Different Stages Of Grain Development Research Articles

Genome-wide identification, abiotic stress, and expression analysis of PYL family in Tartary buckwheat (Fagopyrum tataricum (L.) Gaertn.) during grain development

BackgroundAbscisic acid (ABA) is a plant hormone that plays an important role in plant resistance to drought, salinity, cold, and pathogens. It is also important for regulating plant growth and development. Pyrabactin resistance/pyr1-like/regulatory components of the ABA receptor (PYL/RCAR) are ABA receptor proteins in plants and the core of ABA signal transduction pathways in plant regulatory factors. At present, there are no reports on the PYL family of Tartary buckwheat.ResultsIn this study, 19 paralogous form PYL genes in buckwheat were identified at the whole-genome level and named FtPYL1-FtPYL19 according to their positions on chromosomes. We further analyzed the gene structure, conserved motifs, cis-acting elements, gene duplication, phylogenetic relationships, and expression patterns under different stress treatments and during grain development of the 19 paralogous form PYL genes in Tartary buckwheat. The FtPYL gene exhibits a single exonic gene structure for about 68.4% of the duplicated forms from the total paralogous forms. The remaining subfamilies, such as I and II, contain three exons and two exons (e.g., FtPYL19), respectively. Nineteen FtPYL genes were evenly distributed across the eight chromosomes, with at least one FtPYL gene on each chromosome. In the FtPYL gene family, there was one tandem repeat event and five gene duplication events. We investigated the gene expression levels of FtPYL gene under four abiotic stresses and different stages of grain development. Under drought stress (PEG6000), the relative expression levels of FtPYL14 and FtPYL15 increased by fourfold. Under high temperature stress (38℃), the relative expression level of FtPYL16 dropped to 0.12, and that of FtPYL17 fell to 0.22. At different stages of grain development, the gene expression level of FtPY15 is extremely high at 19 D. The relative expression level of FtPYL7 in roots and stems reaches up to approximately 450, and the relative expression level of FtPYL10 in 13 D also reaches up to 248.SummaryIn this study, the PYL gene family of Tartary buckwheat was identified and analyzed based on the whole genome, and 19 paralogous form FtPYL genes of Tartary buckwheat were bioinformatically analyzed. The expression patterns of 19 paralogous form FtPYL genes in Tartary buckwheat cultivars under different stress treatments and during grain development were analyzed. It was found that the FtPYL gene played an important role in grain development.

Effect of Ultraviolet B Radiation on the Biosynthesis of Carotenoids in Foxtail Millet Grains

Foxtail millet is a vital nutritional cereal. The de-husked grain is usually yellow and mainly contains carotenoids, which directly reflects the millet quality. In this study, the impact of ultraviolet-B(UV-B) on millet color and carotenoid content was determined using two foxtail millet varieties, HuaJinZao (HJZ) and Qinhuang 2 (QH). The b* value at different stages of grain development and the content of carotenoids (primarily lutein and zeaxanthin) in foxtail millet grains decreased when the plants were exposed to low UV-B intensity. A total of 3113 and 96 differentially expressed genes were identified in HJZ and QH, respectively, and were found to be associated with the metabolism of tryptophan, starch, and sucrose as well as the biosynthesis of amino acids, which was relatively consistent with the functional annotation of differential metabolites. Furthermore, we evaluated the changes in the expression of seven and eight genes associated with carotenoid and starch metabolism, respectively, in the kernels of foxtail millet exposed to UV-B and found that appropriate UV-B intensity could promote the expression levels of genes involved in carotenoid synthesis and repress the expression of genes involved in carotenoid degradation. This study lays a theoretical foundation for cultivating new foxtail millet varieties with high carotenoid content.

Fine Mapping and Candidate Gene Analysis of Rice Grain Length QTL qGL9.1.

