The potential role of sucrose transport gene expression in the photosynthetic and yield response of rice cultivars to future CO2 concentration

The metabolic basis for observed differences in the yield response of rice to projected carbon dioxide concentrations (CO₂) is unclear. In this study, three rice cultivars, differing in their yield response to elevated CO₂, were grown under ambient and elevated CO₂ conditions, using the free-air CO₂ enrichment technology. Flag leaves of rice were used to determine (1) if manipulative increases in sink strength decreased the soluble sucrose concentration for the ‘weak’ responders and (2), whether the genetic expression of sucrose transporters OsSUT1 and OsSUT2 was associated with an accumulation of soluble sugars and the maintenance of photosynthetic capacity. For the cultivars that showed a weak response to additional CO₂, photosynthetic capacity declined under elevated CO₂ and was associated with an accumulation of soluble sugars. For these cultivars, increasing sink relative to source strength did not increase photosynthesis and no change in OsSUT1 or OsSUT2 expression was observed. In contrast, the ‘strong’ response cultivar did not show an increase in soluble sugars or a decline in photosynthesis but demonstrated significant increases in OsSUT1 and OsSUT2 expression at elevated CO₂. Overall, these data suggest that the expression of the sucrose transport genes OsSUT1 and OsSUT2 may be associated with the maintenance of photosynthetic capacity of the flag leaf during grain fill; and, potentially, greater yield response of rice as atmospheric CO₂ increases.     

Study on the response of diploid,tetraploid and hexaploid species of wheat to the elevated CO2

Study was done to compare the response of Triticum aestivum (hexaploid), Triticum durum (tetraploid) and Triticum monococcum (diploid) wheat species to the elevated CO₂ using Free Air CO₂ Enrichment (FACE) facility. It was demonstrated that the modern cultivar of wheat Triticum aestivum (hexaploid) was largely sink limited. It appeared to have less photosynthesis per unit leaf area than Triticum monococcum (diploid wheat). While leaf size, grain weight and amylase activity increased with the ploidy level from diploid to hexaploid wheat forms, the photosynthetic rate was reduced significantly. These wheat species responded differentially to the elevated CO₂. The larger leaf area and greater seed weight and presence of 38 KDa protein band caused by elevated CO₂ had additive effect in improving the productivity of hexaploid wheat by changing the source sink ratio. Whereas, such a source sink balance was not induced by elevated CO₂ in diploid wheat. The increasing CO₂ may present opportunities to breeders and possibly allow them to select for cultivars responsive to the elevated CO₂ with better sink potential.Key words: Elevated CO₂, FACE technology, Photosynthesis, Seed weight, Source sink ratio, Triticum     

The effect of source or sink temperature on photosynthesis and 14C-partitioning in and export from a source leaf of Alstroemeria

The influence of source and sink temperature on leaf net C exchange rate (NCER), export, and partitioning in the C₃ monocotyledon Alstroemeria sp. cv. Jacqueline were examined. Leaf (i.e. source) temperature was varied between 12 and 35°C while source leaves were exposed to photorespiratory and nonphotorespiratory conditions during a 2-h steady-state ¹⁴CO₂ labelling period. Between 12 and 20°C, at ambient CO₂ and O₂, leaf NCER and export were similar with maximum rates of 9.71 ± 0.51 and 3.06 ± 0.36 μmol C m^-2 s^-1, respectively. Both NCER and export decreased above 20°C. At 35°C NCER was 30% of the rate at 20°C, but export was totally inhibited. Between 12 and 35°C, at the end of the 2-h feeding period, ¹⁴C was partitioned in the leaf as ethanol insolubles (3–10%), H₂O solubles (88–92%), and chloroform solubles (2–8%). However, above 25°C, less ¹⁴C was recovered in the starch fraction and more in the sugar fractions. At all temperatures, 86 to 94% of the labelled sugars was ¹⁴C-sucrose. In nonphotorespiratory conditions (i.e. 1 800 μI I^-1 CO₂ and 2% O₂). NCER and export were higher than the rates obtained at ambient CO₂ and O₂ at each temperature. Carbon dioxide enrichment sustained high NCER and export rates even at 35°C, Although CO₂ enrichment increased partitioning of ¹⁴C into starch, starch synthesis at 35°C was markedly reduced. Cooling the root-zone mass (i.e. a dominant sink) to 10°C, which simulated the commercial practice used to induce flowering, had no significant effect on source leaf NCER and export rates either during a 2-h steady-state labelling period or subsequently during a 21-h light-dark chase period. Furthermore, partitioning of ¹⁴C among leaf products at the end of the feed-chase period was not affected. Additional pulse and chase experiments using ¹¹CO₂ fed to source leaves of control and root-cooled plants showed that there was no difference in the direction of movement of ¹¹C-assimilates towards the flower or the root zone as a consequence of root cooling. Together, the data indicate that changing source strength, by manipulating photosynthesis and photorespiration, by varying the leaf temperature had a more profound effect on leaf export than manipulating sink activity.     

