods compared to leaves and was reduced by water stress (by 26%) only in leaves. WS pod walls and seeds discriminated less against ¹³CO₂ than did the controls. This response was not observed for leaves as is usually the case. Pod walls and seeds discriminated less against ¹³CO₂ than did leaves in both WW and WS plants. Measurement of carbon isotope composition in pods may be a more sensitive tool for assessing the impact of water stress on long-term assimilation than is the instantaneous measurement of gas exchange rates.     

Influence of water stress and low irradiance on morphological and physiological characteristics of <Emphasis Type="Italic">Picea asperata</Emphasis> seedlings

The combined effects of water stress (WS) and low irradiance (LI) on growth, photosynthesis, osmotic adjustment, and lipid peroxidation were studied in dragon spruce (Picea asperata Mast.) seedlings grown under two water treatments (well watered, 100 % of field capacity, and water stressed, 30 % of field capacity) and two irradiances (HI, 100 % of full sunlight and LI, 15 % of full sunlight). WS reduced growth, chlorophyll (Chl) a and b contents, net photosynthetic rate, transpiration rate, stomatal conductance, and effective quantum yield of photosystem 2 (Y) but increased free proline and malondialdehyde contents. LI increased Chl contents and decreased Y, photochemical quenching (q_P), and non-photochemical quenching (q_N) under both water treatments. Hence the seedlings in the understory were more sensitive to drought than to LI.     

Effect of Cd on growth, photosynthetic gas exchange, and chlorophyll fluorescence of wild and Cd-sensitive mutant rice

Growth, photosynthetic gas exchange, and chlorophyll fluorescence characteristics were investigated in wild type (WT) and Cd-sensitive mutant rice (Oryza sativa L.) plants using 50 μM Cd treatment for 12 d followed by a 3-d recovery. Under Cd stress, net dry mass and pigment contents were significantly lower in the mutant plants than in the WT. The mutant had lower net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) than WT rice, however, it had higher intercellular CO₂ concentration (C _i), indicating that non-stomatal factors accounted for the inhibition of P _N. Maximal photochemical efficiency of photosystem 2 (F_v/F_m), effective quantum yield of PS2 (Φ_PS2), and photochemical quenching (q_P) decreased much in the mutant under Cd stress. Cd content in roots and leaves of the mutant was significantly higher than those in the WT. Hence Cd toxicity was associated with the marked increases in Cd contents of plant tissue. After the recovery for 3 d, the WT rice had higher capacity to recover from Cd injury than the mutant.     

Characterization of photosynthesis of Populus euphratica grown in the arid region

Net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), transpiration rate (E), water use efficiency (WUE), and stomatal limitation (L_s) of Populus euphratica grown at different groundwater depths in the arid region were measured. g _s of the trees with groundwater depth at 4.74 m (D₄) and 5.82 m (D₅) were lower and a little higher than that at 3.82 m (D₃), respectively. Compared with C _i and L_s of the D₃ trees, C _i decreased and L_s increased at 4.74 m, however, Ci increased and L_s decreased at D₅. Hence photosynthetic reduction of P. euphratica was attributed to either stomatal closure or non-stomatal factors depending on the groundwater depths in the plant locations. P _N of the D₃ trees was significantly higher than those at D₄ or D₅. The trees of D₄ and D₅ did not show a significant difference in their P _N, indicating that there are mechanisms of P. euphratica tolerance to mild and moderate drought stress.     

Photosynthetic Performance of Ginkgo biloba L. Grown Under High and Low Irradiance

Diurnal variation in net photosynthetic rate (P _N) of three-year-old plants of Ginkgo biloba was studied under open, O (receiving full sunlight), net-shade, NS (40 % of photosynthetically active radiation, PAR), or greenhouse, G (25 % PAR) conditions. In all three conditions, P _N was higher in morning along with stomatal conductance (g _s), and intercellular CO₂ concentration (C _i), while leaf temperature and vapour pressure deficit were low. The O-plants exhibited a typical decline in P _N during midday, which was not observed in NS-plants. This indicated a possible photoinhibition in O-plants as the ratio of variable to maximum fluorescence (F_v/F_m) and photosystem 2 (PS2) yield (_PS2) values were higher in the NS- and G-plants. On the contrary, stomatal density and index, chlorophyll a/b ratio, leaf thickness, and density of mesophyll cells were greater in O-plants. Further, higher P _N throughout the day along with higher relative growth rate under NS as compared to O and G suggested the better efficiency of Ginkgo plants under NS conditions. Therefore, this plant species could be grown at 40 % irradiance to meet the ever-increasing demand of leaf and also to increase its export potential.     

