Effects of glutelin and globulin on the physicochemical properties of rice starch and flour

The effect of two rice endosperm proteins, glutelin and globulin, on the physicochemical properties of rice starch and flour was investigated. Albumin, globulin, prolamin and glutelin were sequentially extracted from defatted rice flour with de-ionised water, 1.5 M NaCl, propan-2-ol and 0.1 M NaOH, respectively, followed by dialysis and lyophilisation. Globulin and glutelin were then added to pure rice starch at various concentrations, separately and together, and the pasting and textural properties of mixtures were analysed by the Rapid Visco Analyser (RVA) and TA-XT2 textural analyser, respectively. The presence of glutelin in rice starch caused an increase in pasting temperature but a decrease in the viscosity parameters of the starch paste. The concentration of glutelin was also positively correlated with the hardness and adhesive properties of the starch gel. The presence of globulin, on the other hand, resulted in a decrease in all the pasting and textural parameters except gel hardness and the changes were linearly correlated with the concentration of the protein for most of the physical parameters. When the two proteins were added to rice starch together, the outcomes in pasting and textural properties were generally dependent upon the relative concentrations of the two proteins, but were also influenced by the presence of the other two protein fractions, albumin and prolamin. The presence of globulin initially accelerated the rate of water absorption by starch during cooking while the presence of glutelin slowed it down, but in both cases, the ultimate amount of water absorbed was significantly lower than that by pure starch. The contrasting effects of the different protein fractions mean that it might be possible to manipulate the textural properties of rice starch and flour to achieve desirable sensory outcomes by varying the proportions of the protein fractions in product formulations.     

Effects of gamma irradiation on starch granule structure and physicochemical properties of brown rice starch

In the present research effects of gamma irradiation (5–20 kGy) on the physicochemical, antioxidant and thermal properties of brown rice starch were studied. Scanning electron microscopy displayed mostly polyhedral shapes with no surface fractures. XRD displayed characteristic A type pattern and decrease in per cent crystallinity (22.53–20.41%) was observed as the dose increased from 5 to 20 kGy. DPPH % inhibition and FRAP values were increased in a dose dependent manner. Pasting properties (peak, final and setback viscosity) decreased with irradiation. Pasting temperature also significantly decreased with irradiation for all the irradiated starches. The transition temperatures, enthalpy of gelatinization, pH and apparent amylose content of starch decreased significantly with increase in irradiation dose.     

Effect of various drought stresses and subsequent recovery on proline,total soluble sugar and starch metabolisms in Rice (Oryza sativa L.) varieties

ABSTRACT

Rice (Oryza sativa L.) is one of the most important staple foods in the world, however most improved rice varieties are susceptible to drought stress. A two-year study was conducted to explore the effects of various drought stresses and subsequent recovery on the accumulation and degradation of proline, total soluble sugar and starch in different rice varieties at vegetative stage. The results showed that relative water content in the leaves and sheaths of rice varieties significantly decreased under drought stresses, but not at the same rate. Under control and drought conditions, the water content in sheaths was higher than that in leaves. Interestingly, under severe drought stress in 2015, the leaf water content was higher than the sheath water content. The water distribution between leaves and sheaths might be a response of plants to protect leaf system from devastation by drought. Proline was highly accumulated under drought stress but rapidly decreased after re-watering. The drought tolerant variety DA8 expressed higher ability in accumulation of proline than susceptible varieties. In general, total soluble sugar and starch contents in leaves and sheaths of varieties decreased under drought stress conditions. Total soluble sugar and starch content of DA8 were less affected than other varieties under drought conditions. Our study indicated that metabolisms of total soluble sugar and starch in rice were affected by both environmental conditions and characteristics of varieties. Proline accumulation ability of varieties can be used as a useful indicator for drought tolerant potential in rice breeding for water-limited environments.     

