The influence of a spring habit gene, Vrn-D1, on heading time in wheat

The adaptability of wheat cultivars to environmental conditions is known to be associated with a vernalization requirement, that is, spring/winter habit. To clarify the genetic effect of the spring habit gene, Vrn‐D1, on heading time in the field, recombinant inbred lines (RILs) with or without the Vrn‐D1 gene were produced from F₂ plants of the cross between ‘Nanbukomugi’ and ‘Nishikazekomugi’, non‐carrier and carrier cultivars of this gene, respectively. Using growth chambers with a controlled temperature and photoperiod, three components of heading time, i.e. vernalization requirement, photoperiodic sensitivity and narrow‐sense earliness (earliness per se), were evaluated in each RIL. RILs with the Vrn‐D1 gene (E lines) showed greatly reduced vernalization requirements and slightly shorter narrow‐sense earliness than RILs without Vrn‐D1 (L lines), although no difference in photoperiodic sensitivity was observed between the two groups. RILs were planted at four different sites in Japan and examined for their heading time in the field. E lines headed significantly earlier than L lines at all locations, indicating that the earliness of E lines is stable in various environmental conditions. These results indicated that spring habit caused by Vrn‐D1 gene, as well as narrow‐sense earliness, was responsible for heading time in the field.     

Characterization of QEet.ocs-5A.1, a quantitative trait locus for ear emergence time on wheat chromosome 5AL

QEet.ocs‐5A.1, a quantitative trait locus controlling ear emergence time, has been detected on wheat chromosome 5AL using single chromosome recombinant lines (SCRs) developed from a cross between ‘Chinese Spring’ (CS) (‘Cappelle‐Desprez’ 5A) and CS (Triticum spelta 5A). This locus has little influence on grain yield and its components, and thus has breeding potential for changing ear emergence time without yield reduction. To characterize the phenotypic expression of QEet.ocs.1 and to test its interaction with the Vrn‐A1 gene for vernalization response, six near‐isogenic SCRs differing for these two gene regions were grown together with the parental controls under different vernalization and photoperiod regimes. The T. spelta allele of QEet.ocs.1 accelerated heading time when vernalization and photoperiod were satisfied, demonstrating that the function of this QTL is earliness per se. There was no interaction between Vrn‐A1 and QEet.ocs.1.     

The influence of photoperiod on the Vrn-H2 locus (4H) which is a major determinant of plant development and reproductive fitness traits in a facultative × winter barley (Hordeum vulgare L.) mapping population

To determine the effect of Vrn‐H2 locus on plant developmental and agronomic traits, detailed controlled environment tests involving a factorial set of vernalization and photoperiod treatments were carried out using doubled haploid lines developed from a facultative (Vrn‐H2^?) × winter (Vrn‐H2⁺) barley cross. The allele phase at the Vrn‐H2 locus influenced heading date as well as the developmental and agronomic traits. The performance of Vrn‐H2⁺ lines was significantly influenced by vernalization: reproductive fitness traits showed significant decreases without vernalization. However, the effects of alleles at the Vrn‐H2 locus extended beyond simple satisfaction of the vernalization requirement. Vrn‐H2⁺ lines showed increased reproductive fitness compared with Vrn‐H2^? lines when vernalization was followed by a long photoperiod. The responses of the two Vrn‐H2 allele classes to photoperiod duration were quite different in terms of heading date, developmental and agronomic traits. These results suggest that alleles at the Vrn‐H2 locus – and/or tightly linked gene(s) – respond primarily to the exogenous signal of vernalization (temperature), but when the vernalization requirement has been fulfilled, they also respond to photoperiod duration.     

Genetic and Molecular Characterization of Vernalization Genes Vrn‐A1, Vrn‐B1, and Vrn‐D1 in Spring Wheat Germplasm from the Pacific Northwest Region of the U.S.A.

