Basal Spikelets Research Articles

Low sucrose availability reduces basal spikelet fertility by inducing abscisic acid and jasmonic acid synthesis in wheat.

Within a spike of wheat, the central spikelets usually generate three to four fertile florets, while the basal spikelets generate zero to one fertile floret. The physiological and transcriptional mechanism behind the difference in fertility between the basal and central spikelets is unclear. This study reports a high temporal resolution investigation of transcriptomes, number and morphology of floret primordia, and physiological traits. The W6.5-W7.5 stage was regarded as the boundary to distinguish between fertile and abortive floret primordia; those floret primordia reaching the W6.5-W7.5 stage during the differentiation phase (3-9 d after terminal spikelet stage) usually developed into fertile florets in the next dimorphism phase (12-27 d after terminal spikelet stage), whereas the others aborted. The central spikelets had a greater number of fertile florets than the basal spikelets, which was associated with more floret primordia reaching the W6.5-W7.5 stage. Physiological and transcriptional results demonstrated that the central spikelets had a higher sucrose content and lower abscisic acid (ABA) and jasmonic acid (JA) accumulation than the basal spikelets due to down-regulation of genes involved in ABA and JA synthesis. Collectively, we propose a model in which ABA and JA accumulation is induced under limiting sucrose availability (basal spikelet) through the up-regulation of genes involved in ABA and JA synthesis; this leads to floret primordia in the basal spikelets failing to reach their fertile potential (W6.5-W7.5 stage) during the differentiation phase and then aborting. This fertility repression model may also regulate spikelet fertility in other cereal crops and potentially provides genetic resources to improve spikelet fertility.

Gluten subfractions of wheat storage proteins are affected by high night temperature during grain formation

AbstractGluten (gliadin + glutenin) protein in wheat flour is affected by high temperature (day and/or night) resulting in undesirable consequences on dough quality. A study was conducted with early and late‐maturing wheat genotypes, to assess the spatial (superior‐ central and inferior‐ apical and basal spikelets) variation in the composition of gluten subfractions in the developing ear under high night temperature (HNT). We hypothesised that protein content in the superior and inferior grains may show a differential quantitative and qualitative response to HNT. HNT resulted in a significant increase in protein content which exhibited a strong (r = −0.44*) negative correlation with sedimentation volume (SV) that determines baking quality. The late‐maturity genotypes were more responsive to HNT with changes in ω‐5 and γ gliadin subfractions of both superior and inferior spikelets, though a consistent trend was not established. The proportion of high molecular weight (HMW) glutenins increased, whereas low molecular weight (LMW) glutenins reduced in most of the genotypes under HNT. Both HMW and LMW glutenins revealed significant positive (r = 0.43* and r = 0.81***, respectively) correlation with SV. The expression analysis of genes for gluten subfractions showed a significant decrease in transcript abundance of α, ω‐5, γ, HMW, and LMW fractions under HNT.

Delayed development of basal spikelets in wheat explains their increased floret abortion and rudimentary nature.

Large differences exist in the number of grains per spikelet across an individual wheat (Triticum aestivum L.) spike. The central spikelets produce the highest number of grains, while apical and basal spikelets are less productive, and the most basal spikelets are commonly only developed in rudimentary form. Basal spikelets are delayed in initiation, yet they continue to develop and produce florets. The precise timing or the cause of their abortion remains largely unknown. Here, we investigated the underlying causes of basal spikelet abortion using shading applications in the field. We found that basal spikelet abortion is likely to be the consequence of complete floret abortion, as both occur concurrently and have the same response to shading treatments. We detected no differences in assimilate availability across the spike. Instead, we show that the reduced developmental age of basal florets pre-anthesis is strongly associated with their increased abortion. Using the developmental age pre-abortion, we were able to predict final grain set per spikelet across the spike, alongside the characteristic gradient in the number of grains from basal to central spikelets. Future efforts to improve spikelet homogeneity across the spike could thus focus on improving basal spikelet establishment and increasing floret development rates pre-abortion.

