In vitro enzyme activities and products of 14CO2 assimilation in flag leaf and ear parts of wheat (Triticum aestivum L.)

Activities of key enzymes of Calvin cycle and C₄ metabolism, rate of ¹⁴CO₂ fixation in light and dark and the initial products of photosynthetic ¹⁴CO₂ fixation were determined in flag leaf and different ear parts of wheat viz. pericarp, awn and glumes. Compared to the activities of RuBP carboxylase and other Calvin cycle enzymes viz. NADP-glyceraldehyde-3-phosphate dehydrogenase, NAD-glyceraldehyde-3-phosphate dehydrogenase and ribulose-5-phosphate kinase, the levels of PEP carboxylase and other enzymes of C₄ metabolism viz. NADP-malate dehydrogenase, NAD-malate dehydrogenase, NADP-malic enzyme, NAD-malic enzyme, glutamate oxaloacetate transaminase genase, NADP-malic enzyme, NAD-malic enzyme, glutamate oxaloacetate transaminase and glutamate pyruvate transaminase, were generally greater in ear parts than in the flag leaf. In contrast to CO₂ fixation in light, the various ear parts incorporated CO₂ in darkness at much higher rates than flag leaf. In short term assimilation of ¹⁴CO₂ by illuminated ear parts, most of the ¹⁴C was in malate with less in 3-phosphoglyceric acid, whereas flag leaves incorporated most into 3-phosphoglyceric acid. It seems likely that ear parts have the capability of assimilating CO₂ by the C₄ pathway of photosynthesis and utilise PEP carboxylase for recapturing the respired CO₂.     

Monocarpic senescence in wheat: Influence of sterile glumes and ear

The senescence of sterile glumes, flag leaf and the other two leaves below the ear of wheat ( Triticum aestivum L. cv. Sonalika) was studied in relation to grain development and surgical manipulation. The senescence of sterile glumes was faster than that of the leaves in terms of chlorophyll and protein degradation. The flag leaf senesced later than the other two leaves below it. Removal of sterile glumes markedly reduced the harvest index (crop: straw ratio) and average dry weight per grain as compared to removal of the flag leaf. Maximum grain weight was achieved after the glumes had senesced completely. Removal of the ear delayed senescence of all the three leaves. It is concluded that sterile glumes are important suppliers of assimilate for grain filling and that nutrient drainage is the primary cause of the monocarpic senescence in wheat.     

Photosynthetic pigment composition and photosystem II photochemistry of wheat ears.

The characteristics of pigment composition and photosystem II (PSII) photochemistry in the flag leaf and ear parts of wheat (Triticum aestivum L.) grown in the field was compared. At the early stage of flowering, awns and the flag leaf showed the highest values in the maximal efficiency of PSII photochemistry (Fv/Fm), actual PSII efficiency (phi(PSII)), photochemical quenching (qP), and the efficiency of excitation capture by open PSII centres (Fv/F'm), followed by glumes, lemmas, and paleae, respectively except that no differences in F'v/F'm were observed among glumes, leamms, and paleae. With progressing grain filling, there was a change in the photosynthetic pigment stoichiometry. In the ear parts, neoxanthin and antheraxanthin decreased equally with chlorophyll levels. Lutein and zeaxanthin decreased less than chlorophyll levels while beta-carotene and violaxanthin decreased faster than chlorophyll levels. No big differences in pigment composition were observed among different ear parts. For the flag leaf, neoxanthin and beta-carotene decreased concomitantly with chlorophyll, whereas lutein and xanthophyll cycle pigment were less affected, leading to increases in lutein/chlorophyll and xanthophyll cycle pigment/chlorophyll ratios. Fv/Fm, phi(PSII), qP, and F'v/F'm decreased gradually in the flag leaf and ear parts but to different extents. The largest changes were observed in awns, followed by the lemmas of floret 2, the lemmas of floret 1, glumes, and the flag leaf, respectively. The results suggest that during grain filling, a down-regulation of PSII associated with an increase of the de-epoxidation state of the xanthophyll cycle carotenoids occurred in the flag leaf but not in the ear parts.     

Differences in water relations and physiological characteristics in leaves of wheat associated with leaf position on the plant

The seasonal change in leaf water potential and its components, stomatal resistance, specific leaf weight, photosynthesis rate, the activities of ribulose-1,5-bisphosphate carboxylase and nitrate reductase, and soluble proteins were measured in flag leaves (ninth from base in position), seventh and fifth leaves of wheat Triticum aestivum L. cv Kalyansona. Flag leaves had a lower water and solute potential and lower or equal turgor pressure than seventh and fifth leaves. These differences were found to be independent of environment. The rate of photosynthesis and nitrate reductase activity were always lower in fifth and seventh leaves than in flag leaf. The photosynthetic efficiency in flag leaves appeared to be associated with lower stomatal resistance and higher specific leaf weight. The relations between leaf water potential and relative water content showed a change with leaf position. This change possibly allows flag leaf to maintain its functional efficiency despite its lower water potential.     

