Superior Spikelets Research Articles

Post-Anthesis Changes in Concentration of Polyamine in Superior and Inferior Spikelets and Their Relationship with Grain Filling of Super Rice

To reveal the physiological role of polyamines and the mechanism involved in grain-filling of rice ( Oryza sativa L.) superior and inferior spikelets, the relationship between grain-filling rate and polyamine concentrations was analyzed and verified using 4 super rice cultivars and 2 high-yielding cultivars (controls). Compared with the controls, super rice had similar maximum grain-filling rate, mean grain-filling rate, and brown rice weight in superior spikelets and significantly lower values in inferior spikelets. The concentrations of putrescine (Put), spermidine (Spd), and spermine (Spm) varied in single-peak curves in both inferior and superior spikelets during grain-filling period. The peak and mean concentrations of Put were greater in inferior spikelets than in superior spikelets. However, the peak and mean concentrations of Spd and Spm were greater in superior spikelets than in inferior spikelets. The control cultivars had higher concentrations of polyamines than super rice. The mean grain-filling rate and brown rice weight were negatively correlated with Put concentration, and positively correlated with Spd and Spm concentrations and the ratios of Spd/Put and Spm/Put. Spraying of Spd or Spm to panicle at early grain-filling stage significantly enhanced activities of sucrose synthetase, adenine diphosphoglucose pyrophosphorylase, and soluble starch synthetase in inferior spikelets, resulting in improvement of grain-filling rate, seed-setting rate, and grain weight. Application of Put or methylglyoxal-bis (guanylhydrazone) (MGBG, an inhibitor of S-adenosyl- L-methioninedecarboxylase) exhibited an opposite effect. These results indicated a regulative function of polyamines during grain filling. The low grain-filling rate and grain weight in inferior spikelets of super rice are probably in relation to the low concentrations of Spd and Spm and the small ratios of Spd/Put and Spm/Put in inferior spikelets.

Hormones in the grains and roots in relation to post-anthesis development of inferior and superior spikelets in japonica/indica hybrid rice

Grain filling is usually not adequate in later-flowering inferior spikelets in japonica/indica (J/I) hybrid rice (Oryza sativa) although it shows stronger hybrid vigor than indica/indica (I/I) hybrid. This study investigated the potential causes by examining changes in zeatin (Z)+zeatin riboside (ZR), indole-3-acetic acid (IAA), gibberellins (GAs, GA(1)+GA(4)), and abscisic acid (ABA) in spikelets and roots during the grain filling period. The inferior spikelets of J/I hybrid exhibited low rate of endosperm cell division and slow grain filling. During the early grain filing period, they had less Z+ZR, IAA, and ABA, but more GAs, than the earlier-flowering superior spikelets. If compared to the inferior spikelets of the I/I hybrid, the J/I inferior spikelets also had less Z+ZR, IAA, and ABA. Rates of endosperm cell division and grain filling were positively and significantly correlated with Z+ZR and ABA contents in both grains and roots or IAA in grains, whereas not significantly correlated with GAs either in grains or roots or IAA in roots. Applications of kinetin, IAA, or ABA to spikelets, or kinetin and ABA to roots, enhanced cell division and grain filling in the inferior spikelets. Results suggest that low contents of cytokinins and ABA in both grains and roots and low contents of IAA in grains may result in the poor filling of inferior spikelets in the J/I hybrid.

Involvement of polyamines in the post-anthesis development of inferior and superior spikelets in rice

Early-flowered superior spikelets usually exhibit a faster grain filling rate and heavier grain weight than late-flowered inferior spikelets in rice (Oryza sativa L.). But the intrinsic factors responsible for the variations between the two types of spikelets are unclear. This study investigated whether and how polyamines (PAs) are involved in regulating post-anthesis development of rice spikelets. Six rice genotypes differing in grain filling rate were field grown, and PA levels and activities of the enzymes involved in PA biosynthesis were measured in both superior and inferior spikelets. The results showed that superior spikelets exhibited higher levels of free spermidine (Spd) and free spermine (Spm) and higher activities of arginine decarboxylase (ADC, EC 4.1.1.19), S-adenosylmethionine decarboxylase (SAMDC, EC 4.1.1.50) and Spd synthase (EC 2.5.1.16) than inferior spikelets at the early endosperm cell division and grain filling stage. The maximum concentrations of free Spd and free Spm and the maximum activities of ADC, SAMDC and Spd synthase were significantly correlated with the maximum cell division and grain filling rates, maximum cell number and grain weight. Application of Spd and Spm to panicles resulted in significantly higher rates of endosperm cell division and grain filling in inferior spikelets along with the activities of sucrose synthase (EC 2.4.1.13), ADP glucose pyrophosphorylase (EC 2.7.7.27) and soluble starch synthase (EC 2.4.1.21), suggesting that these PAs are involved in the sucrose-starch metabolic pathway. The results indicate that the poor development of inferior spikelets is attributed, at least partly, to the low PA level and its low biosynthetic activity.

