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Original Paper Effects of the Wx Gene on Starch Biosynthesis, Physicochemical Wheat Flour Properties, and Dry Noodle Quality

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Original Paper Effects of the Wx Gene on Starch Biosynthesis, Physicochemical Wheat Flour Properties, and Dry Noodle Quality _ Food Science and Technology Research, 24 (3), 443 453, 2018 Copyright © 2018, Japanese Society for Food Science and Technology http://www.jsfst.or.jp doi: 10.3136/fstr.24.443 Original paper Effects of the Wx Gene on Starch Biosynthesis, Physicochemical Wheat Flour Properties, and Dry Noodle Quality 1* 2 1 Peng QIN , Zhiyou KONG and Yeju LIU 1College of Agronomy and Biotechnology, Yunnan Agricultural University, Kunming 650201, China 2College of Resources and Environment, Baoshan University, Baoshan 678000, China. Received May 24, 2015 ; Accepted October 9, 2015 Wheat starch biosynthesis is influenced by the Wx gene, which may also affect the quality and characteristics of flour and food. In this study, eight near-isogenic Wx wheat lines were bred and used as test materials for analysis of the content and biosynthesis dynamics of starch, and the changes in flour and dry noodle qualities were evaluated. The results showed that the contents and accumulation rates of amylose, amylopectin, and total starch initially increased and then decreased at 20 d after anthesis. Additionally, the three Wx genes examined in this study had varying effects on the amylose content and accumulation rate, in the order of Wx-B1 > Wx-A1 > Wx-D1. Wx genes had only minor effects on amylopectin biosynthesis. Lack of Wx gene expression was expected to affect the physiochemical properties of wheat flour, with the Wx-B1 gene exhibiting the most dramatic effects. Almost all properties except taste, stickiness, and total score of dry noodles decreased following silencing of the Wx-B1 and Wx-D1 genes. Dry noodle qualities deteriorated to different degrees after silencing of one, two, or three Wx genes, and almost no amylose biosynthesis and very poor dry noodle quality were observed in wheat lacking all three Wx genes. Keywords: near-isogenic wheat lines, starch biosynthesis, flour physicochemical characteristics, dry noodle quality Introduction ability of different Wx proteins and the effects of silencing or Starch is the major component in wheat (Triticum aestivum mutation of different Wx proteins on amylose synthesis. For L.) kernels. Wheat starch normally contains 20–30% amylose, example, Miura and colleagues (Mirua et al., 1994; 1999; with the remaining starch content comprised of amylopectin. 2002) found that the presence of a regulator gene that Amylose content plays an important role in the quality of suppresses the activities of Wx genes and the absence of the wheat flour, affecting the texture, stability, and viscosity of Wx-B1 gene or substitution with the corresponding null allele processed foods. Granule-bound starch synthase (GBSS), a key causes dramatic reductions in amylose synthesis. While enzyme involved in the biosynthesis of amylose, is encoded by amylose content is not expected to be increased by double waxy (Wx) genes. Because of its role in amylose biosynthesis, dosages of chromosomes 7A, 4A, and 7D, upregulation of Wx GBSS also influences the texture and quality of wheat flour. proteins is observed in the context of tetrasomic chromosomes. Wx genes include Wx-A1, Wx-B1, and Wx-D1, which are A more comprehensive study of the correlation between located on chromosomes 7AS, 4AL, and 7DS, respectively, amylose content and starch viscosity properties was carried out and mutations in or silencing of one or more Wx genes can using near-isogenic Wx wheat lines (Bhattacharya et al., 2002; cause reduction in endosperm amylose content (Anisworth et Zhao et al., 1998). Nakamura et al. (1995) indicated that there al., 1993; Nakamura et al., 1993; Sun et al., 1998; Yamamori were no significant differences in protein content in eight near- and Endo, 1996). isogenic Wx wheat lines. As compared to the normal line, Some researchers have studied the amylose synthesis lower falling number, glucose yield, and ethanol yield and *To whom correspondence should be addressed. E-mail: [email protected] 444 P. QIN et al. higher grain protein content were observed in double-null flour blend is increased. Moreover, the amylose content of (WxBD) and triple-null (WxABD) near-isogenic lines, starch is positively correlated with hardness and negatively indicating the strong involvement of alleles encoded at Wx-B1 correlated with cohesiveness of cooked white salted noodles and Wx-D1 loci in grain composition (Gadonna-widehem et (Park and Baik, 2004, Baik et al., 2003). Noodles prepared al., 2012). Moreover, waxy wheat starch contains a smaller from waxy wheat flour exhibit the shortest cooking time, proportion of B-type granules, larger average granule diameter, demonstrating the influence of starch amylose content on and higher degree of crystallinity than normal wheat starch; cooking time in noodles (Noda et al., 2001), and have a higher these differences result in higher gelatinization temperature, compressive force than nonwaxy noodles when the strain is transition enthalpy, peak viscosity, breakdown, and swelling greater than 80%, indicating that waxy wheat noodles are soft power and lower peak viscosity temperature and final viscosity, but difficult to cut through completely (Sasakiet al., 2004). suggesting that waxy wheat starch has greater resistance to In this study, we aimed to analyze the influence and retrogradation during cooling and higher water-holding function of all Wx genes. To this end, nonwaxy wheat Ningmai capacity under dry conditions (Zhang et al., 2013; 2014). 