Photosynthesis of Ears and Flag Leaves of Wheat and Barley

Immediately after anthesis ears of spring wheat absorbed lessthan 0.5 mg CO₂, per hour in daylight and later evolved CO₂,in the light and in the dark. The rate of apparent photosynthesisof the combined flag-leaf lamina and sheath and peduncle (collectivelycalled flag leaf) of two spring wheat varieties, Atle and JufyI, was 3–4 mg per hour; the rates of the flag leaf andthe ear of two spring barleys, Plumage Archer and Proctor, wereeach about 1 mg per hour. The gas exchange of ears and flag leaves between ear emergenceand maturity accounted for most of the final grain dry weight.The CO₂, fixed by the wheat ear was equivalent to between 17and 30 per cent of the grain weight, but more than this waslost by respiration, so assimilation in the flag leaf was equivalentto 110–20 per cent of the final grain weight. In barley,photosynthesis in the flag leaf and the net CO₂ uptake by theear each provided about half of the carbohydrate in the grain. Barley ears photosynthesized more than wheat ears because oftheir greater surface, and flag leaves of wheat photosynthesizedmore than those of barley because they had more surface anda slightly greater rate of photosynthesis per dm².     

Physiological Causes of Differences in Grain Yield between Varieties of Barley

In a field experiment on barley at Rothamsted with the highmean yield of 49 cwt. of grain per acre, the varieties Proctorand Herta produced 10—15 per cent, more grain than Plumage-Archeron plots that received no nitrogenous fertilizer. When nitrogenwas applied the difference was increased to about 30 per cent.,because the higher nitrogen supply caused the Plumage-Archercrop to lodge and did not increase its yield, while Proctorand Herta remained standing. The three varieties did not differ in leaf-area index nor innet assimilation rate before ear emergence, so that all hadthe same total dry weight. After ear emergence, the leaf-areaindices of Proctor and Plumage-Archer were nearly equal, butthat of Herta was smaller. Assuming that the photosyntheticefficiency of the leaves continued to be the same in all varieties,the higher grain yields of Proctor and Herta cannot be attributedto greater production of dry matter by the leaves, either beforeor after ear emergence. A pot experiment on plants with shadedears confirmed that the dry matter contributed to grain yieldby unit leaf area was nearly equal in all the varieties. The higher grain yield of Proctor and Herta than of Plumage-Archermust therefore have come from additional photosynthesis in partsof the plant other than the leaves, i.e. in the ears themselves.An attempt to demonstrate this directly in a pot experiment,by comparing the grain yields of plants with shaded or withunshaded ears, was unsuccessful because the varieties behaveddifferently in pots; Proctor and Herta produced only about 6per cent, more grain yield than Plumage-Archer, and though thedecrease in grain yield by shading the ears was slightly greaterfor Proctor and Herta, the differences were not significant. The sum of ear sizes (estimated from length and breadth measurements)per m.1 in the field experiment was greater for Proctor andHerta than for Plumage-Archer. Also the distribution of drymatter between developing ears and shoots apparently differedwith variety, so that at ear emergence the dry weight of earsper m.² was greater in the two higher yielding varieties. Theincreased amount of photosynthetic tissue in the ears of Proctorand Herta, as measured by size or weight, may not wholly explaintheir greater dry-matter production; ears of Herta may alsohave a higher photosynthetic efficiency. No differences in nutrient uptake that could account for thevarietal differences in grain yield were found. Plumage-Archerabsorbed more potassium, and Herta less phosphorus than theother varieties. About a quarter of the final content of nitrogen,and a third of the phosphorus, was absorbed after ear emergence,but the potassium content was nearly maximal at ear emergenceand later decreased. The pot experiment showed that, on the average of all varieties,26 per cent. of the dry matter in the grain at harvest originatedfrom photosynthesis in the ears, including 10 per cent, fromthe awns; 59 per cent, came from photosynthesis in the flag-leaflamina and sheath and peduncle, and 15 per cent, from partsof the shoot below the flag leaf.     

