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.     

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.     

Leaf Regulation of the Nitrogen Concentration in the Grain of Wheat Plants

The regulation of the final grain N concentration in wheat (Triticumaestivum) plants was studied through the alteration of the source/sinkratio. Plants were grown in a greenhouse in pots with soil,and fertilized with a supraoptimal N supply. The plants were divided into six groups. In one treatment, plantsremained untouched as a control (Treatment 1). In another group,all the ears except that of the main tiller were removed atflowering (Treatment 2). All other plants were de-tillered afterthe emergence of the third leaf, leaving only one tiller perplant. At flowering, one plant set was left untouched (Treatment3). In a second group, all the leaves were excised (Treatment4). In another group half the spikelets of the ear were excised(Treatment 5) and in the last group three-quarters of the spikeletswere excised (Treatment 6). Ear excision produced an increase in individual grain weightand the grain N concentration above the normal N concentrationobserved in this cultivar. The final N concentration was correlatedwith the concentration of free amino acids in the flag leaf34 d after flowering. It is concluded that in intact plants grain protein synthesisis substrate-limited by the amino acid export pool in the leaves,and grain excision increases the availability of amino acidsto be transported to the remaining grains. Key words: Amino acids, grain N concentration, nitrogen, remobilization, wheat     

The genetics of nitrogen use in hexaploid wheat: N utilisation, development and yield

A genetic study is presented for traits relating to nitrogen use in wheat. Quantitative trait loci (QTLs) were established for 21 traits relating to growth, yield and leaf nitrogen (N) assimilation during grain fill in hexaploid wheat (Triticum aestivum L.) using a mapping population from the cross Chinese Spring × SQ1. Glutamine synthetase (GS) isozymes and estimated locations of 126 genes were placed on the genetic map. QTLs for flag leaf GS activity, soluble protein, extract colour and fresh weight were found in similar regions implying shared control of leaf metabolism and leaf size. Flag leaf traits were negatively associated with days to anthesis both phenotypically and genetically, demonstrating the complex interactions of metabolism with development. One QTL cluster for GS activity co-localised with a GS2 gene mapped on chromosome 2A, and another with the mapped GSr gene on 4A. QTLs for GS activity were invariably co-localised with those for grain N, with increased activity associated with higher grain N, but with no or negative correlations with grain yield components. Peduncle N was positively correlated, and QTLs co-localised, with grain N and flag leaf N assimilatory traits, suggesting that stem N can be indicative of grain N status in wheat. A major QTL for ear number per plant was identified on chromosome 6B which was negatively co-localised with leaf fresh weight, peduncle N, grain N and grain yield. This locus is involved in processes defining the control of tiller number and consequently assimilate partitioning and deserves further examination. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.     

Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears cultured <Emphasis Type="Italic">in vitro</Emphasis>

A detached culture system and steady-state ¹⁵N labeling technique were used to study the effects of exogenous ABA and ZR on photosynthetic characteristics, nitrogen remobilization and the activities of key enzymes for nitrogen metabolism in detached wheat parts during grain protein accumulation. The differences in net photosynthetic rate, chlorophyll content (SPAD value) and soluble protein content in the flag leaves of detached culture system between the treatments of ABA and ZR showed that ABA facilitates the post-anthesis senescence course compared to the ZR treatment. The differences in the changes of ¹⁵N amount in different organs in the detached culture system between the ABA and ZR treatments showed that nitrogen remobilization from vegetative organs to the grain is accelerated by the ABA treatment but is delayed by ZR. The activities of GS and GPT in grains treated with ABA were significantly higher than those with the control treatment at 5 DAC, but reduced significantly compared with control at 11 DAC. The two enzyme activities in grains were reduced significantly by ZR at 5 DAC and increased significantly at 11 DAC, compared with those treated with ABA. The above changes of enzyme activity showed that the ABA treatment hastens amino acid conversion into grains and protein accumulation in grains, whereas the ZR treatment delays these processes. A significant reduction in grain weight with ABA treatment is associated with the reduction of net photosynthesis, chlorophyll content, and soluble protein content in flag leaves. Compared with the control and ZR treatments, a significant increase in grain protein content with the ABA treatment may result from the accelerating effects of ABA on N remobilization, amino acid conversion into grains and protein accumulation in grains.     

Free Polyamines in Senescing Wheat(Triticum aestivum L.)