Long- and short-term effects of decreased sink demand on carbohydrate levels and photosynthesis in wheat leaves

Wheat (Triticum aestivum L.) ears were removed to investigate long-term regulation of photosynthesis by sink demand at ambient CO₂ and 22 °C. The CO₂ level was also increased to 660 μmol mol^?1 and temperature was lowered to 5 °C to examine short-term responses of photosynthesis to low sink demand. Sink removal inhibited photosynthesis and increased leaf levels of glucose, fructose and ribulose-1, 5-bisphosphate (RuBP), and the glucose-6-phosphate (G6P)/fructose-6-phosphate (F6P) and RuBP/3-phosphoglycerate (PGA) ratios under growth conditions, but had no effect on the activity and activation state of ribulose-1, 5-bisphosphate carboxylase oxygenase (Rubisco) either under growth or short-term conditions, suggesting an inhibition of photosynthesis by decreased in vivo catalysis of Rubisco. Photosynthesis increased similarly in eared and earless shoots after a rise in CO₂ concentration, and the ratio of triose-phosphates (glyceraldehyde 3-phosphate and dihydroxyacetone phosphate, TP) to PGA was similar or higher for removed than intact ears, suggesting that feedback inhibition of photosynthesis was not caused by a limitation of ATP synthesis in chloroplasts. Under short-term conditions (660 μmol mol^?1 CO₂, 5 °C), TP and RuBP levels and the TP/PGA and TP/RuBP ratios were increased by sink removal, indicating an additional limitation of photosynthesis by the rate of RuBP regeneration.     

Long- and short-term responses of leaf carbohydrate levels and photosynthesis to decreased sink demand in soybean

Axillary buds and the apical portion of shoots of soybean [Glycine max (L.) Merr. cultivar Turchina] plants were trimmed to investigate long-term regulation of photosynthesis by sink demand at ambient CO₂ and 22 °C. Also, in intact and trimmed shoots, the CO₂ level was increased to 660 μmol mol^?1 and temperature was lowered to 5°C to examine the superimposed short-term responses of photosynthesis to low sink demand. Under growth conditions, trimming the shoots increased leaf photosynthesis and the levels of sucrose, glucose-6-phosphate (G6P) and 3-phosphoglycerate (PGA), as well as the G6P/fructose-6-phosphate (F6P) and sucrose/starch ratios, while it decreased the level of starch and the triose-phosphate (glyceraldehyde 3-phosphate and dihydroxyacetone phosphate, TP)/PGA ratio. Photosynthesis enhancement was accompanied by increased chlorophyll contents and ribulose-l,5-bisphosphate carboxylase oxygenase (Rubisco) activity. Sink removal consistently increased photosynthesis measured under a variety of conditions (growth CO₂ or a short-term change to 660 μmol mol_-1 CO₂; growth temperature or a short-term change to 5 °C), except when low temperature was combined with ambient CO₂; the increase in photosynthesis was higher under short-term elevated CO₂ than at ambient CO₂. In contrast with its effect at ambient CO₂, shoot trimming increased the levels of TP and ribulose-1,5-bisphosphate (RuBP) and the TP/PGA ratio under high-CO₂ conditions.     

Global terrestrial carbon storage and uncertainties in its temperature sensitivity examined with a simple model