Varietal Differences in Photosynthesis of Ears and Leaves of Barley

Rates of apparent photosynthesis of ears and of the combinedflag leaf and sheath and peduncle of Proctor barley grown inpots or in the field were similar to those of Plumage Archer,or slightly smaller when the dimensions of the ear and leafarea of Proctor were less than those of Plumage Archer. Thephotosynthesis rate of the ear—about 1.0 mg. CO₂ per earper hour—was similar or slightly less than the rate ofthe flag leaf and sheath and peduncle. These rates of photosynthesisindicated that 40-50 per cent, of the carbohydrate in the grainwas provided by photosynthesis in the shoot and about 40 percent, by photosynthesis in the ear. The total CO₂ fixed by theear was equivalent to about 60 per cent, of the grain weight,20 per cent, being lost by respiration. Shading the ear underestimatedthe total amount of CO₂ fixed by the ear and decreased dry weightof grain per ear of both Proctor and Plumage Archer by 26 percent., as in pots. The contribution of ear photosynthesis toyield of grain per acre was greater for Proctor than for PlumageArcher because Proctor had more ears. The rate of apparent photosynthesis per dm.² of leaves of Proctorwas similar to that of Plumage Archer both before and afterear emergence. Before ear emergence, the photosynthesis rateof a particular leaf decreased linearly with time and was slowerfor lower than for higher leaves on the shoot. Respiration ratesper g. dry weight of ears of Proctor and Plumage Archer weresimilar; in one experiment the leaves of Proctor respired slightlyfaster than those of Plumage Archer.     

Photosynthesis of Lamina and Sheath of Barley Leaves

Apparent photosynthesis, in mg. CO₂ absorbed per dm.² per hour,of the sheath and enclosed stem of a barley leaf was about 50per cent. of that of the lamina of the same leaf, when the photosynthesizingarea was measured as one side of the lamina and the outer exposedsurface of the sheath. Apparent photosynthesis of a particularlamina or sheath was about 70 per cent. of that of the one aboveon the same stem. Respiration per dm.², though not per g. dry weight, of sheathwith enclosed stem was greater than of lamina in one experimentdone with low-intensity illumination so that true rates of photosynthesisof lamina and sheath were similar. Differences in respirationrates per unit area of laminae and sheaths probably accountedfor most of the greater apparent photosynthesis of the formerin other experiments done with higher intensity illumination. It is suggested that for growth-analysis studies the size ofthe photosynthetic system of cereals should be measured as thatof one side of the leaf laminae plus the outer surface of thecombined leaf sheaths. In the later stages of growth the surfacearea of exposed stem and peduncle should also be included.     

Comparative Growth-rates of Upland and Swamp Rice Varieties

The growth of swamp rice (Dima) and upland rice (Kindinga) wasstudied under tropical glass-house conditions in Trinidad. Kindinga ultimately attained greater dry weight and height thanDima but the total number of leaves and tillers per plant weregreater in Dima than in Kindinga. Kindinga had a greater numberof leaves, on the main shoot, than Dima. Growth curve of leaf-lamina area per plant was similar to thatof leaf-lamina dry weight in both varieties but leaf area perunit dry weight of lamina and per plant were usually greaterin Kindinga than in Dima. Over the whole period, the mean relative growth rate, leaf weightratio were greater in Kindinga than in Dima but net assimilationrate was greater in Dima than in Kindinga. Total nitrogen content was greater in Dima than in Kindingaduring the tillering period but later the reverse was the case.The rate of nitrogen absorption was high in both varieties duringthe tillering and reproductive phases. Split applications ofnitrogenous fertilizer are therefore suggested. The rate ofnitrogen absorption was, however, greater in Dima than in Kindingaduring the tillering period and latter half of the reproductivephase.     

Water and Nitrogen Uptake in Upland and Swamp Rice

The water uptake per unit dry matter and nitrogen uptake perunit water absorbed of swamp rice (Dima) and upland rice (Kindinga)at three nitrogen levels equivalent to 23.6, 70.8, and 118,kg./ha. (20, 60 and 100 lb./acre) respectively were studiedunder tropical green-house conditions in Trinidad. The water uptake per unit dry matter produced was significantlygreater in Kindinga than in Dima at the three nitrogen levels.Additional nitrogen increased the water requirement of Kindingabut decreased that of Dima, the greater increase in Kindingabeing attributed to increase in leaf area. The efficiency of nitrogen absorption in terms of water transpiredwas greater in Dima than in Kindinga at the three nitrogen levels.     