Hormonal changes are related to the poor grain filling in the inferior spikelets of rice cultivated under non-flooded and mulched condition

Non-flooded plastic mulching cultivation (PM) of rice can save much irrigation but usually exhibits a poor grain filling and low grain weight when compared to traditional flooding cultivation (TF). This study measured the variations of plant hormones during grain filling and investigated whether they were related to the grain-filling problem under mulching in a field experiment. Hormonal levels in the grains and the grain development of both superior (early flowered) and inferior spikelets (later flowered) were monitored continuously during the grain filling period. The contents of indole-3-acetic acid (IAA), zeatin riboside (ZR) and abscisic acid (ABA) were higher in superior grains than in inferior ones at early grain-filling stage. For individual spikelets, the peaks of IAA and ZR contents appeared just before the peak grain-filling rate but the peak ABA content matched it. The earlier flowered superior spikelets did not show much difference in grain filling characteristics and hormonal changes under the two cultivation treatments. Distinctively with the inferior spikelets in a panicle, PM led to lower grain weight, higher grain-filling rate at early stage and shorter active grain-filling period than the TF. PM also resulted in less IAA and ZR but more ABA contents in the grains than TF at early and middle grain filling stages in these inferior spikelets. Peaks of these hormone contents in the PM inferior grains usually appeared earlier and also disappeared more quickly than those in TF inferior grains. When exogenous IAA was applied to plants at the initial grain-filling stage, IAA and ZR were increased and ABA was reduced in the inferior grains. The treatment prolonged the active grain-filing period and reduced the peak grain-filling rate of inferior grains, similarly as the case with TF. As a result, IAA spraying enhanced the weight of PM inferior grains but reduced that of TF ones. Exogenous ABA spraying led to the opposite effects to those of IAA. The results suggest that the shortened grain-filling period in the inferior grains under PM is related to the increased ABA and reduced IAA and ZR in the grains. Regulation of the ratio of ABA to IAA in grains could potentially increase the weight of inferior grains under the water-saving cultivation with film mulching.

Post-anthesis development of inferior and superior spikelets in rice in relation to abscisic acid and ethylene

The purpose of this study was to test the hypothesis that the interaction between abscisic acid (ABA) and ethylene may be involved in mediating the post-anthesis development of spikelets in rice (Oryza sativa L.). Two rice genotypes were field-grown, and the changes of ABA, ethylene, and 1-aminocylopropane-1-carboxylic acid (ACC) levels in spikelets during grain filling and their relationships with endosperm-division and grain-filling rates were investigated. The results showed that earlier-flowering superior spikelets exerted dominance over later-flowering inferior spikelets in endosperm cell-division and grain-filling rates. The two genotypes behaved the same. Later-flowering spikelets had higher levels of ethylene and ACC than earlier-flowering spikelets. The ethylene evolution rate was significantly and negatively correlated with the cell division and grain filling rates. By contrast to ethylene, later-flowering spikelets contained a lower ABA content/concentration and showed a low content ratio of ABA to ACC than earlier-flowering ones. The cell-division and grain-filling rates were significantly and positively correlated with both ABA contents and the ratio of ABA to ACC. Application of cobalt ion (inhibitor of ethylene synthesis) or ABA at an early grain-filling stage significantly increased endosperm cell division rate and cell number, grain-filling rate, and grain weight of inferior spikelets. Application of ethephon (an ethylene-releasing agent) or fluridone (an inhibitor of carotenoid synthesis) had the opposite effect. The results suggest that antagonistic interactions between ABA and ethylene mediate endosperm cell-division and grain-filling in rice. A higher ratio of ABA to ethylene in rice spikelets is required to maintain a faster grain-filling rate.