Enclosed stigma contributes to higher spikelet fertility for rice(Oryza sativa L.) subjected to heat stress

With global warming, rice plants may be subjected to heat stress more regularly during the heatsensitive flowering stage, causing spikelet sterility and grain yield loss.Stigma exsertion is considered to increase pollen reception and promote female reproductive success.The aim of this study was to investigate the role of stigma exsertion on spikelet fertility at high temperatures.Five rice cultivars(Liangyoupeijiu, Shanyou 63, Huanghuazhan, Nagina 22, and IR64) with differing degrees of stigma exsertion were cultivated and exposed to high temperature at anthesis.Heat-tolerant cultivars did not always show a high percentage of spikelets with exserted stigmas, and vice versa.Irrespective of the presence of more pollen grains on exserted stigmas, spikelets with exserted stigmas did not show greater spikelet fertility than spikelets with fewer exserted stigmas or hidden stigmas under heat stress.GA3 application augmented the percentage of spikelets with exserted stigmas;however, it did not increase spikelet fertility under heat stress.Spikelet fertility of whole panicles was negatively correlated with the percentage of spikelets with exserted stigmas, but positively with that with hidden stigmas.Viability of the hidden stigmas was less reduced than that of exserted stigmas under heat stress, suggesting that hidden stigmas have an advantage in maintaining viability.Heat stress delayed anther dehiscence and reduced the viabilities of both exserted stigmas and pollens, thereby causing low spikelet fertility.Together, these results suggest that high spikelet fertility does not depend on stigma exsertion and that enclosed stigma generally contributes to higher spikelet fertility and heat tolerance under high-temperature conditions during flowering in rice.     

Effects of soaking on physicochemical properties of four kinds of rice used in Huangjiu brewing

Rice soaking is an important step in the Huangjiu brewing process. It affects the subsequent fermentation process and the quality of Huangjiu. However, the mechanism and physicochemical variation of rice soaking have been little studied. The physicochemical characteristics of japonica rice (JR), japonica glutinous rice (JGR), indica rice (IR) and indica glutinous rice (IGR) and their properties during the soaking process were studied and compared. The decreased pasting temperature of soaked rice indicated that they were easier to cook and gelatinize after soaking. The four kinds of rice and the soaked water showed different properties after soaking. During soaking, the total acids and amino nitrogen of the four kinds of rice increased slowly at first, then increased rapidly and finally became stable, while the reducing sugar content increased initially and then decreased. The soaking water of JGR showed the highest values of the physicochemical components, showing that it might be more profitable for further Huangjiu fermentation.     

Effect of germination time on physicochemical properties of brown rice flour and starch from different rice cultivars

The present work was designed to obtain information on the effect of germination time on the selected physicochemical properties of brown rice flour and starch prepared from three different rice cultivars. Changes in total starch, amylose and amylopectin contents of flour, amylopectin/amylose ratio and molecular weight of starch, gelatinization, pasting, rheological, and morphological properties of flour and starch during 5 days of germination were investigated. Significant changes of pasting and rheological properties of brown rice flour were found during germination, but only small changes of these properties could be found in isolated starch. Scanning electron micrographs of flour showed that the continuous matrix structure of flour was highly destroyed after germination and scanning electron micrographs of isolated starch showed that after three days of germination, pits and holes were discovered on the surface of some starch granules. Germination had little effect on the average molecular weight of starch, but the polydispersity value in germinated brown rice (2–5 days germination) was higher than that in non-germinated brown rice. The changes observed in physicochemical properties of brown rice flour and starch after germination provided a crucial basis for understanding flour and starch modification mechanisms with potential applications for an industrial scale.     

Stay-Green in Rice (Oryza sativa L.) of Drought-Prone Areas in Desiccated Soils

Abstract

Stay-green in the post-anthesis period is thought to be an efficient drought-tolerance trait in crops, but its effectiveness in rice (Oryza sativa L.) is unknown. Our objectives were to determine whether the stay-green trait exists in rice cultivars in drought-prone areas. Twenty-four cultivars from Japan and Vietnam were grown in pots of 0.08 m in diameter and 1.00 m deep. At heading, irrigation was terminated in half of the pots and continued in the remaining pots. Every four days during the grain-filling period, we measured the leaf green color with a chlorophyll meter (SPAD), the green leaf area (GLA) and the fraction of transpirable soil water (FTSW). The capacity for maintenance of SPAD-value and GLA in desiccated soils was evaluated by determining the ratio of integrated SPAD-value and GLA in desiccated (D) plants to those in irrigated (I) plants (SPAD_d/I or GLA_d/I, respectively). The SPAD_d/I and GLA_d/I in 24 cultivars showed diverse frequency distributions. Cultivars belonging to higher ranges of the distribution in SPAD_d/I and GLA_d/I tended to show higher ratios of plant dry weight at harvest in D to in I plants. SPAD_d/I and GLA_d/I in the grain-filling period were poorly correlated with those in the seedling period in desiccated soils, and hence the capacity for maintenance of green leaves in the grain-filling period would differ from that in the seedling period. These results suggest that the stay-green trait exists as the capacity for maintaining green leaves and benefits dry matter production in desiccated soils in rice cultivars in drought-prone areas.     

