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; )     

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.     

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.     

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.     

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.     

Polyamines and leaf senescence in pyrrolizidine alkaloid-bearing Heliotropium plants

Putrescine, spermidine and spermine were found in leaves and inflorescences of H. angiospermum and H. indicum plants; the levels of these amines declined with leaf age. In addition, homospermidine was identified in the inflorescence axes and youngest leaves of H. indicum. The youngest tissues exhibited the highest levels of both putrescine and pyrrolizidine alkaloids. The detection of homospermidine in the plants supports the theory that the pyrrolizidine moiety is derived from two molecules of putrescine with homospermidine as an intermediate. In the youngest organs, the pyrrolizidines represented over 5% of the total nitrogen content. Their level was 50–100 fold higher than that of the polyamines, including putrescine. When detached and kept in the dark for 100–120 hr, mature older Heliotropium leaves, with a very low polyamine content, exhibited only a weak senescence syndrome. By contrast, in detached, darkened leaves of Avena sativa and Nicotiana alata having high polyamine levels, the chlorophyll and protein degradation and increases in free amino acids were very pronounced.     

Effects of Drought on Partitioning of Nitrogen in Two Wheat Varieties Differing in Drought-tolerance

A model was constructed to describe the translocation and partitioningof nitrogen on the seventh day after anthesis for well-wateredand droughted plants of two wheat varieties (Triticum aestivumL. cv. Warigal and Condor). The glasshouse-grown plants weredetillered so that a simplified model could be derived for themain stem. A 9-d drought treatment was imposed just after anthesisand this coincided with the period of endosperm cell divisionin the grains. Warigal, which had a higher grain yield thanCondor under drought, absorbed up to 15-times more nitrogenand translocated 1.5-fold more nitrogen to the shoot via thexylem. In both varieties, nitrogen redistributed from vegetativeorgans accounted for more than 60 per cent in control and 70per cent in droughted plants of the nitrogen needed for eargrowth. The net loss of nitrogen increased by 4-3 per cent inthe leaves, but decreased by 60 per cent in the stem under drought.Stem and roots appeared to play an important role in the nitrogeneconomy of droughted plants: less nitrogen was translocateddirectly to the grains from the senescing leaves and 40–60per cent more nitrogen was translocated to the roots. Nearlyall the nitrogen reaching the roots in the phloem was reloadedinto the xylem stream and translocated back to the shoot. Thetransfer of nitrogen through the stem was reduced under droughtand this resulted in a constant C:N ratio of the grains whichmay be important in the regulation of endosperm cell division. Triticum aestivum L., wheat, drought, nitrogen, senescence, translocation     

Changes in polyamine pattern mediates sex differentiation and unisexual flower development in monoecious cucumber (Cucumis sativus L.)

Changes in the levels of polyamines are associated with fundamental physiological processes such as embryogenesis, induction of flowering, fruit development and ripening, senescence, and responses to environmental stresses, but the role of polyamines in sex differentiation and unisexual flower development has not been deeply studied. To extend the knowledge on the regulatory mechanisms of flowering in monoecious plant (producing unisexual flowers), we investigated the morphogenesis and free polyamine levels in Cucumis sativus during sex differentiation and unisexual flower development in vitro using histocytological and biochemical methods. As shown in our study, floral development in vitro was undisturbed and flowers of both sexes were produced. Sex differentiation relied on preventing the development of generative organs of the opposite sex, as we observed carpel repression in male flowers and stamen repression in female flowers. Pollen viability was negatively correlated with female flower development on the same node. Biochemical analysis revealed increased accumulation of aliphatic amines (tri, tetra‐amines) in generative (flower buds and flowers) compare to vegetative (axillary buds and leaves) organs. Undifferentiated floral buds contained elevated levels of agmatine, cadaverine, spermidine and spermine. Sex differentiation was associated with significantly decreased levels of agmatine and cadaverine. Our results showed that female flowers contained higher levels of total polyamine than male flowers. The increased level of cadaverine was associated with macrogametogenesis and female flower maturation. Putrescine was important for male flower development. Such results support the hypothesis that aliphatic amines are involved in unisexual flower development.     

