Nana LI,Yanyan PU,Weiguo KONG,Yongchao GONG,Wenbin JIA,Fugang XIN,Yanli YU,Hanfeng DING*

1.Shandong Center of Crop Germplasm Resources,Shandong Academy of Agricultural Sciences,Jinan 250100,China;2.Maize Research Institute,Shandong Academy of Agricultural Sciences,Jinan 250100,China

As a major food crop in China,wheat is an important source of human proteins.The protein quality not only affects the commercial value of wheat,but also lays an important material basis for the processing and production of high-quality foods and directly influences the quality of processed foods[1-2].Wheat quality traits are complex quantitative traits that are commonly influenced by the genotype and the environment,and the influences vary among different influencing factors[3-5].Kanget al.[6]and Xueet al.[7]reported that protein properties of wheat are the most representative traits that contribute greatly to wheat quality traits.Investigating the main components and major influencing factors of protein quality traits of wheat to select important indicators for comprehensive study has important guiding significance to quality improvement and breeding of wheat.

Chinese researchers have begun to investigate the protein content of wheat since the late 1970s and investigate the protein quality of wheat since the 1990s.Wang and Cao[8]described the effects of agricultural technical measureson protein contentand quality of wheat and carried out studies on breeding and improvement of wheatcultivars with good protein quality.Zhaoet al.[9]conducted multipoint tests with 10 winter wheat varieties for many years and found that the environment is a major factor affecting the percentage content of protein and gluten,while starch content,peak viscosity and breakdown value are mainly influenced by the interactions between various factors.Konget al.[10]comprehensively analyzed the genetic effects of protein quality traits and their interactions with the environment with quantitative genetic analysis method to investigate the ef-fects of heritability and environment on different trait indicators.Denget al.[11]carried out multi-point tests with eight winter wheat varieties(lines)for many years and found that the genotype significantly affected GMP (glutenin macro-polymer),high and low molecular weight glutenin subunits,sedimentation value,MPT (midline peak value)and bandwidth at 8 min(8TW);protein traits exhibited certain correlations with other traits;correlation coefficients between various traits varied among different environmental conditions.

In view of the importance of protein quality traits to baking quality and processing quality of wheat,it is very important to investigate protein quality traits of wheat and analyze the major influencing factors.However,most of the above studies are focused on the quality traits of some wheat varieties and the effects of genotype or environmental factors on wheat quality.Little information is available on comprehensive analysis of protein properties and influencing factors of a large population of wheat variety resources in China.In this paper,based on a large number of previous researches,the status,heritability and correlations of various quality traits of wheat varieties in China were analyzed to comprehensively investigate the effects of genotype,environment and G×E interactions on protein quality traits,aiming at providing theoretical basis for genetic improvement,planting structure adjustment,and efficient production of high-quality wheat.

Analysis of Wheat Quality Traits

Status of wheat quality traits

In recent years,in order to improve the quality of wheat,a large number of studies have been carried out on quality traits of different wheat varieties (lines).Zhanget al.[12]analyzed five quality traits of 631 wheat varieties (lines)from Huanghuai and Northern Winter Wheat Region and found that there were 537 mediumgluten wheat varieties(lines),89 highgluten wheat varieties (lines)and five low-gluten wheat varieties(lines).The results indicated that medium-gluten wheat varieties (lines)occupied a dominant position in this region;moreover,the stabilization time and stability were low.Huet al.[13]analyzed the main quality traits of 324 wheat varieties (lines)from Huanghuai Winter Wheat Region by regional trials during 2000-2009 and found that the wheat grains had relatively high protein and wetgluten contents,poorgluten strength,poor protein quality,low stabilization time,sedimentation value,extension area and maximum resistance.Liet al.[14]analyzed the quality traits of 250 wheat varieties from Shaanxi Province and believed that there was a high proportion of medium-gluten wheat varieties and a low proportion of high-quality high-gluten and low-gluten wheat varieties.Zhaoet al.[15]investigated the quality of 89 wheat varieties (lines)from Henan Province by regional trials in 2007 and found that these wheat materials had relatively high protein content,wet gluten content and sedimentation value;moreover,more than 50%of the experimentalmaterials were highgluten wheat varieties with poor general quality due to low water absorption capacity,stabilization time and softening degree.Zhaoet al.[16]reported that wheat varieties newly bred in recentyearshad appropriate bulk density,moderate and high protein content,poorprotein quality and gluten quality,and uncoordinated quality traits,most of which belonged to medium-gluten or mid-low-gluten wheat varieties.Wanget al.[17]analyzed the quality traits of 79 wheat varieties (lines)from seven winter wheat planting regions in China by regional trials during 2001-2002;according to the results,the protein content was 14.1%,sedimentation value was 27.1 ml,dough stabilization time was 3.1 min,indicating that poor gluten protein quality is the main issue in quality improvement of winter wheat in China.