The future of the land carbon sink is a significant uncertainty in global change projections. Here, key controls on global terrestrial carbon storage are examined using a simple model of vegetation and soil. Equilibrium solutions are derived as a function of atmospheric CO₂ and global temperature, these environmental variables are then linked in an idealized global change trajectory, and the lag between the dynamic and equilibrium solutions is derived for different linear rates of increase in atmospheric CO₂. Terrestrial carbon storage is departing significantly from equilibrium because CO₂ and temperature are increasing on a similar timescale to ecosystem change, and the lag is found to be proportional to the rate of forcing. Thus peak sizes of the land carbon sink, and any future land carbon source, are proportional to the rate of increase of CO₂. A switch from a land carbon sink to a source occurs at a higher CO₂ and temperature under more rapid forcing. The effects of parameter uncertainty in temperature sensitivities of photosynthesis, plant respiration and soil respiration, and structural uncertainty through the effect of fixing the ratio of plant respiration to photosynthesis are explored. In each case, the CO₂ fertilization effect on photosynthesis is constrained to reproduce the 1990 atmospheric CO₂ concentration within a closed global model. New literature compilations are presented for the temperature sensitivities of plant and soil respiration. A lower limit, Q₁₀=1.29, for soil respiration significantly increases future land carbon storage. An upper limit, Q₁₀=3.63, for soil respiration underpredicts the increase in carbon storage since the Last Glacial Maximum. Fixing the ratio of plant respiration to photosynthesis (R/P) at 0.5 generates the largest and most persistent land carbon sink, followed by the weakest land carbon source.     

Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat

The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO₂ diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (R_d) was greater than the net photosynthesis rate (P_n) by a factor of 1.74 in the spike, whereas R_d was much smaller, only 22.1, 65.7 and 24.8% of P_n in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO₂ exchange. Results of ¹³C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high R_d values of ears as well as their low diffusive conductance suggest that CO₂ from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO₂ as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear-darkened plants, probably because of some degree of refixation (recycling) of respired CO₂ in the grains.     

Photosynthate supply and utilization in alfalfa : a developmental shift from a source to a sink limitation of photosynthesis

Long-term carbon dioxide enrichment, ¹⁴CO₂ feeding, and partial defoliation were employed as probes to investigate source/sink limitations of photosynthesis during the development of symbiotically grown alfalfa. In the mature crop, long-term CO₂ enrichment does not affect the rates of net photosynthesis, relative growth, ¹⁴C export to nonphotosynthetic organs, or the rates of ¹⁴C label incorporation into leaf sucrose, starch, or malate. The rate of glycolate labeling is, however, substantially reduced under these conditions. When the mature crop was partially defoliated, a considerable increase in net photosynthesis occurred in the remaining leaves. In the seedling crop, long-term CO₂ enrichment increased dry matter accumulation, primarily as a result of increases in leaf starch content. Although the higher rates of starch synthesis are not maintained, the growth enhancement of the enriched plants persisted throughout the experimental period. These results imply a source limitation of seedling photosynthesis and a sink limitation of photosynthesis in more mature plants. Consequently, both the supply and the utilization of photosynthate may limit seasonal photosynthesis in alfalfa.     

The Photosynthetic Role of Ears in C3 Cereals: Metabolism,Water Use Efficiency and Contribution to Grain Yield

This review concerns ear photosynthesis and its contribution to grain filling in C₃ cereals. Ear photosynthesis is quantitatively important to grain filling, particularly in dry areas where source (i.e., assimilate) limitations can occur. Compared to the flag leaf, ear photosynthesis exhibits higher water stress tolerance. Several factors could be involved in the ear's “drought tolerance.” First, although degree of C₄ metabolism in ear parts has been reported, current evidence supports only typical C₃ metabolism. Second, recycling of respired CO₂ (i.e., refixation) could have considerable impact on final crop yield by preventing loss of CO₂. Because refixation of CO₂ is independent of atmospheric conditions, water use efficiency (measured as total ear photosynthesis divided by transpiration) could be higher in the ear than in the flag leaf. Moreover, ear parts (in particular awns) show higher relative water content and better osmotic adjustment under water stress compared to the flag leaf. This capacity, in addition to persistence of photosynthetic components under drought (delayed senescence), might help the ear to continue to fix CO₂ late in the grain filling period.     

Carbon source–sink limitations differ between two species with contrasting growth strategies

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximize crop yield. We investigated how differences in growth rate arise from source–sink limitations, using a model system comparing a fast‐growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow‐growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub‐ambient and elevated CO₂ concentrations ([CO₂]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO₂] indicating that source strength was near maximal at current [CO₂]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO₂] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO₂], and lower non‐structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximizing growth of elite cereals under future elevated [CO₂].     