Grain length (GL) is one of the crucial determinants of rice yield and quality. However, there is still a shortage of knowledge on the major genes controlling the inheritance of GL in japonica rice, which severely limits the improvement of japonica rice yields. Here, we systemically measured the GL of 667 F2 and 1570 BC3F3 individuals derived from two cultivated rice cultivars, Pin20 and Songjing15, in order to identify the major genomic regions associated with GL. A novel major QTL, qGL9.1, was mapped on chromosome 9, which is associated with the GL, using whole-genome re-sequencing with bulked segregant analysis. Local QTL linkage analysis with F2 and fine mapping with the recombinant plant revealed a 93-kb core region on qGL9.1 encoding 15 protein-coding genes. Only the expression level of LOC_Os09g26970 was significantly different between the two parents at different stages of grain development. Moreover, haplotype analysis revealed that the alleles of Pin20 contribute to the optimal GL (9.36 mm) and GL/W (3.31), suggesting that Pin20 is a cultivated species carrying the optimal GL variation of LOC_Os09g26970. Furthermore, a functional-type mutation (16398989-bp, G>A) located on an exon of LOC_Os09g26970 could be used as a molecular marker to distinguish between long and short grains. Our experiments identified LOC_Os09g26970 as a novel gene associated with GL in japonica rice. This result is expected to further the exploration of the genetic mechanism of rice GL and improve GL in rice japonica varieties by marker-assisted selection.

DNA Methylation and RNA-Sequencing Analysis Show Epigenetic Function During Grain Filling in Foxtail Millet (Setaria italica L.)

Grain filling is a crucial process for crop yield and quality. Certain studies already gained insight into the molecular mechanism of grain filling. However, it is unclear whether epigenetic modifications are associated with grain filling in foxtail millet. Global DNA methylation and transcriptome analysis were conducted in foxtail millet spikelets during different stages to interpret the epigenetic effects of the grain filling process. The study employed the whole-genome bisulfite deep sequencing and advanced bioinformatics to sequence and identify all DNA methylation during foxtail millet grain filling; the DNA methylation-mediated gene expression profiles and their involved gene network and biological pathway were systematically studied. One context of DNA methylation, namely, CHH methylation, was accounted for the largest percentage, and it was gradually increased during grain filling. Among all developmental stages, the methylation levels were lowest at T2, followed by T4, which mainly occurred in CHG. The distribution of differentially methylated regions (DMR) was varied in the different genetic regions for three contexts. In addition, gene expression was negatively associated with DNA methylation. Evaluation of the interconnection of the DNA methylome and transcriptome identified some stage-specific differentially expressed genes associated with the DMR at different stages compared with the T1 developmental stage, indicating the potential function of epigenetics on the expression regulation of genes related to the specific pathway at different stages of grain development. The results demonstrated that the dynamic change of DNA methylation plays a crucial function in gene regulation, revealing the potential function of epigenetics in grain development in foxtail millet.

Localization, Gene Expression, and Functions of Glutamine Synthetase Isozymes in Wheat Grain (Triticum aestivum L.).

Glutamine synthetase (GS) plays a major role in plant nitrogen metabolism, but the roles of individual GS isoforms in grains are unknown. Here, the localization and expression of individual TaGS isozymes in wheat grain were probed with TaGS isoenzyme-specific antibodies, and the nitrogen metabolism of grain during the grain filling stage were investigated. Immunofluorescence revealed that TaGS1;1, TaGS1;3, and TaGS2 were expressed in different regions of the embryo. In grain transporting tissues, TaGS1;2 was localized in vascular bundle; TaGS1;2 and TaGS1;1 were in chalaza and placentochalaza; TaGS1;1 and TaGS1;3 were in endosperm transfer cells; and TaGS1;3 and TaGS2 were in aleurone layer. GS exhibited maximum activity and expression at 8 days after flowering (DAF) with peak glutamine content in grains; from then, increased largely from reduction, glutamate dehydrogenase (GDH) aminating activity increased continuously, and the activities of GS and glutamate synthase (GOGAT) decreased, while only TaGS1;3 kept a stable expression in different TaGS isozymes. Hence, GS-GOGAT cycle and GDH play different roles in assimilation of grain in different stages of grain development; TaGS1;3, located in aleurone layer and endosperm transfer cells, plays a key role in Gln into endosperm for gluten synthesis. At 30 DAF, grain amino acids are mainly transported from maternal phloem.

Discovery of genomic regions and candidate genes for grain weight employing next generation sequencing based QTL-seq approach in rice (Oryza sativa L.).

