The effect of sucrose on the levels of abscisic acid, indoleacetic acid and zeatin/zeatin riboside in wheat ears growing in liquid culture

The influence of a varied sucrose supply on grain size and hormonal contents of detached wheat ears ( Triticwn aestivum L. cv. Schirokko) was investigated throughout grain development. In ears led limited amounts, or no sucrose, grain weights in both proximal and distal grain positions of the ear were reduced. Radioimmunoassay for abscisic acid, indoleacetic acid and zeatin/zeatin riboside showed that the changes in the levels of these hormones in grains and bracts were comparable to intact ears when detached ears were well supplied with sucrose. Under conditions of limited sucrose supply, higher abscisic acid levels in the distal and proximal grains of detached ears were found compared to ears supplied with adequate sucrose. Limiting sucrose supply to the ear did not alter the levels of indoleacetic acid or zeatin/zeatin riboside in either the grains or bracts of detached ears.     

Effects of abscisic acid (ABA) on grain filling processes in wheat

The effect of in situ water stress on the endogenous abscisic acid (ABA) content of the endosperm and the in vitro application of ABA on some important yield regulating processes in wheat have been studied. Water stress resulted in a marked increase in the ABA content of the endosperm at the time close to cessation of growth. Application of ABA to the culture medium of detached ears reduced grain weight. Exogenously applied ABA, at the highest concentration (0.1 mM) reduced transport of sucrose into the grains and lowered the starch synthesis ability of intact grains. In vitro sucrose uptake and conversion by isolated grains was stimulated by low ABA concentrations (0.001 mM) in the medium but was inhibited by higher concentrations. ABA application had no effect on sucrose synthase (SS) and uridine diphosphate glucose pyrophosphorylase (UDP-Gppase) activities, whereas adenosine diphosphate glucose pyrophosphorylase (ADP-Gppase), soluble starch synthase (SSS), and granule-bound starch synthase (GBSS) activities were reduced. These results raise the possibility that water stress-induced elevated levels of endogenous ABA contribute to reduced grain growth.     

Changes in Levels of Endogenous Plant Hormones During Floret Development in Wheat Genotypes of Different Spike Sizes

The levels of endogenous plant hormones regulate floret development and degeneration, and thus grain set in flower crops. This study was undertaken to characterize the changes of endogenous hormone levels during floret development in three wheat (Triticum aestivum L.) genotypes: “97J1" with the highest grain set and fertile florets per spike, “H8679" with the lowest grain set and fertile florets per spike, and a medium, “YM158". The results showed that the peak level of ABA appeared between stamen and pistil differentiation and antherlobe formation of floret development, and the timing delayed with the size of spike (earliest in “H8679” and latest in “97J1”). From antherlobe formation to meiosis, the levels of ABA and GA1+3decreased sharply in the ears of “97J1”, while in the ears of “H8679” there was only a slight decrease in ABA, and even an increase in GA1+3. The ratio of isopentenyladenosine (iPA)/ABA and IAA/ABA in the ears of “97J1” increased sharply from antherlobe formation to meiosis, but changed only slightly in the ears of “H8679”. At antherlobe formation, IAA and GA1+3 levels were higher in the ears of “97J1”, but lower in the ears of “H8679”than in the leaves. At meiosis, ABA, GA1+3 and IAA levels in the “97J1” ears were much lower than in the leaves, but similar in “H8679”. These results indicated that the sharp decreases of ABA and GA1+3 in ears from antherlobe formation to meiosis and the lowest maintenance at meiosis may be favorable for development of fertile florets and enhancement of grain set in wheat.     

Effect of abscisic acid on the transport of assimilates in barley

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

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.     

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.     

Phytohormone-mediated transformation of sugars to starch in relation to the activities of amylases, sucrose-metabolising enzymes in sorghum grain

