Foliar Uptake and Movement of Chlormequat in Cereals: Influence of Dose and Addition of Related Chemicals

Increased amounts of chlormequat applied to leaves of barleyand wheat resulted in a greater proportion being taken up andmoving to the tips of treated leaves. Combined foliar applicationwith certain other amines also increased uptake and movementof chlormequat - choline and glycine betaine were particularlyeffective - but a few others had the reverse effect. Ammoniumand sodium salts of acetic, butyric and hydrochloric acids alsoincreased movement of ¹⁴C-chlormequat Hordeum vulgare L., barley, Triticum aestivum L., wheat, chlormequat, foliar uptake     

Effect of L-Methionine Sulphoximine on the Products of Photosynthesis in Wheat (Triticum aestivum) Leaves

Methionine sulphoximine, an inhibitor of glutamine synthetase,caused ammonia accumulation in detached wheat leaves. The ratewas increased by increased oxygen in the atmosphere and by simultaneouslysupplying glycine or giving extra nitrate; it was decreasedby isonicotinyl hydrazide. Ammonia production was light-dependentand continued at a constant rate in air for at least 2 h. Photosynthesiswas progressively inhibited after the first hour; this inhibitionwas not because of increased stomatal resistance. Leaves suppliedwith 30 mol m^–3 ammonium chloride, without methioninesulphoximine, accumulated more ammonia than leaves treated withthe inhibitor but showed less inhibition of photosynthesis.The inhibitor decreased synthesis of [¹⁴C] amino acids from¹⁴CO₂ in the light but increased the synthesis of [¹⁴C] malateand, relatively, the incorporation of ¹⁴C into sugar phosphates.In the absence of inhibitor, nitrate increased and ammoniumion decreased synthesis of malate. Methionine sulphoximine,by causing a shortage of amino acids, probably inhibited photosynthesisin part by decreasing the recycling of carbon from the photorespiratorycycle back to the Calvin cycle. Key words: Photosynthetic ¹⁴CO₂ assimilation, Methionine sulphoximine, Detached wheat leaves     

Tiller Bud Suppression in Reproductive Plants of Lolium multiflorum Lam. Cv. Westerwoldicum

The control of tiller bud growth during reproductive developmentwas investigated in experimental plants ofLolium multiflorumLam. cv. Westerwoldicum that were reduced to a main axis havinga developing but unemerged ear, elongating stem internodes,a series of expanded leaves, slow-growing tiller buds and aroot system. Isolation of the ear by excision of its base, ordecapitation so as to remove the ear together with the upperleaves, promoted the movement of ¹⁴C-assimilates to tiller buds,decapitation being the more effective treatment. Applicationof 0.1 per cent indol–3yl-acetic acid (IAA) to cut tissuesof decapitated plants diverted ¹⁴C-assimilates to upper internodesbut did not reduce import by buds, whereas application of 1.0per cent IAA both diverted labelled assimilates to upper internodesand reduced bud import. Radioactivity from [¹⁴C] IAA appliedto the upper leaves or to the ear base was recovered from budsin very small amounts; larger amounts were recovered from budsfollowing the application of labelled IAA to an elongating internode,especially from the bud at the base of the treated internode.It is suggested that tiller bud suppression may be influencedby the movement of inhibitory levels of auxin into buds fromnearby elongating stem internodes, whose activity in turn maybe controlled by the developing inflorescence and upper leaves.     

Refixation of respiratory CO2 in the ears of C3 cereals

The spatial arrangement of tissues within the ears of cereals,and gas exchange measurements on intact ears of barley and durumwheat suggest that respiratory CO₂ associated with grain-fillingprocesses, may be refixed close to its site of evolution. Apparentrefixation of respiratory CO₂ in intact ears was compared withthat in flag leaves, by feeding both organs with ¹⁴C-labelledsucrose and trapping ¹⁴CO₂ released by respiration. Apparentrefixation in the ears was twice that measured in flag leafblades of durum wheat genotype Durelle. In ears, the capacityof refixation of respiratory CO₂ at 210 mmol mol^–1 O₂ranged from 55% in barley genotype Roxana to 75% in barley genotypeHatif, and 60% in duwm wheat genotype Bidi 17. A low O₂ concentrationincreased refixation of respiratory CO₂ by up to 90%, 92% and82%, respectively. The occurrence of CO₂ refixation in the field,in a set of 12 barley genotypes grown under irrigated and rainfedMediterranean field conditions, was consistent with observedcarbon isotope ratios (     

The Translocation of Sulphonamides in Higher Plants: III. ACETYLATION AND DEACETYLATION OF SULPHANILAMIDE IN BROAD BEANS AND WHEAT

