水稻直链淀粉的影响因素、直链淀粉对加工品质的影响及遗传

综述了影响水稻直链淀粉的因素,直链淀粉对稻米加工品质的影响及直链淀粉的遗传分析。并提出一些问题及看法。     

A novel enzyme-linked immunosorbent assay for the detection of Wx-B1 null wheat lines using a monoclonal antibody and its application for the detection of marker heterogeneity within commercial cultivars

A monoclonal antibody was raised with specificity for the granule bound starch synthase (GBSS1) Wx-B1 homeoallele of wheat (Triticum aestivum L.) using a synthetic peptide immunogen. This was used to develop a simplified enzyme-linked immunosorbent assay (ELISA) for the discrimination of Wx-B1a and Wx-B1b alleles which differ in the starch properties they confer. Discrimination of these alleles is important for the selection of wheat lines for Udon-style noodle production. The simplified ELISA worked in a single antibody (indirect) format and gave improved ease of use, discrimination of alleles and resolution relative to a previously developed 2-antibody (sandwich) ELISA. When the test was validated using breeders seed of a panel of commercial cultivars, heterogeneity of Wx-B1 alleles was observed for a significant proportion of the cultivars tested and this was confirmed using PCR analysis of the Wx-B1 and Wx-A1 genes. This observation has implications for cultivar wheat quality assessment, the application of molecular markers for variety identification purposes and the establishment of mapping populations from commercial cultivars.     

Crucial role of amylose in the rising of gluten- and additive-free rice bread

Gluten- and additive-free rice bread is suitable for consumption by individuals who are sensitive to gluten and prefer to avoid additives. Here, we prepared 100% rice bread, which is a gluten-free rice bread prepared in the absence of additives, using 19 rice flour samples containing amylose contents ranging from 9.6 to 22.3%. The amylose content was positively correlated with the specific volume of the 100% rice bread, whereas no correlation was observed between the protein content and the specific volume. The amylose content was also positively correlated with the specific volume of gluten-free rice bread prepared following protease treatment of the dough. The dough of 100% rice bread prepared from rice flour containing higher amylose contents was better stabilized during leavening than that prepared from rice flour containing a lower amylose content. It was therefore apparent that amylose plays an important role in the preparation of 100% rice bread with a high loaf volume through stabilization of the dough.     

The difference in amylose content within a panicle as affected by the panicle morphology of rice cultivars

The variation of amylose content (AC) among grains within a panicle and its relation to panicle morphology were investigated using 44 japonica rice cultivars differing in grain density (GD), panicle bending degree (BD) and palatability. Morphological traits of the rice panicle were poorly associated with mean grain AC on the average of the whole panicle, but closely related to AC variation and spatial distribution within a panicle, with the compact-panicle cultivars (GD>6.5 grain/cm and BD< 30°) having greater variation than loose-panicle cultivars (GD<6.0 grain/cm and BD>70°). The extent of AC differences among the rachides within a panicle was also cultivar-dependent, and greater differences were found between middle and top rachides for the compact-panicle cultivars, and between middle and bottom rachides for the loose-panicle cultivars. The difference in spatial distribution of AC variation between Bing110 (a compact-panicle cultivar) and XS11 (a loose-panicle cultivar) indicated that grains with low AC were principally distributed on interior rachis of middle branches for Bing110, but on bottom-rachis for XS11. Such a distribution is basically associated with the difference in the timing procedure of floret initiation and development, and final grain weight.     

Starch swelling power and amylose content of triticale and Triticum timopheevii germplasm

Triticale (× Triticosecale Whittmack) and Triticum timopheevii have undergone little selection relative to other grains for quality characters, including starch amylose content. Using starch swelling power (SSP) in water and spectrophotometric analysis of the iodine binding ratio, 247 lines of triticale and 20 lines of T. timopheevii were screened for amylose content. Following this, the expression of the starch-forming protein granule-bound starch synthase (GBSS) in triticale was investigated by SDS-PAGE in the eight highest and eight lowest SSP lines. A strong correlation (R² = 0.8174) was found between iodine binding and SSP. The SSP of T. timopheevii lines ranged from 13.7 to 16.7, indicating an approximate range of amylose content from 28.1 to 33.8%: a small range within typical results from commercial wheat cultivars. The SSP of triticale ranged from 12.5 to 23.6 suggesting amylose content ranged from 12.8 to 35.1%: a much wider range reflecting the contribution of both the wheat and rye genomes. It appeared that expression of GBSS-4A was down-regulated in low amylose lines. Therefore there is significant potential to select for amylose content in triticale to increase quality in both the animal and human feed markets.     

