Subscribe Now Subscribe Today
Research Article

Genetic Diversity of Storage Proteins in Triticum polonicum L.

Dong Pan, Liu Hong , Li Wei , Qi Peng-Fei , Wei Yu-Ming and Y.L. Zheng
Facebook Twitter Digg Reddit Linkedin StumbleUpon E-mail

The gliadins and high molecular weight glutenin subunits (HWM-GSs) variations of 72 Triticum polonicum accessions derived from 23 countries were characterized by A-PAGE and SDS-PAGE, respectively, for the purpose of evaluating the genetic diversity T. polonicum accessions at the level of proteins. Higher genetic variabilities were observed for both gliadins and HWM-GSs. Forty-eight gliadin bands and 65 gliadin patterns were detected. The average of genetic similarity coefficient based on gliadin bands was 0.5123. The cluster analysis indicated that all the accessions could be divided into 5 groups and the gliadin variations among T. polonicum accessions were associated with their geographic origins. Three and five HWM-GS alleles were detected at Glu-A1 and Glu-B1 loci, respectively. A total of 10 HWM-GS combinations were observed. The genetic diversity indices (H) at Glu-B1 loci (0.659) were much higher than that at Glu-A1 loci (0.271).

Related Articles in ASCI
Search in Google Scholar
View Citation
Report Citation

  How to cite this article:

Dong Pan, Liu Hong , Li Wei , Qi Peng-Fei , Wei Yu-Ming and Y.L. Zheng , 2007. Genetic Diversity of Storage Proteins in Triticum polonicum L.. Journal of Plant Sciences, 2: 416-424.

DOI: 10.3923/jps.2007.416.424



Compared with common wheat, Triticum polonicum L. (2n = 28, AABB) has some typical agronomic traits, such as large grains, more spikes and higher tillering ability (Zheng, 1989). T. polonicum is mainly cultivated in Mediterranean and Ethiopia and often mixed grown with durum wheat (Dong and Zheng, 2000). T. polonicum had been widely evaluated from its classification and distribution (Dong and Zheng, 2000; Jin et al., 1996), origin and evolution (Belea, 1971; Belea et al., 1975; Gorgidze and Zhizhilashvili, 1983; Orel et al., 1990; Yang et al., 1992), genetics (Puzyreva, 1971; Prasad, 1972; Zhangaziev, 1990; Watanabe et al., 1998; Efremova et al., 2001; Deng et al., 2005), bio-chemistry marker (Zahoor et al., 1986; Yang et al., 2000; Liu et al., 2001; Sissons and Batey, 2003) and molecular marker (Akond et al., 2007; Liao et al., 2007)

Wheat storage proteins are mainly composed of gliadins and glutenins and the glutenins could be subdivided into high and low molecular weight glutenin subunits (HMW- and LMW- GSs) (Wei et al., 2002). Gliadins encoded by Gli-1 and Gli-2 loci (Lafiandra et al., 1984) are generally considered to contribute to the viscosity and extensibility of gluten (Gianibelli et al., 2001; Khatkar et al., 2002; Clarke et al., 2003). HMW-GSs, controlled by three gene loci identified as Glu-A1, Glu-B1 and Glu-D1 (Payne et al., 1982), have closely associated with bread-making quality (Payne, 1987; Yahata et al., 2006; Chen et al., 2007). In addition, storage proteins are also reliable bio-chemical markers for the evaluation of genetic diversity in wheat (Branlard et al., 2001).

In this study, the genetic variations of gliadins and HMW-GSs in 72 T. polonicum accessions, which were widely collected from various areas, were investigated using A-PAGE and SDS-PAGE methods, respectively. The objective of this study was to evaluate the genetic diversity of T. polonicum accessions at the level of proteins with reference to the improvement of common wheat.


Plant Materials
A total of 72 T. polonicum accessions received from American national resource information net (GRIN) (Table 1) were derived from 6 main regions, including North America, South America, Europe, East Africa and West Asia, Center and South Asia and Australia. Common wheat cultivar, Chines Spring (CS) was used as reference in A-PAGE analysis. The relative motilities of the HMW glutenin subunits were determined by comparison with the references Chinese Spring (1Bx7, 1By8, 1Dx2, 1Dy12), Chuanyu12 (1Ax, 1Bx7, 1By8, 1Dx5, 1Dy10) and Xiaoyan 6 (1Ax1, 1Bx14, 1By15, 1Dx2, 1Dy11).