Detecting ALS and ACCase herbicide tolerant accession of Echinochloa oryzoides (Ard.) Fritsch. in rice (Oryza sativa L.) fields

Inconsistent control of Echinochloa oryzoides has been reported repeatedly by farmers in the major rice growing area of Turkey. Greenhouse studies confirmed the existence of cross and multiple herbicide tolerance of E. oryzoides accessions including acetolactate synthase (penoxsulam, bispyribac-sodium) and acetyl CoA carboxylase (cyhalofob-butyl) inhibiting herbicides. Comparison of 95% lower confidence intervals of ED₉₀ derived from log-logistic dose–response curves, and twice the recommended field rates of the herbicides showed some, but not distinct separation of susceptible and tolerant accessions. We used a novel method to separate heterogeneous data without a priori knowledge of grouping into more than one group. On the basis of the distribution of ED₉₀ it was possible to identify two distinct groups of the 172 accessions tested, 78% were not controlled by ALS inhibitors (penoxsulam, and bispyribac-sodium) at recommended field rates; and 38% were not controlled by the ACCase Inhibitor (cyhalofob-butyl) at twice the field rates. The effective response level of ED₉₀ resulted in 64 and 14 tolerant accessions to ALS and ACCase, respectively. Fourteen accessions showed multiple resistances to ALS and ACCase Inhibitors.Some of the accessions were strongly tolerant to both herbicide modes of action and had 100% survival even at 6 times the recommended rates. Most of these tolerant accessions were from Marmara region, predominantly in Edirne and Balıkesir, which are the regions without any crop rotation.     

Moisture distribution of soaked rice grains observed by magnetic resonance imaging and physicochemical properties of cooked rice grains

Moisture distribution in individual polished rice grains was observed during soaking by magnetic resonance (MR) imaging, and a nuclear magnetic resonance (NMR) signal intensity profile (SI-profile) was generated from the MR image. The water penetration pattern during soaking roughly showed dissimilar trends between different varieties of japonica and japonica-indica hybrid rice. NMR signal intensity at the completion of water absorption varied within each grain; high at the periphery and the central region and low in the area between the periphery and the central region. High moisture content within the central region is due to loosely packed starch granules. The SI-profile was congruent for grains of the same variety harvested in different regions and years and characterized a grain moisture distribution for each variety. Moisture distribution was compared using SI-profiles for varieties with different amylose contents and new varieties bred for specific end-uses in Japan. The NMR signal intensity, which is related to the moisture content, at the surface of soaked grain was negatively correlated to the grain amylose content. The NMR signal intensities at the surface of soaked grain negatively correlated with the overall hardness of the cooked rice grain as measured by the single-grain low-high compression test.     

Investigation on morphological structure and crystal transition of maize starch gelatinized in pure glycerol

The gelatinization phenomena and crystalline structure of maize starch gelatinized in pure glycerol were investigated using confocal laser scanning microscopy (CLSM), scanning electron microscopy (SEM), differential scanning calorimetry (DSC) and X-ray diffraction (XRD). Starch granules were firstly treated in water system, CLSM and SEM micrographs displayed that they were completely broken and the characteristic birefringence of the starch granules disappeared at 70 °C. As for pure glycerol system, the starch granules swelled but maintained granular shape with the increasing of temperature. The crystalline structure of starch granules was partially destroyed at 130 °C and completely destroyed at 140 °C. The DSC thermogram showed that the gelatinization temperature of starch in glycerol started at 123.7 °C, peaked at 128.4 °C, and concluded at 135.2 °C. The X-ray diffractograms indicated that the crystalline structure of maize starch was partially destroyed at 130 °C and completely destroyed at 140 °C. Thus, glycerol served an alternative solvent to destroy crystalline structure of maize starch, which may be helpful for hydrolysis of starch granules by amylase in food industry.     

