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Abstract The composition of high molecular weight glutenin subunits （HMWGS） was analyzed using SDSPAGE with 70 Longdong dryland winter wheat germplasm resources as experimental materials. The results showed that Longdong dryland winter wheat germplasm resources had N， 7+8， 2+12 as the dominant HMWGS composition （42.86%）， while the dominant HMWGS composition in the introduced germplasms was N， 7+9， 2+12 （36.84%）. On the GluB1 locus of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%， and on the GluB1 locus of the introduced germplasms， subunit 7+9 had the highest frequency， accounting for 57.89%. Whether subunit 7+8 is related to drought resistance in varieties still needs further study. On GluD1 locus， highquality subunit 5+10 appeared in the introduced resources for 5 times， accounting for 13.16% of all the introduced lines， and appeared in Longdong dryland winter wheat germplasm resources for 2 times， only accounting for 2.86% of all the resources， suggesting that Longdong dryland germplasms lack highquality subunits and need further improvement.
Key words Germplasm resource of winter wheat; HMWGS; Quality; 5+10 subunit; SDSPAGE
In wheat breeding， the selection of parents is very important， because highquality parent resources facilitate the breeding of good varieties. Wheat HMWGS type and composition are related to quality， and are important factors influencing wheat gluten quality. Different loci and different subunits differ in quality effect. For instance， varieties carrying subunits 1， 2*， 14+15， 17+18 and 5+10 have better gluten strength and bread making quality， while subunit 7+8 has a remarkable positive effect on the improvement of gluten strength in wheat[1-4]. The influences of different subunits on quality mainly depend on the composition of subunits and different abilities of the polymers that they form， and different subunit combinations could change subunit structure and ability to function[5]. Dryland farming refers to rainfed farming in semiarid and semihumid by arid zone with slightly less precipitation without irrigation condition suffering from water stress. Longdong dryland wheat germplasm resources are droughtresistant germplasms bred for dryland agricultural areas， which have watersaving function， and studies on their subunit composition and quality improvement provide highquality droughtresistant resource information for wheat breeding in China. In this study， HMWGS of Longdong dryland winter wheat germplasm resources and introduced resources were studied， and the differences in subunit composition between the two kinds of resources were analyzed， so as to understand subunit composition in Longdong dryland winter wheat germplasm resources. This study provides theoretical basis and basic materials for efficient utilization and quality improvement of droughtresistant varieties. Materials and Methods
Materials
The characteristics of glutenin subunits in Longdong droughtresistant germplasms were studied with 70 droughtresistant winter wheat germplasm resources bred in Longdong dryland as experimental materials， 38 introduced germplasm resources as control materials and Chinese spring （N， 7+8， 2+12） and Zhongyou 9507 （1， 7+9， 5+10） as reference varieties. The emphasis was put on the study on more complicated characteristics of glutenin subunits with Longdong dryland wheat germplasms as experimental materials and introduced germplasms as controls.
Main instruments
DYY12 electrophoresis apparatus and DYCZ24F vertical slab electrophoresis tank （Beijing Liuyi Instrument Factory）; TGL16G centrifuge （Shanghai Anting Scientific Instrument Factory）; WD9409 electric pulverizer （AmonMed Biotechnology Co.， Ltd.）; HH4 thermostat water bath （Jintan Fuhua Instruments Co.， Ltd.）.
Research methods
Highmolecular weight glutelin subunit composition of wheat endosperm was analyzed by sodium dodecyl sulphatepolyacrylamide gel electrophoresis （SDSPAGE） technique. The electrophoresis method for HMWGS referred to the method provided by Zhang et al[6].
Sample extraction
One wheat seed was weighed， ground and transferred to a 0.5 ml centrifuge tube. The ground wheat seed was extracted with 200 μl of 50% isopropanol at 60 ℃ for 20-30 min， with continuous shaking. Centrifugation was performed at 10 000 r/min for 10 min， and the supernatant was discarded. The extraction was repeated for 2-3 times.
Gel casting
Separation gel was casted to the upper tank of the glass slab at the 1 cm position， and the gel surface was sealed with nbutyl alcohol. After the separation gel was solidified， spacer gel was then poured， followed by the inserting of a sample comb.
Sample application
When the gel was solidified， the sample comb was slightly pulled out. During sample application， a proper amount of electrode buffer was added， and a pipettor was used to apply samples at an amount of 9 μl sequentially.