Gliadin proteins were extracted from single seeds with a solution of 25% (v/v) α-ethanol and 0.05% (w/v) methyl green and fractionated by a standard acid-plyacrylamide-ger electrophoresis (A-PAGE) at pH 3.1 (Draper, 1987).

Table 1: The geographical origin and HMW-GSs of 72 T. polonicum accessions
Image for - Genetic Diversity of Storage Proteins in Triticum polonicum L.

The HMW-GS extractions from single seeds were performed as described by Mackie et al. (1996). According to the procedure of Ng and Bushuk (1987), HMW-GSs were separated by polyacrylamide-gelelectrophoresis in the presence of sodium dodecyl sulfate (SDS-PAGE).

Allele Identification
Based on the classification of Payne and Lawrence (1983), the HMW-GSs were identified. The new subunits and alleles were designated according to Wang et al. (2006).

Data Analysis
The gliadin genetic similarity coefficient (GS) among the accessions was calculated by Nei and Li (1979) genetic similarity

GS = 2Nij/(Ni +Nj)

where, Ni is the bands of accession i, Nj is the bands of accession j and Nij is the bands in both accessions i and j. The genetic similarity matrix of all accessions was analyzed by using Unweighted Pair Group Method with Arithmetic Average (UPGMA) algorithm and the result was used to construct a clustering dendrogram under NTSYS-pc2.1.

The gene diversity at Glu-A1 and Glu-B1 loci was calculated by Nei (1973) genetic variation index,

H = 1-∑pi2

where, H is Nei’s genetic variation index and pi is the frequency of a particular allele at that locus.


Gliadin Variations
A total of 48 gliadin bands were detected in the 72 T. polonicum accessions (Fig. 1) and 8 to 24, with an average of 16.2, gliadin bands were identified in each accession. The α, β, ω and γ gliadin zones were identified by 11, 9, 13 and 15 bands, respectively.

Image for - Genetic Diversity of Storage Proteins in Triticum polonicum L.
Fig. 1: Gliadin patterns in the representative T. polonicum accessions. 1 = PI29447, 2 = PI42209, 3 = PI56261, 4 = PI56262, 5 = CItr13919, 6 = CItr14139, 7 = CItr14140, 8 = CItr14803, 9 = Citr14869, 10 = Citr17442, 11 = Citr134945, 12 = PI134945, 13 = PI167622, 14 = PI185309, 15 = PI190951, 16 = 191808, 17 = PI191810, 18 = PI191823

The frequency of each band ranged from 1.4 to 90.3%, with the mean of 33.7%. A total of 65 gliadin patterns were identified from 72 accessions, among which 58 accessions had the unique gliadin genotypes. The gliadin patterns in seven pairs of accessions (i.e., CItr14869 and CItr14892, PI306548 and PI306549, PI384338 and PI384339, PI384337 and PI384342, PI384337 and PI384344, PI384342 and PI384344 and PI629119 and PI608017) were identical, thus, these accessions could not be distinguished by A-PAGE.

The GS among the T. polonicum accessions varied from 0.1429 to 1.000, with the mean of 0.5132, indicating that higher genetic diversity existed in these T. polonicum accessions. The mean GS in the accessions from North America (0.6782) was the highest, followed by Europe (0.5802). The accessions from East Africa and West Asia and Center and South Asia, which were the origin center of wheat, had the higher genetic diversity, with the mean GS of 0.5473 and 0.4165, respectively. Due to the lack of enough accessions, the mean GSs among the accessions from South America and Australia were not calculated.

All the T. polonicum accessions could be divided into 5 groups (Fig. 2). Only one accession PI349051 from Georgia were included in Group I. Twelve accessions from Ethiopia and 1 accession from Egypt were clustered into Group II. Group III consisted of 4 accessions from Ethiopia and Group IV had 5 accessions from Portugal. The remaining 49 accessions were clustered into Group V, which included the all accessions from Hungary, England, the United States, Iraq and Australia and some accessions from European. These results suggested that most of the accessions with close geographic origins had the tendency to cluster together, indicating that the gliadin variations among T. polonicum accessions were associated with their geographic origins.