Waxy gene haplotypes: Associations with apparent amylose content and the effect by the environment in an international rice germplasm collection

Apparent amylose content (AAC), the key determinant of rice end-use quality attributes, is primarily controlled by the Waxy gene which codes for granule bound starch synthase (GBSS). We examined the combination of sequence variation in the Waxy gene and environmental effects, and their associations with AAC using 171 rice accessions originating from 43 countries. The combination of two single-nucleotide-polymorphism (SNP) markers in the Waxy gene allows for the identification of three marker haplotypes in this gene. The first SNP is at the leader intron splice site (In1 SNP), and the second polymorphism is in exon 6. The haplotypes explained 86.7% of the variation in AAC and discriminated the three market classes of low, intermediate and high AAC rice from each other. The environment affected the AAC of all haplotypes. Higher air temperature during grain development associated with a decrease in AAC of low and intermediate AAC-types, but with an increase in AAC of high AAC-type. The association of AAC with several Waxy RM190 microsatellite-(CT_n) alleles in combination with the In1 SNP was also examined. In conclusion, the Waxy haplotypes studied appear to be useful markers for selecting the AAC of breeding lines developed from the world's rice germplasm.     

Physicochemical characterization and in-vitro digestibility of extruded rice noodles with different amylose contents based on rheological approaches

The physicochemical properties and in-vitro digestibility of extruded rice noodles with different amylose contents were characterized from a rheological point of view. Thermo-mechanical measurements showed that the rice flour with higher amylose contents exhibited greater stability to dual-mixing and higher degrees of starch gelatinization and retrogradation. In addition, greater elastic properties were clearly observed in the high amylose rice samples. The use of high amylose rice flour produced noodles with a harder texture, consequently contributing to reduced cooking loss. Furthermore, the rheological changes of extruded rice noodles were monitored in real time during the in-vitro starch digestion. The rice noodle digesta with higher amylose contents exhibited greater viscosities throughout the simulated oral-gastric-intestinal digestion steps. The flow behaviors of the rice noodle digesta consisted of the Power-law region and infinite shear plateau that were satisfactorily characterized by the Sisko model (R² > 0.99).     

Preparation and characterization of gluten-free sheeted doughs and noodles with zein and rice flour containing different amylose contents

This study aimed to explore the feasibility of zein as a gluten alternative in preparing sheeted doughs with rice flours containing different amylose contents (12, 19, and 26%) that were successfully slit into noodle strands. Rice-zein mixtures with lower amylose contents exhibited lower pasting parameters and the high amylose paste samples possessed more elastic properties. Higher water absorption was observed in the low amylose rice-zein mixture whereas the use of zein were effective in increasing the stability of rice dough regardless of amylose content. In case of thermal conductivity, the noodle doughs prepared with high amylose rice-zein mixture had low thermal conductivity, probably implying a long cooking time. The structure of the rice-zein noodles had a tendency to become firmer with increasing levels of amylose by showing greater breaking stress and resistance to extension that could be correlated to reduced cooking loss.     

Impact of amylose content on starch retrogradation and texture of cooked milled rice during storage

Milled rice from 11 varieties, with amylose levels from 1.2 to 35.6% dry base, were collected to study the impacts of amylose content on starch retrogradation and textural properties of cooked rice during storage. The relationship between amylose content and different properties was determined using Pearson correlation. Starch retrogradation enthalpy (ΔH_r) of cooked rice was determined by differential scanning calorimetry. ΔH_r values were found to be positively correlated with amylose content (0.603 ≤ r ≤ 0.822, P < 0.01) during storage. Textural properties were determined by a Texture Analyser. The hardness of cooked rice showed a positive correlation with amylose content (0.706 ≤ r ≤ 0.866, P < 0.01) and a positive correlation with ΔH_r of cooked rice (r = 0.650, P < 0.01) during storage. The adhesiveness showed a negative correlation with amylose content (−0.929 ≤ r ≤ −0.678, P < 0.01) and a negative correlation with ΔH_r of cooked rice (r = −0.833, P < 0.01) during storage. Hardness showed a negative correlation with adhesiveness (r = −0.820, P < 0.01). These results indicated that amylose content has significant effects on starch retrogradation and textural properties of cooked rice. The cooked rice with high amylose content is easy to retrograde, the cooked rice with low amylose content retrograded slowly. Sarch retrogradation contributes to the changes of textural properties of cooked rice during storage.     