Observations on the Distribution of Mineral Elements in the Leaf of Wheat (Triticum aestivum L.), with Particular Reference to Silicon

Wheat (Triticum aestivum L. cv. Wheaton) plants were grown inwater culture or in soil. Basal leaves (B) were harvested after3 weeks from the water culture plants, while flag leaves werecollected from soil-grown material at the time of inflorescenceemergence (E0) and 7 d after emergence (E + 7). Mineral distributionin bulk frozen leaves was investigated using SEM and X-ray microanalysis.The elements detected were silicon, phosphorus, chlorine, sulphur,potassium and calcium. Potassium was present in all cell typesat all harvests, chlorine was almost entirely confined to theadaxial and abaxial epidermi, while sulphur was only rarelydetected in the E0 and E + 7 leaves. Phosphorus was presentat higher levels in the E + 7 leaves than in the B or E0 leaves.At the B harvest calcium was confined to the adaxial epidermalcells, but in the E0 and E + 7 leaves it was present in bothepidermi. Silicon was, initially, mainly detected in the abaxialepidermal cells, but in older (E + 7) leaves it was presentin both epidermi and in some internal tissues. Mineral transportwithin the leaf and ionic environment at silica deposition sitesare discussed. Wheat, Triticum aestivum L., leaf, mineral distribution, X-ray microanalysis, silicon, calcium     

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     

Effects of Abscisic Acid on Growth of Wheat (Triticum aestivum L)

Daily application of abscisic acid (ABA) to growing wheat plants,although initially inhibiting growth, resulted, after a shortlag, in an increase in the number of leaves and tillers. Thismay have been due to reduced apical dominance. At 84 days thetotal dry weight and area of all leaves produced up to thistime was less for the plants treated with ABA than for the controlplants. However, the area of green, living leaves and the dryweight were not significantly affected by the ABA treatment.Further effects of the daily ABA treatment were the inhibitionof transpiration, especially on the abaxial surface, the reductionof leaf size, the promotion of flowering and the stimulationof trichome formation on the leaf surfaces. ABA did not promoteleaf senescence in whole plants and actually increased leaflongevity. Triticum aestivum L., wheat, leaf senescence, transpiration, growth, flowering, abscisic acid     

The Structure of the Mesophyll of Flag Leaves in Three Triticum Species

Flag leaves of Triticum urartu, T. monococcum and T. aestivumcv. Professeur Marchal were examined by light and electron microscopyand by separating cells to determine whether differences inleaf anatomy could be related to known differences in theirlight-saturated rates of photosynthesis. Mesophyll cells fromthe three species were lobed and orientated with their longaxis parallel to the veins. The longest, most-lobed cells flankedthe sclerenchyma associated with the veins. Mean cell dimensionswere greatest in Professeur Marchal, but there was no significantdifference in the ratio of the mesophyll cell surface area tocell volume amongst the three species. Flag leaves of T. urartushowed the highest rates of photosynthesis and were also thethickest, with closely-spaced veins from which many of the mesophyllcells radiated. These flag leaves also had significantly more(21.9 per cent) air-filled space, and the highest ratio (15.2)of mesophyll cell surface exposed to this air-filled space perunit leaf area. Ways in which these anatomical characteristicsmay contribute to the higher rate of photosynthesis are discussed. Triticum urartu, Triticum monococcum, Triticum aestivum, flag leaves, morphology, mesophyll     

Effect of Drought on Metabolism and Partitioning of Carbon in Two Wheat Varieties Differing in Drought-tolerance