Rice (Oryza sativa L.) yield enhancement is one of the prime objectives of plant breeders. Elucidation of the inheritance of grain weight, a key yield component trait, is of paramount importance for raising the yield thresholds in rice. In the present investigation, we employed Next-Generation Sequencing based QTL-seq approach to identify major genomic regions associated with grain weight using mapping populations derived from a cross between BPT5204 and MTU3626. QTL-seq analysis identified three grain weight quantitative trait loci (QTL) viz., qGW1 (35-40Mb), qGW7 (10-18Mb), and qGW8 (2-5Mb) on chromosomes 1, 7 and 8, respectively and all are found to be novel. Further, qGW8 was confirmed through conventional QTL mapping in F2, F3 and BC1F2 populations and found to explain the phenotypic variance of 17.88%, 16.70% and 15.00%, respectively, indicating a major QTL for grain weight. Based on previous reports, two candidate genes in the qGW8 QTL were predicted i.e., LOC_Os08g01490 (Cytochrome P450), and LOC_Os08g01680 (WD domain, G-beta repeat domain containing protein) and through in silico analysis they were found to be highly expressed in reproductive organs during different stages of grain development. Here, we have demonstrated that QTL-seq is one of the rapid approaches to uncover novel QTLs controlling complex traits. The candidate genes identified in the present study undoubtedly enhance our understanding of the mechanism and inheritance of the grain weight. These candidate genes can be exploited for yield enhancement after confirmation through complementary studies.

Hormonal regulation of cereal endosperm development with a focus on rice (Oryza sativa).

The endosperm of cereal grain forms the staple diet for most of the world's population, and feeds much of their stock. Grain size and quality are determined largely by events taking place during coenocytic nuclear division, endosperm cellularisation and cell differentiation, and the production of storage molecules. Thus, understanding the complex signalling processes occurring at each of these steps is essential for maintaining and improving our food supply. Here, we critically review evidence for the effects of phytohormones on grain size, as well as hormone homeostasis, signalling and crosstalk. We focus on rice endosperm due to the importance of rice as a food crop and a model grass, as well as its relative neglect in recent reviews; however, data from other cereals are also discussed due to strong evidence for conserved signalling networks operating during grain development. Discussion is restricted to auxin, cytokinin, ethylene, abscisic acid and gibberellin. Our review highlights the need for accurate hormone determinations combined with information on gene expression. We present evidence for separate, localised signalling roles for auxin at different stages of grain development and highlight key research questions for other hormones where much less data are available.

Metabolomic approach for characterization of phenolic compounds in different wheat genotypes during grain development.

The phenolic-profiling of seven different wheat (Triticum aestivum) genotypes was investigated for the first time during different stages of grain development (milky, softy, physiological maturity and mature). Free and bound phenolic compounds were extracted separately and analyzed by UPLC-QTOF-MSE. Total phenolic content significantly decreased, up to 50% depending on the genotype, towards the maturation of grain. The highest content (free and bound) was observed in the most immature grains, while the lowest level was found in mature grains (408.0 and 165.0 GAE mg/100 g, respectively). Globally, 237 phenolic compounds were identified, divided into 5 classes: flavonoids (85), phenolic acids (77), other polyphenols (51), lignans (16) and stilbenes (8). UPLC-MS results showed a progressively decrease of the number of phenolic identification (ID) all along grain development, milky (213), softy (192), physiological maturity (169) and mature (144). The proportion bound to free phenolic progressively increased, reaching the maximum at physiological maturity, indicating a possible enzymatic reactions and complexation during grain growth. Ferulic acid, diphyllin, 4-hydroxybenzoic acid, ferulic acid isomer, apigenin 7-O-apiosyl-glucoside isomer and myricetin isomer were the most abundant compounds. Chemometric tools showed a clear separation between immature and mature grain for all genotypes. Phenolic profile varied significantly among genotypes, this result can help the selection of varieties towards a higher retention of bioactive compounds. Noteworthy, immature wheat grains can be considered a rich source of phenolic compounds and as an attractive ingredient to incorporate to functional foods.

Differential and temperature dependent regulation of ADP-glucose pyrophosphorylase by specific chromosome in wheat grains