Based on a large number of research achievements,Chinese wheat varieties (lines)exhibit high protein content and generally weak gluten strength,which can not meet the needs of some flour products[18-19].Therefore,the main task in wheat quality improvement currently is to strengthen the improvement of protein quality and gluten quality,thus achieving comprehensive and balanced development of various quality traits.

Protein quality traits of wheat

Only high-quality high-gluten and low-gluten wheat varieties with certain protein content can display the high quality properties well.Huanget al.[20]and Linet al.[21]reported that Protein content is not only highly correlated with the gluten strength but also significantly associated with the appearance performance and total evaluation value of cooked noodles.

Wheat quality mainly depends on the content and quality of gluten.Wet gluten is the material basis of unique foods prepared with flour.Wheat flour can be processed into various types of foods lies in its unique gluten.Gluten plays a decisive role in rheological properties and baking quality of dough[22].According to the content and strength of gluten,various wheat varieties are divided into high-gluten wheat,medium-gluten wheatand low-gluten wheat.The content and quality of gluten in flour are two independent factors that are not necessarily related[14].

Higher sedimentation rate and larger volume measured indicate greatergluten strength and better baking quality of flour.Settlement values include zeleny settlement value and SDS sedimentation value.zeleny method is defined as the standard method by American Association of Cereal Chemists (AACC),which is characterized by small systematic error,accurate and reliable results,and being little affected by temperature and closely related to protein content and quality.Compared with zeleny method,SDS sedimentation method exhibits strong practicality,which can be used for rapid microdetermination.In addition,SDS sedimentation is significantly correlated with processing and baking quality,which is little affected by protein content,with high stability.

Genetic Models for Proteinrelated Traits of Wheat

Investigating the inheritance patterns of wheat grain protein traits has practical significance for understanding the genetic mechanism of proteins in wheat grains and improving the effi-ciency of protein quality improvement and breeding of wheat.The genetic mechanism of wheat grain protein traits is relatively complex.Differences in gene control,complex physiological and biochemical processes lead to differences between various varieties.Moreover,the results of studies on genetic models,heritability and combining ability of wheat exhibit varying degrees of differences due to different experimental conditions.

Genetic models for protein content of wheat

Many studies about the genetic effectofwheatprotein content have achieved inconsistent results.Liet al.[23]reported that the genetic model for protein content is consistent with additive-dominant genetic models,exhibiting primarily additive effects.However,Zhu[24]found that the inheritance of protein content is partially dominant,involving superdominant and epistatic effects.Konget al.[10]conducted multi-point planting trials with parents and F1generations of partial diallel cross and found that the protein content,peak time and right slope are mainly controlled by the dominant effect.Huoet al.[25]reported that the higher value of wheat grain protein content is related with dominant genes and the lower value is related with recessive genes,which is consistent with Mihaljevet al.[26]but contrary to Zhuet al.[27].Liet al.[28]found that the protein content shows intermediate inheritance in F2generations and predicted that protein content-related genes may have bidirectional dominant effects,resulting in varying performances in different combinations.

Genetic models for wet gluten content of wheat

Very few studies have been carried out on the inheritance of wet gluten content of wheat.The inheritance of wheat gluten content and gluten index is generally consistent with additive-dominant genetic models,influenced by additive,dominant effects and nonallelic gene interactions.Tanget al.[29]conducted genetic analysis on wet gluten content and gluten index of 21 hybrid wheat combinations;the results show that the gluten content exhibits medium heritability with the narrow heritability of 44.26%,while gluten index has relatively higher heritability with the narrow heritability of 73.44%.Zhanget al.[30]analyzed glutenin macro-polymer content and other quality traits of 12 parent materials with different quality traits and found that wet gluten content is primarily controlled by additive effects and also affected by non-additive effects.Weiet al.[31]conducted regional trials with 12 wheat varieties(lines)from Guanzhong Plain ofShaanxi Province in 12 points and found that the wet gluten content is extremely significantly affected by year effects,followed by location effects.

