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.     

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.     

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     

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     

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.     

Distribution between parts of the main shoot and the tillers of photosynthate produced before and after anthesis in the top three leaves of main shoots of Hobbit and Maris Huntsman winter wheat

The top three leaves of main shoots in crops of Hobbit and Maris Huntsman winter wheat were exposed to ¹⁴CO₂ at 22 and 16 days before and at 10 days after anthesis in 1978. The distribution of the ¹⁴C recovered in whole plants at anthesis and at maturity was measured. There was negligible loss of ¹⁴C between these two times, but some redistribution. The percentage in the tillers was negligible when the flag leaf (leaf 1) was exposed to ¹⁴CO₂, and otherwise less than 12% except for ¹⁴C absorbed by the third leaf at 16 days before anthesis, when it averaged 26% but was very variable. When ¹⁴C was supplied before anthesis, about 20% reached the grain whichever leaf had been supplied. The ear structures contained about 10% of that absorbed by the third leaf and 35% of that absorbed by the flag or second leaf. When ¹⁴C was supplied after anthesis, the amounts reaching the grain from the different leaves were: flag 82%, second 68%, third 56%. Most of the remainder was in the stem. The exposed leaf never retained more than 6%. The amount of ¹⁴C that moved from the stem to the grain between anthesis and maturity was about 50% greater in the semi-dwarf variety Hobbit than in Maris Huntsman. There was no significant varietal difference in the percentage of post-anthesis ¹⁴C reaching the grain. The ear structures of Hobbit contained about a third more ¹⁴C than those of Maris Huntsman. An additional 90 kg N ha^-1, which increased grain yield by 46%, had negligible effects on the distribution of ¹⁴C.     

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     

Physiological Factors Limiting Grain Size in Wheat

The effects on grain size of changing the supply of assimilates,by thinning before anthesis or by shading the plants or by halvingthe ears either early or late in grain growth, were studiedin two glasshouse experiments with Kleiber spring wheat (Triticumaestivum L.), in 1976 and 1977. Late treatments had no effect,presumably because little grain growth occurred thereafter.Thinning the plants before anthesis increased, and shading theplants soon after anthesis decreased grain size. Halving theears soon after anthesis increased the size of the remaininggrains, but grain weight per ear decreased. The effect on grainsize of halving the ear tended to be smaller under conditionsmore favourable for photosynthesis, except when the plants werethinned before anthesis. Shading decreased the total amountof nitrogen per culm and the proportion of total nitrogen recoveredin the ear. Halving increased the retention of nitrogen in thestem of unshaded shoots and had no effect on nitrogen distributionwithin shaded shoots. In 1977 halving the ear increased the rate of dry matter accumulationin the grain throughout the grain filling period, but in 1976the increase in dry weight was faster in the grains of halvedears only during early grain growth. Later the grains in halvedand intact ears increased in dry weight at the same rate, eventhough the supply of photosynthate and the capacity of the grains(as measured by volume) were greater in the halved ears. Theseresults are discussed in relation to the influence on finalgrain weight of assimilate supply and the storage capacity ofthe grain.     

The tolerance of difenzoquat and diclofop-methyl by winter wheat varieties at different growth stages

Winter wheat varieties were treated with the wild-oat herbicides difenzoquat and diclofop-methyl at several growth stages in a series of four field experiments. High rates of difenzoquat damaged the winter wheat varieties Bouquet, Sportsman, Hobbit and Maris Huntsman but not Hustler. Treatment of the sensitive varieties during tillering stimulated excessive tiller production, resulting in more ears with fewer and smaller grains than on the unsprayed controls; later applications reduced the number of grains and severely depressed yield. In contrast, the early applications of diclofop-methyl, during tillering, were more damaging than the later ones. Bouquet was more tolerant of diclofop-methyl than were the other varieties tested.     

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.     

Incidence and effects of Septoria nodorum on wheat cultivars

Wheat cultivars were grown in two field trials in which half the plots were artificially inoculated with Septoria nodorum. There were differences between cultivars in the degree of 5. nodorum infection on flag leaves and on ears. Hybrid 46, Maris Huntsman and Maris Widgeon were among those with the least infection, and Maris Ranger and Maris Templar among those with the most. Severity of infection was positively correlated with both short stature and earliness of maturity. Infection on leaves was positively correlated with infection on ears. Inoculation reduced yield by decreasing two of its components, grain number per ear and 1000-grain weight. A very early maturing cultivar, Sterling, sustained no yield loss despite severe infection; for the remaining cultivars, yield loss was positively correlated with degree of infection.     

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.     