Electrophoresis and staining
Electrophoresis： With reference to the method of Zhang Xueyong， the control materials were Chinese spring （N， 7+8， 2+12） and Zhongyou 9507 （1， 7+9， 5+10）. After electrophoresis， the power was turned off， and the electrode buffer was poured out; the glass slab was lightly forced open， to remove the spacer gel; and the separating gel was marked， rinsed with distilled water for 2-3 times and placed in a fixation solution， followed by vibrating on a shaker for 2 h. Staining： The fixation solution was poured out， and the gel was rinsed with distilled water for 2-3 times. A staining solution was added to cover the gel， followed by vibrating on a shaker for 12 h.
Destaining
The staining solution was poured out， and the gel was rinsed with distilled water for 2-3 times. A proper amount of destaining solution was added to cover the gel， followed by vibrating for 2-3 h. When the bands were basically clear， the gel was taken out and rinsed with distilled water for 2-3 times， and finally added with a proper amount of distilled water to soak the gel until the bands were completely clear.
Photographing
The gel was placed in a WD9413A gel imaging system produced by Beijing Liuyi Instrument Factory and photographed. Subunits were named according to Payne and Lawrence system[7].
Results and Analysis
HMWGS composition
HMWGS are coded by Glu1 loci located on the long arm of the first homologous group of wheat， i.e.， GluA1， GluB1 and GluD1. It could be seen from Table 1 that the HMWGS compositions of Longdong dryland winter wheat germplasms included N， 7+8， 2+12， N， 7+9， 2+12， N， 7+8， 2+10， N， 7+9， 2+10， N， 7+9， 5+10， 1， 7+8， 2+12， 1， 7+9， 2+12， and 1， 7+9， 2+10， eighty subunit types in total. It could be seen from Table 2 that there were also eight types of HMWGS compositions of the introduced winter wheat germplasms： N， 7+8， 2+12， N， 7+9， 2+12， N， 7+8， 2+10， N， 7+9， 2+10， 1， 7+9， 5+10， N， 7+9， 5+10， 1， 7+8， 2+12， and N， 7+8， 5+10.
It could be seen from Table 3 that each wheat variety contained 3-5 HMWGS generally; and the HMWGS compositions of Longdong dryland winter wheat germplasm resources were mainly N， 7+8， 2+12 （42.86%）， while those of introduced resources were mainly N， 7+9， 2+12 （36.84%）. Overall， Longdong dryland winter wheat germplasm resources and the introduced resource both had 8 types of HMWGS compositions. The highquality HMWGS compositions were 1， 7+9， 5+10， N， 7+8， 5+10 and N， 7+9， 5+10， in total， three subunit compositions which all contained 5+10 subunit. Goodquality subunit combinations 1， 7+9， 5+10 and N， 7+8， 5+10 did no appear in Longdong dryland winter wheat germplasm resources， but showed frequencies accounting for 7.89% and 2.63% in the introduced resources， respectively. N， 7+9， 5+10 only appeared for 2 times in Longdong dryland winter wheat germplasm resources， accounting for 2.86% of the all the lines. The highquality subunit 5+10 appeared in the introduced resources for 5 times， accounting for 13.16% of all the introduced lines， and appeared in Longdong dryland winter wheat germplasm resources for 2 times， accounting for 2.86% of all the resources. Subunits 4+15 and 17+18 did not appear. HMWGS frequency analysis
In the 70 experimental materials and 38 control materials， 8 types of different HMWGS allelic variations were detected. On GluA1， two types of variations， 1 and N appeared. Longdong germplasms had two types of allelic variations， among which subunit N had the highest frequency， accounting for 94.29%， and subunit 1 accounted for 5.71%. There were two types of allelic variations on GluA1 locus， of which subunit N also had the highest frequency， accounting for 89.47%， and subunit 1 accounted for 10.53%. Subunit 2* did not appear in both the two kinds of resources.
On GluB1， there were two types of allelic variations， 7+8 and 7+9. On the GluB1 locus of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%， and subunit 7+9 showed a frequency accounting for 34.29%. On the GluB1 locus of the introduced germplasms， subunit 7+9 showed the highest frequency， accounting for 57.89%， and 7+8 exhibited a frequency accounting for 42.11%. Other several subunits did not appear.