Genetic models for sedimentation value of wheat

Many studies about the inheritance of sedimentation value have achieved significantly inconsistent results.A large number of studies have shown that sedimentation value exhibits high heritability;high sedimentation value is dominant with high heritability and can be easily selected in early generations[32-34].The simple and rapid microdetermination of sedimentation value can reflect genetic differences among genotypes.Therefore,it has aroused wide concern among wheat breeders and cereal chemists.Li[35]believed that the inheritance of sedimentation value is primarily controlled by additive effects,which has high heritability and is little influenced by environmental factors,thereby reflecting the differences in protein quality among various wheat varieties.In addition,Liuet al.[36]reported that sedimentation value is commonly affected by multi-gene additive effects and nonadditive effects.

Based on the results of previous researches,genetic models for wheat grain protein content,wet gluten content,gluten index,sedimentation value and other traits are mainly controlled by additive effects.It is effective to realize the genetic improvement of highquality wheat via parental hybridization and target trait selection.

Main Influencing Factors of Protein Traits of Wheat

Wheat protein traits belong to quantitative traits.A large number of studies have confirmed significant influences of genotype,environment and G×E interactions on protein traits[37-39],and different quality traits are influenced to varying degrees[40-41].Many studies have shown that quality traits are significantly influenced by the interactions between genotype and environment[42-45],indicating that wheat quality traits are resulted from the genetic characteristics of wheat varieties,environment,and their interactions[46].Only under appropriate environmental conditions can high-quality special wheat varieties achieve the best results and play proper value.Therefore,clarifying the effects of genotype,environment and G×E interactions on quality traits of wheat is necessary to wheat quality improvement and highquality wheat production.

The conclusions are not entirely consistent due to varying effects of the environment and genotype on different protein quality traits and differences in sample size and environmental conditions[40-41].Sunet al.[47]and Yuanet al.[48]believed that the genotype exhibits the greatest effect on protein quality traits of wheat,followed successively by the environment and G×E interactions.Liet al.[39]investigated the variation patterns of main quality traits of wheat and found that various quality traits,except grain protein content,are significantly or extremely significantly affected by the interactions between genotype and environment.Wanget al.[43]analyzed the quality traits of eight winter wheat varieties(lines)with various genotypes and found that protein and wet gluten contents are not significantly affected by the environment or G×E interactions.Qiao[49]conducted multi-site regional trials with 13 wheat varieties for many years and drew different conclusions.According to the results, the environment exhibits greater effects on protein content,wet gluten content and zeleny sedimentation value than the genotype;the genotype exhibits greater effects on SDS sedimentation value than the environment.In conclusion,the effects of genotype,environment and G×E interactions on protein quality traits are very important,which should be paid much attention to.

A large number of studies have shown that the environment exhibits greatereffects on protein content than the genotype[47-49].However,Jinget al.[50]and Wanget al.[43]drew different conclusions,who believed that the effects of these three factors on protein contentdemonstrated a descending order of genotype＞environment＞G×E interactions.Most studies indicate that sedimentation value is primarily influenced by the genotype,exhibiting extremely high genetic stability[40,51].Moreover,the environment exhibits greater effects on sedimentation value than on protein content in different locations[52].Guoet al.[53]reported that the effects of these three factors on protein content and sedimentation value demonstrated a descending order of genotype＞environment＞ G×E interactions.

Correlations between Wheat Protein Traits and Other Traits

There are certain correlations between protein traits and other traits of wheat.Therefore,clarifying the correlations among various traits and identifying the main influencing factors contributing greatly to quality trait improvement can provide theoretical basis for genetic improvement,breeding of high-quality wheat varieties,development of cultivation measures,and determination of reasonable breeding programs.

A large number of studies have confirmed the correlations among various quality traits of wheat.It has been reported that there is a significant positive correlation between protein contentand dough rheologicaltraits(dough development time,stabilization time,water absorption capacity,extension area and maximum resistance)[54-56].Wanget al.[57]found that protein content is extremely significantly positively correlated with wet gluten content and zeleny sedimentation value,which is consistent with Zhaoet al.[16];in addition,protein content is extremely significantly negatively correlated with starch content but exhibits no significant correlation with hardness.