The acetylation of both amino groups on sulphanilaniide hasbeen demon strated by paper chromatography of sap expressedfrom broad beans treated with suiphanilamide through the roots.Deacetylation of N¹- and N⁴ mides occurred in the leaves oftreated plants and the N⁴ compound was also acetylated to yieldthe N¹: N⁴-diacetylsulphanilamide. About 30 per cent. of the suiphanilamide entering broad-beanroots was acetylated in the roots, the maximum proportion ofthe N⁴-acetyl compound being reached after 7 days. Deacetylationis a more limited process but reaches a steady state after thesame time. Low levels of acetylation and deacetylation wererecorded in wheat. The amounts of suiphanilamide and N⁴ entering and accumulatingin the various tissues were calculated on a water-uptake basisarid these data show that the main site of acetylation is inthe roots and that deacetyla tion occurs to a limited extentin the roots but predominates in stems and leaves.     

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.     

Uptake and distribution of 65Zn and 54Mn in wheat grownat sufficient and deficient levels of Zn and Mn: I. During vegetative growth

The uptake and distribution of ⁶⁵Zn and ⁵⁴Mn by wheat (Triticumaestivum cv. Aroona) was investigated. Plantswere grown in achelate-buffered nutrient solution with either sufficient Znand Mn, low Zn or low Mn. A single representative seminal rootfrom 14-d-old and 42-d-old plants was dual-labelled with ⁶⁵Znand ⁵⁴Mn. The 14-d-old plants were harvested every 10 min from10–140 min of labelling, whilst the 42-d-old plants wereharvested after 2 h of labelling. At harvest, each plant wasseparated into leaves, main stem, unexposedroots, and tillers.In addition, the crown was separatedfrom the stem in the 14-d-oldplants In the control plants labelled at 14 d, ⁶⁵Zn was firstdetectedand accumulated in the crown of the roots after 40–60min. Labelled Zn was then detected in the stem, followed bythe leaves. The oldest and youngest leaves received less ⁶⁵Znthan the second and third oldest leaves. The plants grown underlow Zn conditions accumulated more ⁶⁵Zn in their older leavesand transferred ⁶³Zn to the unexposed roots. Distribution of⁵⁴Mn was similar in the controls to that of ⁶⁵Zn, except theolder leaves received no HMn, At the second harvest, a similardistribution pattern of ⁶⁵Zn and ⁵⁴Mn was observed with regardto leaf age. Large amounts of ⁶⁵Zn and ⁵⁴Mn were detected withinthe unexposed roots of all treatments. It is suggested thatthe distribution of root-supplied Zn and Mn may be determinedby micronutrient status and its relationship with leaf transpirationrates. Key words: Distribution, manganese, vegetative growth, wheat, zinc     

Variation in Concentration of Ribulose-1, 5-bisphosphate Carboxylase in Leaves of Barley, Wheat and Hybrids of Crested Wheatgrasses

Amounts of the enzyme ribulose-1,5-bisphosphate carboxylasewere estimated in seedling leaves of barley (Hordewn vulgareL.) and flag leaves of wheat (Triticum aestitum L.) by radialimmuno diffusion. A fourfold variation among barley varietiesfor amount of RuBPCase at the seedling stage was observed (c.3.5–15mg g^–1 fr. wt). Range in variation for amountof flag leaf RuBPCase among wheat varieties was 6-09-9.39 mgRuBPCase g^–1 fr. wt. F₁ hybrids from interspecific andintergeneric crosses of crested wheatgrasses (Agropyron andElymus spp.) and their amphidiploid analogues were comparedfor amount of RuBPCase in the most recent fully expanded leavesharvested before seed set. Amount of enzyme varied from 3.4to 77.6 mg g^–1 fr. wt among the hybrids. No effect chromosomenumber on enzyme concentration was observed among 13 hybridsand their amphidiploid counterparts. Key words: RuBPCase, wheatgrasses     

Virulence of Septoria nodorum as Affected by Passage Through Detached Leaves of Wheat and Barley Cultivars1

Abstract Three genetically marked, single–spore isolates of Septoria nodorum from wheat were passed through detached leaves of wheat cvs Blueboy and Coker 747 and the barley cv. Boone to produce three sub–isolates per original isolate. Each sub–isolate was cultured for three pycnidiospore generations on its respective host. Virulence of each sub–isolate on detached leaves of Blueboy, Caldwell, Coker 747, and NK81W701 wheat, and Boone and Surry barley was compared with that of the original single–spore isolate from which it was derived. In most cases, sub–isolates passed through wheat were significantly more virulent than the originals on wheat cultivars. They also were more virulent to barley than the original isolates but they were less virulent to barley than to wheat cultivars. Isolate × cultivar interactions were statistically significant (P < .0001) for isolates passed through wheat or barley and were greater than isolate × cultivar interactions among the original isolates. In seven of eight isolates passed through wheat or barley, only the original genetic marker was recovered after three generations, indicating that cross–contamination could not account for the observed change of virulence. In the single case of apparent contamination, of a sub–isolate, virulence declined.     