Hormones in the grains in relation to sink strength and postanthesis development of spikelets in rice

Inferior spikelets usually exhibit a slower grain filling rate and lower grain weight than superior spikelets in a rice (Oryza sativa L.) panicle. This study investigated whether the variations in grain filling between the two kinds of spikelets were attributed to their sink strength and whether the sink strength was regulated by the hormonal levels in the grains. Using two field-grown rice genotypes, the division rate of endosperm cells, hormonal levels in the grains, and grain weight of both superior and inferior spikelets were determined during the grain filling period. The results showed that superior spikelets had dominance over inferior spikelets in endosperm cell division rate and cell number, grain filling and grain weight. Changes in zeatin (Z) and zeatin riboside (ZR) contents paralleled and were very significantly correlated with the cell division rate and cell number. Cell division rate and the content of indole-3-acetic acid (IAA) in the grains were also significantly correlated. Gibberellin (GAs; GA1+ GA4) content of the grains was high but ABA levels were low at the early grain filling stage. ABA increased substantially during the linear phase of grain growth and was very significantly correlated with grain dry weight during this period. Application of kinetin at 2 through 6 days post anthesis (DPA) significantly increased cell number, while spraying ABA at 11 through 15 DPA significantly increased the grain filling rate. The results suggest that differences in sink strength are responsible for variations in grain filling between superior and inferior spikelets. Both cytokinins and IAA in the grains may mediate cell division in rice endosperm at early grain filling stages, and therefore regulate the sink size of the grain, whereas ABA content correlates with sink activity during the linear period of grain growth.

Effect of irradiance on the partitioning of assimilated carbon during the early phase of grain filling in rice.

Low irradiance in the early phase of grain filling in rice often results in a low grain yield, but its effects on the partitioning of previously or recently assimilated carbon within the plant or panicle have not been seriously examined. The objective of this study was to demonstrate the effect of shading during the different stages in the early phase of grain filling on the partitioning of previously or recently assimilated carbon among constituent organs and into superior and inferior spikelets of the panicle in rice (Oryza sativa L. 'Sasanishiki') plants using 13C as a tracer. Plants were grown either under low (shading) or moderate (non-shading) irradiance (120 and 800 micromol quantum m(-2) s(-1)) for 3 or 4 d before or after the 13CO2 feeding at heading, full-heading or milky stages during the early phase of grain filling. Four days after the 13CO2 feeding, the proportion of labelled (previously assimilated) carbon partitioned into the panicle was 17% higher in plants grown under low irradiance compared with plants grown under moderate irradiance at the full-heading stage (7-11 d after heading), while the proportion partitioned into the culm was 13% lower. The light treatments for 3 d were conducted before the 13CO2 feeding and partitioning of the labelled (recently-assimilated) carbon into spikelets was examined 6 h after feeding. The amount of labelled carbon partitioned into the spikelets of the secondary branch (inferior grains) in the plants grown under low irradiance was only 31% when compared with plants grown under moderate irradiance at the full-heading stage, although the partitioning of labelled carbon into the apical spikelets of the primary branch (superior grains) was not affected by the light treatments. These results clearly indicate that preferential partitioning of assimilated carbon into the panicle occurs under low irradiance at around 7-11 d after heading and that the priority of superior spikelets for assimilated carbon intensifies. This phenomenon is thought to be an important strategy for such rice cultivars as used in this study to achieve a certain proportion of ripened grains even under light limited conditions.

Dynamics of Abscisic Acid Levels during Grain-Filling in Rice : Comparisons between Superior and Inferior Spikelets

In a rice panicle, the superior spikelets display higher growth rates than the inferior spikelets during the initial phase of grain-filling. To better understand the regulatory mechanism of this phenomenon, we examined the dynamics of endogenous abscisic-acid (ABA) levels and the effects of shading on the dynamics during the grain-filling period in superior and inferior spikelets. While ABA content in the superior spikelets increased rapidly after flowering, that in the inferior spikelets increased slowly and reached the maximum later than in the superior spikelets. Shading significantly exaggerated the inherently different ABA dynamics between the superior and inferior spikelets. We concluded that ABA levels are correlated with the dry matter accumulation patterns of spikelets. These results support our hypothesis that ABA is involved in assimilate partitioning among spikelets in a panicle during the initial phase of grain-filling of rice.

Glutelin accumulation and changes in the levels of its mRNA in the superior and inferior spikelets of rice ear during ripening

Glutelin accumulation and changes in the levels of its mRNA in the superior and inferior spikelets of rice ear during ripening

Nitrogen accumulation in the inferior spikelet of rice ear during ripening

Abstract Nitrogen accumulation in the apical spikelet of the top primary branch (superior spikelet) and the second spikelet of the lowest secondary branch (inferior spikelet), of the ear on the main stem of rice plant (Oryza sativa L. var. Sasanishiki) was characterized during grain filling. In the superior spikelet, the accumulation of dry matter and nitrogen which started immediately after lowering, proceeded rapidly, and reached the maturation level at 20 d after heading. In the inferior spikelet, however, the amount of dry matter and nitrogen accumulation was minimal immediately after flowering. It increased when grain filling of the superior spikelet was almost completed. 15N-labeled ammonia was administered to the plants at different stages of ripening and the amount of incorporation in the spikelets was analyzed at harvest. The labeled nitrogen administered at the early stages of ripening was the main source of the labeled nitrogen incorporated in the superior spikelet. However, the labeled nitroge...

