Senescence and net photosynthesis of the flag leaf and grain growth of wheat in response to fungicide

Green leaf area and net photosynthesis of the flag leaves of Avalon and Maris Huntsman winter wheat crops were studied in relation to grain growth following the application of the fungicide propiconazole at flag leaf emergence. Disease levels were low during grain-fill, and were not significantly affected by the fungicide. Propiconazole significantly maintained green leaf area and photosynthesis per unit area during the period of rapid flag leaf senescence, but it had no effect on stem weight, grain growth or yield.     

Seasonal variation in the response of Hagberg falling and liquefaciton numbers to propiconazole fungicide in wheat

The hypothesis was tested that seasonal differences in the response to propiconazole fungicide of hagberg failling and liquefaction numbers are related to grain drying rate, as indicated by potential evaportranspiration. The response of Hagberg failling and liquefaciton numbers to late-season applicaitions of propiconazole was calculated using data from a field experiments in each of the years 1981 to 1990 on winter wheat (Triticum aestivum) cv. Agalong grown with similar husbandry at one site. Cumulative potential evapotranspiration was calculated over seven different periods during grain ripening. No visible sprouting occurred in any experiment and Hagberg falling number was above 250 s in nine of the 10 years. Disease severity on the flag leaf was less than 7% in nine of the 10 years, and diseases were generally controlled well by propiconazole. There were no significant relationships between the response of Hagberg falling number and any of the potential evapotranspiration variables. The response of liquefaction number was exponentially related to each of three cumulative potential evapotranspiration variables which covered periods before the estimated date of 40% moisture content. Years with lower cumulative potential evapotranspiration gave the greater responses, but the relationships were considerably influenced by the two lowest potential evapotranspiration years. It was concluded that these data provide indirect evidence that the response of Hagberg falling number (when transformed to liquefaction number) to fungicide is mediated through grain drying rate, and furthermore that the results are compatible with the hypothesis that pre-maturity alpha-amylase activity is stimulated by slow grain drying.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

Rate and duration of grain filling in relation to flag leaf senescence and grain yield in spring wheat (Triticum aestivum) exposed to different concentrations of ozone

Effects of different concentrations of ozone on grain filling, flag leaf senescence and final grain yield in field‐grown spring wheat (Triticum aestivum L. cv. Dragon) were studied using open‐top chambers. The hypothesis tested was that an ozone‐induced reduction in grain yield is mainly related to an enhanced senescence and a shortening of the grain‐filling period. The plants were exposed to filtered air (F), non‐filtered air without extra ozone (NF) or non‐filtered air with 3 different levels of ozone added (NF1+, NF2+ and NF3+). The mean daytime (08.00–20.00 h) ozone concentrations during the exposure period (31 days) were 7, 20, 34, 48 and 62 nmol mol?1 in F, NF, NF1+, NF2+ and NF3+, respectively. The corresponding ozone doses, expressed as the accumulated exposure over a concentration threshold of 40 nmol mol?1 (AOT40), were 0, 12, 1 989, 5 881 and 10 375 nmol mol?1 h, respectively, and 884, 2 594, 4 557, 6 188 and 7 900 μmol m?2, respectively, expressed as the calculated cumulative flag leaf ozone flux (CFO30). The flag leaves senesced earlier and the grain‐filling duration was significantly shorter at higher ozone exposure compared to F (?5, ?13 and ?18% in NF1+, NF2+ and NF3+, respectively). The relative grain‐filling rate did not differ between the treatments. The 1000‐grain weights were 10, 28 and 37% lower, and the grain yields were 15, 29 and 46% lower than F in NF1+, NF2+ and NF3+, respectively. Ozone exposure had no significant effect on the number of grains per unit ground area or on straw yield, but significantly reduced the harvest index and increased the grain protein concentration in NF2+ and NF3+ compared to F. The grain yield was negatively correlated with the ozone dose, expressed either as AOT40 or as CFO3 with or without an ozone flux threshold. The 1000‐grain weight was positively correlated with the grain‐filling duration (R2=0.998), which in turn was positively correlated with the leaf area duration (R2=0.989).     