Late spring cold reduces grain number at various spike positions by regulating spike growth and assimilate distribution in winter wheat

Late spring cold (LSC) occurred in the reproductive period of wheat impairs spike and floret differentiation during the reproductive period, when young spikelets are very cold-sensitive. However, under LSC, the responses of wheat spikelets at various positions, leaves, and stems and the interactions between them at physiological levels remain unclear. In the present study, two-year treatments at terminal spikelet stage under two temperatures (2 °C, −2 °C) and durations (1, 2, and 3 days) were imposed in an artificial climate chamber to compare the effects of LSC on grain number and yield in the wheat cultivars Yannong 19 (YN19, cold-tolerant) and Xinmai 26 (XM26, cold-sensitive). The night temperature regimes were designed to reproduce natural temperature variation. LSC delayed plant growth and inhibited spike and floret differentiation, leading to high yield losses in both cultivars. LSC reduced dry matter accumulation (DMA, g) in spikes, stems, and leaves, reducing the DMA ratios of the spike to leaf and spike to stem. Plant cell wall invertase (CWINV) activity increased in upper and basal spikelets in YN19, whereas CWINV increased in middle spikelets in XM26. Under LSC, soluble sugar and glucose were transported and distributed mainly in upper and basal spikelets for glume and rachis development, so that spike development was relatively complete in YN19, whereas the upper and basal spikelets were severely damaged and most of the glumes in middle spikelets were relatively completely developed in XM26, resulting in pollen abortion mainly in upper and basal spikelets. The development of glumes and rachides was influenced and grain number per spike was decreased after LSC, with kernels present mainly in middle spikelets. Overall, reduced total DMA and dry matter partitioning to spikes under LSC results in poor spikelet development, leading to high losses of grain yield.

Genetic association of spikelet abortion with spike, grain, and shoot traits in highly-diverse six-rowed barley.

Spikelet abortion is a phenomenon where apical spikelet primordia on an immature spike abort. Regardless of the row-type, both apical and basal spikelet abortion occurs, and their extent decides the number of grain-bearing spikelets retained on the spike-thus, affecting the yield potential of barley. Reducing spikelet abortion, therefore, represents an opportunity to increase barley yields. Here, we investigated the variation for apical spikelet abortion along with 16 major spike, shoot, and grain traits in a panel of 417 six-rowed spring barleys. Our analyses showed a significantly large genotypic variation resulting in high heritability estimates for all the traits. Spikelet abortion (SA) varies from 13 to 51% depending on the genotype and its geographical origin. Among the seven spike traits, SA was negatively correlated with final spikelet number, spike length and density, while positively with awn length. This positive correlation suggests a plausible role of the rapidly growing awns during the spikelet abortion process, especially after Waddington stage 5. In addition, SA also showed a moderate positive correlation with grain length, grain area and thousand-grain weight. Our hierarchical clustering revealed distinct genetic underpinning of grain traits from the spike and shoot traits. Trait associations showed a geographical bias whereby European accessions displayed higher SA and grain and shoot trait values, whereas the trend was opposite for the Asian accessions. To study the observed phenotypic variation of SA explained by 16 other individual traits, we applied linear, quadratic, and generalized additive regression models (GAM). Our analyses of SA revealed that the GAM generally performed superior in comparison to the other models. The genetic interactions among traits suggest novel breeding targets and easy-to-phenotype "proxy-traits" for high throughput on-field selection for grain yield, especially in early generations of barley breeding programs.

High expression of the MADS-box gene VRT2 increases the number of rudimentary basal spikelets in wheat.