The Production and Distribution of Dry Matter in Maize after Flowering

An experiment in which different groups of leaf laminae wereremoved, or ears shaded, shortly after silking showed that mostof the dry-matter increase after flowering was produced by upperleaves. The top five, the middle four, and the bottom six laminaeaccounted, respectively, for 26 per cent, 42 per cent, and 32per cent of the leaf area duration (D) of the laminae afterflowering; the estimated contributions of the three groups todry-matter production by the laminae after flowering were about40 per cent, 35–50 per cent, and 5–25 per cent,respectively. The sheaths provided about one-fifth of the totalleaf area and probably contributed about one-fifth, and laminaefour-fifths, of the total dry matter produced after flowering.The contribution from photosynthesis by the ear was negligible,presumably because its surface area was only 2 per cent of thatof the leaves. Leaf efficiency (dry matter produced per unitarea) decreased greatly from the top to the base of the shoot.When laminae were removed, the grain received a larger fractionof the dry matter accumulated after flowering, less dry matterremained in the stem, and the photosynthetic efficiency of theremaining leaves was apparently increased. When alternate laminae were removed at the time of silking (half-defoliation)D was decreased by 40 per cent, and the subsequent productionof dry matter decreased nearly proportionately, so that netassimilation rate (E) was not affected but grain dry weightwas decreased by only 32 per cent. At the final harvest, thegrain of half-defoliated plants constituted 80 per cent of thedry matter accumulated after flowering, compared with 65 percent for intact plants. Stem weight decreased from two weeksafter flowering in half-defoliated plants, but remained nearlyconstant in intact plants. When pollination was prevented andno grains formed, E during the first month after flowering wasunaffected; the dry matter that would have passed into the grainaccumulated in the stem and husks, not in the leaves. The decrease in stem weight caused by defoliation suggests thatpreviously stored dry matter was moved to the grain. That suchmovement is possible was shown by keeping prematurely harvestedshoots in the dark for two weeks with their cut ends in water;the dry weight of the grain increased and that of the stem,laminae, husks, and core decreased. Nevertheless, dry-matterproduction after flowering was more than sufficient for graingrowth, and previous photosynthesis probably contributed littleto the grain.     

Contributions to Grain Yield from Pre-anthesis Assimilation in Tall and Dwarf Barley Phenotypes in Two Contrasting Seasons

Twenty-four barley genotypes were produced from crosses betweentwo sets of parents each having a different, non-allelic dwarfinggene. Four genotypes were phenotypically tall, twelve were singledwarf and eight were double dwarf. By a combination of carbon-14labelling and growth measurement the contribution to grain drymatter from assimilation before anthesis was estimated for eachgenotype in two years, 1976 and 1977. Pre-anthesis assimilation was estimated to have contributed44 per cent of grain dry matter in 1976, a very dry, hot year,but only 11 per cent in 1977, a wetter, cooler year. These percentageswere equivalent to about 133 and 74 g m^–2 in 1976 and1977. It is shown that some previous estimates of the contributionfrom pre-anthesis assimilation are too low for crops of barleygrown in the field in dry years, while in wet years, the contributionis unlikely to be less than 10 per cent. In neither year did the genotypes differ in the extent to whichpre-anthesis assimilation contributed to grain filling. In bothyears, grain yields of the tall genotypes were 8–9 percent greater than those of the double dwarf ones, and stem weightsat maturity 30 per cent greater. Hordeum vulgare L., barley, grain yield, carbon assimilation     

