Distribution of photosynthate produced before and after anthesis in tall and semi-dwarf winter wheat, as affected by nitrogen fertiliser

In 2 years the distribution of radioactivity recovered in entire shoots of field-grown winter wheat was determined at various times after exposing the top two leaves (flag leaf or second leaf) to ¹⁴CO₂ for 30 s. In 1976 when ¹⁴C was supplied to either leaf 14 days before anthesis, 30% was in the ear at anthesis. Less than 5% was in the leaf exposed to ^I4CO₂. The remainder was equally divided between the stem above and below the flag-leaf node when the flag leaf had been exposed, and was mainly in the lower part of the stem when the second leaf had been exposed. By maturity the proportion in the stem had decreased; 20% of the total activity was in the grain and 30% was still in the ear structures. When ¹⁴C was supplied 10 days after anthesis, the proportion in the ear 24 h later ranged from 42 to 69% of that in the whole shoot when the flag leaf was exposed, and from 6 to 28% when the second leaf was exposed. At maturity these proportions increased to 92 and 85% when the ¹⁴C had been supplied to flag leaves and second leaves respectively. When ¹⁴C was supplied 25 days after anthesis to either flag leaves or second leaves, more than 90% of the activity was in the mature ears. Less than 5% of the ¹⁴C remaining at maturity from any treatment was still in the leaf exposed to ¹⁴CO₂. Between 2 and 6% of ¹⁴C supplied after anthesis was in the non-grain parts of the ear. The proportion of the ¹⁴C in the ear was greatest for the semi-dwarf varieties Maris Fundin and Hobbit, less for Maris Huntsman, and least for Cappelle-Desprez. These varietal differences were large 24 h after exposure to ¹⁴CO₂, especially in 1976. They were small and rarely significant at maturity. Nitrogen fertiliser up to 210 kg N ha^-1 had negligible effects on the distribution of ¹⁴C, although it greatly increased growth and yield, especially in 1975.     

Effects of crop thinning and reduced grain numbers per ear on grain size in two winter wheat varieties given different amounts of nitrogen

To study the importance for final grain size in wheat (Triticum aestivum, L.) of assimilate supply and the storage capacity of the grain, two field experiments were done. In 1976 nitrogen was applied in the range from none to 180 kg ha^-1, part of the crop was thinned, and the top halves of some ears of the short variety Hobbit and of the tall variety Maris Huntsman were removed soon after anthesis. In 1977 ears of Maris Huntsman were halved 5 days after anthesis or at 30 days after anthesis when grain volume was maximum. Thinning the crop from 360 to 180 ear-bearing shoots m^-2 30 days before anthesis increased the number of grains per ear, except in the absence of nitrogen fertiliser, but did not increase grain size, grain dry weight per ear or total dry weight per culm. Removing the upper half of ears of Hobbit 5 days after anthesis increased dry weight per grain, but when this treatment was applied to Maris Huntsman either 5 days after anthesis in 1976 and 1977, or when grain volume was maximal in 1977, the grains failed to increase in dry weight. Non-grain dry weight of both varieties was increased by halving the ear. In both varieties the maximum volume of grains in halved ears was larger than in intact ears. Grain dry weight increased relatively less than volume after halving the ear of Hobbit, and the decrease in volume up to maturity was greater in halved than intact ears of both varieties. The larger grain volume in halved ears of Maris Huntsman in 1977 was associated with more endosperm cells.     

Effects of decreasing the number of grains in ears of cvs Hobbit and Maris Huntsman winter wheat

At 6 days after anthesis, grain numbers in ears of Maris Huntsman and Hobbit winter wheat growing in the field were decreased either by removing the two lower grains in each spikelet (degraining) or by removing the top half of the ear (halving). At maturity, degraining increased the dry weight of the third grains in each spikelet of Maris Huntsman by 11% and of Hobbit by 40%, compared with third grains in intact ears. Halving increased the mean dry weight of all the grains in the lower six spikelets of the ear slightly less; it increased the number of grains in Hobbit but not in Maris Huntsman. The responses to halving in Hobbit were greater with additional nitrogen fertiliser. At 28 days after anthesis in both varieties, degraining increased grain dry weight and the amount of water, reducing sugar, amino acids and total nitrogen in third grains. Effects of halving on these properties of the two lower grains of each spikelet were much less or nil. The increases in nitrogen content of grains at 28 days and at maturity caused by degraining or halving were relatively greater than the increases in dry weight and were similar in the two varieties.     