Growth, Development, and Yield of Spring Wheat in Artificial Climates

Spring wheat was grown to maturity in three growth rooms providing:(a) 18 h of light at 20° C and 6 h of darkness at 15°C (hot long days, HL); (b) 18 h of light at 15° C and 6h of darkness at 15° C (cold long days, CL); (c) 14 h lightat 20° C and 10 h of darkness at 15° C (hot short days,HS). Plants were moved between environments at spikelet initiationand anthesis, so dividing the growth period into three. Meanlengths in days of these periods in the different environmentswere: Period 1: HL 16, CL 18, HS 25; Period 2: HL 42, CL andHS 61; Period3: HL 53, CL 83, HS 63. The length of periods 2and 3 also depended on previous treatments. Grain dry weight was affected by environmental differences inall periods and effects in successive periods were additive.Compared with HL, CL or HS in period I before initiation increasedgrain yield by 6 per cent by increasing grain number per ear,HS in period 2 between initiation and anthesis decreased itby 24 per cent by decreasing the number of grains per spikeletand the proportion of spikelets that contained grain; CL inperiod 2 increased it by 21 per cent by increasing the numberof ears; CL in period 3 after anthesis increased it by 16 percent because leaves died later; HS in period 3 decreased itby 14 per cent because there was less radiation and hence lessphotosynthesis. Dry weight of shoot and root at maturity wasincreased by CL or HS in periods 1 or 2, and increased by CLand decreased by HS in period 3. The effects on final yieldof treatment during periods 1 and 2 were the consequence ofsimilar effects already produced at anthesis, and shoot androot dry weight changed little during period 3. The effects of environmental differences on grain dry weightcould not be explained by differences in leaf-area durationafter anthesis (D₃), except that CL in period 3 increased bothyield and D₃ but not proportionately, so that, as with HS inthe same period, grain: leaf ratio was decreased. Environmentaldifferences in periods 1 and 2 appeared to affect grain weightby altering the capacity of the ear to accumulate carbohydrates,determined by the number of grains per ear, rather than by alteringthe supply of carbohydrates, determined by D₃. There were some interactions between environments in differentperiods which were usually small compared with the main effects.     

Root:shoot ratios of old and modern,tall and semi-dwarf wheats in a mediterranean environment

A field study tested the hypothesis that modern wheat varieties invest a lesser proportion of the total dry matter (root plus shoot) in the root system compared to old varieties. The study was carried out on a duplex soil (sand over clay) at Merredin, Western Australia in a Mediterranean type environment. We also compared the root:shoot dry matter ratios of near-isogenic lines for Rht dwarfing genes.Root:shoot ratios decreased with crop growth stage and were closely related to the developmental pattern of a variety. All varieties appeared to accumulate more dry matter into shoots after the terminal spikelet stage. For the modern variety Kulin this occurred as early as 55 days after sowing (DAS), but did not occur until 90 DAS in the old variety Purple Straw. For all varieties, root dry matter reached its maximum at anthesis, while shoot dry matter continued to increase till maturity. At anthesis there were no significant differences in shoot dry matter between varieties, but from Purple Straw to Kulin root dry matter and thus root:shoot ratio decreased.The tall and dwarf isogenic lines had similar developmental and root:shoot dry matter accumulation patterns.At anthesis, the old variety Purple Straw had significantly higher root dry matter and root length density in the top 40-cm of the profile than modern variety Kulin. There were no varietal differences in rooting depth, water extraction or water use. At maturity about 30% of the total dry matter was invested in the roots among wheat varieties. Grain yield, harvest index (HI) and water use efficiency of grain (WUE_gr) increased from old to modern varieties.The reduced investment of dry matter in the root system and thus the lower root:shoot ratio from early in the growing season may partly explain the increased HI and WUE_gr of modern compared to old varieties.     

Survival of Tillers and Distribution of Dry Matter between Ear and Shoot of Barley Varieties

Plants of Plumage Archer barley grown in pots produced moreshoots than did Proctor, but had fewer ears at maturity, becausesome shoots died about 7 days before ear emergence. The numberand position on the plant of the shoots that died were consistentfor particular growing conditions. Shoots that died were notalways the last to be produced. There were no consistent differencesin dry weight, leaf area, nitrogen content, or apex developmentbetween shoots that did and did not survive; nor did removingthree developing ears on older shoots affect the survival ofother shoots. Growth of Plumage Archer plants was not checkedby death of shoots; dry matter, nitrogen, and leaf area wereredistributed so that increase in surviving shoots compensatedfor losses in the dead shoots. The dry-weight ratio of ear to shoot was smaller for Proctorthan for Plumage Archer at ear emergence and anthesis, but atmaturity it was greater for Proctor, both when ears were shadedto prevent them photosynthesizing and when they were unshaded.The varietal difference in ear: shoot dry-weight ratio was reversedbecause the relative growth-rate of ears of Proctor was greaterand because more dry matter was lost from its shoots. Both changesmay have been caused by translocation to the ear of a greaterproportion of the assimilate from the shoots of Proctor thanof Plumage Archer. Leaves of Proctor appeared to be more efficientin producing dry matter for the ear than those of Plumage Archer.     