Wheat nitrogen metabolism during grain filling: comparative role of glumes and the flag leaf

The mobilization of nitrogen (N) compounds and the roles played by glumes and the flag leaf during grain filling were studied in bread wheat (Triticum aestivum L. cv. Florida) grown under field conditions. Glumes lost twice as much of their total N content as that lost by the flag leaf between the milk and early dough stages. In the flag leaf, glumes and grains, Glu, Asp, Ser and Ala accounted for 85% of all the reductions in the free amino acid pool. Principal component analysis of free amino acid pools separated grains from the glumes and the flag leaf, suggesting grain specific regulations in the use of free amino acids in protein synthesis. In all three organs, no decrease in Gln was detected, probably due to steady glutamine synthetase (GS; EC 6.3.1.2) activities per soluble protein in both the flag leaf and glumes. Compared with the flag leaf, glumes presented relatively smaller amounts of the chloroplast GS associated isoform. This we show is due to a lower relative number of mesophyll cells in glumes as supported by the different anatomy and the cellular pattern of the GS immunolocalization. We argue that cellular distribution plays a key role in supporting metabolism to enable the various functions undertaken by glume tissue.     

Contribution of the wheat (Triticum aestivum L.) flag leaf to grain yield in response to plant growth regulators

In West-Europe, intensive cereal management uses plant growth regulators (PGRs) especially for wheat. A green-house experiment compared the effects of two PGRs on flag leaf characteristics and yield of winter wheat. Chlormequat chloride + choline chloride (CCC) and chlormequat chloride + choline chloride + imazaquin (CCC+I) were applied to winter wheat at growth stage 5 (Feekes Large scale). CCC and CCC+I significantly increased flag leaf surface area at anthesis. Both treatments also enhanced chlorophyll content of the main stem flag leaf. The grain filling period was extended with PGR application by 2 days. CCC and CCC+I significantly increased net CO₂ assimilation rates during the flag leaf life. No effects of PGR spraying were observed on the pattern of ¹⁴C labelled assimilate distribution. Increased grain yield was due to the increase in average grain weight. The results indicate that PGR treatments increased flag leaf contribution to grain filling. The addition of imazaquin (I) to chlormequat (CCC) improved the effects of CCC.     

Genotype and fungicide effects on late-season root growth of winter wheat

The aim of this work was to investigate differences among genotypes in post-anthesis root growth and distribution of modern UK winter wheat cultivars, and the effects of fungicide applications. Post-anthesis root growth of up to six cultivars of winter wheat (Triticum aestivum L.), given either one or three applications of fungicide, was studied in field experiments during two seasons. Total root mass remained unchanged between GS63 (anthesis) and GS85, but root length increased significantly from 14.7 to 31.4 km m⁻² in one season. Overall, there was no evidence for a decline in either root mass or length during grain filling. Root mass as a proportion of total plant mass was about 0.05 at GS85. There were significant differences among cultivars in root length and mass especially below 30 cm. Malacca had the smallest root length and Savannah the largest, and Shamrock had a significantly larger root system below 40 cm in both seasons. Fungicide applied at ear emergence had no significant effect on root mass in either season but increased root length (P<0.01) in the more disease-prone season. By maintaining a green canopy for longer, fungicide applied at flag leaf emergence may have resulted in delayed senescence of the root system and contributed to the post-anthesis maintenance of root mass and length. Section Editor: R. W. Bell     

The nature of the effects of the aphids Sitobion avenae and Metopolophium dirhodum on the growth of wheat

Field-caged, post-anthesis populations of the aphids Sitobion avenae and Metopolophium dirhodum reduced grain weight of wheat by 14 and 7% respectively and induced changes in the senescence of the flag leaf. Spikelet number and grain number were unaffected as they are normally determined by pre-anthesis factors. Percentage grain protein was significantly reduced by both aphid species. The pattern of grain weight reduction within the ear was consistent with known limitations on the distribution of flag leaf assimilates among the grains. The relative effects of the two aphid species apparently resulted from the degree of nutrient drain imposed at particular feeding sites and the reduction in the leaf area duration of the flag leaf.     

Elevated CO2 concentration enhances the role of the ear to the flag leaf in determining grain yield of wheat

Net photosynthetic rate (P _N) of ear and flag leaf during grain filling stage and grain yield of plants with non-darkened or darkened flag leaf or darkened ear were examined in two different CO₂ concentrations: ambient (AC) and AC+200 μmol mol⁻¹ (EC). Ear showed much higher enhancement (56 %) of P _N than flag leaf (23 %) under EC. Moreover, CO₂ enrichment shortened the photosynthetic duration of flag leaf relative to ear. In this way the ratio of ear to flag leaf contribution to grain yield increased from 1.18 (AC) to 1.39 (EC).     

