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 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.     

Persistence of photosynthetic components and photochemical efficiency in ears of water-stressed wheat (Triticum aestivum)

Ear photosynthesis may be an important source of C for grain growth in water-stressed plants of cereals. The main objectives of this work were to determine the stability of the photosynthetic apparatus and the photochemical efficiency of ears in plants subjected to post-anthesis drought. Plants of wheat ( Triticum aestivum L. cv. Granero INTA) were grown in pots under a rain shelter and subjected to water stress (soil water potential around −0.6 to −0.8 MPa) starting 4  days after anthesis. Post-anthesis drought substantially accelerated the loss of chlorophyll, Rubisco and the light-harvesting complex of photosystem II (LHCII) in the flag leaf, but the degradation of these photosynthetic components was much less affected by water deficit in awns and ear bracts. Quantum yield of PSII (Φ_PSII) decreased in leaves of water-stressed plants. In contrast, ear bracts had a higher Φ_PSII than leaves, and Φ_PSII of ear bracts did not decrease at all in response to drought. Removing the grains immediately before fluorescence measurements (less than 30 min) slightly reduced Φ_PSII, indicating that CO₂ supplied by grain respiration may contribute to the high photochemical efficiency of ears in droughted plants. However, other factors may be involved in maintaining high Φ_PSII, since even in the absence of grains Φ_PSII remained much higher in ear bracts than in the flag leaf. The relative stability of ear photosynthetic components and their relatively high photochemical efficiency may help to maintain ear photosynthesis during the grain filling period in droughted plants.     

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.     

Senescence-induced ectopic expression of the A. tumefaciens ipt gene in wheat delays leaf senescence, increases cytokinin content, nitrate influx, and nitrate reductase activity, but does not affect grain yield

The manipulation of cytokinin levels by senescence-regulated expression of the Agrobacterium tumefaciens ipt gene through its control by the Arabidopsis SAG12 (senescence-associated gene 12) promoter is an efficient tool for the prolongation of leaf photosynthetic activity which potentially can affect plant productivity. In the present study, the efficiency of this approach was tested on wheat (Triticum aestivum L.)-a monocarpic plant characterized by a fast switch from vegetative to reproductive growth, and rapid translocation of metabolites from leaves to developing grains after anthesis. When compared with the wild-type (WT) control plants, the SAG12::ipt wheat plants exhibited delayed chlorophyll degradation only when grown under limited nitrogen (N) supply. Ten days after anthesis the content of chlorophyll and bioactive cytokinins of the first (flag) leaf of the transgenic plants was 32% and 65% higher, respectively, than that of the control. There was a progressive increase in nitrate influx and nitrate reductase activity. However, the SAG12::ipt and the WT plants did not show differences in yield-related parameters including number of grains and grain weight. These results suggest that the delay of leaf senescence in wheat also delays the translocation of metabolites from leaves to developing grains, as indicated by higher accumulation of ((15)N-labelled) N in spikes of control compared with transgenic plants prior to anthesis. This delay interferes with the wheat reproductive strategy that is based on a fast programmed translocation of metabolites from the senescing leaves to the reproductive sinks shortly after anthesis.     

Ear photosynthesis, carbon isotope discrimination and the contribution of respiratory CO2 to differences in grain mass in durum wheat

The role of ear photosynthesis in grain filling was studied in a number of durum wheat (Triticum turgidum var durum L.) landraces and varieties from the Middle East, North Africa, and from the collections of ‘Institut National de la Recherche Agronomique’ (INRA, France) and ‘Centro International de Mejora de Maiz y Trigo’ (CIMMYT, Mexico). Plants were grown in the field in a Mediterranean climate. Flag leaves (blade plus sheath) and ears were kept in the dark from 1 week after anthesis to maturity which reduced grain weight by 22.4% and 59.0%, respectively. In a further experiment, the carbon isotope discrimination ratio (Δ) of ear bracts, awns and flag leaves was measured on samples taken at anthesis and on mature kernels. The mean value of Δ for the water soluble fraction of bracts (17.0‰) and awns (17.7‰) were lower than those of leaves (19.5‰) and fairly similar to those of kernels (17.4‰) averaged across all genotypes. Data indicate that most of the photosynthates in the grain come from ear parts and not from flag leaves. In addition, a higher water use efficiency (WUE) of ear parts than of the flag leaf is suggested by their lower Δ values. Gas exchange in ears and flag leaves was measured during grain filling. Averaged over all genotypes, CO₂ diffusive conductance was about five times higher in the flag leaf than in the spike (with distal portions of awns outside the photosynthetic chamber) 2 weeks after anthesis. In absolute terms, the dark respiration rate (R_d) was greater than the net photosynthesis rate (P_n) by a factor of 1.74 in the spike, whereas R_d was much smaller, only 22.1, 65.7 and 24.8% of P_n in blade, sheath and awns, respectively. Data indicate that photosynthesis, and hence the water use efficiency (photosynthesis/transpiration), is greatly underestimated in ears because of the high rates of respiration which diminish the measured rates of net CO₂ exchange. Results of ¹³C discrimination and gas exchange show that genotypes from North Africa have higher WUE than those from the Middle East. The high R_d values of ears as well as their low diffusive conductance suggest that CO₂ from respiration may be used as source of carbon for ear photosynthesis. In the same way, the anatomy of glumes, for example, supports the role of bracts using internal CO₂ as source of photosynthesis. In the first experiment, the Δ in mature grains from culms with darkened ears compared with control culms provided further evidence in support of this hypothesis. Thus, the Δ from kernels of control plants was 0.40 higher than that from ear-darkened plants, probably because of some degree of refixation (recycling) of respired CO₂ in the grains.     

