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Genetic and in silico comparative mapping of the polyphenol oxidase gene in bread wheat (Triticum aestivum L.).

Polyphenol oxidases (PPOs) are involved in the time-dependent darkening and discolouration of Asian noodles and other wheat end products. In this study, a doubled haploid (DH) population derived from Chara (moderately high PPO activity)/WW2449 (low PPO activity) was screened for PPO activity based on L-DOPA and L-tyrosine assays using whole seeds. Both these assays were significantly genetically correlated (r = 0.91) in measuring the PPO activity in this DH population. Quantitative trait loci (QTLs) analysis utilising a skeleton map enabled us to identify a major QTL controlling PPO activity based on L-DOPA and L-tyrosine on the long arm of chromosome 2A. The simple sequence repeat (SSR) marker GWM294b explained over 82% of the line mean phenotypic variation from samples collected in both 2000 and 2003. Four SSR markers were validated for PPO linkage in genetically diverse backgrounds and proven to correctly predict the PPO activity in more than 92% of wheat lines. Physical mapping using deletion lines of Chinese Spring has confirmed the location of the GWM294b, GWM312 and WMC170 on chromosome 2AL, between deletion breakpoints 2AL-C to 0.85. In order to identify functional gene markers, data searches for alignments between rice BAC/PAC clones assembled on chromosome 1 and 4, chromosome 7, and (1) the wheat expressed sequence tags mapped in deletion bin (2AL-C to 0.85) and (2) the coding sequence of a previously cloned wheat PPO gene were made and found significant sequence similarities with the PPO gene or common central domain of tyrosinase. Available PPO gene sequences in the National Centre for Biotechnology Information (NCBI) database have revealed that there is a significant molecular diversity at the nucleotide and amino acid level in the wheat PPO genes.

Alleles↗

[Development and identification of a set of Triticum aestivum-Thinopyrum bessarabicum disomic alien addition lines].

In order to transfer the genes for salt tolerance and disease resistance from Thinopyrum bessarabicum into wheat, the hybrid progenies between T. aestivum cv. Chinese Spring and T. aestivum cv. Chinese Spring-amphiploid Th. bessarabicum were screened. A set of T. aestivum-Th. bessarabicum disomic addition lines was developed with the assistance of mitotic chromosome C-banding and genomic in situ hybridization (GISH), as well as GISH on meiotic M I chromosome preparations. The results indicated that all the wheat chromosomes in the lines remained unchanged karyotypically, while the added Th. bessarabicum chromosomes paired regularly in meiosis. The developed disomic addition lines were designated temporarily as DAJ1, DAJ2, DAJ3, DAJ4, DAJ5, DAJ6 and DAJ7 respectively. Determination of the homoelogous groups of the added Th. bessarabicum chromosomes and localization of the genes for salt tolerance and disease resistance are undergoing.

Adaptation, Physiological↗

Expression of a cold-responsive Lt-Cor gene and development of freezing tolerance during cold acclimation in wheat (Triticum aestivum L.).

Time-courses of the development of freezing tolerance and the expression of a cold-responsive gene wlt10 were monitored during cold acclimation in wheat (Triticum aestivum L.). Bioassay showed that cold acclimation conferred much higher freezing tolerance on a winter cultivar than a spring cultivar. Northern blot analysis showed that the expression of wlt10 encoding a novel wheat member of a cereal-specific LT-COR protein family was specifically induced by low temperature. A freezing-tolerant winter cultivar accumulated the mRNA more rapidly and for a longer period than a susceptible spring cultivar. The increase in the amount of mRNA was temporary but the peak occurred at the time when the maximum level of freezing tolerance was attained. The mRNA accumulated more in the leaves than in the roots, and different light/dark regimes modulated the level of mRNA accumulation. Genomic Southern blot analyses using the nulli-tetrasomic series showed that the wlt10 homologues were located on the homologous group 2 chromosomes.

Acclimatization↗

Alfalfa rapidly remediates excess inorganic nitrogen at a fertilizer spill site.