Starch digestibility and the estimated glycemic score of different types of rice differing in amylose contents

Three types of rice cultivars (indica, japonica and hybrid rice) with four levels of amylose were selected for assessing variability in starch digestibility. A vitro enzymatic starch digestion method was applied to estimate the glycemic index in vivo based on the kinetics of starch hydrolysis in vitro. The results indicated that significant differences in term of glycemic response were observed in three types of rice. Amylose content had an obviously impact on the estimated glycemic score (EGS) value and resistant starch (RS) content. The contents of RS were increased with the increasing amylose in the same type of rice. Japonica rice was significantly lower in RS content compared to indica rice and hybrid rice with similar amylose. The high amylose rice cultivar ZF201, which was characterized by low major RVA parameters, i.e. peak viscosity (PKV), hot paste viscosity (HPV) and cool paste viscosity (CPV), were obviously higher in RS content and lower in EGS. The retrogradation of cooked rice led to a reduction of HI and EGS of all varieties. Starch hydrolysis tends to be more quick and complete for the waxy and low amylose rice than for the intermediate and high amylose rice.     

An SNP downstream of the OsBEIIb gene is significantly associated with amylose content and viscosity properties in rice (Oryza sativa L.)

Allele mining in starch synthesis-related genes (SSRGs) has facilitated the discovery of desired natural sequence variations for eating quality in rice. This study investigated the sequence variations from 10 SSRGs, and further evaluated their relationship with the amylose content (AC) and rapid viscosity analysis profiles in a global collection of rice accessions by association mapping (AM). In total, 83 sequence variations were found in 10 sequenced amplicons, including 73 single nucleotide polymorphisms (SNPs), eight insertion-deletions (InDels) and two polymorphic simple sequence repeats (SSRs). Four subpopulations were identified by population structure analysis based on 170 genome-wide SSR genotypes. AM revealed 11 significant associations between three phenotypic indices and three sequence variations. One SNP with a g/c transversion at the 63rd nucleotide downstream of the OsBEIIb gene termination codon on rice chromosome 2 was significantly associated with multiple trait indices in both the general linear and mixed linear models (GLM and MLM), including the final viscosity (p < 0.001, R² = 23.87%) in both 2009 and 2010, and AC (p < 0.01, R² = 11.25%) and trough viscosity (p < 0.01, R² = 20.43) in 2010. This study provides a new perspective of allele mining for breeding strategies based on marker-assisted selection.     

Effects of added water and retrogradation on starch digestibility of cooked rice flours with different amylose content

Flours derived from rice varieties with different amylose content possess distinct physicochemical and molecular properties. The aim of this study was to determine optimal processing conditions for preparing rice flour-based foods with reduced starch digestibility. To do so, we evaluated the in vitro starch digestibility of rice flours with five varieties. Reducing the amount of water (from 10-fold to 4-fold of rice flour) used for cooking rice flour lowered its starch digestibility, and the magnitude of the decrease was positively correlated with amylose content. When retrogradation of cooked rice flour proceeded for 7 days, the digestibility of high-amylose rice flours declined rapidly in the first 3 days, whereas the digestibility of low-amylose rice flours declined continuously. Our analysis also demonstrated that the chain length distribution of starch molecules and the final and setback viscosity pasting properties were the most important parameters affecting the digestibility of rice flours. Based on our results, it appears possible to reduce rice starch digestibility by establishing optimum processing conditions for different varieties. We suggest a 7-fold addition of water and retrogradation for 1 day for high-amylose rice varieties and a 4-fold addition of water with 3 days of retrogradation for low-amylose rice.     