Photosynthetic and yield responses of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) to different water management strategies in subtropical China

An experiment was performed to study gas exchange and chlorophyll fluorescence responses of rice (Oryza sativa L.) to various regimes, such as flooding–midseason drying–flooding (FDF), flooding–midseason drying–saturation (FDS), and flooding–rain-fed (FR) regimes. Compared to FDF, FR resulted in an obvious decrease in net photosynthetic rate (P_N), due to the decrease in stomatal conductance and the increase in stomatal limitation. In contrast, FDS plants did not suffer stomatal limitation and had comparable P_N with FDF plants. For diurnal light-saturated electron transport rate and saturation irradiance, FDF performed the best, which was followed by FDS and FR successively. FR and FDS plants tended to suffer from midday depression. FDS reduced irrigated water by 17.2% compared to FDF for comparable yields. The results suggested that FDS can be an effective irrigation regime to save water.     

Effect of water stress on leaf photosynthesis,chlorophyll content,and growth of oriental lily

The photosynthetic characterization of the oriental lily (Lilium) cv. Sorbonne and its response to increasing water stress were analyzed based on the net photosynthetic rate (P _n), stomatal conductance (g _s), intercellular CO₂ concentration (C_i), transpiration rate (E), water use efficiency (WUE), and stomatal limitation (Ls) in the Horqin Sandy Land of western China. A photosynthesis-PAR response curve was constructed to obtain light-compensation and light-saturation points (LCP and LSP), the maximum photosynthetic rates (P _max) and dark respiration rates (R _D). The growth of lilies in the pots was analyzed after anthesis. Various intensities of water stress (5, 10, and 20 days without water, and an unstressed control) were applied. The results indicated that drought stress not only significantly decreased P _n, E, g _s, photosynthetic pigment content (Chl a, Chl b, and Chl (a + b)) and increased intrinsic water use efficiency (WUE), but also altered the diurnal pattern of gas exchange. Drought stress also affected the photosynthesis (P _n)-PAR response curve. Drought stress increased LCP and R _D and decreased LSP and P _max. There were both stomatal and nonstomatal limitations to photosynthesis. Stomatal limitation dominated in the morning, whereas nonstomatal limitation dominated in the afternoon. Thus, drought stress decreased potential photosynthetic capacity and affected the diurnal pattern of gas exchange and P _n-PAR response curves, thereby reducing plant quality (lower plant height, flower length, flower diameter, and leaf area). Water stress is likely the main limitation to primary photosynthetic process in the lily. Appropriate watering is recommended to improve photosynthetic efficiency and alleviate photodamage, which will increase the commercial value of the lily in the Horqin Sandy Land.     

Elevated CO<Subscript>2</Subscript> mitigates the adverse effects of drought on daytime net ecosystem CO<Subscript>2</Subscript> exchange and photosynthesis in a Florida scrub-oak ecosystem

Drought is a normal, recurrent feature of climate. In order to understand the potential effect of increasing atmospheric CO₂ concentration (C _a) on ecosystems, it is essential to determine the combined effects of drought and elevated C _a (EC) under field conditions. A severe drought occurred in Central Florida in 1998 when precipitation was 88 % less than the average between 1984 and 2002. We determined daytime net ecosystem CO₂ exchange (NEE) before, during, and after the drought in the Florida scrub-oak ecosystem exposed to doubled C _a in open-top chamber since May 1996. We measured diurnal leaf net photosynthetic rate (P _N) of Quercus myrtifolia Willd, the dominant species, during and after the drought. Drought caused a midday depression in NEE and P _N at ambient CO₂ concentration (AC) and EC. EC mitigated the midday depression in NEE by about 60 % compared to AC and the effect of EC on leaf P _N was similar to its effect on NEE. Growth in EC lowered the sensitivity of NEE to air vapor pressure deficit under drought. Thus EC would help the scrub-oak ecosystem to survive the consequences of the effects of rising atmospheric CO₂ on climate change, including increased frequency of drought, while simultaneously sequestering more anthropogenic carbon.     

Interactions of Elevated CO2 Concentration and Drought Stress on Photosynthesis in Eucalyptus Cladocalyx F. Muell

Response of net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (c _i), and photosynthetic efficiency (F_v/F_m) of photosystem 2 (PS2) was assessed in Eucalyptus cladocalyx grown for long duration at 800 (C₈₀₀) or 380 (C₃₈₀) μmol mol^-1 CO₂ concentration under sufficient water supply or under water stress. The well-watered plants at C₈₀₀ showed a 2.2 fold enhancement of P _N without any change in g _s. Under both C₈₀₀ and C₃₈₀, water stress decreased P _N and g _s significantly without any substantial reduction of c _i, suggesting that both stomatal and non-stomatal factors regulated P _N. However, the photosynthetic efficiency of PS2 was not altered. This revised version was published online in June 2006 with corrections to the Cover Date.     