The objective of this study was to determine the Vrn‐1 allelic composition of spring wheat germplasm from the Pacific Northwest region of the USA. Individual plants from 56 spring wheat lines were crossed to near‐isogenic tester lines carrying the dominant allele Vrn‐A1, Vrn‐B1 or Vrn‐D1. F₂ progeny were evaluated for growth habit in the field and Vrn‐1 allelic composition was determined through chi‐square analysis. Lines also were analysed with DNA sequence‐based Vrn‐1 allele‐specific markers. A majority of the germplasm carried the dominant allele Vrn‐A1a alone or in combination with Vrn‐B1, Vrn‐D1 or Vrn‐B3 alleles. Vrn‐B1 and Vrn‐D1 were almost always associated with other dominant Vrn‐1 allele(s). Based on DNA sequence analysis, a novel Vrn‐B1 allele referred to as Vrn‐B1b, which carried a single nucleotide polymorphism (SNP) and a 36 bp deletion, was identified in cultivar ‘Alpowa’. These results will be useful to wheat breeders for choosing parents with different Vrn‐1 alleles for crossing to maximize diversity at the Vrn‐1 loci with an expectation of identifying superior Vrn‐1 allelic combinations for cultivar improvement.     

Evaluation of cold hardiness in two sets of near-isogenic lines of wheat (Triticum aestivum) with polymorphic vernalization alleles

In wheat, variation at the orthologus Vrn‐1 loci, located on each of the three genomes, A, B and D, is responsible for vernalization response. A dominant Vrn‐1a allele on any of the three wheat genomes results in spring habit and the presence of recessive Vrn‐1b alleles on all three genomes results in winter habit. Two sets of near‐isogenic lines (NILs) were evaluated for DNA polymorphisms at their Vrn‐A1, B1 and D1 loci and for cold hardiness. Two winter wheat cultivars, ‘Daws’ and ‘Wanser’ were used as recurrent parents and ‘Triple Dirk’ NILs were used as donor parents for orthologous Vrn‐1 alleles. The NILs were analysed using molecular markers specific for each allele. Only 26 of 32 ‘Daws’ NILs and 23 of 32 ‘Wanser’ NILs had a plant growth habit that corresponded to the marker genotype for the markers used. Freezing tests were conducted in growth chambers programmed to cool to ?21.5°C. Relative area under the death progress curve (AUDPC), with a maximum value of 100 was used as a measure of death due to freezing. The average relative AUDPC of the spring habit ‘Daws’Vrn‐A1a NILs was 86.15; significantly greater than the corresponding winter habit ‘Daws’Vrn‐A1b NILs (42.98). In contrast, all the ‘Daws’Vrn‐A1bVrn‐B1aVrn‐D1b and Vrn‐A1bVrn‐B1bVrn‐D1a NILs (spring habit) had relative AUDPC values equal to those of their ‘Daws’ sister genotypes with Vrn‐A1bVrn‐B1bVrn‐D1b NILs (winter habit). The average AUDPC of spring and winter habit ‘Wanser’ NILs differed at all three Vrn‐A1, Vrn‐B1 and Vrn‐D1 locus comparisons. We conclude that ‘Daws’ and ‘Wanser’ have different background genetic interactions with the Vrn‐1 loci influencing cold hardiness. The marker for Vrn‐A1 is diagnostic for growth habit and cold hardiness but there is no relationship between the Vrn‐B1 and Vrn‐D1 markers and the cold tolerance of the NILs used in this study.     

The effect of VRN1 genes on important agronomic traits in high‐yielding Canadian soft white spring wheat

For reproductive success, flowering time must synchronize with favourable environmental conditions. Vernalization genes play a major role in accelerating or delaying the time to flowering. We studied how different vernalization (VRN1) gene combinations alter days to flowering and maturity and consequently the effect on grain yield and other agronomic traits. The study focussed on the effect of the VRN1 gene series (Vrn‐A1, Vrn‐B1 and Vrn‐D1) and their combinations. The Vrn gene group Vrn‐A1a, Vrn‐B1, vrn‐D1 was the earliest to flower and mature, while Vrn‐A1b, Vrn‐B1, vrn‐D1 was the latest to flower. Spring wheat lines with vrn‐A1, Vrn‐B1, Vrn‐D1 were the highest yielding and matured at a similar time as those having vernalization genes Vrn‐A1a, Vrn‐B1 and Vrn‐D1. The findings of this study suggest that the presence of Vrn‐D1 has a direct or indirect role in producing higher grain yield. We therefore suggest the introduction of Vrn‐D1 allele into higher‐yielding classes within Canadian spring wheat germplasm.     

