Cell Production and DNA Accumulation in the Wheat Endosperm, and their Association with Grain Weight

Nuclear DNA content was measured in developing endosperm cellsof two wheat varieties, Chinese Spring and Spica. 3C, 6C, 12Cand 24C nuclei were detected, indicating that some form of endoreduplicationand/or endopolyploidization was occurring. The total amountof DNA in the endosperm continued to increase until 24 dayspost anthesis. This accumulation of DNA resulted both from productionof new nuclei and also from increases in the DNA content ofexisting nuclei. Estimates of endosperm cell numbers were made from the totalDNA content per endosperm and the mean DNA content per endospermnucleus for a range of genotypes differing in mature grain weight.Endosperm DNA content and cell number were both positively associatedwith mature grain weight among the genotypes examined. However,not all of the variation in grain weight could be attributedto variation in cell number because of differences in mean dryweight per endosperm cell. The large-grained variety, Spica, had a greater mean weightper endosperm cell than Chinese Spring and this difference aroseafter cell production in the endosperm had ceased. Triticum aestivum, grain weight, cell size, cell number, DNA content     

The Effects of Floret Sterilization on Grain Number and Grain Weight in Wheat Ears

This paper describes the effects of sterilizing certain floretson the development of others within the ears of wheat, cultivarMaris Ranger. Sterilization of all the florets in spikelets2, 4, 6, and 8 (numbered from the base of the ear upwards) ledto more grain setting and greater grain growth in the untreatedspikelets. These compensatory increases were insufficient toprevent a depression in the yield of grain per ear. Sterilizationof more than one of the basal florets of spikelet 8 led to amore frequent setting of the grain in the distal florets onthat spikelet and to the centrally positioned grain becomingheavier. The physiological basis for the inhibitory influences of theolder and developing grain on the unfertilized florets and youngergrain is discussed.     

Interactions Among Number of Spikelets, Number of Grains and Grain Weight in the Spikes of Wheat (Triticum aestivum L.)

In a field experiment, comprising four spring wheat cultivars,the frequency and final weight of the grains developing fromeach individual floret were determined in intact spikes andin spikes of which up to nine spikelets had been removed. Theextent of damage caused by the cutting procedure was estimated. Characteristic distributions of the frequencies and weightsof the individual grains were found for each cultivar. Removalof spikelets resulted, in most cases, in a small increase inthe number of grains and in a considerable increase in the weightof the grains of the remaining spikelets. These increases compensatedonly partially, and differently in the different cultivars,for the loss of the removed spikelets. Defoliation at the timeof earing caused a subsequent reduction in grain yield of intactspikes but no reduction in the yield of spikes from which ninespikelets had been removed. The removal of the upper floretsin each spikelet resulted in a certain increase in the weightof the two basal grains. It is concluded that an increase in the number of spikeletsper spike may reduce grain weight but will nevertheless contributeto yield. The number of grains per spikelet is cultivar dependentbut not causally associated with grain weight. Grain set indistal florets is expected to add rather small grains to thespike's yield. Under conditions of limited supplies it may causea reduction in the weight of the basal grains. Any increasein grain weight is anticipated to contribute to grain yieldand is not liable to affect spikelets per spike or grains perspikelet. Wheat cultivars, Triticum aestivum, growth of inflorescence, grain yield, spikelet number     

大麦胚乳细胞增殖动态及其与粒重的关系

大麦籽粒胚乳细胞数在花后17 d左右就基本决定,增殖动态可用Richards曲线方程拟合,决定系数0.9900以上,达极显著水平.籽粒胚乳细胞数目、单个细胞重量与粒重均存在极显著正相关,r值分别为0.9019**和0.9409**.籽粒胚乳细胞数对粒重影响最大,单个胚乳细胞重次之,胚乳细胞数的多少是决定粒重的主要原因.     

Block-Surface Staining for Differentiation of Starch and Cell Walls in Wheat Endosperm

A staining technique for differentiating starch granules and cell walls was developed for computer-assisted studies of starch granule distribution in cells of wheat [Triticum aestivum L.] caryopses. Blocks of embedded caryopses were sectioned, exposing the endosperm tissue, and stained with iodine potassium iodide (IKI) and Calcofluor White. Excessive tissue hydration during staining was avoided by using stains prepared in 80% ethanol and using short staining times. The IKI quenched background fluorescence which facilitated the use of higher concentrations of Calcofluor White. Cell wall definition was improved with the IKI-Calcofluor staining combination compared to Calcofluor alone. The high contrast between darkly stained starch granules and fluorescent cell walls permitted computer assisted analysis of data from selected hard and soft wheat varieties. The ratio of starch granule area to cell area was similar for both wheat classes. The starch granule sizes ranged from 2.1 μm³ to 22,000 μm³ with approximately 90% of the granules measuring less than 752 μm³ (ca. 11 μm in diameter). Hard wheat samples had a greater number of small starch granules and a lower mean starch granule area compared to the soft wheat varieties tested. The starch size distribution curve was bimodal for both the hard and soft wheat varieties. Three-dimensional starch size distribution was measured for four cells near the central cheek region of a single caryopsis. The percentage of small granules was higher at the ends than at the mid-section of the cells.     