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.     

The analysis of grain yield of wheat in terms of photosynthetic ability and efficiency of translocation

A technique, using ¹⁴CO₂, for measuring the rate of photosynthesis and the distribution of synthesized carbohydrate in the same plant, applied to wheat plants at intervals from 10 days before anthesis until the plants were no longer green, showed that the rate of photosynthesis by the leaves and ears decreased steadily; it was much less for ears than for leaves. The proportion of carbohydrate translocated to the grain was very small at and before anthesis, but increased rapidly afterwards. Integration of these data provided estimates of yield based on physiological components which showed good agreement with measured yields at harvest, though varietal differences in observed yield could not be explained. An experiment in which ears were removed from plants 7 days after anthesis showed that photosynthetic activity was not limited by the size of the ‘sink’ to which photosynthates were translocated.     

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     

Dry Matter Partitioning between Plant Organs in Wheat Cultivars Differing in Productivity and Drought Resistance

In microplot field experiments conducted over the course of 12 years, the accumulation of dry matter was recorded in the leaves, stems, and ears of the primary shoots of plants of four cultivars of spring wheat (Triticum aestivumL.) differing in productivity and drought resistance. The contribution of organs to the accumulation of dry matter by the shoot from emergence to anthesis was calculated, and relative changes in the weight of the ear after anthesis were assessed. In all the cultivars, the contribution of the leaves was the same; however, the share of the ear was greater in short-stem cultivars of the intensive type in which the leaves were more active during the time of grain filling. Furthermore, this load increased, and the relative increment in the ear weight after flowering decreased in the years of drought, because the contribution of the leaves did not depend on water supply during the growing season. During these years, the contribution of the ear increased in the plants of all the cultivars. The authors conclude that, in arid regions, in addition to drought resistance, the cultivar should display such a ratio between the ear weight and the weight of leaves that would still ensure satisfactory grain filling     

Wheat ear carbon assimilation and nitrogen remobilization contribute significantly to grain yield

The role of wheat ears as a source of nitrogen (N) and carbon (C) in the grain filling process has barely been studied. To resolve this question, five wheat genotypes were labeled with ¹⁵N‐enriched nutrient solution. N remobilization and absorption were estimated via the nitrogen isotope composition of total organic matter and Rubisco. Gas exchange analyses showed that ear photosynthesis contributed substantially to grain filling in spite of the great loss of C due to respiration. Of the total kernel N, 64.7% was derived from the N acquired between sowing and anthesis, while the remaining 35.3% was derived from the N acquired between anthesis and maturity. In addition, 1.87 times more N was remobilized to the developing kernel from the ear than from the flag leaf. The higher yielding genotypes showed an increased N remobilization to the kernel compared to the lower yielding genotypes. In addition, the higher yielding genotypes remobilized more N from the ears to the kernel than the lower yielding genotypes, while the lower yielding genotypes remobilized more N from the flag leaf to the kernel. Therefore, the ears contribute significantly toward fulfilling C and N demands during grain filling.     

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.     

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.     