The Stability and Movement of Gibberellic Acid in Pea Seedlings

The stability and movement of gibberellic acid (GA) in intactdwarf pea seedlings growing in the light was studied by meansof both unlabelled GA and GA labelled with isotopic carbon (¹⁴C).After ¹⁴C-GA had been applied to the mature leaves of pea seedlingsmuch remained in association with the treated leaflets, but¹⁴C-GA was also extractable from the young shoots. The yieldwas approximately the same 5 to 96 hours after treatment. GApenetrated leaf surfaces only while the application solventwas moist (about 1 hour), but moved from the treated leafletsinto the shoots for at least 24 hours. Some hours after treatmentthere was an abrupt increase in the growth-rates of the plants,and crude estimates suggest that an effective dose of GA movedto the elongating tissue at about 5 cm/hr. The pattern of distributionof ¹⁴C was examined by autoradiography. The data suggest thatGA which enters the plant is redistributed from maturing leavesto immature leaves, passing through the elongating tissue, foras long as any of the substance is present. The hypothesis remainstenable that GA produces its growth effects by acting only uponexpanding tissue     

The Redistribution of Assimilate in Field-grown Winter Wheat

The long- and short-term movement of carbon in a field cropof winter wheat was investigated with the radioactive tracercarbon-14. The flag leaves of individual plants assimilateda pulse of ¹⁴CO₂ and the plant parts were assayed subsequently.Groups of plants were pulse-labelled four times during the mainperiod of growth—twice before and twice after anthesis.Plants were harvested and assayed twice weekly after labellingand the time-course of the changes in the amount of ¹⁴C recoveredfrom the leaves, stems and ears was observed for each groupof plants. Concurrently with these long-term studies, otherwheat plants were pulse-labelled, then harvested and assayed24 h later. The partitioning of ¹⁴C between leaves, stem, earand in some cases roots, was measured over the period from thestart of stem elongation to the end of grain filling. Two distinct types of relocation of carbon were observed. Carbonassimilated early in the growth of the plant and used in thegrowth of new leaves was seen to be partly relocated to theear. Carbon assimilated 8 d after anthesis was partly storedin the stem, and 15 d later relocated to the ear. This relocationcorresponded to a decrease in stem dry mass seen in growth analysis.Little other change in the ¹⁴C content of the plants occurred,suggesting that most respiration used current rather than storedassimilate. Key words: Carbon assimilate, redistribution, wheat     

Change of virulence of Septoria nodorurn during passage through barley and wheat

The barley biotype of Septoria nodorum was recovered from isolates of the wheat biotype after 2–6 passages through barley. Shifts in the population of the two biotypes during host passage were determined by isolating single spores after passage and observing changes in symptom expression on both hosts. The number of passages required before the barley biotype was recovered varied with the fungus isolate and barley cultivar. Attempts to recover the wheat biotype from isolates of the barley biotype during several passages through wheat were unsuccessful when 10⁵ to 10⁶ conidia m1^-1 were used as inoculum. However, the wheat biotype was recovered from two isolates of the barley biotype during a single passage when a very high concentration of conidia (>10¹² ml^-1) were used as inoculum. From the results of this study and a previous report by the author, the biotype of S. nodorum on barley which occurs in the southeastern United States appears to be largely restricted to barley.     

Assimilate Uptake and Water Loss in Maturing Barley Grains

When detached ears of barley were supplied with [U-¹⁴C] sucrosefor 4 h, ¹⁴C was detected in the endosperms of grains from earsharvested 50 ‘days’ after anthesis, i.e. 10–15‘days’ after cell wall modifications had taken placein the chalazal region of the caryopsis. In 55-‘day’ears which had been supplied with [U-¹⁴C] sucrose for 20 h no¹⁴C was detected in the upper part of the rachis or in grainswith a moisture content of less than about 25%. In the rachisof wheat and barley, the walls of the xylem parenchyma and phloemcells stain strongly with ruthenium red, and in immature rachidesthis stainability is lost when sections are incubated with pectinase.Near the apex of 55-‘day’ ears the pectinaceouswall material in the vascular bundles is resistant to digestionby pectinase, and pectic substances, which are removed by pectinase,fill the lumina of the xylem elements. These observations arediscussed in relation to the dehydration of the maturing grain. Key words: Barley, ¹⁴C-sucrose, Pectic material, Xylem     