Regulation of Grain Weight by Supply of Assimilates and Starch Granule Development in Three Winter Wheat Varieties

On removing the top half of the ear (halving) on several datesafter anthesis, dry weight per grain increased in three winterwheat (Triticum aestivum L.) varieties, in two pot experiments;the increase was greater with early than with late halving.The variety Splendeur had a lower dry weight and water percentagein grains than either Hobbit or Maris Huntsman. The ratio ofthe green area integrated over the post-anthesis period to thenumber of grains per ear (green area per grain) was highestin Splendeur and lowest in Hobbit in the first experiment; inthe second, Splendeur gave a lower ratio than the other twovarieties, which showed similar values. The green area per grainwas greater the earlier the ear was halved. The number of A-typestarch granules per endosperm, but not the volume per A granule(modal volume) and the modal volume of B starch granules, butnot their number, increased to a greater extent with early thanwith late halving. In Splendeur the grains had fewer A starchgranules, although these were of greater modal volume than inHobbit and Maris Huntsman and a number of B starch granulessimilar to the other two varieties, but of smaller modal volume.Maris Huntsman had more A granules than Hobbit, but with smallermodal volume. Dry weight per grain increased linearly with thenumber of A starch granules per endosperm, which in turn increasedasymptotically with green area per grain. The regressions forthe three varieties differed significantly. The influence ofthe supply of assimilates and the capacity for starch granuleformation in the regulation of grain weight is discussed. Key words: Grain weight, starch granules, assimilate supply, variety, wheat     

Measurement of Metabolites Associated with Nonaqueously Isolated Starch Granules from Immature Zea mays L. Endosperm

Starch granules with associated metabolites were isolated from immature Zea mays L. endosperm by a nonaqueous procedure using glycerol and 3-chloro-1,2-propanediol. The soluble extract of the granule preparation contained varying amounts of neutral sugars, inorganic phosphate, hexose and triose phosphates, organic acids, adenosine and uridine nucleotides, sugar nucleotides, and amino acids. Based on the metabolites present and on information about translocators in chloroplast membranes, which function in transferring metabolites from the chloroplast stroma into the cytoplasm, it is suggested that sucrose is degraded in the cytoplasm, via glycolysis, to triose phosphates which cross the amyloplast membrane by means of a phosphate translocator. It is further postulated that hexose phosphates and sugars are produced from the triose phosphates in the amyloplast stroma by gluconeogenesis with starch being formed from glucose 1-phosphate via pyrophosphorylase and starch synthase enzymes. The glucose 1-phosphate to inorganic phosphate ratio in the granule preparation was such that starch synthesis by phosphorylase is highly unlikely in maize endosperm.     

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.     

The Effect of High Temperature at Different Stages of Ripening on Grain Set, Grain Weight and Grain Dimensions in the Semi-dwarf Wheat 'Banks'

Grain number in the wheat cultivar Banks was reduced by up to11 % with a rise in temperature from 21/16 °C to 30/25 °Cover a 10-d period immediately following first anthesis in general,the upper ‘d’ and ‘c’ florets were moreaffected by high temperature than the basal ‘a’and ‘b’ florets within a spikelet and florets fromthe upper spikelets were more sensitive than those lower onthe ear Grain weight and grain length at maturity were most affectedby a 10 d period of high temperature commencing 7–10 dafter anthesis However, if dry-matter accumulation between thestart of a treatment and grain maturity was used as a base forcomparison, the response was more uniform throughout development,with a peak in sensitivity 25 d after anthesis Although grainposition within an ear did not have a large effect on the responseto temperature, grains from the basal spikelets were more sensitivethan those from the apex, and the upper floret grains of a spikeletwere more sensitive to high temperature than those at the base There is a need to obtain, for a range of cultivars, more comprehensivedata on the effect of the timing and degree of temperature stressfollowing anthesis, for use in interpreting the response torising temperatures late in the development of the crop in thefield Triticum aestivum L, wheat, temperature, grain development     