Effect of source-sink relations and nitrogen nutrition on senescence and N remobilization in the flag leaf of wheat

The relation between the source-sink ratio and nitrogen nutrition on grain yield of wheat ( Triticum aestivum L. cv. Klein Chamaco) was studied in a greenhouse experiment. Plants were grown until anthesis in pots with soil fertilized with 0.16 mmol N per plant twice a week. At anthesis, all leaves but the flag leaf were excised in a group of plants. In another group the treatment consisted in a similar defoliation plus the longitudinal excision of half the ear, while a third group was left untouched as a control. At the same time, the N supply to half of the plants in each group was interrupted, while the other half continued receiving 16 m M N. The defoliated plants showed a longer functional life of the flag leaf than the control, retaining the chlorophyll, soluble proteins and total reduced nitrogen for a longer time. The ear-excised plants showed an intermediate behavior. The plants with the interrupted N supply showed a faster leaf senescence than the N supplied ones, and this coincided with an increase in the proteolytic activity and nitrogen transport to the ear. However there were no differences in ear weight between the two nitrogen treatments. It is concluded that leaves and ear compete for the nitrogen, and that a low level of carbohydrates in the flag leaf, due to a low source-sink ratio, delays leaf senescence.     

Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears

The presence of awns doubled the net photosynthetic rate of wheat ears and also increased the proportion of ¹⁴CO₂ assimilated by the ear that moved to the grain. The effect of water supply on photosynthesis and movement of assimilates was greater for leaves than ears, so that drought increased the proportion of assimilate contributed by ear photosynthesis to grain filling from 13% to 24% in the awnless ears, and from 34% to 43% in the awned ears. ¹⁴C assimilated by the ears was most important to the economy of the upper spikelets and to the distal florets in each spikelet, whereas flag leaf assimilate went mainly to the spikelets in the lower half of the ear, and to the proximal florets. Awns increased grain yield in the dry but not in the irrigated treatment, despite the large contribution of awned ears to grain filling. Either the supply of assimilate did not limit grain yield when water supply was not limiting, or there were compensating disadvantages to awns. However, they did not seem to have any adverse effect on the development of the upper florets, nor did they reduce grain number per ear.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.     

Hormonal Regulation of Source-sink Relationship: Effect of Hormones on Excised Source and Sink Organs of Cereals

The regulatory role of plant growth substances in relation tosource and sink activity was studied. Leaching of electrolytesfrom flag leaf cells increased with age. Abscisic acid, however,increased the permeability of leaf cells at 10^–4 M andlower concentrations. Plant growth regulators like auxin, gibberellinand cytokinin at 10^–5 M inhibited the leaching of electrolytesbut slightly enhanced the process at 10^–3 M. The effluxof ¹⁴C-sucrose loaded on the flag leaves was not detectablein the very early stage of panicle development but was prominentin the later stages. While auxin prevented sucrose efflux atall stages of panicle development, gibberellin and cytokininwere effective only in different degrees. All the three growthregulators enhanced the activity of -amylase in the flag leafcells but protease activity was enhanced only by gibberellin.Synthesis of polymers like starch, DNA, RNA and protein in developingrice grains was stimulated by gibberellin, cytokinin and auxinlargely in the order mentioned. (Received September 2, 1986; Accepted May 22, 1987)     

The place of brassinolide in the sequential response to plant growth regulators in elongating tissue

In the sequential response to plant growth regulators in young elongating tissue from peas and wheat the peak of sensitivity to 24-epi-brussinolide (1 μM) occurs after those of gibberellin and cytokinin and begins before that of auxin in isolated wheat ( Triticum vulgare L. ev. Egret) coleoptiles aged from 21-96 h. In dwarf pea ( Pisum sativum L. cv. Greenfeast) segments, the peak of sensitivity also lies between those of gibberellin and auxin, and it also occurs before sensitivity to auxin in sections from first leaves of wheat. All the leaf sections and all but the most mature coleoptiles and pea segments were sensitive to fusicocein (1 μM).     

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.     

Effects of foliar application of BAP on source and sink strength in four six-rowed barley (Hordeum vulgare L.) cultivars

A field experiment was conducted to investigate the effects of foliar application of a synthetic cytokinin (BAP) on source and sink strength of four different six-rowed barley (Hordeum vulgare L.) cultivars. Different spraying treatments consisting of spraying on whole plant, spraying only on leaves and spraying only on ears started at anthesis and continued for 7 days. One additional spraying was carried out on late period of grain filling. Results showed that spraying only on leaves did not affect ear weight, grain yield and 1,000-grain weight, while the two other treatments increased all above mentioned traits. Neither of treatments affected stem weight, biological yield and contribution of stem reserves in grain filling. Exogenous cytokinin did not increase photosynthetic rate and chlorophyll content in treated leaves until late period of grain filling, although there was no significant increase in final grain weight due to late application of BAP. Our results suggested that effects of foliar application of BAP were mostly due to increased sink size soon after anthesis and increased sink demand probably met by current photosynthesis of organs other than leaves, like ear green tissues. An erratum to this article is available at .