There were three types of variations on GluD1， i.e.， 2+12， 2+10 and 5+10. On the GluD1 locus of Longdong dryland germplasms， subunit 2+12 had the highest frequency， accounting for 57.14%， subunits 2+10 and 5+10 showed frequencies accounted for 40.00% and 2.86%， respectively. On the GluD1 locus of the introduced germplasms， subunit 2+12 had the highest frequency， accounting for 68.42%， subunits 2+10 exhibited frequencies accounted for 7.89% and 13.16%， respectively.
Conclusions
The results showed that for Glu1 loci， GluA1， GluB1 and GluD1， Longdong dryland winter wheat germplasm resources had N， 7+8， 2+12 as the dominant HMWGS composition （42.86%）， while the dominant HMWGS composition in introduced germplasms was N， 7+9， 2+12 （36.84%）.
On the GluB1 locus of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%， and on the GluB1 locus of the introduced germplasms， subunit 7+9 had the highest frequency， accounting for 57.89%.
Varieties containing highquality subunit combinations such as 1， 7+9， 5+10， 1， 7+8， 5+10 and N， 7+9， 5+10 belonged to the introduced germplasms， while Longdong dryland germplasms rarely contained highquality subunit combinations， suggesting that Longdong dryland germplasms are not rich in highquality subunit types.
Discussion
Generally， for GluA1 locus， 1 and 2* are better than N; as to GluB1 locus， 7+8， 17+18 and 14+15 are better than other subunits; and in the case of GluD1， 5+10 is better than 2+12， which is better than other subunits. Generally， varieties containing these highquality subunits， especially subunit 5+10 have better quality. However， in this study， Longdong dryland germplasms were detected to have fewer highquality subunits and their combinations， and mostly contain subunits such as N and 2+12. The comparison of HMWGS composition between Longdong dryland winter wheat germplasm resources and introduced germplasms showed that Longdong dryland germplasms had the characteristics of single subunit composition and fewer variation types， while the introduced varieties were richer in subunit type and contained certain quantity of highquality subunits.
On the GluB1 loci of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%. On the GluB1 locus of the introduced germplasms， subunit 7+9 showed the highest frequency， accounting for 57.89%. Field and indoor drought resistance analysis shows that Longdong germplasms are much better than introduced germplasms in drought resistance， while whether subunit 7+8 is related to drought resistance in varieties still needs further study.
Based on the advantages and disadvantages of subunit compositions in Longdong dryland subunits， in future breeding， it is necessary to introduce a large quantity of foreign germplasm resources to broad the genetic basis of local wheat varieties， and breed highquality winter wheat varieties by investigating the positive effect and interaction effect of different subunit compositions and subunit combinations through the directing of highquality subunits， comprehensively considering the effects of quality and quantity of glutenin on the processing quality of wheat and comprehensively optimizing germplasms according to subunit composition characteristics， so as to gradually improve the situation of lacking highquality wheat varieties in Longdong area. How to combine the droughtresistant and watersaving characteristics of Longdong germplasms is the goal in future.
References
[1] LU J， HE ZH， XIA XC， et al. Characterization of Xinjiang local and introduced wheat germplasm for high molecular weight glutenin subunits and qualityrelated genes with molecular markers institute of cereal crops[J]. Acta Agronomica Sinica， 2009， 35（4）： 647-661. （in Chinese）
[2] ZHANG YB， ZHAO HB， SONG QJ， et al. Quality difference of nearisogenic lines with HMW glutenin subunits 7+8* and 17+18 in wheat cultivars Longmai 20[J]. Scientia Agricultura Sinica， 2008， 41（5）： 1536-1541. （in Chinese）
[3] BUTOW BJ， MA WJ， GALE KR， et al. Molecular discrimination of Bx7 alleles demonstrates that overexpression has a major impact on wheat flour dough strength[J].Theoretical and Applied Geneties， 2003， 107： 1524-1532.
[4] LEI ZS， GALE KR， HE ZH， et al. Ytype gene specific markers for enhanced discrimination of highmolecular weight glutenin alleles at the GluB1 locus in hexaploid Cereal Science， 2006， 43（1）： 94-101.