A stock of disomic chromosome substitution (DCS) lines having specific chromosome of wheat variety C591 substituted in the background of rest of Chinese spring chromosomes, were used to analyze grain yield components as a function of enzyme activity of ADP–glucose pyrophosphorylase (AGPase), a starch biosynthesis enzyme in wheat grains. Associations between yield characteristics, grain growth rate (GGR) and AGPase enzyme activity of DCS lines suggested a major involvement of chromosome 3A, 4B, 7D and 2D in a temperature dependent manner. Assessment of AGPase assay at different developmental stages such as 14, 21, 28 days post anthesis (DPA) embodied that gene(s) for this enzyme are present on specific chromosomes and operate at different stages of grain development. The DCS line with 7D chromosome has a major contribution in determining the grain starch content. In this line, AGPase enzyme activity was highest at 21 DPA and was the most crucial determinant in its high GGR. Line 4B performed well at only early stage (14 DPA) suggesting that line 4B AGPase requires a lower temperature range for activation as compared to 7D line. Line 3A had substantially reduced (40%) test weights revealing the presence of few down-regulatory elements on chromosome 3A to reduce the activity of AGPase. The DCS line 2D showed higher test weights and grain number than all other lines ascribed to a consistent AGPase activity along with an efficient mechanism for translocation of photosynthates from source to sink. The chromosome 2D shows positive relation with yield attributes therefore, it can be employed to improve wheat productivity via analytical breeding programme.

Effect of water stress at different stages of grain development on the characteristics of starch and protein of different wheat varieties

The effect of water stress (WS) at 8 and 15 days post anthesis (DPA) on the characteristics of starch and protein separated from C-306, HD-2329, PBW-175, PBW-343 and NI-5439 wheat varieties was studied. WS-induced changes in A-, B- and C-type granules distribution were variety- and stage-dependent. A-type granules increased in response to WS at both stages in all varieties, the extent of increase being greater at 15 DPA. The proportion of B-type granules decreased in all the varieties, except C-306, in response to WS at 15 DPA. C-type granules also decreased in response to 15 DPA in all varieties, except HD-2329. The starch from wheat exposed to WS at 15 DPA showed lower amylose content, lipids content and pasting temperature, and higher peak viscosity, final viscosity and setback. DSC analysis of starches showed two endotherms (associated with the melting of crystallites and amylose–lipid [AML] complexes) during heating, and an exotherm (associated with reforming of AML) during cooling. Transition temperatures ( T o, T p and T c) of AML dissociation and association were lower for starch from wheat exposed to WS, the effect being more at 15 DPA. The changes in pasting and thermal properties of starch caused by WS were observed to be related to lipids, amylose content and distribution of granules. The effect of WS on accumulation of different dimethyl formamide-soluble and insoluble proteins was significant and variety dependent.

Multivariate analysis of 2‐DE protein patterns – Practical approaches

Practical approaches to the use of multivariate data analysis of 2-DE protein patterns are demonstrated by three independent strategies for the image analysis and the multivariate analysis on the same set of 2-DE data. Four wheat varieties were selected on the basis of their baking quality. Two of the varieties were of strong baking quality and hard wheat kernel and two were of weak baking quality and soft kernel. Gliadins at different stages of grain development were analyzed by the application of multivariate data analysis on images of 2-DEs. Patterns related to the wheat varieties, harvest times and quality were detected on images of 2-DE protein patterns for all the three strategies. The use of the multivariate methods was evaluated in the alignment and matching procedures of 2-DE gels. All the three strategies were able to discriminate the samples according to quality, harvest time and variety, although different subsets of protein spots were selected. The explorative approach of using multivariate data analysis and variable selection in the analyses of 2-DEs seems to be promising as a fast, reliable and convenient way of screening and transforming many gel images into spot quantities.

Early prediction of wheat quality: analysis during grain development using mass spectrometry and multivariate data analysis

Matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry and multivariate data analysis have been used for the determination of wheat quality at different stages of grain development. Wheat varieties with one of two different end-use qualities (i.e. suitable or not suitable for bread-making purposes) were investigated. The samples were collected from grains from 15 until 45 days post-anthesis (dpa). Gluten proteins from wheat grains were extracted and subsequently analysed by mass spectrometry. Discrimination partial least-squares regression and soft independent modelling of class analogy were used to determine the quality of new and unknown wheat samples. With these methods, we were able to predict correctly the end-use qualities at every stage investigated. This new fast technique, based on the rapidity of mass spectrometry combined with the objectivity of multivariate data analysis, offers a method that can replace the traditional rather time-consuming ones such as gel electrophoresis. This study focused on the determination of wheat quality at 15 dpa, when the grain is due for harvest 1 month later.