The free polyamine content of flag leaves, peduncles, rachis,glumes, and grains of wheat (Triticum aestivum L., cv. Castell)plants, ripening under field conditions, has been investigatedduring three consecutive growing seasons. Putrescine was quantitativelythe most important of all polyamines detected in these organs.Concentrations were highest in the grains, glumes and flag leaves.No correlation was found between polyamine content and the onsetof senescence of flag leaves and other organs. Excised primaryleaves, however, showed a decrease in polyamine content in thedark and also in light/dark cycles, but in the latter case onlyafter an initial increase. Sink removal of otherwise intactwheat plants caused an accumulation of putrescine in flag leavesat the later stages of senescence, whereas removal of all otherleaves was without any significant effect. Putrescine was alsorecovered in phloem-exudate samples collected throughout theperiod of grain development. In both grains and glumes, peakconcentrations of polyamines were found early during seed development. Key words: Triticum aestivum, polyamines, ripening, senescence     

土壤水分对强筋小麦‘豫麦34’氮素同化酶活性和籽粒品质的影响

 以强筋型小麦(Triticum aestivum)品种‘豫麦34号’为材料,采用盆栽方法研究了土壤水分对氮素同化酶活性及籽粒品质的影响。结果表明：旗叶硝酸还原酶（NR）活性于花后呈下降趋势,且土壤含水量为田间持水量（FC）60%的处理活性最强,其次为40%FC,活性最低的是80%FC。旗叶和籽粒中谷氨酰胺合成酶（GS）活性于开花15 d前均呈下降趋势,15 d后均为上升趋势,各水分处理间酶活性大小关系是：80%FC＞60%FC＞40%FC。各水分处理间旗叶和籽粒谷氨酸合成酶（GOGAT）活性的大小关系同GS。60%FC籽粒产量及品质最优,80%FC产量次之,40%FC产量最低;40%FC品质次之,80%FC品质最低。不同水分处理下籽粒蛋白质含量与叶片NR、GS 和籽粒GOGAT活性均呈正相关,与旗叶GOGAT活性呈负相关。且40%FC和80%FC下籽粒蛋白质含量只与旗叶GS活性相关性达显著水平, 60%FC下蛋白质含量则与旗叶NR和籽粒GS活性均达显著相关,与旗叶GS活性达极显著相关。     

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; )     

Factors Affecting Grain Enlargement in Wheat

When most of the grains were removed from wheat ears the remaininggrains developed increased numbers of aleurone and endospermcells, and the increased grain volume was accompanied by anenlarged endosperm cavity. The giant grains usually synthesizedstarch more rapidly than normal grains. Flag leaf photosynthesiswas unaffected and additional sugar was retained in the culmand glumes. Total nitrogen and free amino acid content increased.Gibberellin and auxin concentrations were greater in the glumesof the partly degrained ears, and auxin in the grains was alsogreater than in equivalent grains from intact ears.     

Regulation by kinetin and abcisic acid of correlative senescence in relation to grain maturation,source-sink relationship and yield of rice (Oryza sativa L.)

When kinetin was applied to the source organ (flag leaf) of rice (Oryza sativa L. cv. Ratna), foliar senescence was delayed and grain yield per plant (as evidenced by grain weight, grain/straw weight ratio and 1,000 grain growth) was increased through the increase of sink activity (increase in dry weight of the grains/plant), duration of sink capacity as well as photosynthetic ability of the glumes (as determined by the chlorophyll content of the glumes of the developing grains). However, application of kinetin to the sink organs (fruits), promoted senescence of the source but increased the yield by increasing the sink capacity and 1,000 grain growth mostly at the earlier stage of reproductive development. Lower sterility percentage was associated with higher grain yield of the plant by kinetin treatments. ABA applied either to the source or the sink promoted leaf senescence and reduced the grain yield by reducing the sink activity, harvest index, sink capacity duration and increasing the sterility percentage. Thousand grain dry weight at harvest did not vary significantly amongst the treatments. It was concluded that nutrient drainage was associated with the correlative influence of fruit on the monocarpic senescence of rice plant and that a competetion for differential allocation of cytokinin and ABA in the source and sink organs initiates this senescence syndrome.     