Genetic Dissection of Grain Size and Grain Number Trade-Offs in CIMMYT Wheat Germplasm

Grain weight (GW) and number per unit area of land (GN) are the primary components of grain yield in wheat. In segregating populations both yield components often show a negative correlation among themselves. Here we use a recombinant doubled haploid population of 105 individuals developed from the CIMMYT varieties Weebill and Bacanora to understand the relative contribution of these components to grain yield and their interaction with each other. Weebill was chosen for its high GW and Bacanora for high GN. The population was phenotyped in Mexico, Argentina, Chile and the UK. Two loci influencing grain yield were indicated on 1B and 7B after QTL analysis. Weebill contributed the increasing alleles. The 1B effect, which is probably caused by to the 1BL.1RS rye introgression in Bacanora, was a result of increased GN, whereas, the 7B QTL controls GW. We concluded that increased in GW from Weebill 7B allele is not accompanied by a significant reduction in grain number. The extent of the GW and GN trade-off is reduced. This makes this locus an attractive target for marker assisted selection to develop high yielding bold grain varieties like Weebill. AMMI analysis was used to show that the 7B Weebill allele appears to contribute to yield stability.     

Effects of Radiation and Temperature on Tiller Survival, Grain Number and Grain Yield in Winter Wheat

Effects of the environment on shoot survival were studied inwinter wheat cv. Avalon grown in microplots at a density of247 plants m^–2. The incident radiation and mean temperaturewere altered during one of three periods of between 14 and 29d duration, the first (P1) starting at the end of tiller productionand the last (P3) finishing near the end of the tiller deathphase, about three weeks before anthesis. Plants were giventemperature and radiation treatments in growth rooms in twoexperiments and extra light outdoors in a third experiment:they were at other times grown outdoors. Increasing radiation by between 60 and 100 per cent during P1had negligible effects on shoot number; during P2 it alwaysdelayed tiller death but increased final ear number in onlyone experiment; during P3 it consistently increased ear numberby up to 100 m^–2. Increased radiation always increasedcrop dry weight immediately after treatment but only sometimesdid this increase persist to maturity. Grain dry weight wasincreased by treatment during P3 of one experiment. Increasingthe temperature by 4 C decreased shoot number, usually onlytemporarily, by hastening death of some tillers. Warmer temperaturesdecreased crop growth after, but not during, treatment and decreasedgrain dry weight. Radiation and temperature rarely interacted. Variation in grain yield within and between experiments relatedwell to variation in number of grains m^–2, which in turnrelated to variation in ear dry weight at anthesis. Triticum aestivum L., wheat, radiation, temperature, tillers, grain yield, grain number     

Copy Number Variation of Cytokinin Oxidase Gene Tackx4 Associated with Grain Weight and Chlorophyll Content of Flag Leaf in Common Wheat

As the main pigment in photosynthesis, chlorophyll significantly affects grain filling and grain weight of crop. Cytokinin (CTK) can effectively increase chlorophyll content and chloroplast stability, but it is irreversibly inactivated by cytokinin oxidase (CKX). In this study, therefore, twenty-four pairs of primers were designed to identify variations of wheat CKX (Tackx) genes associated with flag leaf chlorophyll content after anthesis, as well as grain weight in 169 recombinant inbred lines (RIL) derived from Triticum aestivum Jing 411 × Hongmangchun 21. Results indicated variation of Tackx4, identified by primer pair T19-20, was proven to significantly associate with chlorophyll content and grain weight in the RIL population. Here, two Tackx4 patterns were identified: one with two co-segregated fragments (Tackx4-1/Tackx4-2) containing 618 bp and 620 bp in size (as in Jing 411), and another with no PCR product. The two genotypes were designated as genotype-A and genotype-B, respectively. Grain weight and leaf chlorophyll content at 5~15 days after anthesis (DAA) were significantly higher in genotype-A lines than those in genotype-B lines. Mapping analysis indicated Tackx4 was closely linked to Xwmc169 on chromosome 3AL, as well as co-segregated with a major quantitative trait locus (QTL) for both grain weight and chlorophyll content of flag leaf at 5~15 DAA. This QTL explained 8.9~22.3% phenotypic variations of the two traits across four cropping seasons. Among 102 wheat varieties, a third genotype of Tackx4 was found and designated as genotype-C, also having two co-segregated fragments, Tackx4-2 and Tackx4-3 (615bp). The sequences of three fragments, Tackx4-1, Tackx4-2, and Tackx4-3, showed high identity (>98%). Therefore, these fragments could be considered as different copies at Tackx4 locus on chromosome 3AL. The effect of copy number variation (CNV) of Tackx4 was further validated. In general, genotype-A contains both significantly higher grain weight and flag leaf chlorophyll content at 5~15 DAA than those in genotype-B and genotype-C, among 102 varieties under various environments.     