By 19%, standard remediation techniques had significantly reduced the concentration of nitrate nitrogen (NO3- -N) in local ground water at the site of a 1989 anhydrous ammonia spill, but NO3- -N concentrations in portions of the site still exceeded the public drinking water standard. Our objective was to determine whether local soil and ground water quality could be improved with alfalfa (Medicago sativa L.). A 3-yr study was conducted in replicated plots (24 by 30 m) located hydrologically upgradient of the ground water under the spill site. Three alfalfa entries ['Agate', Ineffective Agate (a non-N2-fixing elite germplasm similar to Agate), and MWNC-4 (an experimental germplasm)] were seeded in the spring of 1996. Corn (Zea mays L.) or wheat (Triticum aestivum L.) was seeded adjacent to the alfalfa each year. Crops were irrigated with N-containing ground water to meet water demand. During the 3-yr period, about 540 kg of inorganic N was removed from the aquifer through irrigation of 4.9 million L water. Cumulative N removal from the site over 3 yr was 972 kg N ha(-1) in Ineffective Agate alfalfa hay, compared with 287 kg N ha(-1) for the annual cereal grain. Soil solution NO3- concentrations were reduced to low and stable levels by alfalfa, but were more variable under the annual crops. Ground water quality improved, as evidenced by irrigation water N concentration. We do not know how much N was removed by the N2-fixing alfalfas, but it appears that either fixing or non-N2-fixing alfalfa will effectively remove inorganic N from N-affected sites.

Agriculture↗

[Localization of DNA probes for human ribosomal genes on barley chromosomes].

To estimate the possibility of plant genome mapping using human genome probes, the probes fluorescent in situ hybridization (FISH) of human 18S-28S rDNA (clon 22F9 from the LA-13NCO1 library) was carried out on chromosomes of the spring barley Hordeum vulgare L. As a control, wheat rDNA probe (clon pTa71) was taken. Hybridization of the wheat DNA probe revealed two major labelling sites on mitotic barley chromosomes 5I (7H) and 6I (6H), as well as several minor sites. With the human DNA probe, signals were detected in the major sites of the ribosomal genes on chromosomes 5I (7H) and 6I (6H) only when the chromosome preparations were obtained using an optimized technique with obligatory pepsin treatment followed by hybridization. Thus, this study demonstrates that physical mapping of plant chromosomes with human DNA probes that are 60 to 75% homologous to the plant genes is possible. It suggests principal opportunity for the FISH mapping of plant genomes using probes from human genome libraries, obtained in the course of the total sequencing of the human genomes and corresponding to the coding regions of genes with known functions.

Chromosomes↗

Characterization of trinucleotide SSR motifs in wheat.

Length differences among trinucleotide-based microsatellite alleles can be more easily detected and frequently produce fewer "stutter bands" as compared to dinucleotide-based microsatellite markers. Our objective was to determine which trinucleotide motif(s) would be the most-polymorphic and abundant source of trinucleotide microsatellite markers in wheat ( Triticum aestivumL.). Four genomic libraries of cultivar 'Chinese Spring' were screened with nine trinucleotide probes. Based on the screening of 28550 clones, the occurrences of (CTT/GAA) (n), (GGA/CCT) (n), (TAA/ATT) (n), (CAA/GTT) (n), (GGT/CCA) (n), (CAT/GTA) (n), (CGA/GCT) (n), (CTA/GAT) (n), and (CGT/GCA) (n) repeats were estimated to be 5.4x10(4), 3.5x10(4), 3.2x10(4), 1.2x10(4), 6.3x10(3), 4.9x10(3), 4.5x10(3), 4.5x10(3) and 3.6x10(3), i.e., once every 293 kbp, 456 kbp, 500 kbp, 1.3 Mbp, 2.6 Mbp, 3.2 Mbp, 3.6 Mbp, 3.6 Mbp and 4.5 Mbp in the wheat genome, respectively. Of 236 clones selected for sequencing, 38 (93%) (TAA/ATT) (n), 30 (43%) (CTT/GAA) (n), 16 (59%) (CAA/GTT) (n), 3 (27%) (CAT/GTA) (n) and 2 (4%) (GGA/CCT) (n) clones contained microsatellites with eight or more perfect repeats. From these data, 29, 27 and 16 PCR primer sets were designed and tested to the (TAA/ATT) (n), (CTT/GAA) (n) and (CAA/GTT) (n) microsatellites, respectively. A total of 12 (41.4%) primers designed to (TAA/ATT) (n), four (14.8%) to (CTT/GAA) (n), and two (12.5%) to (CAA/GTT) (n) resulted in polymorphic markers. The results indicated that (TAA/ATT) (n) microsatellites would provide the most-abundant and the most-polymorphic source of trinucleotide microsatellite markers in wheat.