HWM-GS Variations
The HMW-GSs and their frequencies were identified in 72 T. polonicum accessions (Table 2 and Fig. 3). A total of 8 allelic variants at two Glu-1 loci were observed. At the Glu-A1 locus, only three HWM-GS alleles were detected and the genetic variation index (H) was 0.271. The frequency of Glu-A1c was 84.7%, indicating that Glu-A1c was the most frequent allele. Six accessions had the allele a (subunit 1) at the frequency of 8.3%. A novel subunit (allele Glu-A1-I), moving slightly slower than the subunit 5 but slightly faster than the subunit 2*, was present in five accessions (i.e., PI91808, PI191903, PI272565, PI352488 and CItrl14892) at the frequency of 7.0%.

At the Glu-B1 locus, five alleles were observed and the H (0.659) was much higher than that of Glu-A1 loci. Glu-B1a (subunit 7) and Glu-B1e (subunit 20) were the most frequent subunits with the frequencies of 40.3%, while the other two alleles (Glu-B1b and Glu-B1d) appeared at lower frequencies of 8.3 and 9.7%, respectively. Only one accession, PI19183, had the Glu-B1g (13+19) allele.

A total of 10 HMW-GS combinations (Table 1) were observed at the Glu-A1 and Glu-B1 loci. Two major genotypes (Null + 20 and Null + 7) were observed in higher frequencies with 41.67 and 30.56%, respectively. The remaining 8 patterns were observed at lower frequencies. The HWM-GS combinations among the accessions from different geographic regions were also analyzed. All the HWM-GS combinations detected in this study were also appeared in Europe.

Table 2: Allelic variations and frequencies of HMW glutenin subunits and genetic diversity indices (H) at Glu-1 in 72 T. polonicum accessions
Image for - Genetic Diversity of Storage Proteins in Triticum polonicum L.

Image for - Genetic Diversity of Storage Proteins in Triticum polonicum L.
Fig. 2: Clustering results of 72 T. polonicum accessions based on UPGMA method

Image for - Genetic Diversity of Storage Proteins in Triticum polonicum L.
Fig. 3: HMW-GS patterns in some T. polonicum accessions. 1 = PI191903, 2 = PI272565, 3 = Chinese Spring (CS), 4 = PI191823, 5 = CItr14139, 6 = PI134945, 7 = CItr139919, 8 = PI191826, 9 = Chuan Yu 12

The HMW-GS combination N + 7 was the most frequent combination with the frequency of 30.0%, followed by the combination N + 20 (24.2%). Four HMW-GS combinations were observed among 25 accessions from East Africa and west Asia and the combination N + 20 was also the most frequent combination (76%). In Australia only one subunit combination (N, 6+8) was detected and 2 combination were observed in other three regions.


It has been proposed that gliadin bands are an easy, cheap and dependable marker for the evaluation of wheat genetic resources. Wang et al. (2006) reported that the gliadin patterns could reflect the genetic diversity in durum wheat. Zhang et al. (1995), Hou et al. (2004) and Lan et al. (1999) suggested that A-PAGE method could be used for identifying and evaluating the wheat germplasm resources. In this study, higher variations of gliadins were detected in the T. polonicum. A total of 65 gliadin patterns were identified in 72 T. polonicum accessions.

Li et al. (2002) found that the genetic relationships of wild emmer wheat were associated with their geographical distribution. The similar results were also obtained in T. compactum (Zhang et al., 2005a), T. turanicum (Xu et al., 2005a) and T. durum (Wang et al., 2006). In this study, also it was observed that most of the accessions with close geographic origins had the tendency to cluster together, indicating that the gliadin variations among T. polonicum accessions were also associated with their geographic origins. Meanwhile, cluster analysis indicated that the genetic similarity was the lowest in the West Asia and Center Asia, which were the origin center of wheat.

It was reported that the genes encoding HWM-GS were a resertively restable and dependable marker (Nevo and Payne, 1987). In present study, the subunit null, coded by Glu-A1 locus, was the predominant subunit with the highest frequency of 84.7%, which is similar as in T. astivum (Zeng et al., 2005), T. macha (Xiong et al., 2005), T. turanicum (Xu et al., 2005b), T. durum (Wang et al., 2006) and T. compactum (Zhang et al., 2005b), whereas the predominant subunit in T. dicoccoides (Li et al., 2002), T. turgidum ssp. dicoccum (Li et al., 2006), T. carthlicum (Zhuang et al., 2006) and T. spelt (Xueli et al., 2005) was subunit 1 and subunit 2* was the most frequent subunit in T. turgidum landraces (Zhang et al., 2003). At Glu-B1 loci, subunits 7+8 were the predominant subunits in T. compactum (Zhang et al., 2005b), T. carthlicum (Zhuang et al., 2006), T. macha (Xiong et al., 2005) and T. astivum (Zeng et al., 2005), whereas T. turanicum (Xu et al., 2005b) and T. spelt (Xueli et al., 2005) had the predominant subunits 13+16. In this study, 7 and 20 subunits were the most frequent subunits with the frequency of 40.3%. Therefore, the predominant subunits in different species or subspecies of wheat germplasm were not identical.