Effect of Subsoiling in Fallow Period on Soil Water Storage and Grain Protein Accumulation of Dryland Wheat and Its Regulatory Effect by Nitrogen Application

To provide a new way to increase water storage and retention of dryland wheat, a field study was conducted at Wenxi experimental site of Shanxi Agricultural University. The effect of subsoiling in fallow period on soil water storage, accumulation of proline, and formation of grain protein after anthesis were determined. Our results showed that subsoiling in fallow period could increase water storage in the 0–300 cm soil at pre-sowing stage and at anthesis stage with low or medium N application, especially for the 60–160 cm soil. However, the proline content, glutamine synthetase (GS) activity, glutamate dehydrogenase (GDH) activity in flag leaves and grains were all decreased by subsoiling in fallow period. In addition, the content of albumin, gliadin, and total protein in grains were also decreased while globulin content, Glu/Gli, protein yield, and glutelin content were increased. With N application increasing, water storage of soil layers from 20 to 200 cm was decreased at anthesis stage. High N application resulted in the increment of proline content and GS activity in grains. Besides, correlation analysis showed that soil storage in 40–160 cm soil was negatively correlated with proline content in grains; proline content in grains was positively correlated with GS and GDH activity in flag leaves. Contents of albumin, globulin and total protein in grains were positively correlated with proline content in grains and GDH activity in flag leaves. In conclusion, subsoiling in fallow period, together with N application at 150 kg·hm⁻², was beneficial to increase the protein yield and Glu/Gli in grains which improve the quality of wheat.     

Amino acid metabolism associated with N-mobilization from the flag leaf of wheat (Triticum aestivum L.) during grain development

Field-grown winter wheat (Triticum aestivum L. cv. Castell) was used to study changes in the free amino acid pools of different plant parts and related enzyme activities in the flag leaf throughout the grain-filling period in three consecutive growing seasons. Amino acid analysis data indicated that, during senescence, the nitrogen flow in the flag leaf was directed towards the synthesis of glutamine as a specific nitrogen transport form. Of the enzymes involved, total glutamine synthetase (GS; EC 6.3.1.2) and especially ferredoxin-dependent glutamate synthase (Fd-GOGAT; EC 1.4.7.1) activities declined continuously as senescence progressed. Unlike (chloroplastic) GS₂, (cytosolic) GS₁ was shown to be very persistent suggesting a special role for this isoenzyme in the N-reallocation process. Glutamate-oxaloacetate transaminase (GOT; EC 2.6.1.1), glutamate-pyruvate transaminase (GPT; EC 2.6.1.2) and isocitrate dehydrogenase (IDH; EC 1.1.1.42) showed a characteristic biphasic activity profile after anthesis. It is proposed that these enzymes, for each of which at least two isoenzymes were demonstrated, are involved in glutamate synthesis at the later stages of leaf senescence. Ammonium levels were fairly constant throughout the flag leafs life span, an ultimate rise often following peak values of glutamate dehydrogenase (GDH; EC 1.4.1.4) activity. The enzymology of flag leaf amino acid metabolism during grain development is further discussed in relation to observations of NH₃-volatilization from naturally senescing wheat plants.     

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.     

Mechanism of monocarpic senescence in rice

During grain formation stage (90 to 110 days), the youngest flag leaf of rice (Oryza sativa L. cv. Jaya) remained metabolically most active (as indicated by cellular constituents and enzyme activities) and the third leaf the least active. At the grain development stage (110 to 120 days) the above pattern of age-related senescence of the flag leaf completely changed and it senesced at a faster rate than the second leaf which remained metabolically active even up to grain maturation time (120 to 130 days), when both the flag and the third leaf partially senesced. Removal of any leaf temporarily arrested senescence of the remaining attached leaves, that of flag leaf did not hasten senescence of the second leaf, while that of either the second or the third accelerated senescence of the flag. Removal of the inflorescence after emergence or foliar treatment of intact plant with kinetin equally delayed senescence and produced an age-related, sequential mode of senescence or leaves. Both translocation and retention of ³²P by the flag leaf were maximum at the time of grain formation and that by the second leaf was maintained even up to grain maturation time. The induction of senescence of the flag leaf was preceded by a plentiful transport of ³²P to the grains. Kinetin treatment decreased the transport of ³²P, prolonged its duration, and almost equally involved all of the leaves in this process. The pattern of senescence of isolated leaf tips was similar to that of attached leaves. The level of endogenous abscisic acid-like substance(s) maintained a close linearity with the senescence behavior of the leaves of intact and defruited plants during aging, and the rise in abscisic acid in the flag leaf was also preceded by higher ³²P transport to the grains.     