The Influence of Sucrose and Abscisic Acid on the Determination of Grain Number in Wheat

Using detached ears of wheat cultured in nutrient solution theeffect of sucrose and ABA supply on grain number was examined.In ears detached prior to ear emergence, to correspond withthe time of pollen mother cell meiosis, higher levels of sucroseuptake were associated with higher grain number per ear. ABAindirectly caused a reduction in grain number by decreasinguptake of sucrose through decreasing stomatal conductance andtranspiration. There was possibly also a direct effect of ABAon grain number. At higher levels of ABA the negative effecton grain number was decreased by increasing the sucrose concentration.In ears detached one to three days after anthesis grain numberwas also influenced by the amount of sucrose taken up. althoughgrain number was not as sensitive to the sucrose supply at thistime as it was at meiosis. The influence of ABA following anthesiswas limited to an indirect effect on grain number through reductionsin sucrose uptake. Key words: Wheat, Sucrose, ABA, Grain number     

The Deterioration of Wheat Embryo and Endosperm Function with Age

The rates of senescence of embryos and aleurone tissues of different-agedwheat was compared. Germination behaviour was used as the primarycriterion of embryo senescence and GA₃-induced reducing sugarrelease and -amylase production were the primary criteria ofaleurone senescence. The results suggest that both tissues ageindependently though at identical rates, and that the primaryfactor in the ageing process is probably related to a basicaspect of metabolism or intracellular integrity, rather thanany functional capacity of the cells.     

Leptin基因的PCR-SSCP与牛体重、体尺指标的相关性

利用PCR—SSCP技术研究了南阳牛、秦川牛、郏县红牛、西镇牛、鲁西牛和荷斯坦奶牛6个牛品种539个个体leptin基因的遗传多态性。结果表明,PCR扩增产物大小为330bp,PCR—SSCP分析表现出多态。南阳牛、秦川牛、郏县红牛、西镇牛、鲁西牛和荷斯坦奶牛的A等位基因频率分别为0．558,0．492,0．571,0．658,0．591,0．615;B等位基因频率分别为0．442,0．508,0．429,0．342,0．409,0．385。不同基因型与体重、体尺等生长性状指标相关性分析的结果表明：南阳牛群体内除12月龄的体高和日增重、18月龄的坐骨端宽和日增重外,BB型个体的六月龄、十二月龄、十八月龄、二十四月龄体斜长、胸围、体重、坐骨端宽、体高和日增重均显著的大于AB和AA型个体（P〈0．05）;秦川牛群体内BB基因型个体十字部高上显著高于群体AA、AB型个体（P〈0．05）,即BB〉AA、AB,可作为秦川牛体尺指标（十字部高）候选基因之一,但在体重、胸围、体长指标上均无显著差异（P〉0．05）,所以不宜作为体重、胸围、体长指标候选基因;郏县红牛群体内AB与BB基因型个体在十字部高和坐骨端宽上显著高于群体AA型个体（P〈0．05）,而群体内不同基因型在体重和体尺指标（体高、体斜长、胸围）上无显著差异（P〉0．05）。序列分析表明,leptin基因多态是第66位发生G→T、第67位发生A→C及299位发生新的单核苷酸突变C→T所造成。     

Association Analysis of Genomic Loci Important for Grain Weight Control in Elite Common Wheat Varieties Cultivated with Variable Water and Fertiliser Supply

Grain weight, an essential yield component, is under strong genetic control and markedly influenced by the environment. Here, by genome-wide association analysis with a panel of 94 elite common wheat varieties, 37 loci were found significantly associated with thousand-grain weight (TGW) in one or more environments differing in water and fertiliser levels. Five loci were stably associated with TGW under all 12 environments examined. Their elite alleles had positive effects on TGW. Four, two, three, and two loci were consistently associated with TGW in the irrigated and fertilised (IF), rainfed (RF), reduced nitrogen (RN), and reduced phosphorus (RP) environments. The elite alleles of the IF-specific loci enhanced TGW under well-resourced conditions, whereas those of the RF-, RN-, or RP-specific loci conferred tolerance to the TGW decrease when irrigation, nitrogen, or phosphorus were reduced. Moreover, the elite alleles of the environment-independent and -specific loci often acted additively to enhance TGW. Four additional loci were found associated with TGW in specific locations, one of which was shown to contribute to the TGW difference between two experimental sites. Further analysis of 14 associated loci revealed that nine affected both grain length and width, whereas the remaining loci influenced either grain length or width, indicating that these loci control grain weight by regulating kernel size. Finally, the elite allele of Xpsp3152 frequently co-segregated with the larger grain haplotype of TaGW2-6A, suggesting probable genetic and functional linkages between Xpsp3152 and GW2 that are important for grain weight control in cereal plants. Our study provides new knowledge on TGW control in elite common wheat lines, which may aid the improvement of wheat grain weight trait in further research.     