The fate of carbon in pulse-labelled crops of barley and wheat

Wheat (cv. Gutha) and barley (cv. O'Connor) were grown as field crops on a shallow duplex soil (sand over clay) in Western Australia with their root systems contained within pvc columns. At four stages during growth, the shoots were pulse-labelled for 1.5h with¹⁴CO₂; immediately prior to labelling, the soil was isolated from the shoot atmosphere by pvc sheets. After labelling, the soil atmosphere was pumped through NaOH to trap respired CO₂ and after 2.5, 5, 7.5 and 24 h from the start of labelling, columns were destructively sampled to recover¹⁴C from the roots, soil and shoot.Both species showed similar patterns of¹⁴C distribution and changes in distribution through the growing season. During early tillering, 15–25% of the¹⁴C recovered after 24 h had been respired by the roots and rhizosphere, 17–27% was retained in the roots, 0.4–1.8% was recovered as water-soluble¹⁴C in the soil and the remainder (45–67%) was present in the shoot. These percentages changed during growth so that during grain filling only 2–3% of the¹⁴C recovered after 24 h was as respired CO₂, 2–6% was in the roots, 0.2% was in the soil and over 90% was in the shoot.The distribution of¹⁴C in components of the soil-plant system changed during the 24 h after labelling with the most rapid changes occurring generally during the first 7.5 h after labelling.Using growth measurements from adjacent plots, the amounts of C added to the soil were estimated for the whole season. Carbon input to the soil was about 48 gC m^–2 for wheat and 58 gC m^–2 for barley; the crops produced total shoot dry matter of 494 (wheat) and 735 g m^–2 (barley). Of the C input to the soil, 27.8% (wheat) and 40.3% (barley) was as respired C and only 3.3 (wheat) and 4.1% (barley) was collected as exudate (water-soluble material).     

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     

Comparative mapping of HKT genes in wheat, barley, and rice, key determinants of Na+ transport, and salt tolerance

Salt tolerance of plants depends on HKT transporters (High-affinityK⁺ Transporter), which mediate Na⁺-specific transport or Na⁺-K⁺co-transport. Gene sequences closely related to rice HKT geneswere isolated from hexaploid bread wheat (Triticum aestivum)or barley (Hordeum vulgare) for genomic DNA southern hybridizationanalysis. HKT gene sequences were mapped on chromosomal armsof wheat and barley using wheat chromosome substitution linesand barley–wheat chromosome addition lines. In addition,HKT gene members in the wild diploid wheat ancestors, T. monococcum(A^m genome), T. urartu (A^u genome), and Ae. tauschii (D^t genome)were investigated. Variation in copy number for individual HKTgene members was observed between the barley, wheat, and ricegenomes, and between the different wheat genomes. HKT2;1/2-like,HKT2;3/4-like, HKT1;1/2-like, HKT1;3-like, HKT1;4-like, andHKT1;5-like genes were mapped to the wheat–barley chromosomegroups 7, 7, 2, 6, 2, and 4, respectively. Chromosomal regionscontaining HKT genes were syntenic between wheat and rice exceptfor the chromosome regions containing the HKT1;5-like gene.Potential roles of HKT genes in Na⁺ transport in rice, wheat,and barley are discussed. Determination of the chromosome locationsof HKT genes provides a framework for future physiological andgenetic studies investigating the relationships between HKTgenes and salt tolerance in wheat and barley. Key words: Barley, comparative mapping, HKT, rice, salt tolerance, sodium transport, wheat     

The uptake and the transport of14C-labeled epibrassinolide in intact seedlings of cucumber and wheat

The uptake and the transport of exogenously applied epibrassinolide (EBR) in seedlings of cucumber and wheat were examined by autoradiography using¹⁴C-EBR.¹⁴C-EBR was applied to roots, young and mature leaves, and the shoot apex. When applied to roots,¹⁴C-EBR was readily taken up and was swiftly transported throughout both plant species. When¹⁴C-EBR was applied to the adaxial surface of a young cucumber leaf, it was readily taken up, but was very slowly transported. In cucumber leaves,¹⁴C-EBR was transported throughout the treated leaf after 3 days of treatment, and then it was transported to upper leaves from the treated leaf after 7 days. Some 6.3% of applied¹⁴C-EBR was transported to the newly expanded leaves. In wheat leaves,¹⁴C-EBR was transported only in the apical direction from the treated spot after 3 days of treatment, but it was not transported from the treated leaf to the other leaves or organs even after 7 days. Some 1.3% of applied¹⁴C-EBR was transported to the tip area of the treated leaf. These results indicate that exogenous EBR applied to intact plants is acropetally transported.     