普通野生稻和武育粳8号剑叶光合功能衰退的比较

以武育粳 8号 (Wuyujing8)和普通野生稻 (Oryza.sativaL .f.spontaneaRoschev .)为材料 ,比较了它们剑叶光合功能衰退的进程。结果表明 ,在剑叶一生中 ,武育粳 8号剑叶的叶绿素含量和光合速率都高于普通野生稻 ,光合速率的最高值前者为 2 3 .5 μmolCO2 m 2 ·S 1,后者为 1 5 .5 μmolCO2 m 2 ·S 1。前者剑叶光合速率的高值持续期比后者长近 2 0d ,其叶源量是后者的 2 .8倍。武育粳 8号剑叶叶绿素含量最大值的SPAD值为 5 0 .1 ,而普通野生稻最大值的SPAD值为 42 .7。在可溶性蛋白质的基础上 ,核酮糖 1 ,5 二磷酸羧化酶比活性二者之间没有太大区别。内肽酶活力上升落后于光合功能的衰退。在光合功能衰退的早期 ,内肽酶的活力仅有小幅度提高 ,但在叶片衰老的末期升高幅度则较大。这说明内肽酶主要在光合功能衰退的后期即不可逆衰退期才发挥较大作用。     

Grain filling in relation to monocarpic senescence of wheat in varying source-sink ratios

Flag leaf removal at any stage of grain growth hastened senescence (reduction in chlorophyll content) of the sterile glumes whereas a removal of the latter did not alter senescence of the former. Kernel mass, grain mass per ear, harvest index and sink activity reduced more conspicuously by the removal of glumes than by the flag leaf removal. Removal of grains hastened senescence of the glumes only, although protein content increased at a later grain removal in the majority of source organs. Moisture contents of the developing grains were mostly reduced by the removal of either the flag leaf or the glumes, though flag leaf removal at anthesis affected it the most. Protein levels of the developing grains reduced at harvest only when the glumes were removed at anthesis but sugar levels in later phases always decreased irrespective of the time of removal and nature of the source organs.     

Effect of changed source/sink relations on proteolytic activities and on nitrogen mobilization in field-grown wheat (Triticum aestivum L.)

Nitrogen mobilization and the pattern of proteolytic enzymeswere investigated in leaves and glumes of field-grown winterwheat (Triticum aestivum L.) during maturation. Source/sinkrelations were changed by removal of the ear, the flag leafor the lower leaves shortly after anthesis. Removal of the earwas most effective, resulting in delayed senescence of the flagleaf with the chlorophyll, aminopeptidase and carboxypeptidaseactivities remaining high in contrast to the control, whereasneutral endopeptidase activity increased more slowly. No majorchanges were observed in the second leaf from the top in plantswith either ears or flag leaves removed. Nitrogen mobilizationand proteolytic activities in glumes and the remaining leaveswere influenced only slightly by leaf removal. In earless plants,nitrogen was transported from the second leaf into the leafsheath and stem, but in the flag leaf the total reduced nitrogenremained high and free amino groups increased. The increase in endopeptidase activity was influenced by thesource/sink relations. However, the accumulation of amino groupsand the increasing endopeptidase activity in the flag leaf ofearless plants suggest that the nitrogen sink capacity did notgreatly control protein degradation; it remains to be seen whetherphytohormones, accumulated amino acids or other factors delayedthe increase in endopeptidase activity. (Received September 3, 1979; )     

Variation of RuBP carboxylase activity among anther-derived plants of Solanum phureja

The effect of genotype and ploidy on RuBP carboxylase (EC 4.1.1.39) activity, chlorophyll content, leaf area, chloroplast ultrastructure and not photosynthesis among monoploid. diploid and tetraploid anther-derived plants of Solanum phureja Juz, and Buk. was studied. Within the monoploid group, RuBP carboxylase activity and concentration displayed a significant genotypic effect. For the diploids, variation among genotypes was significant for total protein content and maximum specific activity of RuBP carboxylase, and among the tetraploids for net photosynthesis and specific leaf weight. Ploidy effect was evident regarding net photosynthesis, leaf area and chlorophyll content. The different ploidy groups among the anther-derived plants surpassed the anther donor plant for all characteristics except maximum activity of RuBP carboxylase and net photosynthesis. For the latter only the tetra-ploid group was superior to the anther source plant. However, a monoploid genotype with an increase of 9% in maximum activity of RuBP carboxylase over the anther-donor plant was identified. Segregation of trails rind differential gene expression together with possible mutations during androgenesis are discussed as sources of variation.     