Induction of New Isoforms of Superoxide Dismutase and Catalase Enzymes in the Flag Leaf of Wheat during Monocarpic Senescence

Leaf senescence is a programmed cell death phenomenon and involves oxidative stress. Superoxide dismutase (SOD, EC 1.15.1.1) and catalase (CAT EC 1.11.1.6) activities were studied in the flag leaf of Triticum aestivum cv. Kundan at different stages of grain development. Both SOD and CAT activities showed a decline during monocarpic senescence. Three SOD isozymes were observed in the cytosol, of which one isozyme was observed in the chloroplasts as well. Mitochondria showed the presence of three low abundant SOD isoforms. Inhibitor studies revealed the cytosolic and chloroplastic isoforms to be Cu/Zn SODs. In mitochondria however, two isozymes were MnSOD and one of them appeared to be FeSOD. These isoforms present in the mitochondria increased in activity as senescence progressed. Three isoforms of CAT were observed in peroxisomes which responded differentially during monocarpic senescence. The changes in the kind and pattern of the antioxidant enzymes supported the ordered sequence of events during leaf senescence. This is the first report showing an increase in mitochondrial FeSOD activity during leaf senescence.

Ergosterol concentration in mold-susceptible and mold-resistant sorghum at different stages of grain development and its relationship to flavan-4-ols

Sorghum germplasm accessions (10) and breeding lines (3) exhibiting contrasting reactions to the grain mould complex (including Fusarium moniliforme [Gibberella fujikuroi], Curvularia lunata [Cochliobolus lunatus] and Phoma sorghina) were grown at Patancheru, India. Grains were harvested at different days after flowering (DAF) and analysed for ergosterol. Methanol and acidified methanol extracts of grains were analysed for flavan-4-ols. Ergosterol concn increased with increasing DAF in the mould-susceptible accessions and was 10-fold higher in grains collected at 50 DAF than in the corresponding mould-resistant accessions. It is suggested that ergosterol concn could be used to assess the magnitude of mould damage in sorghum grains. The correlation coeff. between ergosterol and flavan-4-ols concn was significant (P <0.01) and negative in coloured mould-susceptible and mould-resistant accessions that did not have testa, but no significant correlation was observed in white mould-resistant and mould-susceptible sorghum. It is concluded that there must be another genetic trait or mechanism besides flavan-4-ols that is associated with mould resistance

Flavan-4-ols concentration in mold-susceptible and mold-resistant sorghum at different stages of grain development

ADVERTISEMENT RETURN TO ISSUEPREVArticleNEXTFlavan-4-ols concentration in mold-susceptible and mold-resistant sorghum at different stages of grain developmentRamamurthi Jambunathan, Milind S. Kherdekar, and Ranajit BandyopadhyayCite this: J. Agric. Food Chem. 1990, 38, 2, 545–548Publication Date (Print):February 1, 1990Publication History Published online1 May 2002Published inissue 1 February 1990https://doi.org/10.1021/jf00092a046RIGHTS & PERMISSIONSArticle Views60Altmetric-Citations21LEARN ABOUT THESE METRICSArticle Views are the COUNTER-compliant sum of full text article downloads since November 2008 (both PDF and HTML) across all institutions and individuals. These metrics are regularly updated to reflect usage leading up to the last few days.Citations are the number of other articles citing this article, calculated by Crossref and updated daily. Find more information about Crossref citation counts.The Altmetric Attention Score is a quantitative measure of the attention that a research article has received online. Clicking on the donut icon will load a page at altmetric.com with additional details about the score and the social media presence for the given article. Find more information on the Altmetric Attention Score and how the score is calculated. Share Add toView InAdd Full Text with ReferenceAdd Description ExportRISCitationCitation and abstractCitation and referencesMore Options Share onFacebookTwitterWechatLinked InReddit PDF (539 KB) Get e-Alerts Get e-Alerts

Nitrogenous compounds and protease activity in developing testapericarp and endosperm of high and low protein wheat (Triticum aestivum L.)