淡绿叶色标记水稻光温敏核不育系开花习性

以4个淡绿叶粳稻光温敏核不育系和正常叶色粳型不育系浙农大11S(ZAU11S)及籼型不育系培矮64S(PA64S)共6个不育系为材料，在杭州自然条件下观察了上述6个光温敏核不育系的开花习性。结果表明：(1)淡绿叶色不育系同对照正常叶色不育系的叶色差异显著，易于肉眼分离；(2)淡绿叶色不育系的开花天数、开花率、柱头外露率都低于正常叶色不育系，日开花率、颖花张开时间和张开角度易受环境条件影响，表现较大差异；(3)除TS1外，其他三个淡绿叶色不育系在不育期和可育期的开花高峰都位于午前9：00或10：00，表现较强的早花特性。     

Location of a gene for frost resistance on chromosome 5A of wheat

Summary A gene for frost resistance on chromosome 5A of wheat was located using single chromosome recombinant lines from the cross between the substitution line Hobbit (Triticum spelta 5A) and Hobbit. In this sample of recombinant lines the locus for frost resistance, designated Fr1, is completely linked to the locus Vrn1 controlling vernalisation requirement. The results can be explained by a pleiotropic action of the Vrn1 locus or close genetic linkage between Vrn1 and Fr1. Further detailed study is necessary to resolve these alternative hypotheses.     

Comparative Study of CaMsrB2 Gene Containing Drought‐Tolerant Transgenic Rice (Oryza sativa L.) and Non‐Transgenic Counterpart

Concerns regarding the safety of transgenic foods have been raised because of possibility of undesirable effects development during genetic engineering. Analysis of phenotypic traits can increase the likelihoods of identifying those unintended effects in dietary composition of the GM crops. Objective of this study was to compare the transgenic lines with their non‐transgenic counterpart. Different vegetative and reproductive traits as well as antioxidant properties were considered to evaluate the transgenic (HV8 and HV23) lines containing CaMsrB2 gene and their non‐transgenic (Ilmi) parent line. Grain size and weight, seed germination, root length, root and shoot dry weight, length and width of flag leaf, plant height, and ligule, stamen and carpel length were not significantly different. Onset and completion of heading in each line occurred almost during the same period. The antioxidant properties in terms of DPPH (1,1‐diphenyl‐2‐picrylhydrazyl) radical scavenging activity and polyphenol content were not statistically different under same treatment condition. The results suggested that the transgenic rice lines containing CaMsrB2 gene were equivalent to their non‐transgenic counterpart without any visible unintended effects.     

Inheritance of cabbage head formation in crosses of cabbage × ornamental cabbage and cabbage × kale

We investigated the inheritance of head formation in Brassica oleracea by using two crosses, cabbage × ornamental cabbage and cabbage × kale. The degree of head formation (DHF) was classified into nine grades ranging from non‐heading to full heading. DHF in the two F₂ populations showed a continuous distribution. The variance of F₃ offspring selected for full heading or non‐heading was large. The DHF distributions in the F₃ offspring selected for moderate heading or randomly selected F₃ populations were similar to those of the respective F₂ populations, but had smaller average values and variance. However, the realized heritability in F₃ offspring was similar in full‐heading and non‐heading selections. Our findings suggest that head formation is a quantitative trait controlled additively with low dominance effects. In comparisons of leaf developmental patterns among the parents, only cabbage showed a change in leaf shape becoming wider because of the shorter petiole length with increasing leaf position. These findings suggest that cabbage acquired the developmental changes in leaves required to form a head during the process of domestication.     

Variation of PGM and IDH isozymes for identification of alfalfa varieties

Growth habit, heading date and Vrn genotypewere examined for wheat landraces cultivated in China,Korea and Japan, to study their ecogeographicaldifferentiation in east Asia. Spring type landracesaccounted for 43.6% of the whole, and the frequencyvaried between the localities, being closely relatedto the degree of winter coldness. Spring typelandraces mainly adapted to north and south Chinawhere average January temperature is under –7 ^°Cand over 4 ^°C, respectively. On the contrary,winter type adapted to areas of average Januarytemperature from –7 ^°C to 4 ^°C. As toheading date, significant difference was not observedbetween spring and winter type landraces but betweenlocalities, and those cultivated in north China weresignificantly later in heading. It is thereforeindicated that spring type mainly adapts to areaswhere wheat is sown in spring to avoid frost injury,and where winter temperature is not low enough tovernalize winter type wheat. Genetic analysis forspring type landraces showed that the relativefrequency of four Vrn genes was different witheach other. Vrn3 was most widely and frequentlyfound among the four genes, followed by Vrn1 andVrn2. Only seven landraces proved to be thecarrier of Vrn4. The frequency was alsodifferent between localities. Genotype with Vrn1plus other dominant gene(s) adapted to spring sowingto avoid severely cold winter in north China, whilegenotype with only Vrn3 adapted to winter sowingin south China and southwest Japan. It is thereforeconcluded that at least three ecotypes, differing ingrowth habit and Vrn genotype, areallopatrically distributed in east Asia, as a resultof adaptation to winter coldness in each locality.     