Spikelets are the fundamental building blocks of Poaceae inflorescences, and their development and branching patterns determine the various inflorescence architectures and grain yield of grasses. In wheat (Triticum aestivum), the central spikelets produce the most and largest grains, while spikelet size gradually decreases acropetally and basipetally, giving rise to the characteristic lanceolate shape of wheat spikes. The acropetal gradient corresponds with the developmental age of spikelets; however, the basal spikelets are developed first, and the cause of their small size and rudimentary development is unclear. Here, we adapted G&T-seq, a low-input transcriptomics approach, to characterize gene expression profiles within spatial sections of individual spikes before and after the establishment of the lanceolate shape. We observed larger differences in gene expression profiles between the apical, central, and basal sections of a single spike than between any section belonging to consecutive developmental time points. We found that SHORT VEGETATIVE PHASE MADS-box transcription factors, including VEGETATIVE TO REPRODUCTIVE TRANSITION 2 (VRT-A2), are expressed highest in the basal section of the wheat spike and display the opposite expression gradient to flowering E-class SEPALLATA1 genes. Based on multi-year field trials and transgenic lines, we show that higher expression of VRT-A2 in the basal sections of the spike is associated with increased numbers of rudimentary basal spikelets. Our results, supported by computational modeling, suggest that the delayed transition of basal spikelets from vegetative to floral developmental programs results in the lanceolate shape of wheat spikes. This study highlights the value of spatially resolved transcriptomics to gain insights into developmental genetics pathways of grass inflorescences.

Effect of Zn and sucrose supply on grain Zn, Fe and protein contents within wheat spike under detached-ear culture

Wheat (Triticum aestivum L.) is one of most widely consumed cereal crops globally. More than two billion people are affected by nutritional deficiencies caused by zinc (Zn) and iron (Fe) deficiency. Therefore, biofortification is a strategy supposed to be economical, sustainable and easily implemented to increase the micronutrient content in the crops. However, micronutrient distribution, especially Zn and Fe, within the spike has been rarely reported. For this, different concentrations of Zn (0, 30, 60, 90 and 120 μmol L–1) and sucrose (2%, 4%, 6%) were supplied at Zn concentrations of 30 and 90 μmol L–1 under detached-ear culture. The results showed that grain weight and grain Zn, Fe and protein contents were higher in superior grains than in inferior grains, and higher in central spikelets than in basal and apical spikelets. Zn supply significantly improved grain Zn, Fe and protein content. Increasing sucrose concentration at low and high Zn supply levels significantly increased grain weight, and grain Zn, Fe and protein contents. The percentage increase in dry weight and nutrient contents of superior and inferior grains was higher in central spikelets than in apical and basal spikelets. It is concluded that grain position significantly affected the grain weight and nutrient content, and these effects could be improved by application of Zn and sucrose. These findings suggest Zn and sucrose application for increasing mineral nutrients contents in wheat grain, and mechanisms of nutrient accumulation in relation to external sucrose and Zn supply should be elucidated in the future.

Identification of novel QTLs for grain fertility and associated traits to decipher poor grain filling of basal spikelets in dense panicle rice

High grain number is positively correlated with grain yield in rice, but it is compromised because of poor filling of basal spikelets in dense panicle bearing numerous spikelets. The phenomenon that turns the basal spikelets of compact panicle sterile in rice is largely unknown. In order to understand the factor(s) that possibly determines such spikelet sterility in compact panicle cultivars, QTLs and candidate genes were identified for spikelet fertility and associated traits like panicle compactness, and ethylene production that significantly influences the grain filling using recombinant inbred lines developed from a cross between indica rice cultivars, PDK Shriram (compact, high spikelet number) and Heera (lax, low spikelet number). Novel QTLs, qSFP1.1, qSFP3.1, and qSFP6.1 for spikelet fertility percentage; qIGS3.2 and qIGS4.1 for panicle compactness; and qETH1.2, qETH3.1, and qETH4.1 for ethylene production were consistently identified in both kharif seasons of 2017 and 2018. The comparative expression analysis of candidate genes like ERF3, AP2-like ethylene-responsive transcription factor, EREBP, GBSS1, E3 ubiquitin-protein ligase GW2, and LRR receptor-like serine/threonine-protein kinase ERL1 associated with identified QTLs revealed their role in poor grain filling of basal spikelets in a dense panicle. These candidate genes thus could be important for improving grain filling in compact-panicle rice cultivars through biotechnological interventions.