This study was supported by the National High Technology Research and Development Program of China (863 program 2006AA10Z179 and 2006AA10Z1F8) and the FANEDD project (200357 and 200458) from Ministry of Education, China. Dr. Y.M. Wei was supported by the Program for New Century Excellent Talents in University of China. Prof. Y.L. Zheng was supported by the Program for Changjiang Scholars and Innovative Research Teams in University of China (IRT0453).


  1. Akond, M.A., N. Watanabe and Y. Furuta, 2007. Exploration of genetic diversity among Xinjiang triticum and Triticum polonicum by AFLP markers. J. Applied Genet., 48: 25-33.
    Direct Link  |  

  2. Belea, A., 1971. Are these other possibilities for the origin of T. spelta or T. aestivum? Yakubtsiner, M: Is T. carthlicum the product of T. durum H T. aestivum. Acta Agron. Acad. Sci. Hung., 20: 227-229.

  3. Belea, A., D. Fejer and K.K. Ghosal, 1975. Genetic analysis of Triticum ispahanicum Heslot hybrids. Nucl. India, 18: 50-53.

  4. Branlard, G., M. Dardevet, R. Saccomano, F. Lagoutte and J. Gourdon, 2001. Genetic diversity of wheat storage proteins and bread wheat quality. Euphytica, 119: 59-67.
    CrossRef  |  

  5. Chen, J., M. Ding and X. Chen, 2007. Analysis of the primary structure of high molecular weight blutenin subunits in wheat. J. Triticeae Crops, 27: 67-74.

  6. Clarke, B.C., T. Phongkham and M.C. Gianibelli, 2003. The characterisation and mapping of a family of LMW-gliadin genes: Effects on dough properties and bread volume. Theor. Applied Genet., 106: 629-935.
    Direct Link  |  

  7. Deng, X.F., Y.H. Zhou and R.W. Yang, 2005. Chromosomal location of genes for spike length in dwarfing polish wheat by monosomic analysis. J. Sichuan Agric. Univ., 23: 12-14.
    Direct Link  |  

  8. Dong, Y.S. and D.S. Zheng, 2000. Genetic Resource of Chinese Wheat. Chinese Agriculture Press, Beijing, pp: 46-47

  9. Draper, S.R., 1987. 1987 ISTA and 1987 Variety committee: Report of the working group for biochemical tests for cultivar identification 1983-1986. Seed Sci. Technol., 15: 431-434.
    Direct Link  |  

  10. Efremova, T.T., L.I. Laikova and V.S. Arbuzova, 2001. Use of aneuploid analysis in chromosomal localisation and wheat genes mapping. Proceedings of the 11th European Wheat Aneuploid Conference, July 24-28, 2001, Novosibirsk, Russia, pp: 115-118

  11. Gianibelli, M.C., O.R. Larroque, F. MacRitchie and C.W. Wrigley, 2001. Biochemeical, genetic and molecular characterization of wheat glutenin and its component subunits. Cereal Chem., 78: 635-646.
    Direct Link  |  

  12. Gorgidze, A.D. and K.M. Zhizhilashvili, 1983. Phylogeny of the wheat Triticum polonicum L. Bull. Acad. Sci. Georgian SSR, 109: 381-383.

  13. Hou, Y.C., Y.L. Zheng and Y.M. Wei, 2004. Analysis of genetic diversity of hordein in wild relatives of barley from Qing Zang plateau. Southwest China J. Agric. Sci., 17: 545-551.

  14. Jin, S.B., 1996. The Wheat of China. Chinese Agriculture Press, Beijing, pp: 259

  15. Khatkar, B.S., R.J. Fido, A.S. Tatham and J.D. Schofield, 2002. Functional properties of wheat gliadin: I. Effects on mixing characteristics and bread making quality. J. Cereal Sci., 35: 299-306.
    Direct Link  |  

  16. Lafiandra, D., D.D. Kasarda and R. Morris, 1984. Chromosomal assignment of genes coding for the wheat gliadin protein components of the cultivars Cheyenne and Chinese Spring by two dimensional (two-pH) electrophoresis. Theor. Applied Genet., 68: 531-539.
    CrossRef  |  

  17. Li, B.Y., C.J. Xie, M.S. You, S.F. Mao and G.T. Liu, 2002. Diversity of high molecular weight glutenin subunits (HMW-GS) in wild emmer. J. Triticeae Crops, 22: 21-24.