Ribulose-1,5-bisphosphate Carboxylase/Oxygenase Content and Degradation in Diploid,Tetraploid, and Hexaploid Wheat Species during Monocarpic Senescence

Wheat provides a unique genetic system in which variable sink size is available across the ploidies. We characterized monocarpic senescence in diploid, tetraploid, and hexaploid wheat species in flag leaf from anthesis up to full grain maturity at regular intervals. Triticum tauschii Acc. cv. EC-331751 showed the fastest rate of senescence among the species studied and the rate of loss per day was highest in terms of photosynthesis rate, ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) content, and flag leaf N content coupled with a higher rate of gain in grain N content. Cultivars Kundan and HD 4530 maintained high flag leaf N content throughout grain filling as compared to the diploids and showed a slower rate of senescence. RuBPCO content was higher in the diploids as compared to Kundan and HD 4530 at anthesis. However, the rate of decline in RuBPCO content per day was also higher in the diploids. This degradation in RuBPCO was mediated by high endoproteolytic activities in the diploids which in turn supported its higher rate of N mobilization as compared to the tetraploid and hexaploid wheat. Acidic endopeptidases were responsible for the mobilization of flag leaf nitrogen in wheat across ploidy levels (r=–0.582, p<0.01).     

Leaf-applied sodium chloride promotes cadmium accumulation in durum wheat grain

Cadmium (Cd) accumulation in durum wheat grain is a growing concern. Among the factors affecting Cd accumulation in plants, soil chloride (Cl) concentration plays a critical role. The effect of leaf NaCl application on grain Cd was studied in greenhouse-grown durum wheat (Triticum turgidum L. durum, cv. Balcali-2000) by immersing (10 s) intact flag leaves into Cd and/or NaCl-containing solutions for 14 times during heading and dough stages. Immersing flag leaves in solutions containing increasing amount of Cd resulted in substantial increases in grain Cd concentration. Adding NaCl alone or in combination with the Cd-containing immersion solution promoted accumulation of Cd in the grains, by up to 41%. In contrast, Zn concentrations of grains were not affected or even decreased by the NaCl treatments. This is likely due to the effect of Cl complexing Cd and reducing positive charge on the metal ion, an effect that is much smaller for Zn. Charge reduction or removal (CdCl₂⁰ species) would increase the diffusivity/lipophilicity of Cd and enhance its capability to penetrate the leaf epidermis and across membranes. Of even more significance to human health was the ability of Cl alone to penetrate leaf tissue and mobilize and enhance shoot Cd transfer to grains, yet reducing or not affecting Zn transfer.     

Photosynthesis of Ears and Flag Leaves of Wheat and Barley

Immediately after anthesis ears of spring wheat absorbed lessthan 0.5 mg CO₂, per hour in daylight and later evolved CO₂,in the light and in the dark. The rate of apparent photosynthesisof the combined flag-leaf lamina and sheath and peduncle (collectivelycalled flag leaf) of two spring wheat varieties, Atle and JufyI, was 3–4 mg per hour; the rates of the flag leaf andthe ear of two spring barleys, Plumage Archer and Proctor, wereeach about 1 mg per hour. The gas exchange of ears and flag leaves between ear emergenceand maturity accounted for most of the final grain dry weight.The CO₂, fixed by the wheat ear was equivalent to between 17and 30 per cent of the grain weight, but more than this waslost by respiration, so assimilation in the flag leaf was equivalentto 110–20 per cent of the final grain weight. In barley,photosynthesis in the flag leaf and the net CO₂ uptake by theear each provided about half of the carbohydrate in the grain. Barley ears photosynthesized more than wheat ears because oftheir greater surface, and flag leaves of wheat photosynthesizedmore than those of barley because they had more surface anda slightly greater rate of photosynthesis per dm².     

Factors of between- and within-plant distribution of Metopolophium dirhodum (Hom., Aphididae) on small grain cereals

Abstract: The relationship between abundance of rose‐grain aphid Metopolophium dirhodum (Walker) and leaf chlorophyll content of spring wheat, spring oats and winter barley was investigated. Within production stands of each crop 18–25 plots were established, located at places with different plant quality where aphids were counted on particular leaves of 50–100 tillers, and the leaf chlorophyll content and area were determined. In all stands aphid numbers × tiller^?1 increased exponentially with chlorophyll content (r² = 0.783–0.933). This parallel variation may be explained by increased nitrogen content and assimilate production of vigorous chlorophyll‐rich plants. The aphid numbers on leaves of particular order (within‐plant distribution) were also correlated with leaf chlorophyll content (r² = 0.373–0.827). However, in oats and barley the analysis of variance of residuals of log aphids × leaf^?1 versus leaf chlorophyll regression revealed a significant effect of leaf position (the order of the leaf from the top of the plant). The magnitude of residuals was positively related to leaf size and may be related to the intensity of phloem transport from the source leaves to sink organs.     