Zhanget al.[58]found that wet gluten content of wheat grains exhibits extremely significant positive correlations with hardness and sedimentation value,but has no significant correlation with protein content.Liet al.[59]found that wet gluten content and dry gluten content exhibit extremely significant negative correlations with gluten index,while gluten index is significantly or extremely significantly positively correlated with dough development time,stabilization time and farinograph index,and extremely significantly positively correlated with protein content.Wanget al.[54]drew inconsistent conclusions with Liet al.[59],who reported that wet gluten content exhibits extremely significant negative correlations with dough development time and flour whiteness.Wanget al.[57]found that wet gluten content of wheat is extremely significantly positively correlated with zeleny sedimentation value and hardness and exhibits extremely significant negative correlations with starch content.

The sedimentation value plays an important role in wheat quality improvement,which is not only closely related with gluten content and quality,but also significantly positively correlated with wheat protein content,bread volume and farinograph-related traits,thereby comprehensively reflecting the content and quality of proteins[60].Sedimentation value issignificantly positively correlated with protein content,wet gluten content,dough development time and stabilization time,negatively correlated with protein content and yield,but not necessarily related to grain yield[32,35,16,61].Wanget al.[57]also found that there is an extremely significant positive correlation between zeleny sedimentation value and hardness.Therefore,sedimentation value can effectively reflect the processing quality of wheat varieties.

There are various quality traits of wheat,and the inheritance of each trait is controlled by complex factors,leading to difficulties in selecting offspring with a variety of excellent quality traits.The protein content,wet gluten content and sedimentation value exhibit significant correlations with other traits.Therefore,protein content,wet gluten content and sedimentation value can be used as main indicators for highquality wheat breeding in early generations.Moreover,these three protein quality traits can be improved to enhance the overall quality of wheat.

Main Existing Problems

In recent years,through the combined efforts of the breeders,great progresses have been made in analysis of wheat quality traits and breeding of high-quality wheat varieties.However,the levelofhigh-quality wheat breeding in China still lags behind foreign countries.Due to narrow hereditary basis and lack of high-quality germplasm resources,no breakthrough has been achieved so far in wheat quality improvement in China.During 2003-2005,National Center for Cereal Quality Supervision and Inspection of the Ministry of Agriculture(Beijing)conducted wheat quality survey for three consecutive years[61].The survey results indicate that most of wheat varieties in China are mediumgluten wheat with moderate and high protein and gluten contents,and 60%of samples reached 14.0%;however,wet gluten content and stabilization time are relatively poor,and very few samples have achieved the national standard for high-gluten wheat,which may be due to the poor protein quality.

A large number of studies have been carried out on genetic models,main influencing factors and correlations among various traits of wheat quality.However,no systematic and complete study has been conducted.Most of the studies are focused on a single trait of a particular aspect,and the results are inconsistent.It is very important to combine these inconsistent findings and apply the theoretical results to guiding high-quality wheat breeding and production practice based on the correlations among various quality traits,genetic models for quality traits and effects of genotype,environment and G×E interactions.

Conclusion

By analyzing the present situation and existing problems,combined with production practice investigation and requirements for high-quality wheat in food processing industry, several countermeasures are proposed as follows.

Strengthening utilization of highquality wheat germplasm resources,broadening breeding approaches

Strengthen the searching and introduction of excellent wheat germplasm resources,creating new high-quality germplasm resources via hybridization,mutagenesis,molecular marker-assisted selection and other means,and breaking through the bottlenecks in the lack of high-quality wheat breeding materials,low genetic diversity and single parental source to enrich high-yield high-quality wheat varieties and broaden sources of breeding materials is the key to successful breeding of high-quality wheat.

Appropriately selecting crossing parents,emphasizing comprehensive improvement of various quality indicators

Parents should be selected basedon the geneticmodelsfor wheat quality traits.Wheat quality is not only determined by the genetic basis of wheat varieties,but also affected by the environmental conditions and interactions between genotype and environment[63-67].Therefore,it is necessary to optimize the breeding methods,cultivation areas and cultivation methods of high-quality wheat,thus improving the success rate of quality improvement.

Perfecting the evaluation system of wheat quality

Wheatvarieties with different quality levels have different commercial uses.Thus,accurately evaluating the appropriate food products prepared by the wheat varieties,such as bread,noodles or cakes,is conducive to developing great commercial value.Therefore,it is necessary to further investigate the correlations between high-gluten,medium-gluten and lowgluten wheat varieties and final products,and establish quality evaluation system of wheat varieties with different gluten levels,thereby providing scientific basis for breeding,production and processing of high-quality wheat.

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