Cell cycle events and expression of cell cycle regulators are determining factors in differential grain filling in rice spikelets based on their spatial location on compact panicles.

Rice being a staple crop for human, its production is required to be increased significantly, particularly keeping in view the expected world's population of 9.6 billion by the year 2050. In this context, although the rice breeding programs have been successful in increasing the number of spikelets per panicle, the basal spikelets remain poorly filled, undermining the yield potential. The present study also found the grain filling to bear negative correlation with the panicle grain density. The poorly filled basal spikelets of the compact-panicle cultivars showed a lower endosperm cell division rate and ploidy status of the endosperm nuclei coupled with no significant greater expression of CYCB;1 and CYCH;1 compared with the apical spikelets, unlike that observed in the lax-panicle cultivars, which might have prevented them from overcoming apical dominance. Significantly greater expression of CYCB2;2 in the basal spikelets than in the apical spikelets might also have prevented the former to enter into endoreduplication. Furthermore, expression studies of KRPs in the caryopses revealed that a higher expression of KRP;1 and KRP;4 in the basal spikelets than in the apical spikelets of the compact-panicle cultivars could also be detrimental to grain filling in the former, as KRPs form complex primarily with CDKA-CYCD that promotes S-phase activity and G1/S transition, and thus inhibits endosperm cell division. The study indicates that targeted manipulation of expression of CYCB1;1, CYCB2;2, CYCH1;1, KRP;1 and KRP4 in the basal spikelets of the compact-panicle cultivars may significantly improve their yield performance.

Physicochemical and milling properties of rice kernels from upper, middle, and basal spikelets of hybrid and inbred lines at early and ideal harvesting stages

AbstractBackground and objectivesGrains from a hybrid (Inov CL) and an inbred (Irga 424 RI) line were studied as a function of harvesting time (supposed early and ideal maturity stages) and panicle position (upper, middle, and basal).FindingsIn general, greater head rice yield (HRY) and thousand grain weight were observed when the kernels were harvested with 22% moisture content. At this moisture content, HRY varied from 52.3% to 63.8% for Inov CL and from 68.2% to 69.8% for Irga 424 RI, depending on panicle position. All studied starches showed similar molecular weight distribution. Relative crystallinity varied from 18.7% to 22.0%. The protein content and extractability at different media differed just for kernels from Inov CL at the 2nd harvesting time.ConclusionsTo achieve better industrial performance, the rice kernels may be harvested at the ideal maturity stage of 22% moisture content or closer. Amylose content and starch structure of the Inov CL and Irga 424 RI cultivars did not differ among the studied harvesting times.Significance and noveltyObtained results improve the scientific database on factors affecting the grain quality of hybrid and inbred rice cultivars, addressing gaps that may be considered for farmers, industries, as well as for rice breeders in future rice breeding programs.

Differential grain filling in apical and basal spikelets of compact panicle rice is associated with difference in expression of miRNAs targeting gene products involved in grain filling