Journal Article↗

[Structural changes of 4V chromosome of Haynaldia villosa induced by gametocidal chromosome 3C of Aegilops triuncialis].

Chromosome 3C of Aegilops triuncialis was discovered with ability to be transferred preferentially in the case of its monosomic status in wheat background, whereas, those gametes without 3C would result in chromosome structural changes including deletions and translocations. In the present study, Triticum aestivum-Haynaldia villosa substitution line 4V(4D) developed in our laboratory, was crossed to T. aestivum c.v. Norin 26-Aegilops triuncialis 3C addition line, and the hybrids F1 were then backcrossed with common wheat in order to induce structural changes of 4V. Both chromosome C-banding and genomic in situ hybridization was applied to search such chromosome variations. In this case, total genomic DNA of Haynaldia villosa was labelled by Biotin-11-dUTP as probes and total genomic DNA of Chinese Spring as the block. Moreover, several chromosome changes within common wheat such as isochromosome 1BL.1BL(B39-2) and others were also revealed. The result indicated that two translocation lines T4VL.3AS(A47-10-3) and T4VS.4DL(A47-25-4), two telocentric chromosome lines A47-7-2(4VS) and A47-32-2(4VL), and two isochromosomes including 4VS.4VS(A47-23) and 4VL.4VL(A412-5-4) were identified from BC1F2 or BC1F3. This result indicated that gametocidal chromosome 3C of Aegilop triuncialis could effectively induce structural changes of both chromosome 4V of Haynaldia villosa and chromosomes of wheat.

Chromosome Aberrations↗

Fusarium head blight resistance in hexaploid wheat (Triticum aestivum)-Lophopyrum genetic lines and tagging of the alien chromatin by PCR markers.

The objective of this research was to identify Fusarium head blight (FHB) resistance in wheat ( Triticum aestivum)- Lophopyrum genetic lines that might complement FHB resistance in common wheat; and to identify DNA markers that can be used to tag the resistance gene in the alien chromatin (E or el(2) genome) for the development of improved wheat cultivars. FHB resistance was evaluated in 19 Chinese Spring- Lophopyrum elongatum (EE) substitution lines, two Thatcher -L. ponticum (el(1) and el(2)) substitution lines, and four Thatcher- L. ponticum translocation lines. Significant resistance was identified in the substitution lines 7E(7A), 7E(7B), and 7E(7D). The homoeologous chromosome, 7el(2),()also showed resistance in the Thatcher genetic background. Both the Thatcher-7el(1) substitution and translocation lines were susceptible, like Thatcher, indicating that there is no resistance gene on the 7el(1) chromosome. Simple sequence repeat (SSR) and cleaved amplified polymorphic sequences (CAPS) in homoeologous group 7 chromosomes were used to identify DNA markers located on 7E and 7el(2). As expected, the transferability of wheat SSR markers to Lophopyrum is low. Of the 52 SSR markers that we tested, only five were found to be co-dominant on 7E of L. elongatum versus 7A, 7B, and 7D, one of which is also positive on 7el(2). A CAPS marker, derived from the RFLP probe PSR129, can serve as a dominant marker for 7el(2) chromatin.

Fusarium↗

Escape protein for beef cows: II. Source and level in ammoniated wheat straw-corn silage diets.

Two experiments, using Angus x Hereford spring-calving beef cows in mid- or late lactation nursing Simmental-sired calves, were conducted to evaluate the relative value of a corn gluten meal-blood meal mixture (CGM-BM; 50% of supplemental protein from each source). In Exp. 1 (78 d), cows in late lactation were assigned to one of three treatments: control at 8.2% CP (C), soybean meal at 10.4% CP (SBM), or CGM-BM at 10.3% CP. Diets were calculated to be isocaloric at 55% TDN. In Exp. 2 (65 d), cows in mid-lactation were assigned to four treatments: urea, SBM, low CGM-BM (LM), and high CGM-BM (HM). Diets in Exp. 2 were isonitrogenous (9.5% CP) and isocaloric (55% TDN). Diets in both experiments were based on ammoniated wheat straw and corn silage. Weight gains of cows and cow-calf pairs were greater (P less than .06) when protein was supplemented in Exp. 1. Gains were lower for cows fed urea (P less than .03) in Exp. 2 but were similar when cows were supplemented with SBM vs either the low or the high level of CGM-BM. Performance of calves did not differ among dietary treatments.