[5] KANG ZY， WANG JJ， SHANG XW. Establishing the score system of HMWGS on alveogram characters in wheat[J]. Journal of Triticeae Crops， 2007， 27（6）： 1029-1033. （in Chinese）
[6] ZHANG XY， DONG YC， YOU GX， et al. Allelic variation of GluA1， GluB1 and GluD1 in Chinese commercial wheat varieties in the last 50 years[J]. Scientia Agricultura Sinica， 2001， 34（4）： 355-362. （in Chinese）
[7] PANYNE PI， LAWRENCE GJ. Catalogue of alleles for the complex geneloci， GluA1， GluB1 and G luD1， which codefor high molecular weight subunits of glueenin in hexaploid wheat[J]. Cereal Res Comn， 1983， 11： 29-35.
Key words Germplasm resource of winter wheat; HMWGS; Quality; 5+10 subunit; SDSPAGE
In wheat breeding， the selection of parents is very important， because highquality parent resources facilitate the breeding of good varieties. Wheat HMWGS type and composition are related to quality， and are important factors influencing wheat gluten quality. Different loci and different subunits differ in quality effect. For instance， varieties carrying subunits 1， 2*， 14+15， 17+18 and 5+10 have better gluten strength and bread making quality， while subunit 7+8 has a remarkable positive effect on the improvement of gluten strength in wheat[1-4]. The influences of different subunits on quality mainly depend on the composition of subunits and different abilities of the polymers that they form， and different subunit combinations could change subunit structure and ability to function[5]. Dryland farming refers to rainfed farming in semiarid and semihumid by arid zone with slightly less precipitation without irrigation condition suffering from water stress. Longdong dryland wheat germplasm resources are droughtresistant germplasms bred for dryland agricultural areas， which have watersaving function， and studies on their subunit composition and quality improvement provide highquality droughtresistant resource information for wheat breeding in China. In this study， HMWGS of Longdong dryland winter wheat germplasm resources and introduced resources were studied， and the differences in subunit composition between the two kinds of resources were analyzed， so as to understand subunit composition in Longdong dryland winter wheat germplasm resources. This study provides theoretical basis and basic materials for efficient utilization and quality improvement of droughtresistant varieties. Materials and Methods
Materials
The characteristics of glutenin subunits in Longdong droughtresistant germplasms were studied with 70 droughtresistant winter wheat germplasm resources bred in Longdong dryland as experimental materials， 38 introduced germplasm resources as control materials and Chinese spring （N， 7+8， 2+12） and Zhongyou 9507 （1， 7+9， 5+10） as reference varieties. The emphasis was put on the study on more complicated characteristics of glutenin subunits with Longdong dryland wheat germplasms as experimental materials and introduced germplasms as controls.
Main instruments
DYY12 electrophoresis apparatus and DYCZ24F vertical slab electrophoresis tank （Beijing Liuyi Instrument Factory）; TGL16G centrifuge （Shanghai Anting Scientific Instrument Factory）; WD9409 electric pulverizer （AmonMed Biotechnology Co.， Ltd.）; HH4 thermostat water bath （Jintan Fuhua Instruments Co.， Ltd.）.
Research methods
Highmolecular weight glutelin subunit composition of wheat endosperm was analyzed by sodium dodecyl sulphatepolyacrylamide gel electrophoresis （SDSPAGE） technique. The electrophoresis method for HMWGS referred to the method provided by Zhang et al[6].
Sample extraction
One wheat seed was weighed， ground and transferred to a 0.5 ml centrifuge tube. The ground wheat seed was extracted with 200 μl of 50% isopropanol at 60 ℃ for 20-30 min， with continuous shaking. Centrifugation was performed at 10 000 r/min for 10 min， and the supernatant was discarded. The extraction was repeated for 2-3 times.
Gel casting
Separation gel was casted to the upper tank of the glass slab at the 1 cm position， and the gel surface was sealed with nbutyl alcohol. After the separation gel was solidified， spacer gel was then poured， followed by the inserting of a sample comb.
Sample application
When the gel was solidified， the sample comb was slightly pulled out. During sample application， a proper amount of electrode buffer was added， and a pipettor was used to apply samples at an amount of 9 μl sequentially.
Electrophoresis and staining
Electrophoresis： With reference to the method of Zhang Xueyong， the control materials were Chinese spring （N， 7+8， 2+12） and Zhongyou 9507 （1， 7+9， 5+10）. After electrophoresis， the power was turned off， and the electrode buffer was poured out; the glass slab was lightly forced open， to remove the spacer gel; and the separating gel was marked， rinsed with distilled water for 2-3 times and placed in a fixation solution， followed by vibrating on a shaker for 2 h. Staining： The fixation solution was poured out， and the gel was rinsed with distilled water for 2-3 times. A staining solution was added to cover the gel， followed by vibrating on a shaker for 12 h.