The effects of puroindoline b on the ultrastructure of endosperm cells and physicochemical properties of transgenic rice plant

Endosperm texture is an important factor governing the end-product quality of cereals. The texture of wheat (Triticum aestivum L.) endosperm is controlled by puroindoline a and b genes which are both absent in rice (Oryza sativa L.). It has been reported that the endosperm texture of rice can be modified by puroindoline genes. The mechanism, however, by which puroindolines affect the ultrastructure of rice endosperm cells remains to be investigated. In this study, we observed the ultrastructure of endosperm cells and the morphology of isolated starch granules of the transgenic rice expressing the puroindoline b gene. SEM and TEM observations indicated that compound starch granules were embedded within the matrix material in non-transgenic rice, Nipponbare, whereas they were surrounded by spaces in the transgenic rice. The morphology and size of each starch granule were not different between non-transgenic and the transgenic rice. However, the transgenic rice flour showed smaller particle size, higher starch damage, and lower viscosity during gelatinization than that of non-transgenic rice. These results confirm that puroindoline b reduces the grain hardness in rice. Moreover, the results also suggest that puroindoline b functions at the surface of compound starch granules, and not on polygonal starch granules in rice endosperm.     

Effects of Water-Saving Irrigation and Nitrogen Fertilization on Yield and Yield Components of Rice (Oryza sativa L.)

Abstract

The effects of nitrogen (N) application (32, 72 and 112 kg N ha-¹ in 2000, and 32, 92 and 152 kg N ha-¹ in 2001) and water-saving irrigation and their interaction on grain yield and yield components of the rice cultivar Champa-Kamphiroozi, which is a local cultivar in a semi-arid area in the south of Islamic Republic (I.R.) of Iran, were investigated. The plants were cultivated under sprinkler irrigation (1.0 ET_p and 1.5 ET_p), intermittent flooding (1-day and 2-day intervals) and continuous flooding (control). The experiments were conducted on a clay loam-clay soil under a semi-arid environment using four replications in a split plot design with irrigation method as main plots and N levels as subplots. The results indicated that intermittent flooding irrigation at 2-day intervals was as effective as continuous flooding for grain yield, showing high water-use efficiency (WUE). The soil moisture tension in the root zone before each irrigation under this condition was –300 to –400 cm. Sprinkler irrigation and intermittent flooding increased WUE by 20 to 60%, compared with continuous flooding, and the increase in N application rate to 112-152 kg ha-¹ increased grain yield under any irrigation condition. Under sprinkler irrigation, grain yield was low and percentage of unfilled grain was high, although WUE was high. However, by adopting sprinkler irrigation, the amount of nitrogen fertilizer application necessary for cultivation was reduced. Furthermore, when nitrogen application must be limited due to groundwater pollution, the amount of nitrogen fertilizer necessary for cultivation can be reduced.     

Effects of Silicon on Transpiration and Leaf Conductance in Rice Plants (Oryza sativa L.)

Summary

To characterize the protection effect of silicon against excessive transpiration in rice plants (Oryza sativa L.), we investigated the factors responsible for transpiration, such as cuticular conductance (gc), leaf surface wax content (WC), and stomatal movement in response to environmental stimuli, using leaves of plants grown in nutrient solution with silicon (+Si) and without silicon (–Si). The rate of transpiration of whole plants (E) under field conditions was clearly reduced by application of silica. The gc of –Si leaves under precisely regulated conditions was 33% higher than that of +Si leaves, reflecting higher adaxial gc. Silicon has little effect on WC, a factor responsible for regulation of cuticular resistance to water flux in rice leaves. These results suggest that silicon is one of the main factors responsible for regulation of gc, but that its effect was not due to an increase in the WG. Leaf conductance (gl) of –Si leaves tended to decrease more slowly than that of +Si leaves over time after changes from high to low light intensity. Moreover, it was always higher over the entire duration of the experiment, irrespective of irradiation intensity. The gl of –Si leaves was higher at lower relative humidity (RH) and the initial response to a subsequent increase in RH was slower than that of +Si leaves. A comparison of gl and gc between +Si and –Si leaves showed that the higher gl of –Si leaves was due to higher stomatal conductance. These results indicated that the reduction in E by application of silica was mainly attributable to reduction in the rate of transpiration through stomatal pores.     