Animal Feed↗

[Developing waxy wheat with backcrossing approach and molecular markers-assisted selection].

A series of Chinese Spring Wx loci near-isogenic lines were used to identify the specific bands of 6 STS markers and 1 CAPS marker of Wx genes. After optimizing PCR amplification and separation of PCR products, 3 co-dominant and dominant STS-markers were identified at the Wx-A1, Wx-D1 loci and Wx-B1 loci, respectively, which were used in molecular markers-assisted selection of waxy wheat in our breeding program. A cross between waxy wheat "98Y1441" and variety "Chuanyu 12" was made, and I2-KI dying method was used to select full waxy genotype individual from F2 seeds. The full waxy genotype individual was crossed with recurrent parent "Chuanyu 12". After 5 generations of self-cross and backcross, BC5F2 segregating population was obtained. Using the markers above, 8 Wx genotypes were developed from the BC5F2 population, which follows Mendelian segregation. Among them, there were 2 aabbdd waxy plants whose amylose content in the BC5F3 seeds were 1.81% and 0.82%, respectively. In addition, there were partial waxy plants (AAbbdd and aabbDD) whose amylose content in the BC5F3 seeds were 15.24 % and 17.57%, respectively. Most agricultural traits of the BC5F2 segregating population resembled those of the recurrent parent "Chuanyu 12", and superior to waxy wheat parent "98Y1441". This shows that backcross approach in combination with molecular markers-assisted selection of waxy genes is helpful to develop partial and full waxy wheat with good traits in the waxy wheat breeding program.

Breeding↗

Construction of comparative genetic maps of two 4Bs.4Bl-5Rl translocations in bread wheat (Triticum aestivum L.).

The physical length of the rye segment of a 4BS.4BL-5RL translocation derived from the Cornell Wheat Selection 82a1-2-4-7 in a Triticum aestivum 'Chinese Spring' background was measured using genomic in situ hybridization (GISH) and found to be 16% of the long arm. The size of this translocation was similar to previously published GISH measurements of another 4BS.4BL-5RL translocation in a Triticum aestivum 'Viking' wheat background. Molecular maps of both 4BS.4BL-5RL translocations for 2 different wheat backgrounds were developed using RFLP analysis. The locations of the translocation breakpoints of the 2 4BS.4BL-5RL translocations were similar even though they arose in different populations. This suggests a unique property of the region at or near the translocation breakpoint that could be associated with their similarity and spontaneous formation. These segments of rye chromosome 5 also contain a gene for copper efficiency that improves the wheat's ability to cope with low-copper soils. Genetic markers in these maps can also be used to screen for copper efficiency in bread wheat lines derived from the Cornell Wheat Selection 82a1 2-4-7.

Chromosome Mapping↗

Plant contents of magnesium, calcium and potassium in relation to ruminant nutrition.

Grass tetany and wheat pasture poisoning are metabolic diseases of mature, lactating beef cattle. In some other countries, similar problems occur with dairy cattle. In grass tetany, the animals generally are grazing cool-season forages in which Mg concentration or bioavailability of plant Mg is low. Levels of Mg in the blood serum also generally are low. Grass tetany can occur in beef cattle fed hay made from grass, small grains or alfalfa Medicago sativa L.). In wheat pasture poisoning where animals are grazing small grains forages, blood levels of Ca often are low, and blood Mg also may be low. Both grass tetany and wheat pasture poisoning occur when plants are growing rapidly in the spring, at the time of heavy lactation demand by ruminants for Mg and Ca. When the temperature increases and plants start to grow rapidly in the spring, concentrations of K, N, organic acids, and the ratio of K/(Ca + Mg) all increase, and the percent dry matter decreases. Much research remains to be done to understand all the soil, plant, and animal aspects of grass tetany and wheat pasture poisoning.

Animal Nutritional Physiological Phenomena↗

Dissecting a wheat QTL for yield present in a range of environments: from the QTL to candidate genes.