Development of rice cultivars bearing numerous spikelets by breeding approach to increase production of the crop to feed the ever increasing human population accompanies poor filling of grains in their basal spikelets preventing achievement of the yield potential. To understand the reason of such poor grain filling at the level of the initial biochemical events, 5 RACE library was prepared taking RNA extracted from the spikelets of two indica rice cultivars, a compact-panicle Mahalaxmi bearing numerous spikelets per panicle (>350) and a lax-panicle Upahar bearing fewer spikelets (<250) per panicle, and the presence of the miRNA cleaved gene products involved in grain filling was identified by running PCR using gene-specific primers. Bioinformatics analysis was conducted to find if the grain filling enzymes were targeted by miRNAs. The effort led to identification of three enzymes, namely granule bound starch synthase, invertase and sucrose phosphate synthase, the transcripts of which were targeted by the miRNAs osa-miR160a-5p, osa-miR166e-3p and osa-miR156b-5p, respectively. The expressions of the three miRNAs were confirmed by Northern in the apical and basal spikelets of the two cultivars. The expression analysis of the three miRNAs further revealed that these were expressed significantly more in the basal spikelets than in the apical ones of Mahalaxmi, but not of Upahar. Real time PCR of the target genes showed an opposite result, i.e., these were expressed significantly less in the basal spikelets than in the apical ones of Mahalaxmi, but mostly not of Upahar. The finding led to the conclusion that the grain filling in rice could be regulated at the level of miRNAs, and manipulation of their expression could be proved an important tool in improving grain filling in compact-panicle rice cultivars.

Radiotracer and radiation application for rapid measurement of contribution of stem assimilates towards grain filling and for alleviating terminal heat stress in wheat

Contribution of stem photoassimilates towards grain filling may have significance under stress, since it may force the plant to initiate an early leaf senescence, thereby limiting the availability of flag leaf photosynthates for grain filling. Radiotracer feeding of 14C-sucrose to the internode by stem injection method at 20 days stage after anthesis and measurement of 14C activity in grains at 5 days after foliar application of potassium iodide, meant to inhibit current photosynthesis by inducing leaf senescence, showed a gradual decline in flag leaf chlorophyll but an increase in accumulation of 14C particularly in the basal spikelet grains. However, a reduction in grain filling at all the sink positions within the wheat ear was measured with hastened leaf senescence. On the other hand, abiotic stresses such as high temperature stress, is known to cause an abortion of the basal florets in wheat and induce an early flag leaf senescence. We have earlier shown that seed gamma irradiation, at 0.1 kGy causes an increase in grain mass at middle and basal spikelet positions. In this paper we report that the reduction in grain yield in response to terminal heat stress can be compensated by gamma irradiation, owing to better stem reserve mobilization into the basal and middle spikelet under condition of stress induced senescence.

A genome-wide associate study reveals favorable alleles conferring apical and basal spikelet fertility in wheat (Triticum aestivum L.)

Kernel number per spike (KNPS) is one of the key factors affecting wheat yield, which can be significantly reduced by lower fertility or sterility of the apical and basal spikelets. In this study, the spikelet number per spike (SNPS), thousand kernel weight (TKW), KNPS, total grain numbers of the top three apical spikelets (GNAS), and total grain numbers of the bottom three basal spikelets (GNBS) of 212 wheat lines were recorded from five different environmental conditions. These 212 accessions were genotyped using the 9K iSelect SNP Beadchip. A total of 3269 SNP markers were used for genome-wide association analysis (GWAS). One hundred twelve significant marker-trait associations (MTAs) were identified. Twenty-two MTAs were identified in at least two environments and two of them showed association with two or more grain setting properties. Different loci showed an additive effect with both GNAS and GNBS being much higher in the lines with more favorite alleles. Two SNP loci, wsnp_Ex_c31799_40545376 and wsnp_BF293620A_Ta_2_3, showed the largest effects on increasing KNPS through improved fertility of apical and basal spikelets, respectively. These MTAs have the potential to be used in future marker-assisted selection.