A model was constructed to describe the partitioning of carbonon the third and seventh day from anthesis for well-wateredand droughted plants of two wheat varieties (Triticum aestivumL. cv. Warigal and Condor). The glasshouse-grown plants weredetillered so that a simplified model could be derived for themain stem. The 9-d drought treatment, imposed just after anthesisduring the period of cell division in the grains, reduced grainyield by 18 per cent in Warigal and 30 per cent in Condor. Netcarbon fixation was up to 60 per cent higher in Warigal thanCondor towards the end of the drought period and this correlatedwith better osmotic adjustment in the flag leaf. Carbon partitioningbetween plant organs responded to water deficit more rapidlythan net carbon fixation. On day 3, carbon allocation to theroots of droughted plants was maintained in Condor and increasedby 14 per cent in Warigal, whereas carbon allocation to theear decreased in both varieties. However the roots did not competewell with the ear when the water deficit became more severe.Warigal accumulated 3 times more stem reserves than Condor underdrought. In the roots, the pattern of carbon allocation betweenrespiration and carbon accumulation changed soon after impositionof drought. Although total root respiration decreased underdrought it became more energy efficient, particularly for Warigal,as less respiration took place via the alternative pathway.On day 3, the larger carbon allocation to the roots and thelower root respiration accounted for the 4-times larger sugaraccumulation in droughted roots of Warigal compared with thoseof Condor. Osmotic adjustment in mature leaves and roots maybe of importance for the maintenance of vital processes andfor recovery after drought. Triticum aestivum L., wheat, drought, carbon, partitioning     

The Effects of Dwarfing Genes Rht1 and Rht2 on Cellular Dimensions and Rate of Leaf Elongation in Wheat

The gibberellin insensitivity genes, Rht1 and Rht2, reducedepidermal cell lengths in leaves of isogenic lines of field-and laboratory-grown wheat (Triticum aestivum L.). Rht dosagesof zero (wild type), two (semi-dwarf) and four alleles (doubledwarf) had a linear negative effect on cell length in flag leavesof field-grown plants, and in the sheaths and blades of leafnumber 1 in laboratory grown plants. Decrease in cell length,rather than reduced cell number, accounted for most to all ofthe reduction in blade and sheath length. In sheaths, cell widthincreased with Rht dosage, but not sufficiently to compensatefor decreased length in determining average projected surfacearea. Rates of extension of leaf number 1 in laboratory-grownplants were negatively and linearly correlated with Rht dosage.Maximal growth rate was maintained longer in wild type thanin double dwarf, but the total duration of measurable extensionin leaf number 1 was not affected by Rht dosage. Cell size, elongation, Rht, wheat, Triticum aestivum L     

Copper Supply in Relation to Content and Redistribution of Copper Among Organs of the Wheat Plant

The relationship of copper supply to the content and movementof copper among organs of wheat plants was examined at sevenstages in their growth from seedlings to maturity on a copperdeficient sand. In the absence of copper (Cu₀), plants becameseverely copper deficient and produced no grain; developmentof tillers, leaves, stems, and inflorescences was delayed andgrowth of roots strongly depressed; leaf senescence was retardedand tiller growth was prolonged. Application of a marginal supplyof copper (Cu₁) overcame all symptoms and promoted growth andgrain production. Increasing copper supply eightfold (Cu₂) didnot change vegetative or grain production. Copper concentrations in stems, individual leaves, and wholetops were highest and responded most strongly to copper supplywhen they were young. As they aged, Cu₁ and Cu₂ leaves lostcopper rapidly; the first Cu₀ leaves retained their copper andremained healthy for more than 7 weeks even though younger leavesdeveloped severe copper deficiency. In all treatments, lossof copper from the oldest leaf paralleled senescence and theloss of nitrogen. It is suggested that copper does not move out of plant leavesuntil they lose organic nitrogen compounds. As a result, copperbehaves in non-senescent leaves as if it is not mobile in plantphloem. But under conditions favouring senescence, copper ishighly mobile: in the present experiment, 67 per cent of thecopper present in vegetative organs of the Cu₂ primary shootat flowering moved from them during grain development and thiscould account for all of the copper found in the grain at maturity. The retention of copper by leaves before senescence, its rapidloss during senescence, and the effect of copper deficiencyin delaying senescence resulted in the oldest leaf of severelydeficient Cu₀ plants in the present experiment having a highercopper concentration than that of copper adequate Cu₁ and Cu₂plants. This behaviour could account for the many reports ofanomalous C-shaped ‘Piper-Steenbjerg’ curves inthe relationship of yield to copper concentrations in planttops. The coupling of copper movement from leaves to nitrogenmovement can also account for the unusually high values reportedfor critical concentrations of copper in tops of plants givenhigh levels of nitrogen fertilizers. Old organs should not be included in samples for diagnosis ofcopper deficiency. Only young organs should be used. In thepresent experiment, the copper concentration of young leavesgave a good indication of the copper status of wheat: a valueof 1 µg g^–1 in young leaves indicated copper deficiency. copper, nitrogen, phloem transport, mineral transport, deficiency diagnosis, wheat, Triticum aestivum L.     