水稻叶片的衰老与超氧物歧化酶活性及脂质过氧化作用的关系

研究了从抽穗开花到籽粒成熟过程中,水稻植株顶部三片叶子的超氧物歧化酶(SOD),脂质过氧化产物丙二醛(MDA)含量及二磷酸核酮糖羧化酶活性的变化。实验结果表明:叶片的衰老伴随着 SOD 活性、RuBP 羧化酶活性及叶绿素含量的降低、丙二醛含量显著增高。分离了三个 SOD 的同工酶,证明为 Cu—Zn SOD。观察了 SOD 同工酶在叶片老化及酶液存放不同时间中的变化。讨论了叶片衰老过程中氧自由基对酶及质膜的损伤影响。     

Photosynthesis and Transpiration in Leaves and Ears of Wheat and Barley Varieties

Carbon exchange rate (CER) and transpiration were measured inflag leaves, whole ears, glumes (referring to the total areaof glumes and lemmas) and awns, in six hexaploid spring wheats(Triticum aestivum L.), three cultivated tetraploid spring wheats(T. turgidum L.), four wild tetraploid wheats (T. dicoccoides),eight six-rowed barleys (Hordeum vulgare L.) and five two-rowedbarleys (H. vulgare L.). Differences between varieties and between species in total earCER and transpiration were associated largely with differencesin ear surface area rather than with rates per unit area. Ratesof CER and transpiration per unit area of ears were 40–80%of those of flag leaves, depending on the species. However, since ear surface area was greater than flag leaf areaby a factor of 1.1, 3.9, 5.5 and 4.4, in hexaploid wheat, tetraploidwheat, six-rowed barley, and two-rowed barley, respectively,total ear CER reached up to 90% of that of the flag leaf. The contribution of awns to total ear CER depended largely ontotal awn surface area per ear, rather than on CER per unitawn area. Awns contributed about 40–80% of total spikeCER, depending on the species, but only 10–20% of spiketranspiration. The disproportionately small contribution ofawns to ear transpiration was caused by the very low rate oftranspiration per unit area of awns. Thus, while transpirationratio (CER/transpiration) was about the same in flag leavesand glumes, it was higher by several orders of magnitude inthe awns. A large amount of awns in the ear is therefore a drought adaptiveattribute in these cereals, for which tetraploid wheat exceededhexaploid wheat and six-rowed barley exceeded two-rowed barley. Key words: Carbon exchange rate, Transpiration, Barley, Wheat     

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.     

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 (ψ₁) 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 ψ₁between 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.     

Carbon isotope ratios in ear parts of triticale : influence of grain filling

Four triticale (×Triticosecale Wittmack) genotypes were grown under rainfed conditions with limited irrigation support in Lleida in northeast Spain. For each variety, samples consisting of 10 tillers with half-sterilized spikes were taken three times from anthesis to maturity. Carbon isotope ratios (δ¹³C) were then determined in water extracts from ear bracts (glumes, paleas, and lemmas), awns and flag leaves, and in powdered kernels. For the half-sterilized spikes, carbon isotope analysis was carried out separately in bracts and awns from fertile and nonfertile spikelets. The δ¹³C in the water-soluble fraction of awns, glumes, and glumells from fruitless spikelets was significantly higher than that from fertile spikelets sampled at mid-grain filling. Differences in δ¹³C among sterile and fertile spikelets were not significant in samples taken a few days after anthesis or at maturity. These results are in accordance with some degree of refixation by awns and ear bracts of the CO₂ respired by grains during grain filling. There was progressively higher δ¹³C from flag leaf blades to awns, glumes, and glumells. This variation in δ¹³C along plant parts may be caused by differences in the ratio of assimilation rate to CO₂-diffusive conductance. Values of δ¹³C of mature kernels were between the values at anthesis and mid-grain filling for the water-soluble fraction of flag leaves and inner bracts and were fairly similar to those of glumes and awns.     