Destaining
The staining solution was poured out， and the gel was rinsed with distilled water for 2-3 times. A proper amount of destaining solution was added to cover the gel， followed by vibrating for 2-3 h. When the bands were basically clear， the gel was taken out and rinsed with distilled water for 2-3 times， and finally added with a proper amount of distilled water to soak the gel until the bands were completely clear.
Photographing
The gel was placed in a WD9413A gel imaging system produced by Beijing Liuyi Instrument Factory and photographed. Subunits were named according to Payne and Lawrence system[7].
Results and Analysis
HMWGS composition
HMWGS are coded by Glu1 loci located on the long arm of the first homologous group of wheat， i.e.， GluA1， GluB1 and GluD1. It could be seen from Table 1 that the HMWGS compositions of Longdong dryland winter wheat germplasms included N， 7+8， 2+12， N， 7+9， 2+12， N， 7+8， 2+10， N， 7+9， 2+10， N， 7+9， 5+10， 1， 7+8， 2+12， 1， 7+9， 2+12， and 1， 7+9， 2+10， eighty subunit types in total. It could be seen from Table 2 that there were also eight types of HMWGS compositions of the introduced winter wheat germplasms： N， 7+8， 2+12， N， 7+9， 2+12， N， 7+8， 2+10， N， 7+9， 2+10， 1， 7+9， 5+10， N， 7+9， 5+10， 1， 7+8， 2+12， and N， 7+8， 5+10.
It could be seen from Table 3 that each wheat variety contained 3-5 HMWGS generally; and the HMWGS compositions of Longdong dryland winter wheat germplasm resources were mainly N， 7+8， 2+12 （42.86%）， while those of introduced resources were mainly N， 7+9， 2+12 （36.84%）. Overall， Longdong dryland winter wheat germplasm resources and the introduced resource both had 8 types of HMWGS compositions. The highquality HMWGS compositions were 1， 7+9， 5+10， N， 7+8， 5+10 and N， 7+9， 5+10， in total， three subunit compositions which all contained 5+10 subunit. Goodquality subunit combinations 1， 7+9， 5+10 and N， 7+8， 5+10 did no appear in Longdong dryland winter wheat germplasm resources， but showed frequencies accounting for 7.89% and 2.63% in the introduced resources， respectively. N， 7+9， 5+10 only appeared for 2 times in Longdong dryland winter wheat germplasm resources， accounting for 2.86% of the all the lines. The highquality subunit 5+10 appeared in the introduced resources for 5 times， accounting for 13.16% of all the introduced lines， and appeared in Longdong dryland winter wheat germplasm resources for 2 times， accounting for 2.86% of all the resources. Subunits 4+15 and 17+18 did not appear. HMWGS frequency analysis
In the 70 experimental materials and 38 control materials， 8 types of different HMWGS allelic variations were detected. On GluA1， two types of variations， 1 and N appeared. Longdong germplasms had two types of allelic variations， among which subunit N had the highest frequency， accounting for 94.29%， and subunit 1 accounted for 5.71%. There were two types of allelic variations on GluA1 locus， of which subunit N also had the highest frequency， accounting for 89.47%， and subunit 1 accounted for 10.53%. Subunit 2* did not appear in both the two kinds of resources.
On GluB1， there were two types of allelic variations， 7+8 and 7+9. On the GluB1 locus of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%， and subunit 7+9 showed a frequency accounting for 34.29%. On the GluB1 locus of the introduced germplasms， subunit 7+9 showed the highest frequency， accounting for 57.89%， and 7+8 exhibited a frequency accounting for 42.11%. Other several subunits did not appear.
There were three types of variations on GluD1， i.e.， 2+12， 2+10 and 5+10. On the GluD1 locus of Longdong dryland germplasms， subunit 2+12 had the highest frequency， accounting for 57.14%， subunits 2+10 and 5+10 showed frequencies accounted for 40.00% and 2.86%， respectively. On the GluD1 locus of the introduced germplasms， subunit 2+12 had the highest frequency， accounting for 68.42%， subunits 2+10 exhibited frequencies accounted for 7.89% and 13.16%， respectively.