Relationships Between Grain Weight and the Size of Floret Cavity in the Wheat Spike

The volume of the floret cavity at different floret positionsalong the spike and within a single spikelet was estimated in10 Triticum aestivum and three T. durum lines by injecting floretcavities with liquid silicone rubber which solidified thereafter.Highly significant correlation coefficients, ranging from 0.40to 0.76 were found between floret volume and grain weight; inmost lines the basal florets had a higher correlation than theterminal ones. The relationships between floret volume and grainweight were characterized by an intercept of about 30–60per cent of the mean grain weight (heavy-grained lines havinga larger intercept) and a slope of about 1 mg µI^–1.Differences in grain size and shape, both within spike as wellas among lines, closely reflected the variation in the sizeof floret cavity. The data support the hypothesis that grainweight is partly determined by the volume of the floret cavity. Grain weight, grain volume, floret cavity     

Changes in Cell Number and Cell Division Activity during Endosperm Development in Allohexaploid Wheat, Triticum aestivum L.

Nuclear or cell number, and the mitotic index, were recordedin endosperms of Triticum aestivum cv. Mardler to test if aparticular stage of endosperm development was critical in determiningthe final grain weight. The basal four florets of emasculatedspikelets (controls), and the third and fourth florets of spikeletswhere the two basal ovaries were removed (ovary-removed), weresampled at various times up to 360 h after hand-pollination.The coenocytic phase in endosperms ended about 84 h after pollinationregardless of both grain position and the treatment. The onsetof the cellular stage was characterized by the final large fluctuationsin the mitotic index reflecting the culmination of the synchronousnuclear division of the coenocytic stage. Thereafter, the mitoticindex fluctuated with smaller amplitudes and, by 216 h afterpollination, was < 1%. Neither floret position in the spikeletnor the treatment affected the pattern of alteration to themitotic index. However, ovary removal from first and secondflorets resulted in significantly heavier grains and higherendosperm cell number in the 3rd and 4th florets compared withthe controls. In all florets, mean endosperm cell number peakedat 280 h but decreased by 360 h after pollination. At this time,the mean cell numbers in endosperms of the 3rd and 4th floretsof ovary-removed spikelets were significantly higher than inthe corresponding endosperms in the controls. Thus, a key contributoryfactor in determining the final endosperm cell number may bethe number of cells which are lost during the late period ofthe cellular stage of endosperm development. Key words: Endosperm cell number, florets, grain weight, mitotic index, Triticum aestivum     

三种粒型小麦品种胚乳细胞增殖动态研究

以三种粒型小麦品种(系)为材料,观察了不同品种和同一品种不同粒位籽粒胚乳细胞增殖动态。结果表明,用Richards方程能较好地模拟胚乳细胞增殖动态。强势粒胚乳细胞分裂起始势高,达到最高增殖速率的时间短,活跃分裂期长,可分裂出更多的胚乳细胞。弱势粒胚乳细胞增殖起始势低,细胞分裂速率变化缓慢,其最终胚乳细胞数显著低于强势拉。不同品种间胚乳细胞数有一定的差异,表现为大粒饱满品种(鄂思1号)>不饱满品系(95A-10)>小粒饱满品种(华麦8号)。胚乳细胞增殖速率变化为单峰曲线,强势粒胚乳细胞增殖速率曲线偏左,弱势粒胚乳细胞增殖速率曲线偏右。     

The Development and Abnormality of Embryo and Endosperm in Trigrain Wheat

The development processes of embryos and endosperm of trigrain wheat were observed by using paraffin section method and intact dissection. The results were as follows: 1. Fertilization: the development and progression of embryo and endosperm were similar to those as common wheat. 2. The grain come from the primary pistil embryo developed more early among the three-grains. 3. Many abnormal structures, such as Conversion of position between the top and base, back and belly of embryos, horizontal embryos, embryos moved on top, polyembryony and endosperm deficiency etc, appeared in additional pistils. All abnormalities accounted for 88 percent in whole additional pistils.