Previous studies with 95 bread wheat doubled haploid lines (DHLs) from the cross Chinese Spring (CS)xSQ1 trialled over 24 yearxtreatmentxlocations identified major yield quantitative trait loci (QTLs) in homoeologous locations on 7AL and 7BL, expressed mainly under stressed and non-stressed conditions, respectively. SQ1 and CS contributed alleles increasing yield on 7AL and 7BL, respectively. The yield component most strongly associated with these QTLs was grains per ear. Additional results which focus on the 7AL yield QTL are presented here. Trials monitoring agronomic, morphological, physiological, and anatomical traits revealed that the 7AL yield QTL was not associated with differences in flowering time or plant height, but with significant differences in biomass at maturity and anthesis, biomass per tiller, and biomass during tillering. In some trials, flag leaf chlorophyll content and leaf width at tillering were also associated with the QTL. Thus, it is likely that the yield gene(s) on 7AL affects plant productivity. Near-isogenic lines (NILs) for the 7AL yield QTL with CS or SQ1 alleles in an SQ1 background showed the SQ1 allele to be associated with >20% higher yield per ear, significantly higher flag leaf chlorophyll content, and wider flag leaves. Epidermal cell width and distance between leaf vascular bundles did not differ significantly between NILs, so the yield-associated gene may influence the number of cell files across the leaf through effects on cell division. Interestingly, comparative mapping with rice identified AINTEGUMENTA and G-protein subunit genes affecting lateral cell division at locations homologous to the wheat 7AL yield QTL.

Alleles↗

Identification of SNPs and development of allele-specific PCR markers for gamma-gliadin alleles in Triticum aestivum.

The coding regions of 28 entries of hexaploid wheat gamma-gliadin genes, gene fragments or pseudogenes in GenBank were used for nucleotide alignment. These sequences could be divided into nine subgroups based on nucleotide variation. The chromosomal locations of five of the seven unassigned subgroups were identified through subgroup-specific polymerase chain reactions (PCR) using Chinese Spring group-1 nulli-tetrasomic lines. Multiple single nucleotide polymorphisms (SNPs) and small insertions/deletions were identified in each subgroup. With further mining from wheat expressed sequence tag databases and targeted DNA sequencing, two SNPs were confirmed and one SNP was discovered for genes at the Gli-A1, Gli-B1 and Gli-D1 loci. A modified allele-specific PCR procedure for assaying SNPs was used to generate dominant DNA markers based on these three SNPs. For each of these three SNPs, two allele-specific primer sets were used to test Chinese Spring and 52 commercial Australian wheat varieties representing a range of low-molecular-weight (LMW) alleles. PCR results indicated that all were positive with one of the primer sets and negative with the other, with the exception of three varieties containing the 1BL/1RS chromosomal translocation that were negative for both. Furthermore, markers GliA1.1, GliB1.1 and GliD1.1 were found to be correlated with Glu-A3 a, b or c, Glu-B3 b, c, d or e and Glu-D3 a, b or e LMW glutenin alleles, respectively. Markers GliA1.2, GliB1.2 and GliD1.2 were found to be correlated with the Glu-A3 d or e, Glu-B3 a, g or h and Glu-D3 c alleles, respectively. These results indicated that the gamma-gliadin SNP markers could be used for detecting linked LMW glutenin subunit alleles that are important in determining the quality attributes of wheat products.

Alleles↗

[Isolation and characterization of two closely linked phenylalanine ammonia-lyase genes from wheat].

Two closely linked, highly homologous phenylalanine ammonia-lyase (PAL) genes located in one phage clone, PAL1 and PAL2, were isolated from a wheat genomic library by using a polymerase chain reaction (PCR) fragment of a wheat PAL gene as probe. The two PAL genes were located approximately from 7 kb apart and displayed 93% identity with the same orientation. Southern blot analysis with a PAL1 specific fragment as probe showed the presence of a multiple gene family of PAL in wheat. Northern analyses demonstrated a differential expression of PAL in two Chinese Spring near-isogenic lines upon infection with stem rust fungus Puccinia graminis. In the resistant Chinese Spring Sr11 isogenic line with the resistance gene Sr11 that is known to interact with the avirulence gene P11 from the stem rust fungus, an induced expression of PAL was observed 4 days post inoculation (d.p.i.), and a massive induction was evident 8 d.p.i. By contrast, in the susceptible Chinese Spring sr11 line lacking the resistance gene, induction of PAL was seen 6 d.p.i. and the expression level at 8 d.p.i. was similar to that observed in the resistant line at 6 d.p.i. In wheat suspension culture cells, treatment with either an elicitor isolated from the stem rust fungus or chitin oligomers could activate PAL gene expression within 2 h. The fungal elicitor appeared to be more active in early activation of the PAL gene than were chitin oligomers. These results demonstrated that wheat PAL played an important role in the induced resistance response upon infection with stem rust fungus at transcription level.