不同水分管理下优质稻花后植株碳氮流转与籽粒生长及品质的相关性

以桂华占、八桂香为材料,在干湿交替灌溉、亏缺灌溉、淹水灌溉3种水分条件下,研究优质稻花后植株碳氮流转与籽粒生长及品质的相关性。结果表明:不同水分管理下,桂华占和八桂香花后碳氮流转与籽粒的生长间存在密切相关。主要表现在:(1)茎鞘和叶片干物质转运对籽粒干物质积累的贡献率为16.86%～25.68%,花后茎叶干物质运转速度和运转率与籽粒起始灌浆势呈显著甚至极显著正相关;籽粒最大灌浆速率、活跃灌浆期、持续灌浆时间与叶片干物质运转速度和运转率呈极显著正相关,与茎鞘干物质运转速度和运转率呈极显著负相关;(2)茎鞘碳同化物转运对籽粒的产量和淀粉产量的贡献率则为干湿交替灌溉>亏缺灌溉>淹水灌溉;但叶片碳同化物转运对籽粒的产量和淀粉产量的贡献率则为淹水灌溉>亏缺灌溉>干湿交替灌溉;茎叶可溶性糖积累量的减少和籽粒直链淀粉含量和积累量增加是同步的,且茎叶可溶性糖积累量快速递减期(花后3～12d)与直链淀粉含量和积累量快速递增期(花后6～12d)同步;(3)茎鞘和叶片氮素转运对籽粒氮素积累的贡献率为44.05%～117.66%,叶片总氮转运对籽粒氮素积累的贡献率大于茎鞘,茎鞘和叶片氮同化物对籽粒氮素的贡献率以淹水灌溉处理的最大,亏缺灌溉处理的次之,干湿交替灌溉处理的最小。     

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.     

籼稻桂华占、八桂香花后营养器官干物质流转与籽粒灌浆关系

以桂华占和八桂香2个籼稻品种为材料,研究籼稻花后不同部位器官物质积累、运转与籽粒生长的动态特征及相互关系。结果表明:(1)叶、叶鞘、节间干物质流转存在一定差异,倒2叶鞘对籽粒的贡献超过倒3叶鞘和倒1叶鞘,倒3节间对籽粒的贡献超过倒2节间和倒1节间;(2)不同部位籽粒的灌浆速率和拐点粒重呈现UPG(上部籽粒)MPG(中部籽粒)BPG(下部籽粒)变化趋势,拐点时间和活跃灌浆时间及持续灌浆时间均呈现BPGMPGUPG变化规律,UPG启动早,灌浆速率大,BPG的灌浆速率小,灌浆时间滞后,籽粒粒重呈现UPGMPGBPG;(3)叶片、叶鞘及节间干物质运转速度和运转率都与籽粒起始灌浆势呈正相关,其中器官间与起始灌浆势的相关系数大小表现为节间叶鞘叶片,不同叶位间与起始灌浆势的相关系数大小表现为倒2叶倒3叶倒1叶,其中节间干物质运转对籽粒生长的作用大于叶鞘,叶片干物质运转与籽粒生长的相关性最小。倒1节鞘物质输出与BPG生长时间上同步,倒2节鞘与MPG生长同步,倒3节鞘与UPG生长同步。     

Starch storage in the stems of wheat plants: localization and temporal changes

Background and Aims

Carbohydrate temporarily accumulates in wheat stems during the early reproductive growth phase, predominantly as water soluble carbohydrate (WSC), and is subsequently remobilized during grain filling. Starch has also been reported as a minor storage carbohydrate component in wheat stems, but the details are lacking.

Methods

The accumulation and localization of starch in wheat stem and leaf sheath tissue over a developmental period from 6 d before anthesis to 35 d after anthesis was investigated.

Key Results

The region of the peduncle enclosed by the flag-leaf sheath, and the penultimate internode were the main tissues identified as containing starch, in which the starch grains localized to the storage parenchyma cells. In contrast, the exposed peduncle lacked starch grains. Starch grains were also found in the flag-leaf and second-leaf sheath. Plants grown in low-nitrogen conditions exhibited increased storage of both starch and WSC compared with plants grown in high-nitrogen supply.

Conclusions

The major accumulation and decrease of starch occurred temporally independently to that for WSC, suggesting a different functional role for starch in wheat stems. Starch reutilization concomitant with peduncle growth, and the early development of the reproductive structures, suggested a role in provision of energy and/or carbon scaffolds for these growth processes.Key words: Carbohydrate partitioning, peduncle, starch, wheat stem, storage parenchyma, Triticum aestivum     

Bioregulation of carbohydrate metabolism in relation to source-sink operation during grain-filling phase of growth in wheat