Dwarfing effect associated with the threshability gene Q on wheat chromosome 5A

A dwarfing effect of the 44.1 cM chromosomal region between the threshability gene Q and Xfba068 on the long arm of hexaploid wheat chromosome 5A has been reported. To clarify whether Q or its adjacent region is responsible for regulating culm elongation, two precise genetic stocks of near‐isogenic lines (NIL), a single chromosome substitution line, ‘Chinese Spring’ (CS; ‘Cappelle‐Desprez’ 5A) (NIL‐Q) and a single chromosome recombinant substitution line (NIL‐q) were used. The target segment of NIL‐q included the q allele and QEet.ocs‐5A.1, an earlinessper se gene, from spelt wheat in the CS genetic background. They were grown under 16‐h day length with and/ or without vernalization treatment. Being independent of heading date, NIL‐Q showed shorter elongation in lower internodes and decreased internode differentiation in comparison with NIL‐q. The culm‐length reduction associated with Q was confirmed in the recombinant F5 population derived from the cross between NIL‐Q and NIL‐q. Vernalization promotion had a tendency to reduce this dwarfing effect.     

Genetic control of pre-heading phases in the Steptoe × Morex barley population under different conditions of photoperiod and temperature

Although there are numerous studies on the genetic control of flowering time in barley, little is known on the genetic control of duration of different particular pre-heading phases. Extending the stem elongation phase (SE), without modifying total time to heading, has been proposed as a trait to raise yield potential. Moreover, studying the genetic control of pre-heading phases would be of interest for a better understanding of crop phenology which is crucial for adaptation. We studied the genetic control of the leaf and spikelet intitiation phase (LS), the stem elongation phase, and within this, from the onset of jointing to flag leaf (J-FL), and from then to heading (FL-HD), in the Steptoe × Morex population, which is known to segregate for some major developmental genes, under different environmental conditions. After a preliminary greenhouse study in which the appropriateness of the population was tested, 130 double haploid lines and the two parents were grown under four field environments that differed in photoperiod and temperature conditions. Amongst all QTLs detected (13), only three were significant for HD and for both LS and SE and with the same allele direction (although with greater effects in one phase than the other in some cases). Genotype by environment interactions for LS and HD were due to both photoperiod and other factors as temperature or its interaction with photoperiod, while for SE responses to only photoperiod were negligible. QTL × E interactions were important for some QTLs, and either cross-over (e.g. Ppd-H1) or quantitative (e.g. Ppd-H2). However, heritability across field environments for the ratio SE/LS was high (0.8) and several of the QTL effects that were significant for only LS or SE, were conserved across different environments, that is, they were significant with the increasing allele derived from the same parent in all or most environments.     

Chromosomal effects in the endogenous contents of non-structural carbohydrates and proteins measured in wheat substitution lines