Adaptation of rice (Oryza sativa L.) genotypes in the rainfed lowlands of Lao PDR

Genotype by environment (G×E) interactions for grain yield were investigated in 14 rice genotypes across eight rainfed lowland field environments in Lao PDR, in order to identify stable adapted cultivars for improved farmer livelihood and food security. G×E accounted for 20.3% of the total variance, with three vectors from ordination analysis accounting for 75.1% of the G×E-SS, in 6 genotype?×?6 environment groups. PCA1 indicated water-limited yield potential, PCA2 pre-flowering stress and PCA3 post-flowering stress. Genotype groups (G1–G6) differed in adaptation to these environments. G5 (VT450-2 and TSN9) were widely adapted and high-yielding. G6 (TDK11 and TDK37) were also high-yielding, topping the rankings in three environment groups, but yielded less in Phalanxay 2012 and Phalanxay 2011, where their phenology was unstable under stress. Other genotype groups showed specific adaptations, but failed to exceed yields of G5 and G6. Hence, VT450-2 and TSN9 (G5) were the preferred genotypes for rainfed lowland in southern Lao PDR, due to their high and stable grain yields. Stability in flowering time and high yield in rainfall deficit were desirable traits for improved farmer livelihood and food security.     

High temperature and low solar radiation during ripening differentially affect the composition of milky-white grains in rice (Oryza sativa L.)

ABSTRACT

Temperatures and solar radiation during ripening critically affect grain appearance in rice (Oryza sativa L.). Climatic factors to induce chalky grains were analyzed under the experimental conditions of high-temperature and shading treatment and also under the ambient condition in a high-temperature-prone region of Japan. The frequency of white-back (WB) and basal-white (BW) grains correlated with temperature and solar radiation, whereas that of milky-white (MW) grains was not correlated, suggesting that complicated climatic factors are involved in the formation of MW grains. Further investigation was carried out to identify the parameters that distinguish perfect and MW grains grown in high-temperature versus those grown in low-solar-radiation conditions. As reported previously, the chalk phenotypes in the transverse section of the MW grains were quite different between environments: oval-shaped chalk for MW grains grown in low-solar-radiation condition and center chalk for MW grains grown in high-temperature condition. Grain hardness and amylopectin chain-length distribution did not explain the difference in MW grains between environments. MW grains subjected to high temperatures had a lower protein content without a consistent reduction in the single-grain weight, whereas those from the low-solar-radiation condition had a lower amylose content with a consistent reduction in the single-grain weight, when compared with perfect grains that developed in either environmental condition. Overall, our results suggest that MW grains are formed through different physiological mechanisms with altered starch and protein synthesis under high-temperature and low-solar-radiation conditions.

Abbreviations: BW: basal-white; DAH: days after heading; DP: degree of polymerization; MW; milky-white WB: white-back     

Water mobility,rheological and textural properties of rice starch gel

Three rice starches from indica (TNuS19), japonica (TNu67) and waxy (TCW70) were used as samples to investigate the water mobility, viscoelasticity and textural properties of starch gels using pulsed nuclear magnetic resonance (PNMR), dynamic rheometer and texture analyzer. The spin–spin relaxation time (T₂), showed water mobility of starch gels was detected with starch concentrations 10–30%. Generally, the TNuS19 and TNu67 at ≥20% showed two components (T_2a and T_2b) in water mobility, where T_2a and T_2b related to solid-like and liquid-like water molecules in starch gels, respectively. However, the TCW70 over the concentrations examined had only T_2b component, higher than those of corresponding TNuS19 and TNu67. The storage (G′) and loss (G″) moduli of starch gels were in the order of TNuS19 > TNu67 > TCW70. Texture analyzer analysis indicated that TNuS19 had higher hardness, stickiness and adhesiveness than did the TNu67 and TCW70, and changed significantly with the starch concentration increase. The value of T_2b was highly correlated with physical properties of starch gels, especially with dynamic rheological parameters. It is suggested that amylose content may play a major role to influence the water mobility of starch gels which affects the specific viscoelasticity and textural properties of starch gels.     