Mobilization of free sugars from vegetative tissues to grain and their transformation to starch in relation to activities of some relevant enzymes during growth and development were investigated in wheat (Triticum aestivum L.). Vegetative tissues, viz. flag-leaf, flag-leaf sheath, nodes and internodes contained high concentration of free sugars from 70 DAS to 18 DPA and that was in the order of accumulation--flag-leaf sheath> flag-leaf and internodes > nodes. In these tissues, major portion of 14C appeared in endogenous sucrose, irrespective of the nature of (U-14C]-sugars supplied. In photosynthetic structures above flag-leaf node, namely peduncle, rachis and bracts, the free sugar make-up was maximum at anthesis (90 DAS). Activity of soluble acid invertase (EC 3.2.1.26) was high in these tissues during early stages of grain growth but reverse was true for soluble neutral invertase (EC 3.2.1.27) activity. In apical and basal portions of grain, free sugars were more or less similarly distributed in concentration. Linear and rapid accumulation of starch in endosperm paralleled with a decline in accumulation of this polymer in pericarp-aleurone. In the latter tissue, the activities of starch hydrolyzing enzymes, i.e alpha- and beta-amylases (3.2.1.1 and 3.2.1.2) were high during initial stages of grain growth. During active grain-filling, alkaline inorganic pyrophosphatase (EC 3.6.1.1) seemed to play a vital role during starch accumulation in endosperm, whereas the involvement of 3-PGA phosphatase (EC 3.1.3.38) was almost confined to pericarp-aleurone. Impairement of ear head photosynthesis by shading depressed starch synthesis (approximately 50%) indicating, thereby, the significant role of current photosynthates during grain-filling. The results suggested that grain growth in wheat was influenced by an efficient operation of source as well as regulatory factors, including enzymes, constituting intrinsic potential of grain sink.     

Genotypic variation in partitioning of dry matter and manganese between source and sink organs of rice under manganese stress

Key message

Genetic variability in dry matter and manganese partitioning between source and sink organs was the key mechanism for Mn efficient rice genotypes to cope with Mn stress.

Abstract

Considerable differences exist among cereal genotypes to cope manganese (Mn) deficiency, but the underlying mechanisms are poorly understood. Minimal information regarding partitioning and/or remobilization of dry matter and Mn between source and sink organs exists in rice genotypes differing in Mn efficiency. The present study was aimed to assess the growth dynamics in terms of dry matter and Mn remobilization in the whole plant (leaves and tillers as source and panicles and grains as sink) during the grain development in diverse rice genotypes. The efficient genotypes accumulated higher dry matter than inefficient genotypes under low Mn level. The translocation index i.e., uptake in grain/total uptake was 0.11 in efficient genotype (PR 116) and 0.04 in inefficient genotypes (PR 111). The efficient genotype had higher grain Mn utilization efficiency of 0.71 in comparison to 0.48 of inefficient genotype indicating that in efficient genotype, Mn in grain produces more dry matter than inefficient genotypes. The efficient genotypes also had higher flag leaf area and nitrate reductase activity. The source of efficient genotypes contributed to a greater extent to developing sink but further mobilization to grain was hindered by panicle. The panicle of inefficient genotypes had higher per cent of Mn uptake than efficient genotypes indicating that Mn was least mobilized from panicle to grain in inefficient genotypes. The lower per cent uptake of Mn in efficient genotypes indicated that Mn was mobilized from panicle to developing grain and this led to higher Mn translocation index in grain of efficient genotypes. The uptake partitioning revealed that source of all genotypes mobilized the Mn towards the sink to almost same extent but it was the panicle where highest per cent uptake per plant was in inefficient genotypes and lowest in efficient genotypes. The lowest per cent uptake in panicle of efficient genotypes revealed that it supported developing grain to have highest translocation index.     

The Developmental Anatomy of Leaves in Lolium temulentum

Measurements of the epidermal cells in Lolium temulentum andassessments of cell number indicate that differences in bladewidth are due mostly to variations in cell number, whereas changesin blade and sheath length result mainly from differences incell length. A marked increase in length of the flag-leaf aheath,however, was related to an increase in cell number. Considerable changes in the relative proportions of leaf bladeand sheath were observed in the flag leaf, and the leaves immediatelypreceding it, associated with inflorescence initiation. As oneconsequence of this the area of the flag leaf, the largest onthe shoot, is virtually constant under different environmentalcondi-tions. It is suggested that this aspect of correlateddevelopment is related to the nutrition of the panicle, sincein annual grasses such as the cereals the flag leaf may be responsiblefor producing up to 30 per cent. of the starch in the grain.     