Effects of fully open-air [CO2] elevation on leaf ultrastructure, photosynthesis, and yield of two soybean cultivars

The objective of this study was to investigate the effect of elevated (550 ± 17 ??mol mol^?1) CO₂ concentration ([CO₂]) on leaf ultrastructure, leaf photosynthesis and seed yield of two soybean cultivars [Glycine max (L.) Merr. cv. Zhonghuang 13 and cv. Zhonghuang 35] at the Free-Air Carbon dioxide Enrichment (FACE) experimental facility in North China. Photosynthetic acclimation occurred in soybean plants exposed to long-term elevated [CO₂] and varied with cultivars and developmental stages. Photosynthetic acclimation occurred at the beginning bloom (R1) stage for both cultivars, but at the beginning seed (R5) stage only for Zhonghuang 13. No photosynthetic acclimation occurred at the beginning pod (R3) stage for either cultivar. Elevated [CO₂] increased the number and size of starch grains in chloroplasts of the two cultivars. Soybean leaf senescence was accelerated under elevated [CO₂], determined by unclear chloroplast membrane and blurred grana layer at the beginning bloom (R1) stage. The different photosynthesis response to elevated [CO₂] between cultivars at the beginning seed (R5) contributed to the yield difference under elevated [CO₂]. Elevated [CO₂] significantly increased the yield of Zhonghuang 35 by 26% with the increased pod number of 31%, but not for Zhonghuang 13 without changes of pod number. We conclude that the occurrence of photosynthetic acclimation at the beginning seed (R5) stage for Zhonghuang 13 restricted the development of extra C sink under elevated [CO₂], thereby limiting the response to elevated [CO₂] for the seed yield of this cultivar.     

Isolation and photosynthetic characteristics of mesophyll cells from developing leaves of soybean

Mesophyll cells were isolated from developing sink leaves (25 to 30 mm in length) of soybean, Glycine max (L.) Merr. cv. Will. Leaf strips were incubated for two h in a buffered medium containing osmoticum and 0.2% Pectolyase Y-23. Gently stirring the leaf strips released from 7 to 16% of the total leaf mesophyll cells. Other pectinase enzymes, effective in releasing cells from mature source leaves (70 to 75 mm in length), did not release cells from sink leaves. Sink and source cell preparations were about 50 and 95% intact, respectively, based on the exclusion of Evans Blue dye. Intact cells could not be separated from broken cells on Ficoll or metrizamide density gradients. Total protein and catalase, glyceraldehyde-3-phosphate dehydrogenase, glycolate oxidase, phosphoenolpyruvate carboxylase, and ribulose 1,5-bisphosphate carboxylase activities on a chlorophyll basis were about 50% lower in sink mesophyll cells than in sink leaf homogenates indicating that broken sink cells lost soluble protein to the medium. Source cells and source leaf homogenates had comparable amounts of protein and enzymatic activities. Enzymatic activities on a chlorophyll basis were similar in source and sink leaves with the exception of phosphenolpyruvate carboxylase, which was two times higher in sink leaves. This enzyme was also exceptionally low in source and sink cells being only 61 and 23%, respectively, of whole leaf activities. Sink cell rates of ¹⁴CO₂ fixation were only 7% of source cell rates and sink cells did not show light-dependent O₂ evolution. Both cell preparations had photosystem II activiteis which were comparable to rates of ¹⁴CO₂ fixation at satuarating light and CO₂ concentration. It was concluded that the reduced photosynthetic rate of sink cells was limited by the low photochemical capacity rather than a limitation of Calvin cycle enzymes.     

Lack of C4 photosynthetic metabolism in ears of C3 cereals

There is continuing controversy over whether a degree of C₄ photosynthetic metabolism exists in ears of C₃ cereals. In this context, CO₂ exchange and the initial products of photosynthesis were examined in flag leaf blades and various ear parts of two durum wheat (Triticum durum Desf.) and two six-rowed barley (Hordeum vulgare L.) cultivars. Three weeks after anthesis, the CO₂ compensation concentration at 210 mmol mol^?1 O₂ in durum wheat and barley ear parts was similar to or greater than that in flag leaves. The O₂ dependence of the CO₂ compensation concentration in durum wheat ear parts, as well as in the flag leaf blade, was linear, as expected for C₃ photosynthesis. In a complementary experiment, intact and attached ears and flag leaf blades of barley and durum wheat were radio-labelled with ¹⁴CO₂ during a 10s pulse, and the initial products of fixation were studied in various parts of the ears (awns, glumes, inner bracts and grains) and in the flag leaf blade. All tissues assimilated CO₂ mainly by the Calvin (C₃) cycle, with little fixation of ¹⁴CO₂ into the C₄ acids malate and aspartate (about 10% or less). These collective data support the conclusion that in the ear parts of these C₃ cereals C₄ photosynthetic metabolism is nil.     

Differential CO2 effect on primary carbon metabolism of flag leaves in durum wheat (Triticum durum Desf.)