Understanding the mechanism of change in morphological structures,visualization features,and physicochemical characteristics of adlay seeds (Coix lacryma-jobi L.): The role of heat soaking

Heat soaking is an essential factor deciding the final quality of hard-to-cook grains. In this study, the morphological structures, visualization features, and physicochemical characteristics of adlay seeds soaked at 30–70 °C were investigated compared with waxy rice (easy to cook). The morphology of macromolecules and section cracks were observed by scanning electron microscopy and light microscopy as these are the crucial factors for penetrate water penetration, which showed that starch granules were slightly eroded when adlay seeds were soaked at 70 °C. The MRI images showed that at higher soaking temperatures, the moisture of soaked adlay seeds was distributed more evenly across a gradient. Meanwhile, heat soaking has a minor effect on starch and protein content of adlay seeds, and the proportion of bound water of adlay seeds slightly declined after heat soaking. Moreover, hardness analysis and pasting measurements exhibited structural changes and viscosity transition in the grains. However, the contribution of heat soaking to the infrared spectrogram and relative crystallinity of adlay seeds was insignificant. This study provides effective methods to evaluate changes in adlay seeds during heat soaking and explains why it is hard to cook.     

Response of the leaf photosynthetic rate to available nitrogen in erect panicle-type rice (Oryza sativa L.) cultivar,Shennong265

Increasing the yield of rice per unit area is important because of the demand from the growing human population in Asia. A group of varieties called erect panicle-type rice (EP) achieves very high yields under conditions of high nitrogen availability. Little is known, however, regarding the leaf photosynthetic capacity of EP, which may be one of the physiological causes of high yield. We analyzed the factors contributing to leaf photosynthetic rate (P_n) and leaf mesophyll anatomy of Nipponbare, Takanari, and Shennong265 (a EP type rice cultivar) varieties subjected to different nitrogen treatments. In the field experiment, P_n of Shennong265 was 33.8 μmol m^?2 s^?1 in the high-N treatment, and was higher than that of the other two cultivars because of its high leaf nitrogen content (LNC) and a large number of mesophyll cells between the small vascular bundles per unit length. In Takanari, the relatively high value of P_n (31.5 μmol m^?2 s^?1) was caused by the high stomatal conductance (g_s; .72 mol m^?2 s^?1) in the high-N treatment. In the pot experiment, the ratio of P_n/C_i to LNC, which may reflect mesophyll conductance (g_m), was 20–30% higher in Nipponbare than in Takanari or Shennong265 in the high N availability treatment. The photosynthetic performance of Shennong265 might be improved by introducing the greater ratio of P_n/C_i to LNC found in Nipponbare and greater stomatal conductance found in Takanari.     

Effects of prolamin on the textural and pasting properties of rice flour and starch

Prolamin is a major class of rice proteins but its influence on the physicochemical properties of rice is not clear. Rapid Visco Analyser (RVA) and TA-XT2 TPA textural analyses were performed on rice starch with the addition of prolamin extracted from three rice cultivars (Hitomebore, M103 and Amaroo), and on rice flour with the prolamin removed by propan-2-ol extraction. Addition of prolamin to rice starch was found to cause a significant (P<0.05) increase in RVA breakdown viscosity but significant (P<0.05) decreases in hardness, adhesiveness and gumminess of the starch gel. Similarly, when prolamin was removed from rice flour, exactly the opposite effect was observed. Addition of prolamin to rice starch also caused it to absorb water faster during cooking but the gelatinised starch absorbed less water compared with control samples without prolamin.     

Characteristics of grain quality and starch fine structure of japonica rice kernels following preharvest sprouting

Preharvest sprouting (PHS) is one of the most serious defects that impacts rice production and grain quality. Knowledge about the effects of PHS on the eating and cooking quality (ECQ) of milled rice is limited. Here, we selected four japonica rice varieties to study the influences of PHS on grain quality. The results showed that PHS strongly led to poor grain appearance and the development of small starch granules whose surfaces were eroded. Analysis of starch fine structure revealed that PHS resulted in amylose (AM) degradation; in particular, PHS caused a decrease in the content of long AM chains. Moreover, PHS led to a decrease in the content of short amylopectin (AP) chains in the varieties Yandao 815 (YD815), Wuyugeng 27 (WY27) and Yangeng 13 (YG13), while the variety Sidao 785 (SD785) displayed the opposite trend. We propose that the main reasons for the decrease in starch crystallinity and the pasting profiles of the germinated rice were due to both the tendency of the AM content to decrease and the degree of this decrease in the content of the different AM chains along with changes in AP and other major components. All these changes caused by PHS led to a decrease in both ECQ and palatability of milled rice.