缺氮和复氮对菘蓝幼苗生长及氮代谢的影响

对基质育苗后水培的菘蓝进行缺氮与复氮处理,分析其生长情况及氮代谢产物含量的变化,探讨缺氮和复氮对菘蓝幼苗生长及氮代谢的影响,以提高菘蓝产量和品质以及栽培过程中的氮素利用效率。结果显示:(1)正常供氮条件下,菘蓝幼苗的叶绿素含量、谷氨酰胺合成酶(GS)活性、硝态氮含量、靛玉红含量为最高,而其株高、主根直径、根的鲜重与干重、叶的鲜重与干重、根系活力均最小。(2)缺氮处理增加了菘蓝幼苗的主根直径和根干重,提高其根系活力和硝酸还原酶(NR)活性,促进游离氨基酸在叶中的积累;但降低了GS的活性,也降低了叶中硝态氮、可溶性蛋白、靛玉红及根中游离氨基酸的含量;缺氮对叶中靛蓝的含量无明显影响。(3)复氮处理增加了菘蓝幼苗的株高、主根长、根鲜重、叶鲜重、叶干重,提高了其根系活力,降低了NR和GS的活性;与对照相比,复氮降低了叶中硝态氮含量,提高了叶中可溶性蛋白、靛蓝及根中游离氨基酸的含量,但对叶中游离氨基酸和靛玉红含量影响较小。研究表明,缺氮后再复氮有利于菘蓝幼苗叶的生长,同时有利于增加其叶内靛蓝含量,从而提高其产量和品质。     

RuBP carboxylase activity in wheat following ear emergence and evaluation of the role of ear

Ribulose bisphosphate carboxylase activity and chlorophyll content were measured in the leaves and reproductive parts of two wheat cultivars after ear emergence. The chlorophyll content of the flag leaf was mostly higher than that of the awns and glumes. Awns had the highest chlorophyll content among reproductive parts. Light transmission to the lower leaves was higher in the dwarf cultivar Moti than in the medium tall cultivar Kalyansona. RuBP carboxylase activity in Kalyansona leaves was higher than in Moti leaves. In postanthesis stages there was no difference in RuBP carboxylase activity in the flag leaf and lower leaves between cultivars. Awns had the maximum activity of RuBP carboxylase followed by glumes and grains among ear parts. The relative capacity for photosynthesis in the ear parts was several times higher than in the flag leaf on a unit chlorophyll basis. It is suggested that in a crop canopy in the field, the spike(ear) may have a greater importance in grain development than has been previously estimated.     

氮素水平对花生氮素代谢及相关酶活性的影响

 在大田高产条件下研究了氮素水平对花生(Arachis hypogaea)可溶性蛋白质、游离氨基酸含量及氮代谢相关酶活性的影响, 结果表明, 适当提高氮素水平既能增加花生各器官中可溶性蛋白质和游离氨基酸的含量, 又能提高硝酸还原酶、谷氨酰胺合成酶和谷氨酸脱氢酶等氮素同化酶的活性, 使其达到同步增加; 氮素水平过高虽能提高硝酸还原酶和籽仁蛋白质含量, 但谷氨酰胺合成酶(GS)和谷氨酸脱氢酶(GDH)的活性下降; N素施肥水平不改变花生植株各器官中可溶性蛋白质、游离氨基酸含量以及硝酸还原酶(NR)、谷氨酰胺合成酶、谷氨酸脱氢酶活性的变化趋势, 但适量施N (A2和A3处理)使花生各营养器官中GS、GDH活性提高; 氮素水平对花生各叶片和籽仁中GS、GDH活性的高低影响较大, 但对茎和根中GDH活性大小的影响较小。     

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.     

夏谷籽粒蛋白质积累和旗叶蛋白质周转的关系

谷子灌浆期旗叶蛋白水解酶活性、籽粒和旗叶游离氨基酸含量均呈双峰曲线变化。第一峰在花后１３天;第二峰分别在２５、２８天。籽粒和旗叶游离氨基酸峰稍滞后,且后者不明显。籽粒总蛋白积累和干重增加均为Ｓ型变化,与旗叶总蛋白和可溶性蛋白变化趋势相反。花后２２─２５天上述物质和酶活变化均出现─转折。转折点之前,蛋白质合成和周转活跃;之后,由叶片蛋白酶活性升高引起的旗叶氮素撤离对籽粒蛋白质积累仍有一定贡献（约１０％）。