  18. Li, Q.Y., Y.M. Yan, A.L. Wang, X.L. An, Y.Z. Zhang, S.L.K. Hsam and F.J. Zeller, 2006. Detection of HWM glutenin subunit variations among 205 cultivated emmer accessions (Triticum turgidum ssp. Dicoccum). Plant Breed., 125: 120-124.

  19. Liao, J., R. Yang, Y. Zhou and H. Tsujimoto, 2007. FISH analysis of 45srDNA and 5srDNA genes in Triticum polonicum L. and T. turgidum L.W. Ailanmai. Genetics, 29: 449-454.

  20. Liu, X., A. Guo and Y. Li, 2001. An improved acid polyacylamid gel electrophoresis method and it's application in germ plasm resources analysis. J. Northwest Sci-Tech Univ. Agric. For. Nat. Sci., 29: 17-20.

  21. Mackie, A.M., E.S. Lagudah, P.J. Sharp and D. Lafiandra, 1996. Molecular and biochemical characterisation of HMW glutenin subunits from T. tauschii and the D Genome of hexaploid wheat. J. Cereal Sci., 23: 213-225.
    Direct Link  |  

  22. Nei, M., 1973. Analysis of gene diversity in subdivided populations. Proc. Natl. Acad. Sci. USA., 70: 3321-3323.
    PubMed  |  Direct Link  |  

  23. Nei, M. and W.H. Li, 1979. Mathematical model for studying genetic variation in terms of restriction endonucleases. Proc. Natl. Acad. Sci. USA., 76: 5269-5273.
    CrossRef  |  PubMed  |  Direct Link  |  

  24. Nevo, E. and P.I. Payne, 1987. Wheat storage proteins: diversity of HMW glutenin subunits in wild emmer from Israel. TAG Theor. Applied Genet., 74: 827-836.
    CrossRef  |  Direct Link  |  

  25. Ng, P.K.W. and W. Bushuk, 1987. Glutenin of Marquis wheat as a reference for estimating molecular weights of glutenin subunits by sodium dodecyl sulfate-polyacrylamide gel electrophoresis. Cereal Chem., 64: 324-327.
    Direct Link  |  

  26. Orel, L.I., R.A. Udachin and M.A. Agafonova, 1990. Morphological and karyological features of Triticum prtropavlovskyi Poaceae. Botanicheskii Zhurnal., 75: 37-44.

  27. Payne, P.I., L.M. Holt, A.Y. Worland and C.N. Law, 1982. Structural and genetical studies on the high-molecular-weight subunits of wheat, Part 3. Telocentric mapping of the subunit genes on the long arms of homoelogous group 1 chromosomes. Theory Applied Genet., 63: 129-138.

  28. Payne, P.I. and G.J. Lawrence, 1983. Catalogue of alleles for the complex gene loci, Glu-A1, Glu-B1 and Glu-D1 which code for high molecular-weight subunits of glutenin in hexaploid wheat. Cereal Res. Commun., 11: 29-35.

  29. Payne, P.I., 1987. Genetics of wheat storage proteins and the effect of allelic variation on bread-making quality. Annu. Rev. Plant Physiol., 38: 141-153.
    CrossRef  |  Direct Link  |  

  30. Prasad, M.V.R., 1972. Studies on induced mutants with reference to species relationships in some tetraploid Ttiticums. Theor. Applied Genet., 42: 160-167.

  31. Puzyreva, V.A., 1971. Segregation in interspecific hybrids of wheat. Nauchtr Tyumen skh Int., 9: 18-24.

  32. Sissons, M.J. and I.L. Batey, 2003. Protein and starch properties of some tetraploid wheat. Cereal Chem., 80: 468-475.
    Direct Link  |  

  33. Wang, H., Y. Wei, Z. Yan and Y. Zheng, 2006. Genetic variations of gliadin and HMW glutenins subunits in durum wheat. J. Agric. Biotechnol., 14: 721-727.