Indole-acetic acid (IAA) and abscisic acid (ABA) were fed throughcomplete liquid medium (containing 2, 4, 8% sucrose) to detached earheads of sorghum. The effect of these phytohormones on interconvertion ofsugarsand their transformation to starch in relation to the activities of-, -amylases, sucrose-synthase (synthesis), sucrose-phosphatesynthase and soluble invertases was studied in the grain. This effect on theuptake of (U-¹⁴C) sucrose by detached ear heads and incorporation of¹⁴C into free sugars and starch of grain and into free sugars ofinflorescence parts was also studied. At concentrations of up to 4%sucrose in the culture medium, IAA increased the content of total free sugarsinthe grain. However, accumulation of starch and activities of - and-amylases increased when lAA was present even beyond the 4%sucroseconcentration in the culture medium. At all sucrose concentrations, the effectsof ABA and IAA were opposite. With 4% sucrose, both phytohormones causedmaximum accumulation of starch in the grain. ABA enhanced the relativeproportion of sucrose in the sugar pool with a concomitant reduction in theactivities of soluble acid (pH 4.8) and neutral (pH 7.5) invertases. Incontrast, IAA decreased the sucrose proportion of grain sugars with asimultaneous elevation and reduction in the activities of invertases andsucrose-phosphate synthase, respectively. Irrespective of sucrose concentrationin the culture medium, the activity of sucrose synthase (synthesis) wasenhancedwith IAA as well as ABA at their 10 M concentration. IAA alsoenhanced incorporation of ¹⁴C from (U-¹⁴C) sucrose intothe EtOH extract (principally constituted by free sugars) and starch of thegrain, but ABA caused the reverse effect. Based on the results, it is suggestedthat IAA and ABA have contrasting effects on the transformation of sucrose tostarch in sorghum grain where its capacity to synthesise starch is modulatedpositively by IAA and negatively by ABA.     

Limitations to the Accumulation of Starch in the Developing Wheat Grain

The postulate that photosynthetic capacity limits the rate ofaccumulation of starch and dry matter in the developing wheatgrain has been tested by defoliating plants growing in the field,and by culturing detached ears on solutions of sucrose. In mostcases complete defoliation did not reduce the amounts of starchor dry matter produced 4 to 10 days later, or affect the amountsof sucrose in the grain. Increasing the potential supply ofsucrose above normal levels by culturing did not lead to increasesin either the deposition of starch or the quantity of sucrosein the grain. Removing leaves caused small depressions in thegain in fresh weight of the grain, and where defoliation didreduce the accumulation of starch there was no comparable effecton the amount of sucrose in the grain. Accordingly, it is concludedthat the postulate stated above is invalid. The case in support of the concept that it is the mechanismregulating the passage of sucrose into the grain which imposesa limitation on the accumulation of starch is presented.     

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.     

Long distance transport of potassium in cereals during grain filling in detached ears

Ears of wheat plants ( Triticum aestivum L. cv. Kolibri), which were given different and uniform K⁺-nutrition in two experiments, were cut at 2, 4 and 6 weeks after anthesis at 15 cm below the ear. These detached ears were fed 30 m M (experiment 1) or 15, 30, 60 or 90 m M ⁸⁶Rb-K₂ malate (experiment 2) and 146 m M [¹⁴C]-sucrose. After a pulse period of 6 and 4 h, respectively, the ears were transferred to identical non-labeled solutions for additional 0, 4, 8 or 20 h.
About 50% of the K⁺ and sucrose supplied was absorbed by detached ears. This rate declined with plant age and decreasing transpiration. Within the 6 and 4 h uptake period less than 7% of the absorbed K⁺, but 20% of the sucrose taken up were incorporated into the grain. During the chase period labeled K⁺ in the grain increased to 15% and ¹⁴C even to 50% of total tracer uptake. Incorporation of labeled K⁺ into the grain was not affected by the previous K⁺ nutrition of the plant and was proportional to the K⁺ concentration in the uptake solution. Transition of K⁺ from xylem into phloem during its acropetal transport is assumed. No evidence was found that the grain itself could control its uptake of K⁺.     

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 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.     

Endogenous abscisic acid levels are not linked to methyl jasmonate-promoted senescence of detached rice leaves

Both abscisic acid (ABA) and jasmonates are known to promote leaf senescence. Since ABA and jasmonates have both chemical and physiological similarities, we are interested to know whether senescence of detached rice leaves induced by methyl jasmonate (MJ) is mediated through an increase in endogenous ABA levels. In darkness, the endogenous level of ABA in detached rice leaves remained unchanged in the first day of incubation in water and increased about 5 times its initial value in the second day. However, the pattern of senescence, as judged by protein loss, was rapid during the first day. MJ significantly promoted senescence of detached rice leaves. Contrary to our expectation, endogenous ABA levels decreased in MJ-treated detached rice leaves. Similar to the effect of MJ, endogenous ABA levels decreased in detached rice leaves which were induced to senesce by treatment with NH₄Cl. These results suggest that endogenous ABA levels are not linked to MJ-induced senescence of detached rice leaves.     