Biochemical and molecular characterisation of exogenous cytokinin application on grain filling in rice

BackgroundPoor filling of grains in the basal spikelets of large size panicles bearing numerous spikelets has been a major limitation in attempts to increase the rice production to feed the world’s increasing population. Considering that biotechnological intervention could play important role in overcoming this limitation, the role of cytokinin in grain filling was investigated based on the information on cell proliferating potential of the hormone and reports of its high accumulation in immature seeds.ResultsA comparative study considering two rice varieties differing in panicle compactness, lax-panicle Upahar and compact-panicle OR-1918, revealed significant difference in grain filling, cytokinin oxidase (CKX) activity and expression, and expression of cell cycle regulators and cytokinin signaling components between the basal and apical spikelets of OR-1918, but not of Upahar. Exogenous application of cytokinin (6-Benzylaminopurine, BAP) to OR-1918 improved grain filling significantly, and this was accompanied by a significant decrease in expression and activity of CKX, particularly in the basal spikelets where the activity of CKX was significantly higher than that in the apical spikelets. Cytokinin application also resulted in significant increase in expression of cell cycle regulators like cyclin dependent kinases and cyclins in the basal spikelets that might be facilitating cell division in the endosperm cells by promoting G1/S phase and G2/M phase transition leading to improvement in grain filling. Expression studies of type-A response regulator (RR) component of cytokinin signaling indicated possible role of OsRR3, OsRR4 and OsRR6 as repressors of CKX expression, much needed for an increased accumulation of CK in cells. Furthermore, the observed effect of BAP might not be solely because of it, but also because of induced synthesis of trans-zeatin (tZ) and N6-(Δ2-isopentenyl)adenine (iP), as reflected from accumulation of tZR (tZ riboside) and iPR (iP riboside), and significantly enhanced expression of an isopentenyl transferase (IPT) isoform.ConclusionThe results suggested that seed-specific overexpression of OsRR4 and OsRR6, and more importantly of IPT9 could be an effective biotechnological intervention towards improving the CK level of the developing caryopses leading to enhanced grain filling in rice cultivars bearing large panicles with numerous spikelets, and thereby increasing their yield potential.

Ethylene production and constitutive expression of ethylene receptors and ethylene signal transduction during grain filling in apical and basal spikelets of compact-and lax-panicle rice (Oryza sativa) cultivars

Ethylene production and constitutive expression of ethylene receptors and ethylene signal transduction during grain filling in apical and basal spikelets of compact-and lax-panicle rice (Oryza sativa) cultivars

MicroRNA172 plays a crucial role in wheat spike morphogenesis and grain threshability.

Wheat domestication from wild species involved mutations in the Q gene. The q allele (wild wheats) is associated with elongated spikes and hulled grains, whereas the mutant Q allele (domesticated wheats) confers subcompact spikes and free-threshing grains. Previous studies showed that Q encodes an AP2-like transcription factor, but the causal polymorphism of the domestication traits remained unclear. Here, we show that the interaction between microRNA172 (miR172) and the Q allele is reduced by a single nucleotide polymorphism in the miRNA binding site. Inhibition of miR172 activity by a miRNA target mimic resulted in compact spikes and transition from glumes to florets in apical spikelets. By contrast, overexpression of miR172 was sufficient to induce elongated spikes and non-free-threshing grains, similar to those observed in three Q loss-of-function mutations. These lines showed transitions from florets to glumes in the basal spikelets. These localized homeotic changes were associated with opposing miR172/Q gradients along the spike. We propose that the selection of a nucleotide change at the miR172 binding site of Q contributed to subcompact spikes and free-threshing grains during wheat domestication.

Eleocharis pedrovianae, a new species of Cyperaceae from Northern Brazil (Serra dos Carajás, Pará State)

Eleocharis pedrovianae is a new species of Cyperaceae from the Serra dos Carajás (Pará State, Brazil), herein described and illustrated. This species lives as a submerged herb in wetlands, marshes and margins of temporary lakes and lagoons with ferric soil (canga). It is morphologically similar to Eleocharis angustispicula, differing by its stoloniferous stem system, the presence of chasmogamous basal spikelets, a perianth of 4(5) bristles, an ellipsoid achene, and a stylopodium that is brown pyramidal, and separated from the achene body by a constriction.