  34. Watanabe, N., Y. Yotani and M. Anada, 1998. Inheritance and the effects of a gene for long glume: A key character for taxonomy. Proceedings of the 3rd International Triticeae Symposium, Aleppo, Syria, May 4-8, 1998, IEEE Xplore, London, pp: 103-108

  35. Wei, Y., Y. Zheng, D. Liu, Y. Zhou and X. Lan, 2002. HWM-glutenin and gliadin variations in Tibetan weedrace, Xinjiang rice wheat and Yunnan hulled wheat. Genet. Resour. Crop Evol., 49: 327-330.

  36. Xiong, L., Y. Zheng, Y. Wei and W. Li, 2005. High-molecular-weight glutenin subunits variation in Triticum macha dek. et men. J. Sichuan Agric. Univ., 23: 264-267.
    Direct Link  |  

  37. Xu, L., Z. Yan, Y. Wei and Y. Zheng, 2005. Analysis of HWM glutenin subunits in Triticum turanicum Jakubz. J. Sichuan Agric. Univ., 23: 137-141.

  38. Xu, L., W. Li, Y. Wei and Y. Zheng, 2005. Genetic diversity of Triticum turanicum jakubz. based on gliadin analysis. J. Plant Genet. Resour., 6: 195-199.

  39. Xueli, A., L. Qiaoyun, Y. Yueming, X. Yinghua, S.L.K. Hsam and F.J. Zeller, 2005. Genetic diversity of European spelt wheat (Triticum aestivum ssp. Spelta L. em. Thell.) revealed by glutenin subunit variations at the Glu-1 and Glu-3 loci. Euphytica, 146: 193-201.
    Direct Link  |  

  40. Yahata, E., W. Maruyama-Funatsukiw, Z. Nishio Y. Yamamoto and A. Hanaoka et al., 2006. Relationship between the dough quality and content of specific glutenin proteins in wheat mill streams and its application to making flour suitable for instant Chinese noodles. Biosci. Biotechnol. Biochem., 70: 788-797.
    Direct Link  |  

  41. Yang, W.Y., C. Yen and J.L. Yang, 1992. Cytogenetic study on the origin of some special Chinese landraces of common wheat. Wheat Inf. Serv., 75: 14-20.

  42. Yang, R W., Y.H. Zhou, Y.L. Zheng and C. Hu, 2000. Genetic differences and the relationship of gliadin between T. polonicum and T. petropavlovskyi. J. Triticeae Crops, 20: 1-5.
    Direct Link  |  

  43. Zahoor, A., R. Habibur and M. Tahir, 1986. Histochemical distribution of proteins in wheat and its relationship with important genetic traits. Pak. J. Agric. Res, 6: 82-85.

  44. Zeng, X., W. Ji, X. Qiang and Y. Tang, 2005. Analysis on the component of HWM-GS of Tibet wheat variety resources. J. Triticeae Crops, 25: 98-101.

  45. Zhang, X., X. Yang and Y.S. Dong, 1995. Genetic analysis of wheat germplasm by acid polyacrylamide gel electrophoresis of gliadins. Sci. Agric. Sin. A, 28: 25-32.
    Direct Link  |  

  46. Zhang, D.F., Y.L. Zheng, Y.M. Wei, Z.H. Yan and Z.Q. Zhang, 2003. Composition analysis of high molecular weight glutenin subunits in Triticum Turgidum L. ssp. Turgidum. J. Triticeae Crops, 23: 29-32.

  47. Zhang, X., Z. Guo, Z. Yan, Y. Wei and Y. Zheng, 2005. Composition of the subunits at Glu-1 composition loci in Triticum compactum. Acta Agric. Bureali-Occidentalis Sin., 14: 25-29.

  48. Zhang, X., W. Li, Y. Wei and Y. Zheng, 2005. Genetic Variability of gliadin in Triticum compactum host. Southwest China J. Agric. Sci., 18: 680-683.

  49. Zhangaziev, A.S., 1990. Phylogenetic relationship in tetroploid wheat species. Problemy teoreticheskoi I prikladnoi genetiki v Kazakhatane: Materialy respublikanskoi komferentsii. Alma Ata, 25: 18-22.

  50. Zheng, D., 1989. The species and subspecies in China. Crop Germplasm Resour., 2: 1-3.

  51. Zhuang, P., Z. Guo, Z. Yan, Y. Wei and Y. Zheng, 2006. Analysis of HWM glutenin subunits in Triticum carthlicum nevski. Southwest China J. Agric. Sci., 9: 5-9.

©  2022 Science Alert. All Rights Reserved