Studies on the Developmental Physiology in Rice Pollen : III. Effects of excessive nitrogen top-dressed at meiotic stage on the amino acid metabolism and the development of pollens

These experiments were carried out to throw light upon a phenomenon of the increased sterility by top-dressing of excessive nitrogen at meiotic stage (hereinafter referred to as nitrogen treatment) in rice plants from the view of functions and structural characteristics of pollens as well as proline metabolism in anthers. It was found that the nitrogen treatment increased the sterility in both superior spikelets (spikelets of the 4th, 5th and 6th position on primary branches) and inferior ones (spikelets of the 2nd and 3rd position on secondary branches) to three times as much as control. On and after trinuclear period (III period), the content of proline in anthers was decreased conspicuously with nitrogen treatment, especially in the anthers of inferior spikelets in which the sterility reached about 40 percent. Morphological observations on the development of tapetum after nitrogen treatment indicated that there were no distinctive differences in behaviors of tapetum between control and treated anthers. In the late III period of pollen development, there were a lot of pollens without deformation of the vegetative nucleus and with a higher staining ability of the nucleoli by aceto-carmine in the treated anthers. These pollens were evidently smaller in diameter than the control. This supressed development was a consequence of the reduced growth rate of pollens in III period. Additionally, the nitrogen treatment also reduced the pollen germination. These results lead to the conclusion that the increased sterility of spikelets by nitrogen treatment belongs to the male sterility which may be caused by a disturbance of the developmental pattern of pollen at III period, when being under the control of two sperm nuclei.

Studies on Tissue of Endosperm and Developmental Process in Opaque Kernel

In this studies we observed on opaque kernel, especially green opaque kernels. The results are summarized as follows: 1) Opaque kernels are classiffied in several types according to its shapesas green opaque kernel (type 6, type 7-1, type 7-2, ), white opaque kernel, small translucent green kernel, and monstrous kernel. Tissue and outlook of green opaque kernel are shown in Figure 1 to Figure 12. Type 6 and type 7 showing green opaque kernel, have immature parts (starch-less tissue) in endosperm as in Figure 4, 6, 8, 10, 12. 2) Developmental process of kernels are classified into 6 types showing in Table 1. According to Table 1, early growth of opaque kernels are suppressed by the competition of superior kernels. And in the late ripening stage at 17 days after flowering, opaque kernels grow comparatively rapid. 3) From the incressing curve of 1000 kernels weight of inferior spikelet, we observed the competition of growth of kernel among inferior spikelet and superior spikelet as in Figure 9. 4) Protein content of opaque kernels are always richer than ripened kernels. 5) If we could promote growth of opaque kernel in late stage of ripening, some of these might be grown to ripennd kernel in harvesting time, then we consider that above results show its possibility.

Studies on Positions in Panicle of Ventral white, Basal white, Milky white like white Core and Milky white kernels, and shapes of White opaque parts in these kernels.

From these studies investigated on positions in panicle of ventral white, basal white, milky white like white core and milky white kernels, we considered on relations between shapes of white opaque parts and positions in panicle of these kernels. Based above consideration, we intend to make clear up the mechanisms of development of these kernels. Results are summarized as follows; 1) Milky white kernels developed at inferior positions in panicle as in Figure 1, and shapes are shown in Figure 2 to 13. White opaque parts of milky white kernels have many different shapes as shown in these figures. 2) Ventral white kernels developed at superior position in panicle as in Figure 1, and shapes are shown in Figure 14 and Figure 15. 3) Milky white like white core kernels developed in high prcentages in the second and the third grains of secondary branch of middle parts in panicle as in Figure 1, and shapes are shown in Figure 16 and Figure 17. 4) Basal white kernels developed in high percentages in the second grain of upper branches in panicle as in Figure 1, and shapes are shown in Figure 18 and Figure 19. 5) We make diagram of developmental process of superior medium and inferior spikelet as Figure 20. This figure is made by the consideration that inferior spikelets develop very slowly and need long days for full ripening. As shown in Figure 20, the most rapidly development time of superior spikelet corresponds to the early development time of medium spikelet (about several days after flowering). As same as above mentioned, the most rapidly development time of medium spikelet corresponds to the middle development time of inferior spikelet (about twenty days after flowering).