In hexaploid bread wheat, Triticum aestivum (2n = 6x = 42), little work has been carried out to study the genetic control of the synthesis of reduced, non‐reduced and total non‐structural carbohydrates and soluble proteins in aerial and rooting structures. The aim of this paper was to determine the chromosomal location of genes determining carbohydrate and protein synthesis that could be used for diagnostic selection in segregating breeding populations. A set of wheat intervarietal chromosome substitution lines [‘Chinese Spring’ (CS) × synthetic wheat (Triticum diccocoides×Aegilops squarrosa) (Syn)], was used. Plants were cultivated in hydroponic solutions to the fully expanded third leaf stage. Carbohydrate and protein contents and dry matter were determined for aerial and root parts. The root dry weight did not show significant differences between the parental varieties and the substitution lines, except for 5A, 2B and 6B, which had significantly lower dry weights. The aerial dry weight was significantly higher for Syn and the 2A substitution line. The ratio aerial dry weight/root dry weight was significantly higher in Syn, 1A, 2A and 4B. The protein content of the plant showed highly significant differences between both parental lines but 6A and 1D of the substitution lines showed highly significant differences, with contents as high as that for Syn. Syn produced significantly lower total aerial carbohydrates. The substitution lines 2A, 5A, 6A, 7A, 2B, 3D, 5D and 6D showed highly significant total carbohydrate content increases in the aerial parts compared with both parental lines. The non‐reduced carbohydrate contents showed a pattern similar to that of the total carbohydrates. Syn had a lower reduced carbohydrate content than CS. Only the 5A, 2B, and 1D substitution lines had a highly significantly different content of reduced carbohydrates than CS. In roots, Syn produced the lowest values for every type of sugar. The highest significant values for total carbohydrates were found in substitution lines 2B, 4B, 5B, 6B, 1D and 6D. The non‐reduced carbohydrate levels were significantly higher than CS in 2B, 5B, 6B and 6D substitution lines. Only the substitution lines 3B and 1D showed a significantly higher reduced carbohydrate content in roots compared with CS. The photoassimilate partitioning in Syn, 1 A, 2A and 4B favoured the aerial parts but, in contrast, higher partitioning to the roots was found in the 7B, 1D and 3D substitution lines. Both groups appear to carry interesting patterns worth incorporating in wheat cultivars.     

Effect of alien 5R(5A) chromosome substitution on ear-emergence time and winter hardiness in wheat-rye substitution lines

Ear emergence time and response to vernalization were investigated in 12 alien substitution lines in which a pair of chromosomes 5A of recipient spring wheat cultivars was replaced by a pair of chromosomes 5R of Siberian spring rye ‘Onokhoiskaya’. The recipients were 12 spring cultivars of common wheat, each carrying different Vrn genes. Spring rye ‘Onokhoiskaya’ had the Sp1 (now called Vrn-R1) gene for spring growth habit located on chromosome 5R, but its expression was weaker. The Vrn-R1 gene had no effect on growth habit, ear emergence time and response to vernalization in wheat-rye substitution lines. Ears emerged significantly later in the 5R(5A) alien substitution lines than in the recipient wheat cultivars with the Vrn-A1/Vrn-B1/vrn-D1 or Vrn-A1/vrn-B1/Vrn-D1 genotypes. No difference in ear emergence time was found between most of the 5R(5A) alien substitution lines and the cultivars carrying the recessive vrn-A1 gene. The presence of the Vrn2a and Vrn2b alleles at the Vrn2 (now called Vrn-B1) locus located on wheat chromosome 5B was confirmed.The replacement of chromosome 5A by chromosome 5R in wheat cultivars ‘Rang’ and ‘Mironovskaya Krupnozernaya’, which carries the single dominant gene Vrn-A1, converted them to winter growth habit. In field studies near Novosibirsk the winter hardiness of 5R(5A) wheat–rye substitution lines of ‘Rang’ and ‘Mironovskaya Krupnozernaya’ was increased by 20–47% and 27–34%, respectively, over the recurrent parents.     

Wheat Rht‐B1 Dwarfs Exhibit Better Photosynthetic Response to Water Deficit at Seedling Stage Compared to the Wild Type

Wheat reduced height (Rht) genes encode modified DELLA proteins, which are gibberellin insensitive, accumulate under stress, restrain growth and affect plant stress response. The seedling reaction to soil water deficit regarding leaf gas exchange and chlorophyll fluorescence was compared in near‐isogenic lines carrying the alleles Rht‐B1a (tall), Rht‐B1b (semi‐dwarfing) and Rht‐B1c (dwarfing) and was related to leaf water content and anatomy. Under drought, Rht‐B1c line was characterized by less decreased CO₂ assimilation, delayed non‐stomatal limitation of photosynthesis and higher instantaneous water use efficiency. The functional state of its photosynthetic apparatus was better preserved as evidenced by the less decreased actual quantum yield (Φ_PSII) and potential maximum quantum yield (F_v/F_m) of PSII, and the less increased quantum yield of non‐regulated energy dissipation (Φ_NO). Rht‐B1b line also tended to perform better than Rht‐B1a, but differences were less pronounced. Although the leaves of both dwarf lines were smaller, thicker and more pubescent, their water content was not higher in comparison with the tall line. Nevertheless, in Rht‐B1c, leaf thickness was less decreased and mesophyll cells were less shrunk under drought. The more effective performance of the photosynthetic machinery of dwarf lines under water deficit could be explained by a combination of morpho‐anatomical and metabolic characteristics.     