Growth, yield and grain quality of barley (Hordeum vulgare L.) in response to nitrogen uptake. II. Plant development and rate of germination

A nutrient culture system supporting stands of barley was used to examine the effect of nitrogen supply on plant development and germination rate. Two contrasting cultivars of barley were grown for two seasons with nitrogen supplied at an optimal rate and at one-third that rate. Mature grain was dissected from ears by grain position; stored until out of dormancy, weighed, and the time from inhibition to germination estimated for different ear positions. A parallel set of samples was analysed for grain nitrogen concentrations. A systematic increase in germination time was observed from the bottom to the top of the ear. This effect was in addition to and of the same order as the effect of grain size on germination time. Although there was significant variation in nitrogen concentration by grain position, these effects were smaller than the variations in grain weight within the ear and were not related to germination rate. These results are discussed in relation to plant development and events at the onset of germination.Keywords: Barley, germination, grain size, ear position, nitrogen.      

Fluoride-induced alteration of carbon and nitrogen metabolism in developing wheat grains

Effect of fluoride (10 and 50 mM) on the activities of sucrose metabolizing enzymes, alkaline inorganic pyrophosphatase, and transaminases in relation to the accumulation of free sugars, starch, and soluble protein was studied in detached ears of wheat cultured in a liquid medium. Culturing for 5 d in the presence of fluoride reduced the amount of grain starch whereas contents of total free sugars, particularly sucrose, and soluble protein increased. Fluoride inhibited the activities of soluble acid and neutral invertases, as well as sucrose synthase acting in the cleavage direction. Uptake of uniformly labelled ¹⁴C-sucrose or fructose was also drastically reduced by fluoride. Glutamate-oxaloacetate transaminase (GOT) and glutamate-pyruvate transaminase (GPT) activities also increased with fluoride addition in correspondence with an increase in soluble protein. Apparently, the wheat grain responds to fluoride-mediated disruption of carbon metabolism by a compensatory effect on nitrogen metabolism. This revised version was published online in July 2006 with corrections to the Cover Date.     

Manipulation of xylem transport affects Zn and Mn transport into developing wheat grains of cultured ears

Zinc and manganese loading into developing wheat grain is little understood at present. The objective of this work was to investigate factors that may affect the rate of transport of Zn and Mn into developing wheat grain of cultured ears. Ears 18-22 days post-anthesis were cultured in solutions containing labelled Zn and Mn. The effect of additions of Cu, Fe, citrate, malate and EDTA to the culture solution was observed. The effect of humidity and awn removal as well as the sucrose status of the ears on Zn and Mn transport was also investigated. The effect of high concentration of Zn and Mn on [¹⁴C]-sucrose transport was determined. High humidity almost completely blocked transport of Zn and Mn into the grain. Awn removal reduced the transport of Zn and Mn to the lemma but not the grain. When the ears were depleted of sucrose (by maintaining them in the dark prior to labelling) transport of Zn and Mn to the grain was reduced compared to ears cultured with sucrose. The presence of Cu reduced the loading of Zn into the grain. There was little effect of Cu on Mn transport or Fe on either Zn or Mn transport. High concentrations of Zn and Mn in the culture solution did not affect [¹⁴C]-sucrose loading into the grain but loading of Zn and Mn was limited at high concentrations suggesting membrane saturation. This study demonstrates that sucrose status and humidity clearly influence the transport of Zn and Mn into the grain, and that other ions may influence Zn and Mn transport.     

Effect of sucrose level on rice grain deposition in detached panicle system

In detached IR36 rice panicles incubated in liquid medium for 10 days in 1.0–2.0% sucrose and 0.6% glutamine, final panicle weight increased but percentage and weight of grain protein decreased with 1.0–1.75% sucrose. Soluble sugars increased in stem and hull but not in developing grain. With five other rice panicles and IR36 panicles differing in grain size incubated in liquid culture containing 0.075% glutamine for 7 days, percentage grain protein was again lower in 1.5% sucrose than in 1.0% sucrose, with correspondingly heavier grain weight in four cases. Free-sugar levels of developing grains were lower in detached panicles than in the field grain samples in both experiments. Thus, sucrose level has a depressing effect on protein accumulation in the developing rice grain. Lysine content of grain protein decreased with increase in protein content.