C sink/source balance and N assimilation have been identified as target processes conditioning crop responsiveness to elevated CO₂. However, little is known about phenology‐driven modifications of C and N primary metabolism at elevated CO₂ in cereals such as wheat. Here, we examined the differential effect of elevated CO₂ at two development stages (onset of flowering, onset of grain filling) in durum wheat (Triticum durum, var. Sula) using physiological measurements (photosynthesis, isotopes), metabolomics, proteomics and ¹⁵N labelling. Our results show that growth at elevated CO₂ was accompanied by photosynthetic acclimation through a lower internal (mesophyll) conductance but no significant effect on Rubisco content, maximal carboxylation or electron transfer. Growth at elevated CO₂ altered photosynthate export and tended to accelerate leaf N remobilization, which was visible for several proteins and amino acids, as well as lysine degradation metabolism. However, grain biomass produced at elevated CO₂ was larger and less N rich, suggesting that nitrogen use efficiency rather than photosynthesis is an important target for improvement, even in good CO₂‐responsive cultivars.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Comparison of the Drought Stress Responses of Tolerant and Sensitive Wheat Cultivars During Grain Filling: Changes in Flag Leaf Photosynthetic Activity,ABA Levels,and Grain Yield

Water status parameters, flag leaf photosynthetic activity, abscisic acid (ABA) levels, grain yield, and storage protein contents were investigated in two drought-tolerant (Triticum aestivum L. cv. MV Emese and cv. Plainsman V) and two drought-sensitive (cvs. GK élet and Cappelle Desprez) wheat genotypes subjected to soil water deficit during grain filling to characterize physiological traits related to yield. The leaf water potential decreased earlier and at a higher rate in the sensitive than in the tolerant cultivars. The net CO₂ assimilation rate (P _N) in flag leaves during water deficit did not display a strict correlation with the drought sensitivity of the genotypes. The photosynthetic activity terminated earliest in the tolerant cv. Emese, and the senescence of flag leaves lasted 7 days longer in the sensitive Cappelle Desprez. Soil drought did not induce characteristic differences between sensitive and tolerant cultivars in chlorophyll a fluorescence parameters of flag leaves during post-anthesis. Changes in the effective quantum yield of PSII (Φ_PSII) and the photochemical quenching (qP) depended on the genotypes and not on the sensitivity of cultivars. In contrast, the levels of ABA in the kernels displayed typical fluctuations in the tolerant and in the sensitive cultivars. Tolerant genotypes exhibited an early maximum in the grain ABA content during drought and the sensitive cultivars maintained high ABA levels in the later stages of grain filling. In contrast with other genotypes, the grain number per ear did not decrease in Plainsman and the gliadin/glutenin ratio was higher than in the control in Emese during drought stress. A possible causal relationship between high ABA levels in the kernels during late stages of grain filling and a decreased grain yield was found in the sensitive cultivars during drought stress.     

Higher than expected CO2 fertilization inferred from leaf to global observations

Several lines of evidence point to an increase in the activity of the terrestrial biosphere over recent decades, impacting the global net land carbon sink (NLS) and its control on the growth of atmospheric carbon dioxide (c_a). Global terrestrial gross primary production (GPP)—the rate of carbon fixation by photosynthesis—is estimated to have risen by (31 ± 5)% since 1900, but the relative contributions of different putative drivers to this increase are not well known. Here we identify the rising atmospheric CO₂ concentration as the dominant driver. We reconcile leaf‐level and global atmospheric constraints on trends in modeled biospheric activity to reveal a global CO₂ fertilization effect on photosynthesis of 30% since 1900, or 47% for a doubling of c_a above the pre‐industrial level. Our historic value is nearly twice as high as current estimates (17 ± 4)% that do not use the full range of available constraints. Consequently, under a future low‐emission scenario, we project a land carbon sink (174 PgC, 2006–2099) that is 57 PgC larger than if a lower CO₂ fertilization effect comparable with current estimates is assumed. These findings suggest a larger beneficial role of the land carbon sink in modulating future excess anthropogenic CO₂ consistent with the target of the Paris Agreement to stay below 2°C warming, and underscore the importance of preserving terrestrial carbon sinks.     