Effect of Indol-3-yl Acetic Acid on Photosynthetic Characteristics of Wheat Flag Leaf During Grain Filling

Area and fresh and dry masses of flag leaf show two phases of development during grain filling in Triticum aestivum. The initial large increase in leaf size is mainly due to water intake. Contents of chlorophylls and carotenoids, reducing sugars, and sucrose, Hill reaction rate, and photosynthetic activity increased during leaf growth, but a noticeable decline in these parameters followed throughout leaf senescence. The maximum accumulation of polysaccharides and proteins occurred at the beginning of grain set, but a continuous decline in their absolute values was manifested during grain filling. Grain priming with indol-3-yl acetic acid (IAA) at 25 mg kg^-1 stimulated the flag leaf growth, namely its fresh and dry masses and its area. Furthermore, the stimulatory effect was mainly due to the increase in the pigment formation that in turn increased the photosynthetic activity of flag leaf during grain filling. On the other hand, the highest dose of IAA (50 mg kg^-1) attenuated the growth and physiological activity of flag leaf through its inhibitory action on leaf fresh and dry masses, leaf area, pigments, saccharides and protein formation, as well as its effect on ¹⁴CO₂ assimilation.     

Wheat ear carbon assimilation and nitrogen remobilization contribute significantly to grain yield

The role of wheat ears as a source of nitrogen (N) and carbon (C) in the grain filling process has barely been studied. To resolve this question, five wheat genotypes were labeled with ¹⁵N‐enriched nutrient solution. N remobilization and absorption were estimated via the nitrogen isotope composition of total organic matter and Rubisco. Gas exchange analyses showed that ear photosynthesis contributed substantially to grain filling in spite of the great loss of C due to respiration. Of the total kernel N, 64.7% was derived from the N acquired between sowing and anthesis, while the remaining 35.3% was derived from the N acquired between anthesis and maturity. In addition, 1.87 times more N was remobilized to the developing kernel from the ear than from the flag leaf. The higher yielding genotypes showed an increased N remobilization to the kernel compared to the lower yielding genotypes. In addition, the higher yielding genotypes remobilized more N from the ears to the kernel than the lower yielding genotypes, while the lower yielding genotypes remobilized more N from the flag leaf to the kernel. Therefore, the ears contribute significantly toward fulfilling C and N demands during grain filling.     

Remobilization of cadmium in maturing shoots of near isogenic lines of durum wheat that differ in grain cadmium accumulation.

Cadmium accumulation in grain of durum wheat (Triticum turgidum L. var. durum) represents a concern to consumers. In an effort to understand the regulation of Cd accumulation in maturing grain, the remobilization of 109Cd applied to stem and flag leaves was examined in two near-isogenic lines that differ in grain Cd accumulation. Absorbed 109Cd was primarily retained in the labelling flap (50-54% and 65-80% for stem and flag leaves, respectively). Cadmium exported from the stem flap initially (3 d) accumulated in the stem in a declining gradient towards the head. Subsequent remobilization of Cd deposited in the stem was associated with Cd accumulation in the grain. Cadmium exported from the flag leaf flap was primarily directed to the grain. Little (<1%) Cd accumulated in the glumes or rachis, and transport of Cd to shoot tissues below the flag leaf node was low (<1%). On average, 9% and 17% of absorbed 109Cd accumulated in the grain 14 d after labelling the stem and flag leaf, respectively. Irrespective of labelling position, the low Cd-accumulating isoline averaged 1.5-2-fold lower Cd accumulation per grain and Cd concentration in the grain than the high Cd-accumulating isoline. Cadmium accumulation in the grain was inversely correlated with Cd retention in the stem (stem labelled) and labelling flap (flag leaf labelled) for both isolines. Cadmium translocation to the grain was not inhibited by Zn when both were applied simultaneously (50 pM 109Cd; 0.5 microM 65Zn) to the flag leaf. These results show that elevated remobilization of Cd from the leaves and stem to the maturing grain may be partially responsible for the high accumulation of Cd in durum wheat grain.     