Glycolate Pathway and Serine Synthesis in Relation to CO2 Concentration in Tomato Leaves

Isotopic trapping of the carbon flowing through the glycolatepathway by exogenous glycolate, glycine and L-serine was investigatedduring ¹⁴CO₂ photosynthesis at different CO₂ concentrationsin tomato leaves. L-Serine markedly trapped the carbon flowingfrom ¹⁴CO₂. The amounts of ¹⁴C incorporated into serine decreasedat a high CO₂ concentration, but increased with an increasein the CO₂ concentration in the presence of exogenous serineduring 10-min photosynthesis in ¹⁴CO₂. When ¹⁴CO₂ was fed for5 to 40 sec at 1300 ppm CO₂ to tomato leaves which had beengiven L-serine, an increase in the accumulation of ¹⁴C-serinebegan after 20 sec, and the ¹⁴C-serine molecules formed at 20and 40 sec were labeled uniformly. In the presence of exogenousserine during 10-min photosynthesis in 1300 ppm CO₂, isonicotinicacid hydrazide increased the incorporation of ¹⁴CO₂ into glycinewith a corresponding decrease in the accumulation of ¹⁴C-serine,but it did not inhibit serine accumulation completely; an evidencefor that some serine was formed by a pathway other than theglycolate pathway. The effect of the CO₂ concentration on theglycolate pathway is discussed in terms of serine synthesisin the presence of exogenous serine. (Received June 1, 1981; Accepted September 30, 1981)     

A simple method for the isolation of intact mesophyll protoplasts from cereal plants

Intact mesophyll protoplasts from cereal plants were easilyprepared by incubating leaves with the abaxial epidermis peeledoff at 20–25?C for 2–3 hr in 0.6 M mannitol containing1% cellulase at pH 5.6. From one gram (fresh weight) of leaves1.5–6?10⁶ protoplasts, more than 90% of which were morphologicallyintact, could be obtained. Protoplasts isolated from wheat,oat, corn and barley were efficiently infected with brome mosaicvirus (BMV), and supported viral multiplication. (Received June 21, 1977; )     

Photosynthesis by Flag Leaves of Wheat in Relation to Protein, Ribulose Bisphosphate Carboxylase Activity and Nitrogen Supply

The effects of nitrate supply on the composition (cell numbers,protein and chlorophyll contents) of flag leaves of winter wheatgrown with two amounts of N fertilizer and of spring wheat grownin the glasshouse under controlled nitrate supply are describedand related to photosynthesis. Nitrogen deficiency decreasedthe size of leaves, mainly by reducing cell number and, to asmaller extent, by decreasing cell volume. Protein content perunit leaf area, per cell and per unit cell volume was largerwith abundant N. Total soluble protein, ribulose bisphosphatecarboxylase-oxygenase (RuBPc-o) protein and chlorophyll changedin proportion irrespective of nitrogen supply and leaf age.Photosynthesis per unit area of flag leaf and carboxylationefficiency in both winter and spring wheat were proportionalto the amount of total soluble protein up to 7.0 g m^–2and to the amount of RuBPc-o protein up to 4.0 g m^–2.However, photosynthesis did not increase in proportion to theamount of total soluble or RuBPc-o protein above these amounts.In young leaves with a high protein content the measured ratesof photosynthesis were lower than expected from the amount andactivity of RuBPc-o. Carboxylation per unit of RuBPc-o protein,measured in vitro, was slightly greater in N-deficient leavesof winter wheat but not of spring wheat. RuBPc-o activity perunit of RuBPc-o protein was similar in winter and spring wheatleaves and remained approximately constant with age, but increasedin leaves showing advanced senescence. RuBPc-o protein fromN-deficient leaves migrated faster on polyacrylamide gels thanprotein from leaves with high N content. Regulation of the rateof photosynthesis in leaves and chloroplasts with a high proteincontent is discussed. The conductance of the cell to the fluxof CO₂ from intercellular spaces to RuBPc-o active sites iscalculated, from cell surface areas and CO₂ fluxes, to decreasethe CO₂ partial pressure at the active site by less than 0.8Pa at an internal CO₂ partial pressure of 34 Pa. Thus the decreasein partial pressure of CO₂ is insufficient to account for theinefficiency of RuBPc-o in vivo at high protein contents. Otherlimitations to the rate of photosynthesis are considered. Key words: Wheat, photosynthesis, nitrogen, ribulose, bisphosphate carboxylase