Low temperature increased the biosynthesis of 2-AP,cooked rice elongation percentage and amylose content percentage in rice

Rice fragrance, cooked rice elongation and amylose content are some of the traits used in determining rice grain quality. 2-acetyl-1-pyrroline (2-AP) is the major compound that is responsible for fragrance in rice.This experiment was conducted with two indica rice cultivars: Meixiangzhan 2 and Xiangyaxiangzhan. Three temperature-controlled chambers: High (32 °C from 06:00 to 17:59 and 26 °C from 18:00 to 05:59), medium (27 °C from 06:00 to 17:59 and 21 °C from 18:00 to 05:59) and low (22 °C from 06:00 to 17:59 and 16 °C from 18:00 to 05:59) were used for the study. At the onset of heading, both cultivars were transferred to the three temperature-controlled chambers. Our results showed that in both cultivars, low temperature growth had the highest 2-AP content, while high temperature growth recorded the lowest 2-AP content. Principal component analysis showed remarkable differences in the metabolomes of rice grown in high, medium and low regimes. In the grains of both cultivars, we observed significant reduction in 2-AP content after 6 and 9 months of storage, however; 2-AP retention differed greatly in both cultivars. We also reported that low temperature growth increased cooked rice elongation percentage and the biosynthesis of amylose in rice. In conclusion, our findings will provide more information to breeders working on the improvement of rice grain quality.     

Association mapping of grain quality and flowering time in elite japonica rice germplasm

Grain quality traits play an important role in the economic prosperity of commercial rice markets. The objective of our research was to identify candidate molecular markers associated with three grain quality and flowering traits in a collection of elite rice japonica inbred lines evaluated in five U.S. states. Candidate marker effects were associated with the traits mapped within regions reported from previous QTL analyses while several new allelic interactions were also detected. Common markers for each trait were observed across two or more locations, and two-way interactions unique to a single location were also found. Significant genotype × location interactions were detected while broad-sense heritability estimates were low for all characters. All but one selected marker effect was associated with a reduction in apparent amylose content. A reduction in heading date at three locations was observed with one marker as a main effect or as a component of two-way interactions that mapped ∼ 5 cM from the Hd3a flowering locus. The majority of selected effects for head rice were associated with modest to substantial increases in value. Marker loci and their interactions identified in this study highlighted targeted regions for future association studies and marker-assisted breeding efforts of grain quality traits.     

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.     

Potential of Waxy gene microsatellite and single-nucleotide polymorphisms to develop japonica varieties with desired amylose levels in rice (Oryza sativa L.)

The Waxy (Wx) gene is responsible for the synthesis of amylose, a key determinant of the cooking and processing qualities of rice. Polymorphisms of CT-microsatellite and G–T single-nucleotide polymorphism (SNP) in the Wx gene and their relationship to amylose content (Ac) were explored using 178 non-waxy rice genotypes. Nine Wx microsatellite alleles, namely (CT)10 and 11, and (CT)14–20 were identified and 11 haplotypes were recognised by different combinations of CT-microsatellite and G–T SNP. Amylose content analysed in a random set of 39 genotypes was correlated with different microsatellite alleles/haplotypes. The highest Ac levels (>30%) correlated with (CT)10 and 16, high (26–30%) with (CT)11, 15 and 20, and intermediate (21–25%) with (CT)14, in all cases with G at the G–T SNP. The CT-classes (CT)17 and 18 (mean Ac value of 21%), could be subdivided into low amylose haplotypes (16–20%) for 17T and 18T and intermediate amylose haplotypes (21–25%) for 17G and 18G. The use of haplotypes proved to discriminate between intermediate and low amylose accessions within the same microsatellite class. Analyses of a segregating population of a cross between low and high Ac parents showed that CT-microsatellite may help to classify breeding lines and identify pollen contamination. We suggest that CT-microsatellite together with G–T SNP may be used as molecular marker by breeders to develop varieties with desired amylose levels.     