Crassulacean acid metabolism in the epiphytic fern <Emphasis Type="Italic">Patycerium bifurcatum</Emphasis>

The epiphytic fern Platycerium bifurcatum grows in different habitats characterized by drought and high irradiance stress. The plant shows diurnal malate oscillations, indicative for CAM expression only in cover leaves, but not in sporotrophophyll. In P. bifurcatum cover leaves exposed to high irradiance and desiccation, the decrease in both CO₂ assimilation (P _N) and stomatal conductance (g _s) was accompanied with occurrence of diurnal malate oscillations. Exogenously applied abscisic acid (ABA) induced the decrease in P _N and g _s, but no clear change in malate oscillations. The measurements of the maximum quantum efficiency of photosystem 2 (F_v/F_m) under high irradiance showed distinct photoinhibition, but no clear changes in F_v/F_m due to desiccation and ABA-treatment were found.     

Diurnal variation of gas exchange, chlorophyll fluorescence, and xanthophyll cycle components of maize hybrids released in different years

Diurnal variation of gas exchange, chlorophyll (Chl) fluorescence, and xanthophyll cycle components of three maize (Zea mays L.) hybrids released in different years, i.e. Baimaya (1950s), Zhongdan2 (1970s), and Nongda108 (1990s), were compared. On cloudless days, the newer hybrids always had higher net photosynthetic rate (P _N), especially at noon, than the older ones. At noon, all the hybrids decreased their maximal yield of photosystem 2 (PS2) photochemistry (F_v/F_m) and actual quantum yield of PS2 (Φ_PS2), the newer ones always showing higher values. Generally, the newer hybrids displayed higher photochemical quenching of Chl (q_P) and lower non-photochemical quenching (NPQ). The interhybrid differences in P _N may be owing to their differential photochemical efficiency. A midday depression in P _N occurred in all hybrids, which might be caused by serious photoinhibition or by decreased stomatal conductance. However, midday depression in P _N was more obvious in the older hybrids, especially when leaves were senescent. The higher de-epoxidation state of the xanthophylls was noted in older hybrids, which was confirmed by their larger NPQ. The newer maize hybrids did not need a strong de-epoxidation state since they had a better photosynthetic quantum conversion rate and a lower NPQ.     

Modelling Diurnal Courses of Photosynthesis and Transpiration of Leaves on the Basis of Stomatal and Non-Stomatal Responses,Including Photoinhibition

A mathematical model for photoinhibition of leaf photosynthesis was developed by formalising the assumptions that (1) the rate of photoinhibition is proportional to irradiance; and (2) the rate of recovery, derived from the formulae for a pseudo first-order process, is proportional to the extent of inhibition. The photoinhibition model to calculate initial photo yield is integrated into a photosynthesis-stomatal conductance (g _s) model that combines net photosynthetic rate (P _N), transpiration rate (E), and g _s, and also the leaf energy balance. The model was run to simulate the diurnal courses of P _N, E, g _s, photochemical efficiency, i.e., ratio of intercellular CO₂ concentration and CO₂ concentration over leaf surface (C _i/C _s), and leaf temperature (T ₁) under different irradiances, air temperature, and humidity separately with fixed time courses of others. When midday depression occurred under high temperature, g _s decreased the most and E the least. The duration of midday depression of g _s was the longest and that in E the shortest. E increased with increasing vapour pressure deficit (VPD) initially, but when VPD exceeded a certain value, it decreased with increasing VPD; this was caused by a rapid decrease in g _s. When air temperature exceeded a certain value, an increase in solar irradiance raised T ₁ and the degree of midday depression. High solar radiation caused large decrease in initial photon efficiency (). P _N, E, and g _s showed reasonable decreases under conditions causing photoinhibition compared with non-photoinhibition condition under high irradiance. The T ₁ under photoinhibition was higher than that under non-photoinhibition conditions, which was evident under high solar irradiance around noon. The decrease in C _i/C _s at midday implies that stomatal closure is a factor causing midday depression of photosynthesis.     

Effect of drought on ear and flag leaf photosynthesis of two wheat cultivars differing in drought resistance

We investigated net photosynthetic rate (P_N) of ear and two uppermost (flag and penultimate) leaves of wheat cultivars Hongmangmai (drought resistant) and Haruhikari (drought sensitive) during post-anthesis under irrigated and non-irrigated field conditions. The P_Nof ear and flag leaf were significantly higher and less affected by drought in Hongmangmai than in Haruhikari. The rate of reduction in stomatal conductance (g_s) was similar for the two cultivars, but intercellular CO₂concentration (C_i) in the flag leaf of Hongmangmai was lower than that of Haruhikari in non-irrigated treatment. No differences were observed in leaf water potential (₁) and osmotic adjustment of the flag leaf of the cultivars. These results imply that differences in photosynthetic inhibition on the flag leaf at low leaf ₁between the cultivars were primarily due to non-stomatal effects. Hence the main physiological factor associated with yield stability of Hongmangmai under drought stress may be attributed to the capacity for chloroplast activity in the flag leaf, which apparently allows sustained P_Nof flag leaf during grain filling under drought stress. The higher P_Nof ear in Hongmangmai under drought could also be related to its drought resistance.This revised version was published online in March 2005 with corrections to the page numbers.