Genetic Control of Vernalization, Day-length Response, and Earliness per se by Homoeologous Group-3 Chromosomes in Wheat

To identify homoeologous group-3 chromosomes that carry genes for vernalization, day-length responses, and earliness per se, a series of aneuploid lines (mono-somics and tetrasomics) and chromosome-substitution lines in ‘Chinese Spring’ (CS) were surveyed under different vernalization and day-length regimes in controlled environments. The results indicated that genes on all three chromosomes of group 3 can have striking effects on ear-emergence time. The replacement of CS 3B by its homologues in ‘Lutescens 62’ and ‘Cheyenne’ produced an increased insensitivity to vernalization, while 3B homologues from ‘Ceska Presivka’ gave CS a remarkable sensitivity to vernalization. This provided evidence for multiple allelism at a new Vrn locus on chromosome 3B. A negative association between gene dosage and day-length response was found in CS 3D which was thought to carry a gene for promoting insensitivity to day-length. The behaviour of CS monosomic 3A and CS (Timstein 3A), in reducing numbers of days to heading independently of environmental stimuli, suggested the presence of earliness per se genes on this chromosome.     

Identifying AFLP and microsatellite markers for vernalization response gene Vrn-B1 in hexaploid wheat using reciprocal mapping populations

Marker‐assisted selection may be useful for combining specific vernalization response (Vrn) alleles into a single wheat genotype for yield enhancement; however, DNA markers are only available for two of the three genes identified to date. The objectives of this study were to investigate reciprocal effects on days to heading using F₂ populations generated by cross‐hybridizing near‐isogenic lines (NILs) carrying spring (Vrn‐B1; TDB) and winter (vrn‐B1; TDC) alleles, and to identify markers linked to Vrn‐B1 through genetic linkage analysis. Heading data were recorded for 91 and 89 progeny from reciprocal mapping populations TDB/TDC and TDC/TDB, respectively, and significant (P < 0.0001) reciprocal and dominance effects were detected. Among 207 amplified fragment length polymorphisms primer pairs and seven wheat microsatellite markers screened, two and one, respectively, were linked distally to Vrn‐B1 on wheat chromosome 5BL. Microsatellite Xgwm408 was most closely linked to Vrn‐B1 at 3.9 and 1.1 cM in the TDB/TDC and TDC/TDB map, respectively. Reciprocal differences in recombination distances emphasize the importance of female parent choice when generating mapping populations. Molecular markers are now available for three Vrn loci in wheat.     

Adaptation and ecological differentiation in wheat with special reference to geographical variation of growth habit and Vrn genotype

Geographical variation of growth habit was studied for 749 landraces from various parts of the world, with special reference to their adaptation and ecogeographical differentiation. The total frequency of spring‐type landraces was 49.9%, and varied between localities. Spring‐type landraces were frequent in two distinct areas where the average January temperature was either below ‐7°C or above 4°C, with winter‐type landraces in areas from ‐7°C to 4°C. These results indicated that geographical variation of growth habit is closely related to the degree of winter coldness. An analysis of the Vrn genotype for 216 spring‐type landraces demonstrated the uneven distribution of four Vrn genes, with Vrn4 being the least frequent. The adaptive Vrn genotype was different between localities. Genotypes carrying Vrn‐A1 and additional Vrn gene(s) were frequent in two distinct areas where the average January temperature was either below ‐7°C or over 10°C, while genotypes with any of three Vrn genes, except Vrn‐A1, adapted to areas with temperatures from 4°C to 10°C. Therefore, it was concluded that the adaptability of wheat landraces differed depending on their growth habit and Vrn genotype, and that ecotypes with different Vrn genotypes were allopatrically distributed as a result of adaptation to different winter temperature. However, the differential distribution of Vrn‐B1, Vrn‐D1 and Vrn4 could not be explained by their adaptability, and might reflect the polyphyletic origin of common wheat.