Conclusions
The results showed that for Glu1 loci， GluA1， GluB1 and GluD1， Longdong dryland winter wheat germplasm resources had N， 7+8， 2+12 as the dominant HMWGS composition （42.86%）， while the dominant HMWGS composition in introduced germplasms was N， 7+9， 2+12 （36.84%）.
On the GluB1 locus of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%， and on the GluB1 locus of the introduced germplasms， subunit 7+9 had the highest frequency， accounting for 57.89%.
Varieties containing highquality subunit combinations such as 1， 7+9， 5+10， 1， 7+8， 5+10 and N， 7+9， 5+10 belonged to the introduced germplasms， while Longdong dryland germplasms rarely contained highquality subunit combinations， suggesting that Longdong dryland germplasms are not rich in highquality subunit types.
Discussion
Generally， for GluA1 locus， 1 and 2* are better than N; as to GluB1 locus， 7+8， 17+18 and 14+15 are better than other subunits; and in the case of GluD1， 5+10 is better than 2+12， which is better than other subunits. Generally， varieties containing these highquality subunits， especially subunit 5+10 have better quality. However， in this study， Longdong dryland germplasms were detected to have fewer highquality subunits and their combinations， and mostly contain subunits such as N and 2+12. The comparison of HMWGS composition between Longdong dryland winter wheat germplasm resources and introduced germplasms showed that Longdong dryland germplasms had the characteristics of single subunit composition and fewer variation types， while the introduced varieties were richer in subunit type and contained certain quantity of highquality subunits.
On the GluB1 loci of Longdong dryland germplasms， subunit 7+8 had the highest frequency， accounting for 65.71%. On the GluB1 locus of the introduced germplasms， subunit 7+9 showed the highest frequency， accounting for 57.89%. Field and indoor drought resistance analysis shows that Longdong germplasms are much better than introduced germplasms in drought resistance， while whether subunit 7+8 is related to drought resistance in varieties still needs further study.
Based on the advantages and disadvantages of subunit compositions in Longdong dryland subunits， in future breeding， it is necessary to introduce a large quantity of foreign germplasm resources to broad the genetic basis of local wheat varieties， and breed highquality winter wheat varieties by investigating the positive effect and interaction effect of different subunit compositions and subunit combinations through the directing of highquality subunits， comprehensively considering the effects of quality and quantity of glutenin on the processing quality of wheat and comprehensively optimizing germplasms according to subunit composition characteristics， so as to gradually improve the situation of lacking highquality wheat varieties in Longdong area. How to combine the droughtresistant and watersaving characteristics of Longdong germplasms is the goal in future.
References
[1] LU J， HE ZH， XIA XC， et al. Characterization of Xinjiang local and introduced wheat germplasm for high molecular weight glutenin subunits and qualityrelated genes with molecular markers institute of cereal crops[J]. Acta Agronomica Sinica， 2009， 35（4）： 647-661. （in Chinese）
[2] ZHANG YB， ZHAO HB， SONG QJ， et al. Quality difference of nearisogenic lines with HMW glutenin subunits 7+8* and 17+18 in wheat cultivars Longmai 20[J]. Scientia Agricultura Sinica， 2008， 41（5）： 1536-1541. （in Chinese）
[3] BUTOW BJ， MA WJ， GALE KR， et al. Molecular discrimination of Bx7 alleles demonstrates that overexpression has a major impact on wheat flour dough strength[J].Theoretical and Applied Geneties， 2003， 107： 1524-1532.
[4] LEI ZS， GALE KR， HE ZH， et al. Ytype gene specific markers for enhanced discrimination of highmolecular weight glutenin alleles at the GluB1 locus in hexaploid Cereal Science， 2006， 43（1）： 94-101.
[5] KANG ZY， WANG JJ， SHANG XW. Establishing the score system of HMWGS on alveogram characters in wheat[J]. Journal of Triticeae Crops， 2007， 27（6）： 1029-1033. （in Chinese）
[6] ZHANG XY， DONG YC， YOU GX， et al. Allelic variation of GluA1， GluB1 and GluD1 in Chinese commercial wheat varieties in the last 50 years[J]. Scientia Agricultura Sinica， 2001， 34（4）： 355-362. （in Chinese）
[7] PANYNE PI， LAWRENCE GJ. Catalogue of alleles for the complex geneloci， GluA1， GluB1 and G luD1， which codefor high molecular weight subunits of glueenin in hexaploid wheat[J]. Cereal Res Comn， 1983， 11： 29-35.