14 and waxy wheat Wx 9 were hybridized and continuously Several studies have shown that waxy wheat behaves backcrossed, generating near-isogenic Wx wheat lines by differently from wild-type and partial-waxy wheat in terms of marker-assisted selection. The effects of various Wx genes on physicochemical characteristics. Waxy wheat flours exhibits wheat starch biosynthesis and the quality of wheat flours and low falling numbers that are independent of α-amylase levels dry noodles were investigated. Our results provide important and lower setback, pasting, and peak temperatures than normal insights into the effects of Wx genes on starch quality, and partial-waxy wheat flour (Graybosch et al., 2000; Kim et agronomic traits, wheat flour properties, and food quality in al., 2003; Park et al., 2001; Sharma et al., 2002). In addition, winter wheat and may facilitate further studies and applications waxy wheat flour exhibits higher breakdown, peak viscosity, of Wx genes in wheat. gelatinization temperature, protein content, water absorption, SDS-sedimentation volume, starch damage, swelling power, Materials and Methods and alkaline water retention capacity as well as a greater degree Wheat samples The waxy wheat line Yangfan 9, provided of crystallization and an absence of amylose-lipid complex by Professor Shunhe Cheng, Lixiahe Agricultural Research compared with nonwaxy wheat (Chakraborty et al., 2004; Guo Institute, Jiangsu Province, lacking the Wx-A1, Wx-B1 and Wx- et al., 2003; Morita et al., 2002; Park.and Baik, 2004; Sasaki et D1 proteins, was used as the waxy gene donor. The wild-type al., 2004). Furthermore, waxy wheat starches show greater wheat genotype, Ningmai 14, was used as the recurrent parent refrigeration and freeze-thaw stabilities than nonwaxy starches to create sets of NILs with null alleles at one, two, or all three (Abdel-Aal et al., 2002). Thus, it is clear that Wx genes have Wx loci. Both Ningmai 14 and Yangfan 9 are hard red wheat substantial effects on the texture and qualities of wheat flour. cultivars with medium gluten content and good agronomic Dry noodles are widely consumed in China and can be traits. The eight near-isogenic Wx wheat lines with the Ningmai influenced by the physicochemical properties of wheat flour. 14 background were developed after seven backcross Both grain hardness and water absorption are negatively related generations with an individual heterozygous plant at three Wx to the color, appearance, smoothness, and taste of cooked dry loci. Individual plants from backcross progenies were selected noodles (Liu et al., 2003). Flour ash content and polyphenol by microscopic examination after pollen staining with 0.07% oxidase (PPO) have moderate negative effects on noodle color, I2-KI; plants with one-eighth yellow pollens and seven-eighths while protein content is negatively associated with the black pollens were used as male parents, while plants with all appearance and smoothness of noodles. The associations of dry black pollens were eliminated. The BC7F2 populations were noodle score with SDS-sedimentation volume, stability time, obtained by individual plant selfing of BC7F1; the absence of and maximum resistance can be fit to a quadratic regression the waxy protein at various loci was confirmed by sodium model, and the peak viscosity and breakdown contribute dodecyl sulfate polyacrylamide gel electrophoresis (SDS- positively to dry noodle qualities (He et al., 2004). Importantly, PAGE, for Wx-A1) and agarose gel electrophoresis (AGE, for waxy flour has little or no amylose in its starches; thus, Wx-B1 and Wx-D1) (Table 1) (Kim et al., 2003). amylose content will be decreased when waxy flour is added to The results of agarose gel electrophoresis showed that the normal wheat flours. Some studies have described the Wx-A1 and Wx-D1 genes were codominant, whereas the Wx- production of noodles made with waxy flour blends. For B1 gene was dominant. 336 bp (lanes 9, 10 and 11) and 317 bp example, adding 20–30% waxy flour to wild-type flour (lanes 1, 2, 3, and 4) for Wx-A1 (Figure 1), 840 bp (lanes 1, 2, 3, improves the quality of white salted noodles, and the optimal and 4) and 260 bp (lanes 9, 10, 11, and 12) for Wx-D1 (Figure amylose content has been reported to be 21–24% (Noda et al., 3) were indicated dominant and recessive respectively. 259 bp 2001). White salted noodles produced from blends of regular and 217 bp (lanes 1, 2, 3, 4, and 5) were indicated recessive, and waxy wheat flours become softer as the proportion of waxy that 259 bp, 217 bp, and 187 bp were homozygous or wheat flour is increased, even when the protein content of the heterozygous dominant (Figure 2), which should be confirmed Effects of Wx Gene on Starch, Flour Properties, Dry Noodle Quality 445 Table 1.
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