不同供水条件下冬小麦叶与非叶绿色 器官光合日变化特征

为揭示小麦叶片与非叶绿色器官的光合活性在一日中的变化特性及其在器官间的差异性,探讨群体及不同器官光合日变化对不同供水条件的响应特征,在田间设置生育期不灌水(I0)、灌2水(I2,拔节水＋开花水)和灌4水(I4,起身水+孕穗水+开花水+灌浆水)3个处理,于灌浆期测定了群体光合与呼吸速率的日变化, 旗叶片、叶鞘、穗、穗下节间各器官光合速率、蒸腾速率、气孔导度及叶绿素荧光参数的日变化。结果表明,灌浆期小麦穗和穗下节间光合速率日变化呈单峰曲线,而旗叶叶片与旗叶鞘光合速率均呈双峰型,表现出不同程度的午休。随着灌水次数减少,各器官光合速率降低,叶片对严重水分亏缺的反应大于各非叶器官。器官光合速率的日变化与F_v/F_m变化相一致,而与气孔导度日变化有较大差异。各器官上午的累积光合量均高于下午,上午光合量占日总光合量的比例为51％-62％,随着灌水次数减少而增大。不同灌水处理群体光合速率、呼吸速率日变化均未出现午休现象。春季浇2水处理与春浇4水处理相比,灌浆期群体光合速率及日光合积累量没有显著差异。综合研究认为,小麦叶与非叶器官光合性能及其日变化特征有较大不同,非叶光合对水分亏缺的敏感性低于叶片,生育期浇2水可以获得与浇4水相似的群体日光合积累量。     

The effect of reducing growth in winter wheat on the population dynamics of the grain aphid Sitobion avenae (F.)

1 The effect of reducing the growth of winter wheat on population size and development of the grain aphid Sitobion avenae was studied. 2 Automatic, mobile, crop shading devices, which reduced radiation comparable to levels found on a cloudy day, whilst minimizing other climatic changes, were used to reduce crop growth. Shading was applied between either GS 31 and 39 (1st node detectable → flag leaf ligule just visible) or GS 39 and 55 (flag leaf just visible → 50% of inflorescence emerged). 3 Sitobion avenae populations were initiated at GS 55 and highest populations were subsequently observed on plants shaded between GS 31 and 39. 4 Individual aphids confined within clip cages were observed to measure development from birth to adult moult, adult lifespan and fecundity. Results indicated that adult lifespan was shortest on plants shaded between GS31 and 39, whilst there were no differences in development time or fecundity. 5 Possible explanations for these observations are discussed.     

Effects of elevated carbon dioxide and ozone on the growth and yield of spring wheat (Triticum aestivum L.)

Spring wheat cv. Minaret was grown under three carbon dioxide(CO₂) and two ozone (O₃) concentrations from seedling emergenceto maturity in open-top chambers. Under elevated CO₂ concentrations,the green leaf area index of the main shoot was increased, largelydue to an increase in green leaf area duration. Biomass increasedlinearly in response to increasing CO₂ (ambient, 550 and 680ppm). At anthesis, stem and ear dry weights and plant heightwere increased by up to 174%, 5% and 9 cm, respectively, andbiomass at maturity was 23% greater in the 680 ppm treatmentas compared to the ambient control. Grain numbers per spikeletand per ear were increased by 0.2 and 5 grains, respectively,and this, coupled with a higher number of ears bearing tillers,increased grain yield by up to 33%. Exposure to a 7 h daily mean O₃ concentration of 60 ppb inducedpremature leaf senescence during early vegetative growth (leaves1–7) under ambient CO₂ concentrations. Damage to the mainshoot and possible seedling mortality during the first 3 weeksof exposure altered canopy structure and increased the proportionof tillers 1 and 2 which survived to produce ears at maturitywas increased; as a result, grain yield was not significantlyaffected. In contrast to the older leaves, the flag leaf (leaf8) sustained no visible O₃ damage, and mean grain yield perear was not affected. Interactions between elevated CO₂ andO₃ influenced the severity of visible leaf damage (leaves 1–7),with elevated CO₂ apparently protecting against O₃-induced prematuresenescence during early vegetative growth. The data suggestthat the flag leaf of Minaret, a major source of assimilateduring grain fill, may be relatively insensitive to O₃ exposure.Possible mechanisms involved in damage and/or recovery are discussed. Key words: Carbon dioxide, ozone, spring wheat (cv. Minaret), leaf damage, tiller, yield