Nitrogenous compounds such as total protein, true protein, soluble protein, non-protein nitrogen, total amides, ammonium, nitrate, nitrite, free amino acids and neutral and acid protease activities were estimated in a high protein wheat cv Shera (12·8% protein) and a relatively low protein wheat cv C-306 (10·4% protein) at different stages of grain development. All the nitrogenous compounds studied, as well as protease activities, were located in both testa-pericarp and endosperm. Developmental patterns and relative levels were, more or less, similar when expressed on per organ basis, or on dry weight basis. In the testa-pericarp; dry weight, total protein, soluble protein, true protein, amide, ammonium, nitrate and nitrite contents increased during development, and were higher in Shera, compared with cv C-306. Non-protein nitrogen content decreased during pericarp development, and was higher in cv Shera, compared with cv C-306. Free amino acid content and protease activities decreased in developing pericarp, and were lower in cv Shera, compared with cv C-306. Similar developmental patterns and relative levels of nitrogenous compounds and protease activities were found in endosperm. It is suggested that both testa—pericarp and endosperm make qualitatively similar contributions to the accumulation of nitrogenous compounds in developing wheat grain; higher protease activity in the endosperm of C-306 is not responsible for lower protein accumulation, and nitrate and nitrite are assimilated mainly in the green pericarp of the wheat grain. It appears that differences in protein contents of Shera and C-306 wheat arise primarily from differences in translocation of nitrogenous solutes from the phloem sap to the peduncle and pericarp, enroute endosperm.

Factors affecting dry weight accumulation in developing barley endosperm

Some factors that may be concerned in determining final grain weight in barley (Hordeum vulgare L. var. distichum) have been investigated. Variation in endosperm fresh and dry weight, volume and starch content have been recorded at different stages of grain development between anthesis and harvest‐ripeness for two barleys, cvs Kym and Golden Promise, differing in final grain weight. Results were recorded under both field and glasshouse conditions. The results suggest that the higher final dry weight of Kym, in comparison with Golden Promise, is a function of both rate and duration of grain filling. Only at later stages of endosperm development did the differences in volume become significant and the Kym endosperms continued to increased rapidly in volume for two to three days after endosperm volume had reached a maximum in Golden Promise. The rates of starch accumulation in both cultivars were very similar but starch deposition continued in Kym endosperms for four to five days after deposition in Golden Promise endosperms had slowed down.

Assimilate supply in relation to grain weight in wheat (Triticum aestivum L)

The contents of various free sugars in large and small wheat grains were compared with those in normal grains at different stages of grain development. Their content paralleled grain weight only upto 14 days after anthesis (A+14). On a dry weight basis, no positive correlations were observed between the concentrations of various free sugars and grain weight at any of the stages. Differences in grain weight were dependent mainly on the number of cells formed in the endosperm which in turn seemed to be regulated by the assimilate supply available to the grain during the first 14daa.

Effects of drought on yield and on grain and malt characters in spring barley

AbstractThe effects on yield and on grain and malt characters, of drought stress applied at different stages of grain development and ripening were studied in three varieties of spring barley grown in a glasshouse. Differences between variety means were found for most of the characters studied, but only for wort filtration time was there a significant interaction between variety and irrigation treatment. Grain yield was reduced by all the treatments involving drought stress and the reduction was greatest when stress was applied at heading and maintained for at least 14 days. A similar response was found for the number of ears per plant. Number of grains per main‐shoot ear was reduced by drought stress applied at heading but not when the stress was applied from 32 days after heading until harvest. Grain size was significantly reduced by all treatments. Raw‐grain characters known to be correlated with malt extract were found to be affected by the treatments: grain nitrogen content, barley extract viscosity and the rate of sedimentation of barley flour in ethanol were all increased by drought stress, the degree of response varying with the length and timing of the period of drought. Malt extracts were reduced by drought stress whether this was applied early or late in grain development. Malting loss and germinative energy were reduced, and wort filtration time increased by late stress. α‐Amylase activity in the germinating grain was unaffected by any of the treatments.

Free sugars and their relationship with grain size and starch content in developing wheat grains

AbstractFree sugars in wheat varieties, at different stages of grain development and differing in grain size and starch content per grain, were analysed to determine any relationship with grain size and starch content. Fresh and dry weight (mg per grain) of developing grain increased with maturity until 35 days after anthesis. Fresh weight declined after 35 days but dry weight remained almost constant. Water content increased until the 21 and/or 28 day stage and then decreased. Active starch synthesis started from 14 days onwards and continued until 35 days after anthesis. Free sugars (total sugars, reducing sugars, non‐reducing sugars and sucrose) per grain followed closely the changes in the rate of starch accumulation and, in general, were maximum at the 14 day stage followed by a decline up to 28 days, with no significant alteration afterwards. Presence of adequate amounts of soluble sugars in grain even when starch synthesis had stopped suggested that the supply of sugar precursors did not limit starch synthesis. None of the sugars was found to have a significant relationship with either grain size or starch content. A hypothesis explaining the termination of starch accumulation in developing cereal grains has been proposed.