Photosynthetic characteristics and enzymatic antioxidant capacity of flag leaf and the grain yield in two cultivars of Triticum aestivum (L.) exposed to warmer growth conditions

Using late sowing practice, the reproductive growth (anthesis and kernel filling) phase of two wheat cultivars, HD 1553 and HD 2307 was exposed to warmer growth conditions, and the effect on grain yield was examined. The grain weight declined in late-sown plants of both cultivars, but the number of grains per spike decreased drastically in HD 1553 plants. In this cultivar exposure to warmer temperature during reproductive phase led to 67% fewer grains per spike. Examination of photosynthetic and enzymatic antioxidant capacity in flag leaves of late-sown plants revealed a marked reduction in chlorophyll and carotenoid pigmentation in addition to a decline in the activity of H2O2 metabolising enzymes in HD 1553 cultivar. The photo-oxidative pigment loss due to warmer growth conditions in late-shown HD 1553 plants could lead to a reduction in flag leaf photosynthesis and contribute to poor grain yield.     

Dorsoventral asymmetry of photosynthesis and photoinhibition in flag leaves of two rice cultivars that differ in nitrogen response and leaf angle

Rice is believed to show photosynthetic symmetry between adaxial and abaxial leaf sides. To verify this, we re‐examined dorsoventral asymmetry in photosynthesis, chlorophyll fluorescence and anatomical traits in flag leaves of two Oryza sativa cultivars that differ in nitrogen (N) response and in leaf angle: ‘Akenohoshi’, a cultivar that can adapt to low‐N (LN), with low leaf angle (more erect leaves), and ‘Shirobeniya’, a cultivar that is unable to adapt to LN, with higher leaf angle. Plants were grown under standard‐N (SN) and LN conditions. LN leaves of both cultivars became more erect than SN, but LN Akenohoshi still had more erect ones than Shirobeniya. Contrary to results of previous studies, leaves of both cultivars showed an asymmetry in photosynthetic rate between adaxial and abaxial sides (higher on the adaxial side) under SN. SN leaves of both cultivars showed lower susceptibility to photoinhibition on the adaxial side than on the abaxial side. However, leaves of Akenohoshi showed less asymmetry in these traits under LN than under SN, whereas leaves of Shirobeniya had similar degrees of asymmetry in these traits under both SN and LN. Both cultivars also showed dorsoventral asymmetry in anatomical traits of mesophyll tissue regardless of N level, but the degree of asymmetry was lower in LN Akenohoshi. These data reveal that rice leaves exhibit dorsoventral asymmetry in photosynthetic and anatomical features, and that the degree of asymmetry varies with cultivar and N level. It is suggested that lower leaf angles (particularly in Akenohoshi) in the presence of LN represent a light acclimation to prevent photoinhibition.     

Photosynthesis and nitrogen relationships in leaves of C3 plants

Summary The photosynthetic capacity of leaves is related to the nitrogen content primarily bacause the proteins of the Calvin cycle and thylakoids represent the majority of leaf nitrogen. To a first approximation, thylakoid nitrogen is proportional to the chlorophyll content (50 mol thylakoid N mol^-1 Chl). Within species there are strong linear relationships between nitrogen and both RuBP carboxylase and chlorophyll. With increasing nitrogen per unit leaf area, the proportion of total leaf nitrogen in the thylakoids remains the same while the proportion in soluble protein increases. In many species, growth under lower irradiance greatly increases the partitioning of nitrogen into chlorophyll and the thylakoids, while the electron transport capacity per unit of chlorophyll declines. If growth irradiance influences the relationship between photosynthetic capacity and nitrogen content, predicting nitrogen distribution between leaves in a canopy becomes more complicated. When both photosynthetic capacity and leaf nitrogen content are expressed on the basis of leaf area, considerable variation in the photosynthetic capacity for a given leaf nitrogen content is found between species. The variation reflects different strategies of nitrogen partitioning, the electron transport capacity per unit of chlorophyll and the specific activity of RuBP carboxylase. Survival in certain environments clearly does not require maximising photosynthetic capacity for a given leaf nitrogen content. Species that flourish in the shade partition relatively more nitrogen into the thylakoids, although this is associated with lower photosynthetic capacity per unit of nitrogen.     

Photosynthetic capacity and dry mass partitioning in dwarf and semi-dwarf wheat (Triticum aestivum L.).