Abscisic Acid Translocation and Influence of Water Stress on Grain Abscisic Acid Content

The movement of foliar applied [1-¹⁴C]abscisic acid (ABA) inwheat plants (Triticum aestivum L., cv. Kolibri) was investigatedat two stages of grain development (1000 grains, weight 19 and24 g dry matter). [1–¹⁴C]ABA seemed to be readily translocated within 12h into the developing grains as well as in other plant parts.A subsequent rapid metabolism took place leading to a decreasedactivity of the ABA-containing chromatogram fraction in theyounger plants 48 h after application. The metabolism seemodto be less intensive in the older grains, where the activityrunning with the ABA increased over 64 h. Treating the leaves of barley plants (Hordeum vulgare, L., cv.Union) 2 weeks after anthesis with a gentle stream of warm air(36° C) resulted in a significant increase in the ABA contentof all parts of the ear. The results mentioned above indicatethat this may be partially due to translocation from other partsof the plant such as the leaves.     

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.     

Nitrogen Redistribution during Grain Growth in Wheat (Triticum aestivum L.) : IV. Development of a Quantitative Model of the Translocation of Nitrogen to the Grain

Translocation of nitrogen was measured in wheat (Triticum aestivium L. cv SUN 9E) plants grown without an exogenous supply of nitrogen from the time that the flagleaf began to emerge, and a model of nitrogen translocation was constructed to describe translocation on one day during the linear period of grain growth. Nitrogen for grain development was derived entirely by the redistribution of nitrogen from vegetative organs. Leaves contributed 40%, glumes 23%, stem 23%, and roots 16% of the nitrogen incorporated by the grains on the fifteenth day after anthesis. Less than 50% of the nitrogen exported from leaves was translocated directly to the grain via the phloem, the rest was translocated to the roots and was cycled in the roots and exported to the shoot in the transpiration stream. Nitrogen imported by leaves and glumes via the xylem was not accumulated in these organs but was transferred to the phloem for reexport from the organs. A large proportion (60%) of the nitrogen in the transpiration stream was cycled in the glumes. The glumes were also a major source of nitrogen for grain development. It was considered likely that this organ always plays an important role in nitrogen metabolism in wheat.     

Enhancement in Growth and Yield of Wheat (Triticum aestivum L.) by Pre-sowing Chill Treatment to Grains

Pre-sowing chill treatments to grains of Wheat (Triticum aestivumL. cvs Kalyan sona and cv. 4530) induce enhancements in growth,tillering and yield. Further, levels of several biochemicalcomponents are also higher in the grains harvested from plantsgrown from chill-treated grains. This method applied in thelaboratory and on a farm scale, effective also in other plants,can be conveniently used for yield enhancement and improvementof seed quality. Triticum aestivumL., wheat, chill treatment     

Changes in polyamine concentration during seed germination

Phaseolus mungo seeds 0 to 10 days after germination contained putrescine, spermidine, spermine, cadaverine, agmatine and tyramine. The rate of biosynthesis of total polyamines, proteins and RNA in the developing seeds follows similar profiles, reaching maxima 3 hr from germination. Putrescine, cadaverine, spermidine, spermine and agmatine were the major amines found in Pisum sativum 0–7 days after germination. RNA and proteins seem to follow the same pattern as polyamines during the first 12 hr in the developing pea seeds. RNA reaches a peak at 15 hr and polyamines and proteins peak 24 hr after germination. A rise to total polyamine concentration was also observed in seeds of Tragopogon porrifolius, Zea mays and Triticum aestivum 2–12 hr after germination.