Changes in the properties of rice varieties with different amylose content on dry heat parboiling

Dry heat parboiling is a unique paddy processing technique that has been scarcely exploited. Dry heat parboiling at high temperature for short time and low temperature for long time on physical and physicochemical properties of three rice varieties differing in amylose content were studied. Hardness of the kernels increased from 66.4 N, 68.8 N and 59.8 N in raw samples to 89.1 N, 86.9 N and 59.8 N in parboiled high amylose, low amylose and waxy rice samples respectively. Rapid migration and evaporation of water from severely heated kernels caused cavity formation at the centre. Irreversible damage of amylopectin structures to leachable fractions caused continuous rise of the pasting curve. Crystallinity was thereby reduced. Parboiled high amylose samples gave X-ray diffraction patterns with peaks characteristic of A, B and V-type starch crystallinity. Crystalline starch-lipid complexes were observed in low amylose and waxy rices. The significant increase in the amount of rapidly digestible starch from 56.7%, 61.7% and 66.6% in raw samples to 92.1%, 90.8% and 94.8% respectively in severely processed rice samples and subsequent reduction in resistant starch from 24.5%, 21.2% and 18.4% to 0.4%, 1.9% and 0.1% indicated possibilities for targeted food use of the dry heat parboiled samples.     

Effects of Variations in Starch Synthase on Starch Properties and Eating Quality of Rice

Abstract

We evaluated the effects of functional variation in three starch synthases in rice (Oryza sativa L.)?granule-bound starch synthase I (GBSSI, wx), starch synthase I (SSI, SSI), and starch synthase IIa (SSIIa, alk)?between indica cultivar Kasalath and japonica cultivar Nipponbare on starch properties and eating quality. We used three near-isogenic lines?NIL(Wx^a), NIL(SSI ^k), and NIL(Alk)?containing chromosomal segments of Kasalath on a Nipponbare genetic background. The Wx^a allele explained most of the difference in amylose content between the two cultivars, and decreased the peak viscosity and breakdown to less than half of those of Nipponbare. These changes reduced the quality of cooked rice both just after cooking and after storage at 5ºC. The variation in SSIIa also affected the eating quality after storage of cooked rice at 5ºC : NIL(Alk) became harder and less sticky than Nipponbare, although the rices were comparable just after cooking. Differential scanning calorimetry revealed faster retrogradation of the once-gelatinized starch in NIL(Alk). The variation in SSI alleles hardly affected these properties.     

Changes in gelatinization and rheological characteristics of japonica rice starch induced by pressure/heat combinations

Rice starch suspensions of 10% dry matter (DM) were treated by heat (0.1 MPa at 20–85 °C) or pressure/heat combinations (100–600 MPa at 20, 40 and 50 °C) for 15 min to investigate their gelatinization and rheological characteristics. The maximum swelling index of about 12 g water per gram of DM was obtained by thermal treatment at 85 °C, meanwhile, that of 7.0 g was observed by 600-MPa pressurization at 50 °C. The higher temperatures or pressures resulted in the higher degrees of gelatinization. Furthermore, treatments of 0.1 MPa at 85 °C, 500 MPa at 50 °C and 600 MPa at various temperatures caused complete gelatinization of rice starch. The consistency index (K) and storage modulus (G′) dramatically increased from 70 °C or 400 MPa. The G′ values were higher in pressure-treated samples than those in thermal-treated samples. Therefore, an application of pressure/heat combinations as a processing method to improve the quality of rice starch products would be possible.     

An SNP downstream of the OsBEIIb gene is significantly associated with amylose content and viscosity properties in rice (Oryza sativa L.)

Allele mining in starch synthesis-related genes (SSRGs) has facilitated the discovery of desired natural sequence variations for eating quality in rice. This study investigated the sequence variations from 10 SSRGs, and further evaluated their relationship with the amylose content (AC) and rapid viscosity analysis profiles in a global collection of rice accessions by association mapping (AM). In total, 83 sequence variations were found in 10 sequenced amplicons, including 73 single nucleotide polymorphisms (SNPs), eight insertion-deletions (InDels) and two polymorphic simple sequence repeats (SSRs). Four subpopulations were identified by population structure analysis based on 170 genome-wide SSR genotypes. AM revealed 11 significant associations between three phenotypic indices and three sequence variations. One SNP with a g/c transversion at the 63rd nucleotide downstream of the OsBEIIb gene termination codon on rice chromosome 2 was significantly associated with multiple trait indices in both the general linear and mixed linear models (GLM and MLM), including the final viscosity (p < 0.001, R² = 23.87%) in both 2009 and 2010, and AC (p < 0.01, R² = 11.25%) and trough viscosity (p < 0.01, R² = 20.43) in 2010. This study provides a new perspective of allele mining for breeding strategies based on marker-assisted selection.