Efficient use of space and high yields are critical for long-term food production aboard the International Space Station. The selection of a full dwarf wheat (less than 30 cm tall) with high photosynthetic and yield potential is a necessary prerequisite for growing wheat in the controlled, volume-limited environments available aboard long-term spaceflight missions. This study evaluated the photosynthetic capacity and carbon partitioning of a full-dwarf wheat cultivar, Super Dwarf, which is routinely used in spaceflight studies aboard U.S. space shuttle and NASA/Mir missions and made comparisons with other dwarf and semi-dwarf wheat cultivars utilized in other ground-based studies in plant space biology. Photosynthetic capacity of the flag leaf in two dwarf (Super Dwarf, BB-19), and three semi-dwarf (Veery-10, Yecora Rojo, IBWSN 199) wheat cultivars (Triticum aestivum L.) was assessed by measuring: net maximum photosynthetic rate, RuBP carboxylation efficiency, chlorophyll concentration and flag leaf area. Dry mass partitioning of carbohydrates to the leaves, sheaths, stems and ear was also assessed. Plants were grown under controlled environmental conditions in three replicate studies: slightly enriched CO2 (370 micromoles mol-1), high photosynthetic photon flux (1000 micromoles m-2 s-1; 58 mol m-2 d-1) for a 16 h photoperiod, 22/15 degrees C day/night temperatures, ample nutrients and water provided by one-half strength Hoagland's nutrient solution (Hoagland and Arnon, 1950). Photosynthetic capacity of the flag leaf was determined at anthesis using net CO2 exchange rate versus internal CO2 concentration curves measured under saturating light (2000 micromoles m-2 s-1) and CO2 (1000 micromoles mol-1). Dwarf wheat cultivars had greater photosynthetic capacities than the taller semi-dwarfs, they averaged 20% higher maximum net photosynthetic rates compared to the taller semi-dwarfs, but these higher rates occurred only at anthesis, had slightly greater carboxylation efficiencies and significantly increased chlorophyll concentrations per unit leaf area. The reduced-height wheat had significantly less dry mass fraction in the stem but greater dry mass partitioned to the ear than the taller semi-dwarfs (Yecora rojo, IBWSN-199). Studies with detached heads confirm that the head is a significant sink in the shorter wheat cultivars.     

三种栽培模式下不同基因型冬小麦旗叶衰老代谢比较

为了探索冬小麦在不同栽培模式下功能叶片衰老代谢的生理机制,以cp02(213)、cp99(1)和陕农512为材料,比较研究了常规栽培、覆草栽培、地膜覆盖3种栽培模式下小麦旗叶衰老代谢特性.结果表明,覆草栽培叶面积、旗叶功能期、叶绿素含量、叶片保护酶活性(SOD、POD、CAT)显著高于常规栽培,膜脂过氧化程度较低,叶片衰老速度缓慢,代谢强度旺盛,有利于籽粒灌浆和光合产物的积累.灌浆前期,地膜覆盖叶面积、旗叶功能期、叶绿素含量、叶片保护性酶活性(SOD、POD、CAT)显著高于常规栽培,膜脂过氧化程度低于常规栽培;灌浆后期,叶绿素含量急剧下降,叶片衰老速度加快,膜脂过氧化程度加剧.参试品种(系)中陕农512叶片衰老速度缓慢,保绿性好.     

水稻叶片生育过程中RubisCO活性与光合、光呼吸的关系

水稻生育过程中,ＲｕＢＰ羧化酶活性与光合速率、ＲｕＢＰ加氧酶活性与光呼吸速率、ＲｕＢＰ羧化酶活性与加氢酶活性以及光合速率与光呼吸速率之间是相关的。籼型品种与粳型品种间酶活性的高低及光合、光呼吸速率的高低基本一致,籼型三系杂交稻（Ｆ１）无明显的光合优势。酶的羧化活性的高低只在一定范围内与光合速率的高低平行。在正常生育条件下,酶蛋白的数量不是水稻光合速率的限制因子。     

Influence of reproductive organs on plant senescence in rice and wheat

The chlorophyll and protein contents of the flag, second and third leaves gradually decreased during the reproductive development of rice (Oryza sativa L. cv. Rasi) and wheat (Triticum aestivum L. cv. Sonalika) plants, whereas proline accumulation increased up to the grain maturation stage and slightly decreased thereafter. In rice plant, the rate of decrease in chlorophyll and protein and increase in proline level were higher in the flag leaf than in the second leaf. It was opposite in wheat plant. The export of [³²P]-phosphate from leaves to grains gradually increased reaching a maximal stage at the grain development stage, and then declined. The export of this radioisotope was greater in rice than in wheat. Removal of panicle at the anthesis and grainfilling stages delayed leaf senescence of rice plant, while in wheat the ponicle removal at any stage did not have a marked effect on delaying leaf senescence. The contents of chlorophyll and protein of glumes were higher in wheat than in rice. The variation of such source-sink relationship might be one of the possible reasons for the above effect on leaf senescence.