Dark Respiration of Several Varieties of Winter Wheat Given Different Amounts of Nitrogen Fertilizer

Carbon dioxide production in the dark by ears and by the restof the shoot of winter wheat grown in the field was measuredin 2 years during grain growth. The respiration rate per g d.wt of the ears was increased by nitrogen fertilizer. Ears ofthe semi-dwarf varieties Maris Fundin and Hobbit respired moreslowly than ears of Maris Huntsman and Cappelle-Desprez. Respirationrates of the rest of the shoot were unaffected by nitrogen orvariety. The amount of carbohydrate required to provide the CO₂ respiredduring the whole period of grain growth varied from 163 to 443g m^–2, or 42 to 76 per cent of the dry weight of the grain.More than half the CO₂ lost was respired by the ear. The additionof 180 kg N ha^–1, which increased grain yield by 78 percent in 1975, almost trebled the amount of CO₂ lost by the ears.The semi-dwarf varieties lost less CO₂ from ears and shootsthan did the taller ones, and had larger yields of grain. Respiration was also estimated from the difference between the¹⁴C contents of shoots sampled immediately after a 30 s exposureto ¹⁴CO₂ and at maturity. When 14C was supplied 10 days afteranthesis, the loss by maturity amounted to 16–28 per centof that initially absorbed by flag leaves and 40 per cent ofthat absorbed by the leaf below the flag leaf. Most of the lossoccurred in the first day. The loss of 14C by maturity was significantlyincreased by nitrogen fertilizer in 1975. Triticum aestivum L., wheat, respiration, nitrogen supply, fertilizer treatment     

Characterisation of grain yield and nitrogen level in relation to flag leaf senescence of wheat varieties

Three wheat cultivars ( Triticum aestivum L.), Splendeur, Hobbit and Maris Huntsman grown in pots were compared. Especially when compared to Splendeur, the flag leaf senesced most rapidly in Maris Huntsman, which presented the most rapid loss of moisture, chlorophyll and nitrogen. The uptake of exogenous nitrogen during the post-anthesis period was lower in the rapidly than in the slowly senescing variety. A higher concentration of free amino nitrogen in the flag leaf at a given sampling date was associated with a lower percentage decrease of soluble proteins at the following date. Acid proteinase activity in the flag leaf was inversely related to moisture percentage and free amino nitrogen level, but unrelated to the nitrogen loss of the flag leaf. Acid proteinase activity in the flag leaf was directly related to grain nitrogen percentage, but inversely related to grain yield. Grain yield was also directly related to the mean soluble protein content of the flag leaf through senescence.     

Effects on dry weight and nitrogen content of grains of semi-dwarf and tall varieties of winter wheat caused by decreasing the number of grains per ear

The number of grains in ears of six varieties of winter wheat growing in the field was decreased by removing the top half of the ear 6 days after anthesis (halving). In the semi-dwarf varieties Hustler, Sentry and Hobbit, and in the taller varieties Armada and Flanders, the mean dry weight per grain in the lower six spikelets of halved ears was about 23% greater than the dry weight of grains in the comparable part of intact ears. In Maris Huntsman the increase in grain size amounted to only 13%. Halving increased the number of grains in the semi-dwarf varieties but not in the others. Consequently, the increases caused by halving in the total weight of grain in the lower six spikelets ranged from 41% in Hustler and Hobbit to 7% in Maris Huntsman. Halving increased the amount of nitrogen in the grain of the lower half of the ear similarly in all varieties, and relatively more than it increased dry weight. So nitrogen per cent dry weight of grain was increased, especially in Armada, Flanders and Maris Huntsman which responded least in dry weight. The uptake of nitrogen into the stem plus ear after anthesis was unaffected by halving. Most of the nitrogen that normally moved to the upper grains accumulated in the lower grains of halved ears. More nitrogen moved into the grain of intact ears of tall than of semi-dwarf varieties after anthesis, because there was greater uptake of nitrogen into the shoot from the roots.     