Wheat Grain Yield Increase in Response to Pre-Anthesis Foliar Application of 6-Benzylaminopurine Is Dependent on Floret Development.

Wheat yield is largely determined during the period prior to flowering, when the final numbers of fertile florets and grains per spike are established. The aim of this study was to assess the dynamics of floret primordia development in winter wheat in response to pre-anthesis application of a synthetic cytokinin, 6-benzylaminopurine (6-BA). We conducted an experiment in which two foliar spray treatments were applied (water or 6-BA) to Chinese winter wheat at 25 days after jointing during two growing seasons (2012–2013 and 2013–2014). Both the final grain number per spike and grain yield at maturity exhibited remarkable increases in response to the 6-BA treatment. Application of 6-BA increased the number of fertile florets in basal spikelets and, to a greater extent, in central spikelets. The mechanism by which 6-BA application affected the final number of fertile florets primarily involved suppression of the floret abortion rates. Application of 6-BA considerably reduced the abortion rates of basal, central and apical spikelet florets (by as much as 77% compared with the control), as well as the degeneration rates of basal and central spikelet florets, albeit to a lesser degree. The effect of 6-BA application on the likelihood of proximal florets being set was limited to the distal florets in the whole spike, whereas obvious increases in the likelihood of grain set under 6-BA treatment were observed in distal florets, primarily in central spikelet positions. The results of this study provide important evidence that 6-BA application to florets (final fertile floret production) results in an increased grain yield.

1-MCP treatment enhanced expression of genes controlling endosperm cell division and starch biosynthesis for improvement of grain filling in a dense-panicle rice cultivar

High ethylene production in dense-panicle rice cultivars impacts grain filling. 1-MCP (ethylene action inhibitor) treatment increased assimilates partitioning, cell number and size and expression of starch synthesizing enzyme genes of developing caryopses mostly in the basal spikelets of panicle at early post-anthesis stage. The gain in cell number was less compared to the increase of size. High ethylene production in spikelets matched with greater expression of ethylene receptor and signal transducer genes. Genes encoding cell cycle regulators CDK, CYC and CKI expressed poorly on 9 DAA. 1-MCP treatment enhanced their expression; the increase of expression was higher for CDKs and lower for CKIs in basal compared to apical spikelets. Greater expression of CDKB2:1 might have lifted cytokinesis of nascent peripheral cells of endosperm, while promotion of CDKAs, CYCD2:2 and inhibition of CYCB2:2 expression contributed to endoreduplication of central cells increasing cell size and DNA ploidy level. It is concluded that the process of endoreduplication, which begins at mid-grain filling stage, is crucially linked with the final caryopsis size of rice grain. The enhanced endosperm growth brought about by repressed ethylene action during the first few days after anthesis seems to be associated with the overall increased cell cycle activity and sink strength.

Dynamics of Expression of Ethylene Signal Transducer and Starch Synthesizing Enzyme Genes in Developing Spikelets of Rice Panicle Contrasting for Grain Growth

Spikelets are individual flower units of the complex panicle type inflorescence of rice. Each spikelet is genetically competent to bear a well-filled grain at maturity. However, heterogeneous grain development underscores this potential and spikelets located on the basal part of panicle do not bear grains suitable for human consumption. The problem of poor grain filling is more acute in the newly introduced large-panicle super rice. My research was intended to find out the chemical routes for enhancing grain filling in the inferior basal spikelets. The research ruled out discrimination in photosynthetic assimilate supply, but identified ethylene as being responsible for variation in grain filling capacity among spikelets of rice panicle. Inferior spikelets produce more ethylene impeding the activities of starch synthesizing enzymes to the detriment of grain filling, as a result of which assimilates not used in growth accumulate in the developing kernel. In contrast, superior spikelets located on apical branches of panicle produce less ethylene and fill faster because of metabolic dominance in starch biosynthesis.