Basidiomycota↗

Mitotic instability in wheat x Thinopyrum ponticum derivatives revealed by chromosome counting, nuclear DNA content and histone H3 phosphorylation pattern.

To evaluate the mitotic stability of Triticum aestivum x Thinopyrum ponticum derivatives (BC(2)F(7) and BC(2)F(5) doubled haploids), chromosome counting by both conventional and immunostaining techniques, and measurement of DNA content were performed. The wheat progenitor line, PF 839197, the wheat recurrent parent CEP 19 and the control Chinese Spring were also investigated. In the hybrid derivatives, chromosome number ranged from 2n=36 to 60, with a predominance of chromosome numbers higher than 2n=42, that was confirmed by determination of nuclear DNA content. Chinese Spring' and PF 839197 were stable, but CEP 19 showed chromosome number variation (20%). Analyses of non-pretreated cells revealed the presence of anaphase bridges, lagging chromatids, chromosome fragments and micronuclei. Immunostaining with an antibody recognizing histone H3 phosphorylated showed dicentric chromatids forming anaphase bridges and pericentromeric phosphorylation at centric chromosome fragments but not at lagging chromatids. The possible causes of the observed mitotic instability are discussed.

Cell Nucleus↗

Gene-dosage compensation of endosperm proteins in hexaploid wheat Triticum aestivum.

Several aneuploid lines and one intervarietal substitution line of the hexaploid wheat Triticum aestivum (2n = 6x = 42; genomes AABBDD) cv. Chinese Spring were used to study the effects of different doses of chromosomes 1B, 1D, or 1A on the amount of the high molecular weight ("HMW") glutenins and gliadins of endosperm. These homeologous chromosomes carry HMW glutenin and gliadin gene clusters on their long and short arms, respectively. Increasing the dosage of chromosome 1B of Chinese Spring in plants having in their 3n endosperm zero or the normal three doses of the homeologue 1D, as well as in plants carrying in their endosperm one dose of 1B of the cultivar Timstein, had a dual effect: on one hand, a nonlinear increase in the amount of each subunit encoded by the chromosome whose dosage was elevated and, on the other hand, a compensating nonspecific decrease in the amount of other HMW glutenin and gliadin subunits encoded either by the homeoalleles on 1D or by the homoalleles on 1B of Timstein, respectively. Deletion of chromosome arm 1BL, which carries only a few HMW glutenin genes, had no significant effect on the amount of HMW glutenins encoded by 1DL and HMW gliadins encoded by 1DS and 1BS. However, deletion of 1BS or 1DS, each carrying many gliadin genes, caused a significant but nonspecific increase in the HMW glutenins and gliadins encoded by the remaining arms of 1B and 1D. The possible mechanism and evolutionary implications of gene-dosage compensation in polyploid wheat are discussed.

Journal Article↗

[Quality variation of wheat planted in different regions of Tibet and Beijing].

In order to supply experimental basis for developing high quality and special use wheat for Tibet, multi-varieties experiments in different regions were carried out to study the quality variation of wheat varieties during 1998 to 2001. Field trials of sowing in spring and in autumn were conducted in Linzhi, Rikaze, Lasa and Beijing, and 5 indices including grain protein content, wet gluten content, sedimentation value, falling number and grain hardness were determined. The results showed that the quality characters were significantly different among the varieties, and extremely significant among the sites of same variety. The protein content of grains had a positive correlation to the other 4 indices, and could be a core index in quality characters of wheat. The parameters of quality characters were higher in inland varieties than in Tibet ones, the latter appearing weak or medium gluten, and those of the same varieties were higher in Beijing than in Tibet. When inland varieties were planted in Tibet, the parameters of quality characters would be decreased. The same variety in 3 sites of Tibet had different quality characters. There was a positive correlation between ecological height (latitude x elevation) and growth-development duration of varieties, but a negative correlation between growth-development duration and protein content, and between ecological height and protein content. Medium gluten varieties should be mainly planted in Tibet, and some good inland varieties could be introduced and planted according to the quality regionalization.

China↗