Nitrogen Uptake and Accumulation in Grains of Three Winter Wheat Varieties with Altered Source--Sink Ratios

In two pot experiments, removal of the top halves of ears (halving)of three winter wheat (Triticum aestivum L.) varieties at varioustimes after anthesis increased the nitrogen content in the grainsof the lower half of the ear. The increase was greater withearly (anthesis and 5 d later) than late (15 and 25 d post-anthesis)removal in the Splendeur and Hobbit varieties, but there wereno significant differences among halving times in Maris Huntsman.Halving also increased nitrogen as a percentage of dry weightof grain. The percentage of nitrogen in the grain decreasedas the time of halving was delayed in Splendeur (expt. 1) andHobbit, but was unaffected by the time of halving in Splendeur(expt. 2) and Maris Huntsman. Nitrogen uptake of shoots afteranthesis decreased with halving. Early halving decreased nitrogenuptake to a lesser extent than did late halving in Splendeurin expt. 1 and in Hobbit, while it decreased nitrogen uptakemore than late halving in Splendeur in expt. 2 and in Mans Huntsman. Key words: Grain nitrogen, nitrogen uptake, source-sink, wheat, variety     

Effect of Nitrogen Fertilizer on Photosynthesis of Several Varieties of Winter Wheat

The rates of gross photosynthesis of the flag leaf and the nextleaf below (second leaf) in crops of winter wheat were estimatedfrom the ¹⁴C uptake of the leaves after exposure to short pulsesof ¹⁴CO₂. The photosynthetic rates of both leaves during thegrain-filling period decreased with increase in nitrogen fertilizerbecause the intensity of photosynthetically active radiationwas less at the surface of the leaves in the dense crops withadditional nitrogen. In addition, the rate of photosynthesisat saturating light intensity was slightly decreased by nitrogen.The effects of nitrogen, in decreasing the rate of photosynthesisper unit area of leaf and in increasing the leaf-area indexof the top two leaves, were such that the photosynthetic productivityper unit area of land of the flag leaf was increased by nitrogenbut the productivity of the second leaf was unaffected. Applying180 kg N ha^–1 increased the productivity of the top twoleaves by a factor of 2.3 but increased grain yield by only1.8. The photosynthetic productivity of the second leaf duringthe grain-filling period was about half that of the flag leaf. There was no difference in photosynthetic rate per unit areaof leaves of Cappelle-Desprez and Maris Huntsman which couldaccount for the larger yield of the latter cultivar. There wasa slight indication that the leaves of the semi-dwarf cultivarsMaris Fundin and Hobbit photosynthesized faster than those ofMaris Huntsman. Triticum aestivum L., winter wheat, photosynthesis, nitrogen fertilizer     

Regulation of Grain Weight by Supply of Assimilates and Starch Granule Development in Three Winter Wheat Varieties