Studies on the Germination of Cereals: I. The Germination of Wheat Grains in the Ear during Development, Ripening, and After-ripening

The ability of individual grains to germinate in the ears ofa red and a white wheat variety has been determined at differentperiods after anthesis, and at different moisture contents,before the stage of full maturity. No grains germinated while active growth was taking place, butafter desiccation during ripening, 88·5 per cent. ofthe white grains and 7 per cent. of the red grains were ableto germinate in the ear; the percentage germination of the redgrains increased to 83 per cent., when further desiccation occurredduring the first 5 weeks of after-ripening, but some grainsin the basal spikelets of the ears of both varieties failedto germinate until they had been subjected to the same desiccationfrom 13 to 23 weeks after anthesis. The ability of the grains to germinate has been correlated withtheir desiccation at different stages during maturation, andthe effect of certain factors, which inhibit the germinationof immature grains, are discussed in relation to varietal differencesin the colour of the grains and their position in the ear.     

IAA level and dry matter accumulation at different positions within a wheat ear

Dry matter accumulation and IAA level of grains in various positions within the wheat ( Triticum aestivum L. cvs Solo and Kolibri) ear were investigated during the grain filling period. Main shoot ears were partitioned into basal, central and apical regions and further into proximal and distal grains. Results from a two-year experiment with two wheat cultivars of different grain size are presented. The IAA level increased rapidly from about 8 until 20 to 30 days after anthesis, and then decreased dependent upon year and cultivar. The differences in IAA levels, both among spikelets in different regions of the ear (i.e. basal, central and apical) and also among florets within a spikelet (i.e. proximal and distal), were positively correlated with the differences in dry matter accumulation. Although the correlations held true within cultivars in a particular year, differences in dry matter between cvs Solo and Kolibri or differences from year to year were not well correlated with the corresponding changes in IAA levels. Possible interactions between dry matter accumulation and IAA levels are briefly discussed.     

Changes in growth-substance contents during growth of wheat grains

Samples of developing wheat grains were extracted at weekly intervals from ear emergence until maturity and the growth substances estimated by bioassay. Immature grains contained two cytokinins; one was similar to zeatin and another, with more cytokinin activity, had different properties. Ovules contained only small amounts of growth substances but at the end of anthesis the grains had a maximum content of cytokinin. The gibberellin content increased until 3 wk after anthesis then decreased; their auxin content increased until 4 wk after anthesis but decreased as the grains ripened and lost fresh weight. The husks contained smaller amounts of growth substances than the grains they surrounded. Exudates from young stems contained cytokinins and these may originate in the roots and move to the ears through the stems. The auxin in the grains was identified as indole-3yl-acetic acid and may be derived from the phenols present reacting with tryptophan.     

Effect of variation in ear temperature on gibberellin content of wheat ears

When the ears of wheat were grown after anthesis at temperatures different from the rest of the plant the gibberellin content of the ears increased more rapidly at higher ear temperatures, whilst the plant temperature had little effect. This greater rate of gibberellin accumulation was associated with a greater rate of ear dry weight increase. Wheat ears detached from the plant increased their gibberellin content over 4 days when provided with sucrose and ewfkaurene.     

The Effect on Wheat Grain Growth of the Removal or ABA Treatment of Glumes and Lemmas

When the glumes and lemmas were removed from wheat spikeletsduring the main grain filling period the uptake of sucrose intothe subtended grains was reduced in a few hours, but the abilityto synthesize starch was not affected for several days, by whichtime a marked loss of water content had occurred. When the deglumedears were covered to maintain high humidity, the grains hada weight and water content similar to those from unprotecteddeglumed ears. The application of ABA to the glumes occasionallyhad a small short-term effect on the uptake of sucrose intothe grain, but there were no long-term effects on dry matteraccumulation despite increased glume senescence. When the glume and lemma were removed shortly after anthesisthe subtended grain grew much less than grains from intact florets,with fewer and smaller aleurone cells. The removal of the lemmafrom the third floret of the spikelet had a similar effect onthe third grain, which could be partly reversed by removal ofthe grains in the lower florets. The role of the glumes andlemmas in grain development is discussed in the light of theseresults.