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.     

Experiments with CCC on wheat: effects of spacing, nitrogen and irrigation

In experiments with spring and winter wheat at Rothamsted and Woburn during 4 years CCC increased yield at close spacing (4 in) (10 cm) more than at usual spacing (8 in) (20 cm), but there was no interaction between spacing and yield. Some experiments tested up to 2·4 cwt/acre (300 kg/ha) N to see whether yields continued to increase with more than usual amounts of N, when CCC prevented lodging. There was no evidence of this. When a short dry spell occurred at ear emergence, yield of spring wheat was increased by 6 cwt/acre (750 kg/ha) by CCC and 10 cwt (1250 kg) by irrigation. CCC probably improves yield in these conditions because the larger root system it causes enables more ear-bearing shoots to survive. CCC increases yield in two ways, either by increasing ears or grain per ear. In an unlodged crop CCC usually makes the grains smaller, but by preventing lodging it can also increase size. Usually CCC decreases the leaf area per shoot. The flag leaf may be smaller, unchanged or larger than those of untreated plants. There was no obvious connexion between flag-leaf area and grain yield; when CCC decreased flag-leaf area duration by 25 %, grain yield was unchanged. The results suggest that using CCC gives a more than even chance of a profitable yield increase.     

Ontogenetic changes in photosynthetic capacity and dry matter production of flag wheat leaves during the grain filling period

The relationships between photosynthetic capacity and dry matter accumulation during the grain filling period have been studied in flag leaves of Triticum aestivum L., cv. Kolibri grown in Mediterranean field conditions. Particular importance has been given to assimilate accumulation in relation to the onset of senescence. During grain filling, the time course of specific dry weight (SDW) was similar in the blade and in the sheath. Variations in SDW were about six times larger in the sheath than in the blade. Minimum blade SDW values occurred during heading and at anthesis. Maximum blade SDW values were observed two weeks after anthesis. After this, SDW values decreased sharply. The dry matter increase per grain in the period from two weeks after anthesis to the end, was only about 25% of final grain dry weight. The importance of environmental constraints on maximum SDW values are discussed. Maximum SDW values occurred at the beginning of the period of rapid decline in blade net CO₂ assimilation rate and leaf nitrogen content, that is, at the beginning of senescence. On the other hand, the stomatal resistance to CO₂ and the development of senescence are not apparently related. The maximum blade dry weight increase (considering a value of zero at heading) was about 60 mg dry weight per g fresh weight. The possible relationships between dry matter accumulation and senescence onset are discussed.     

Effects of Age and Environment on Net Assimilation Rate of Barley

To distinguish between the effects of age and environment onnet assimilation rate (E) of barley (var. Brant) grown in theopen in pots at Ottawa, E was measured outdoors (treatment N)and on similar plants transferred to a constant environmentfor the 2 weeks during which E was determined (treatment T).During June and July, E of treatment N decreased by 77 per cent.;for treatment T this fall increased to 90 per cent, of the initialvalue. E and average day and night temperatures were greaterin the constant environment than outdoors in early June, andin July they were greater outdoors. Incident light energy wasalways greater outdoors. Thus, the fall in E with age was partiallymasked outdoors by an increase caused by the improvement inenvironmental conditions during the experiment. Plants grown continuously in the constant environment had lowerE, greater leaf area and dry weight, and ears emerged earlierthan plants of similar age that had been transferred for 2 weeksto the constant environment from outdoors. E of plants growncontinuously in the constant environment decreased with time,both when the pots were moved away from the light panel to maintaina constant light intensity at the base of the youngest leafand when the pots remained stationary so that the plants grewtowards the lights. After ears emerged, E for treatments N and T was measured onplants whose ears were shaded to prevent them photosynthesizing.Photosynthesis in the ear accounted for 19 per cent, of thefinal ear dry weight during the first 2 weeks after emergence,12 per cent, during the 3rd and 4th, and 3 per cent, duringthe 5th and 6th weeks. Shading had no effect on weight of plantparts other than ears and had similar effects outdoors and inthe constant environment.