On removing the top half of the ear (halving) on several datesafter anthesis, dry weight per grain increased in three winterwheat (Triticum aestivum L.) varieties, in two pot experiments;the increase was greater with early than with late halving.The variety Splendeur had a lower dry weight and water percentagein grains than either Hobbit or Maris Huntsman. The ratio ofthe green area integrated over the post-anthesis period to thenumber of grains per ear (green area per grain) was highestin Splendeur and lowest in Hobbit in the first experiment; inthe second, Splendeur gave a lower ratio than the other twovarieties, which showed similar values. The green area per grainwas greater the earlier the ear was halved. The number of A-typestarch granules per endosperm, but not the volume per A granule(modal volume) and the modal volume of B starch granules, butnot their number, increased to a greater extent with early thanwith late halving. In Splendeur the grains had fewer A starchgranules, although these were of greater modal volume than inHobbit and Maris Huntsman and a number of B starch granulessimilar to the other two varieties, but of smaller modal volume.Maris Huntsman had more A granules than Hobbit, but with smallermodal volume. Dry weight per grain increased linearly with thenumber of A starch granules per endosperm, which in turn increasedasymptotically with green area per grain. The regressions forthe three varieties differed significantly. The influence ofthe supply of assimilates and the capacity for starch granuleformation in the regulation of grain weight is discussed. Key words: Grain weight, starch granules, assimilate supply, variety, wheat     

Effects of Nitrogen Fertilizer on the Distribution of Photosynthate during Grain Growth of Spring Wheat

The distribution of photosynthate labelled with ¹⁴C was studiedin spring wheat grown with different amounts of nitrogen fertilizerin the three years 1972–4, after exposing the flag leafor the leaf below the flag leaf to ¹⁴CO₂ at 6–10 or 19–26days after anthesis. The movement of ¹⁴C to ears was unaffectedby nitrogen fertilizer except after early exposure in 1973,when nitrogen increased the retention of ¹⁴C in stems at maturity The concentration of sugar in the top part of the shoot at theend of the day was unaffected by nitrogen in 1973, but at 22days after anthesis in 1974 the concentration of sucrose inthe glumes and rachis, and in the flag leaf lamina was increasedby nitrogen. Loss of sugar by translocation and respirationduring the night may explain why this increase in concentrationwas not reflected in the ¹⁴C distribution 24 h after supplying¹⁴C. The proportion of the total ¹⁴C content of the shoot that wasin the ear at maturity ranged from 68 to 95 per cent dependingon when and to which leaf the ¹⁴CO₂ was supplied. Less than5 per cent remained in the leaf exposed to ¹⁴CO₂. The proportionof the final ear weight contributed by the leaf below the flagleaf was about half that contributed by the flag leaf. In 1974 about 24 per cent of the ¹⁴C absorbed by the flag leaf,and 56 per cent of that absorbed by the second leaf, was lostby maturity, presumably by respiration. Most loss occurred inthe first 24 h.     

The Influence of Leaf Removal upon the Development of the Grain of Winter Wheat

The paper describes the effects on grain development of theremoval of half the flag leaf, the entire flag leaf and allthe leaves from selected main shoots of a field crop of wheat,cultivar Maris Ranger, 7 days after anthesis. None of the treatmentsreduced grain weights in the 14 days following defoliation.Even in the next 14 days only the more severe defoliations reducedgrain weight, with the most severe treatment causing the greatestreduction. Ultimately the rate of grain growth in plants onwhich only a half of the flag leaf had been removed also fellbehind that of the controls. Final grain yields were in theorder, control plants, > plants with half the flag leaf removed,> the entire flag leaf removed, > all the leaves removed.The significance of these results in interpreting the factorslimiting grain growth in wheat is discussed.     

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.     

Translocation of photosynthetic assimilates in wheat

When ¹⁴CO₂ was supplied to three varieties of wheat in the field translocation of carbohydrates took place freely throughout young plants, but after the stems elongated the tillers became autotrophic. The efficiency with which carbohydrate was translocated to the grain increased during the first 4 weeks after anthesis and then fell. Translocation from the glumes and flag leaves was almost entirely towards the grain; that from the second and third leaves was partly towards the grain and partly downward, with evidence of varietal differences in behaviour.     

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.     