Photosynthesis, photorespiration and productivity of wheat and soybean genotypes

The results of the numerous measurements obtained during the last 40 years on gas exchange rate, photosynthetic carbon metabolism by exposition in ¹⁴CO₂ and activities of primary carbon fixation enzyme, ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RuBPC/O), in various wheat and soybean genotypes grown over a wide area in the field and contrasting in photosynthetic traits and productivity are presented in this article. It was established that high productive wheat genotypes (7–9 t ha^?1) with the optimal architectonics possess higher rate of CO₂ assimilation during the leaf ontogenesis. Along with the high rate of photosynthesis, high values of photorespiration are characteristic for the high productive genotypes. Genotypes with moderate (4–5 t ha^?1) and low (3 t ha^?1) grain yield are characterized by relatively low rates of both CO₂ assimilation and photorespiration. A value of photorespiration constitutes 28–35% of photosynthetic rate in contrasting genotypes. The activities of RuBPC and RuBPO were changing in a similar way in the course of the flag leaf and ear elements development. High productive genotypes are also characterized by a higher rate of biosynthesis and total value of glycine–serine and a higher photosynthetic rate. Therefore, contrary to conception arisen during many years on the wastefulness of photorespiration, taking into account the versatile investigations on different aspects of photorespiration, it was proved that photorespiration is one of the evolutionarily developed vital metabolic processes in plants and the attempts to reduce this process with the purpose of increasing the crop productivity are inconsistent.     

Sucrose-phosphate synthase responds differently to source-sink relations and to photosynthetic rates: Lolium perenne L. growing at elevated pCO2 in the field

Lolium perenne, a main component species in managed grassland, is well adapted to defoliation, fertilization, and regrowth cycles; and hence, to changes in the assimilatory carbon source‐sink ratio. In the Swiss Free Air CO₂ Enrichment experiment the source‐sink ratio is (i) increased by elevated partial pressure of CO₂ (p_CO2), (ii) decreased by enhanced carbon use under high N fertilization, and (iii) gradually increased during regrowth after defoliation. Since sucrose synthesis plays a central role in leaf carbohydrate metabolism in this fructan‐accumulating species, we investigated how sucrose‐phosphate synthase (SPS) responds to the differing assimilatory carbon fluxes and source‐sink ratios in the field. Assimilatory carbon flux, as estimated by leaf gas exchange, strongly depended on p_CO2. Surprisingly, the SPS content per leaf area did not increase with p_CO2, but increased with N fertilization. During later regrowth, when a dense canopy had formed, the SPS content decreased; in particular, SPS was decreased at high N under elevated p_CO2. Further, the higher assimilatory carbon flux through SPS at elevated p_CO2 was accompanied by a higher activation state of SPS. The SPS content correlated very strongly with the ratio of free sucrose to free amino acid in leaves, which represents the carbon source‐sink ratio. Hence, SPS content in L. perenne appears to be regulated by the current, strongly nitrogen‐dependent, source‐sink relation.     

Variation in leaf respiration in relation to growth and photosynthesis of Lolium

Six Lolium genotypes with contrasting apparent photorespiration and CO_a compensation concentration, [C0₂]_c, were compared for net photosynthesis, dark respiration, leaf starch accumulation, rate of leaf expansion and shoot regrowth. Plants were grown in day/night temperatures of 15/10 and 25/20 ^oC. There were significant (P < 0–05) differences between the genotypes in all these parameters. At 25/20 ^oC apparent photorespiration was correlated with [CO₂]_c. Correlation coefficients, pooled from both temperature regimes, revealed that genotypes with high rates of net photosynthesis accumulated more leaf starch during light periods than genotypes with slow photosynthesis, but rates of leaf expansion and dry matter increase were only correlated, negatively, with dark respiration. Apparent photorespiration was negatively correlated with dark respiration. These findings suggest that attributes related to photorespiration such as [CO₂]_c and O₂ uptake from CO₂-free air in the light are unlikely to be useful selection criteria for growth of C₃ grasses, that net photosynthesis was probably not limiting growth and that maintenance respiration may have been an important determinant of genotypic differences in growth rate. Selections for slow and fast rates of dark respiration of mature leaves were therefore made at 8 and at 25 ^oC from within two different populations of L. perenne, S.23. This characteristic showed repeatabilities (broad-sense heritability) of from 0–41 to o-66. Six independent comparisons of simulated swards of the slow- and fast-respiring selections were made under periodic cutting regimes, either in a growth room at 25 ^oC or in a glasshouse from August to May. Growth of all plots of slow-respiring genotypes was consistently more rapid than that of the fast-respiring, at 25 ^oC in the growth room, and during autumn and spring in the glasshouse. There was no difference in winter growth. The implications of these results for the use of gas exchange measurements as selection criteria in plant breeding programmes are discussed.