Photosynthetic and translocation pattern in contrasting winter wheat varieties

The rates of photosynthesis and patterns of translocation of ¹⁴C from the flag leaves of tall and semi-dwarf wheat varieties, with and without awns, were compared in a field experiment. The rates of photosynthesis of the awnless varieties fell during the 14 days after anthesis but those of the awned varieties did not change. Wide differences in translocation pattern of ¹⁴C supplied at anthesis were also found between varieties, but the differences in distribution pattern of ¹⁴C supplied 14 days later were less. At anthesis, the high yielding semi-dwarf selection translocated most carbohydrate to the ears and awned selections least. It is suggested that varietal differences in translocation pattern are determined by differences in the capacity of the grain to store carbohydrate.     

Translocation from the Flag Leaf of Winter Wheat in the Field

Translocation of assimilate from the flag leaf of winter wheat(Triticum aestivum cv. Maris Huntsman) was studied in the fieldby monitoring the export of photo-assimilated [^l4C]carbon dioxidewith a Geiger-Miiller counter placed under the fed area of leaf.The resulting export curve was analysed as a sum of two exponentialterms, and interpreted as a two-pool compartmental system. Therate constant for export from the leaf increased slightly frommaximum elongation to anthesis, then declined to almost halfits peak value just before the leaf lost all visible chlorophyll.The inter-pool transfer rate constants did not change significantlyover the same period, but all rate constants varied with timeof day. Short-term changes in the environment of the flag leaf had nodiscernible influence on translocation in the field. The timeconstants of the two pools of assimilate agreed with those forother species reported in the literature. These results areconsistent with the suggestion that sucrose is stored in thevacuole of mesophyll cells. The variations in rate constants with time of day, and deviationsof the export data from the two-pool model, suggest that exportand inter-pool transport have saturation kinetics. A model withMichaelis-Menten kinetics was formulated, and simulations ofthis model showed similar deviations from a simple two-poolsystem to those seen in our data.     

Effect of abscisic acid on the transport of assimilates in barley

The effect of abscisic acid (ABA) on assimilate transport in barley was investigated in two parallel experiments. First, the effect upon [¹⁴C]sucrose transport from the flag leaf to the ear of a single ABA application made at different stages of growth of the fruits was investigated; the effect was measured 24 h after treatment. Second, the effect of a single application of ABA made at the same stages of growth as above on grain weight of the mature plant was investigated. In both types of experiments ABA was applied once to the ear of different plants as an aqueous solution (10^-3–10^-5 M), one to five weeks after anthesis. [¹⁴C] sucrose was applied by means of agar blocks. Parallel to these experiments, the endogenous content of ABA was investigated in the developing grains. When ears were treated with ABA two or four weeks after anthesis, an increase of up to 70% in the ¹⁴C-transport from the flag leaf to the ear was observed within a 24-h period after treatment (short duration experiments). At these growth stages the endogenous concentrations of ABA were low. In sharp contrast, ABA, especially in a concentration of 10^-3 M, decreased ¹⁴C-import from the flag leaf when applied three weeks after anthesis. At this stage the endogenous ABA content had reached its maximum. Long duration experiments with a single application of ABA to the car two weeks after anthesis resulted in a marked increase of weight per thousand kernels. ABA applications made earlier or later than two weeks after anthesis either reduced the grain weight or had no effect. It is concluded that ABA is involved in the regulation of assimilate transport from the leaves to the grains, possibly by influencing the unloading of sieve tubes in the ears. Promotion or inhibition of assimilate import by exogenously applied ABA may depend on the developmental stage of the grains and on the endogenous ABA level.Abbreviations ABA abscisic acid - TKW weight per thousand kernels     

Effects of Nitrogen Fertilizer on Growth and Yield of Spring Wheat

Nine amounts of nitrogen fertilizer, ranging from 0 to 200 kgN ha^–1, were applied to spring wheat cv. Kleiber in the3 years 1972-1974. In 1972 grain dry weight with 125 kg N ha^–1or more was 100 g m^–2 (23 per cent) greater than withoutnitrogen. Grain yield was unaffected by nitrogen in the otheryears. Leaf area at and after anthesis was increased throughoutthe range of nitrogen tested, most in 1972 and least in 1973.Consequently, the addition of 200 kg N ha^–1 decreasedthe amount of grain produced per unit of leaf area by approximately25 per cent in all years. The dry weight of leaves and stems at anthesis and maturitywas increased by nitrogen in all years, similarly to leaf area.However, the change in stem dry weight between anthesis andmaturity was not affected by nitrogen; stems increased in dryweight for about 20 days after anthesis and then decreased tovalues similar to those at anthesis. The uptake of CO₂ per unit area of flag leaf or second leaf(leaf below the flag leaf) was slightly decreased by nitrogenwhen the increase in leaf area caused by nitrogen appreciablydecreased the light intensity at the surface of these leaves.In spite of such decreases the CO₂ absorbed by flag and secondleaves per unit area of land was always increased by nitrogen,and relatively more than was grain yield. It is suggested that increases in respiratory loss of CO₂ withincreasing nitrogen fertilizer may explain why nitrogen increasedvegetative growth and leaf area relatively more than grain yield.     

Effect of assimilate utilization on photosynthetic rate in wheat

Summary Two weeks after anthesis, when the grain is filling rapidly, the rate of photosynthesis by flag leaves of wheat cv. Gabo was between 20 and 30 mg CO₂ dm^-2 leaf surface hour^-1 under the conditions used. About 45% of flag-leaf assimilates were translocated to the ear, and only about 12% to the roots and young shoots.On removing the ear, net photosynthesis by the flag leaves was reduced by about 50% within 3–15 hours, and there was a marked reduction in the outflow of ¹⁴C-labelled assimilates from the flag leaves.Subsequent darkening of all other leaves on plants without ears led to recovery of flag-leaf photosynthesis, as measured by gas analysis and ¹⁴CO₂ fixation, and to increased translocation of assimilates to the roots and young shoots. Minor changes in the rates of dark respiration accompanied these major, reversible changes in photosynthetic rate.After more than a week in continuous, high-intensity light, the rate of photosynthesis by flag leaves of intact plants had fallen considerably, but could be restored again by a period in darkness, or by inhibiting photosynthesis in the ears by spraying them with DCMU. The inhibition of ear photosynthesis increased translocation of labelled assimilates from the flag leaf to the ears, without affecting leaf sugar levels.The application of TIBA to the culm below the ear inhibited auxin movement throught the culm, but had no influence on flag-leaf photosynthesis.These results suggest that, at least in this system, photosynthesis by the flag leaf is regulated directly by the demand for assimilates from the flag leaf and not indirectly through action in the leaf of auxins produced by the sink organs.     

Dry weight accumulation and starch-synthesis enzymes in grain of cultured ears of three winter wheat varieties

Detached ears of three winter wheat ( Triticum aestivum L.) varieties were cultured in solution for 12 days with sucrose levels varying from 36.5 to 292 m M. The dry weight and starch content of grains increased asymptotically with the sucrose level in the solution. At 4 days of culture, glucose phosphate isomerase (EC 5.3.1.9) activity grain⁻¹ was lower with 36.5 m M than with higher sucrose levels in the medium; at 8 days, adenosinc diphosphoglucose pyrophosphorylase (EC 2.7.7.27) and (soluble plus bound) starch synthase (EC 2.4.1.21) activities grain⁻¹ were higher with 146 and 292 m M sucrose than with 36.5 and 73 m M sucrose. The multiple regression of starch content over these enzyme activities showed that starch synthase was relatively more important as an independent variable. The dry weight and starch content of grains were higher in the variety Maris Huntsman than in Splendeur and Hobbit. The water content of grains was lower in Splendeur than in the other two varieties. At 4 days the glucose phosphate isomerase, adenosine diphosphoglucose pyrophosphorylase and starch synthase activities grain⁻¹ were smaller in Splendeur than in Hobbit and Maris Huntsman and al 8 days they were higher in Maris Huntsman than in Hobbit and Splendeur. The varietal differences in starch content of grains were related to the activities of glucose phosphate isomerase and especially of starch synthase.