The physiology of variation in the time of ear emergence among wheat varieties from different regions of the world

Summary Differences in response to photoperiod and vernalization and genetic variation independent of photoperiod and vernalization (earliness per se), affecting time of ear emergence of wheat, were identified in controlled environment experiments with 33 varieties of diverse geographical origin. The results were compared with an analysis of time of ear emergence of 10409 T. aestivum accessions from the USDA Small Grain Collection grown from autumn sowings in Pendleton, Oregon, and spring sowings in Fargo, North Dakota. The effect of differences in photoperiod and vernalization sensitivity on time of ear emergence was similar to the effect of earliness per se, both under controlled environment conditions and in the field. Most of the accessions from low latitude regions reached ear emergence rapidly owing to their insensitivity to photoperiod and vernalization and earliness per se factors accelerating ear emergence. Lateness was common among accessions from Northern Europe, Afghanistan and Turkey, which was due to sensitivity to photoperiod and vernalization, and to earliness per se factors delaying ear emergence. The physiological basis of earliness per se is discussed.     

Temperature as a component of the expression of developmental responses in wheat

Summary Studies were made of days to ear emergence under the constant temperatures of 9, 14, 19 and 25°C and 16 h photoperiod in three sets of wheat lines each possessing genotypes differing for developmental responses.Days to ear emergence in three near-isogenic lines of the wheat cultivar Triple Dirk, which differed for vernalization response, increased as the strength of the response increased. At the four temperatures Triple Dirk D (Vrn 1 vrn 2) was not significantly different from normal Triple Dirk (Vrn 1 Vrn 2) but Triple Dirk B (vrn 1 Vrn 2) was significantly (P=0.01) later than normal Triple Dirk at each temperature. This indicates that the vrn 1 allele confers stronger vernalization response than vrn 2 over the range of temperatures (9–24°C). However, Triple Dirk C (vrn 1 vrn 2) failed to head after 120 days at each temperature indicating strong interaction between vrn 1 and vrn 2 with each other (and possibly the Triple Dirk back-ground) to give a much stronger vernalization response than predictions from additivity of their individual effects.The second set comprised the four Chinese Spring/Thatcher chromosome substitution lines CS/T 3B, 6B, 7B and 5D, plus Chinese Spring and Thatcher, and were grown in the unvernalized condition. CS/T 5D was similar in days to ear emergence as Chinese Spring at all four temperatures but the other three lines were earlier to ear emergence, particularly as the temperature increased. Days to ear emergence was fastest at 14°C in all lines, except CS/T 3B, in which it decreased progressively from 9 to 24°C.The third set of Chinese Spring and Thatcher and the homoeologous group 2 chromosomes of Thatcher substituted in Chinese Spring, the group which is considered to be involved in the control of photoperiod sensitivity. The three substitution lines responded differently to temperature compared with Chinese Spring and with each other, with chromosome 2D being the least, and chromosome 2B the most, responsive to temperature.     

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.     

Validation of the two-gene epistatic model for vernalization response in a winter × spring barley cross

A two gene epistatic model in which a dominant “winter growth habit” allele at Vrn-H2 encodes a repressor with a corresponding binding site in a recessive vrn-H1 allele explains the vernalization response phenotypes in an array of barley germplasm. In order to validate the model genetically, we developed an F ₂ population (and F ₂-derived F ₃ families) from the cross of Hardy (winter) × Jubilant (spring). Using gene-specific primers, we determined the Vrn-H1 and Vrn-H2 allele architecture of each F ₂ plant and we measured the growth habit phenotype of each F ₂ plant via phenotyping of its F ₃ progeny under controlled environment conditions. We used a set of treatments involving plus/minus vernalization under long photoperiod and vernalization under short photoperiod. Alleles at the two loci showed expected patterns of segregation and independent assortment. Under long day conditions, the two Vrn genes were the primary determinants of heading date, regardless of the vernalization treatment. Under short photoperiod, the effects of these loci were not significant. There was incomplete dominance at Vrn-H1: heterozygotes were significantly later to head than Vrn-H1Vrn-H1 genotypes. Vrn-H2 genotypes were also significantly later to head, even when plants were vernalized. These results validate the two-gene epistatic model for vernalization response under long-day conditions. The results under short photoperiod, and the variance in flowering with vernalization, confirm that while the two Vrn genes are the primary determinants of vernalization response, they are part of a larger interactome that determines the timing of the vegetative to reproductive transition.     

Detection of an earliness per se quantitative trait locus in the proximal region of wheat chromosome 5AL

A homoeologous quantitative trait locus to that of eps5L on barley chromosome 5H was identified in a syntenic region of wheat chromosome 5A. Wheat single chromosome recombinant lines (SCRs) were developed from a cross between ‘Chinese Spring’(‘Cappelle-Desprez’ 5A) and ‘Chinese Spring’(Triticum spelta 5A), these were grown together with the parental controls under different vernalization and photoperiod regimes. The variation for ear emergence time accelerated heading induced by the T. spelta segment indicated an effect associated with the Xcdo412-Xbcd9 interval. Since no differences between the SCRs and controls in responses to vernalization and photoperiod treatments were detected, this effect was identified as an earliness per se gene, Q Eetocs-5 A.2, which may be homoeologous to the eps5L quantitative trait locus of barley. Xbcd926 has been found to be closely linked to the rice flowering time quantitative trait loci, QHd9a or FLTQ2, on chromosome 9, suggesting possible relationships among the quantitative trait loci across wheat, barley and rice genomes.     

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.     

Genetic control of photoperiodic sensivity and maturity in spring wheat within narrow limits of adaptation

Summary Photoperiodic respose, as assessed by a regression technique, exhibited complete dominance averaged over the crosses of an eight parent diallel in the vernalized condition. Photoperiodic response as final leaf number for the vernalized 8-hour photoperiod diallel was closely related to photoeriodic response of the regression method. However, the diallel analyses of both sets of data showed little agreement in terms of respectieve array positions.The inheritance of photoperiodic response in diallels using regression values showed little agreement between the vernalized and unvernalized conditions. This difference was postulated to be due to interaction of vernalization and photoperiodic response in the unvernalized situation. In the unvernalized condition photoperiodic response exhibited non-allelic interaction, attributable mainly to the cultivar Pinnacle in general behaviour in its crosses. Its removal gave a situation of high average dominance for photoperiodic response with a clear indication that high photoperiodic sensitivity was dominant to comparative insensitivity.Days to ear emergence (vernalized and 18-hour photoperiod) exhibited non-allelic interaction in its expression, due mainly to the general behaviour of the cultivar Pinnacle in its crosses. Removal of its array gave a situation of a moderately strong degree of overdominance in the expression of days to ear emergence. Maturity differences amongst parents and F₁'s, vernalized and under 18-hour photoperiod, are postulated to be due to a factor other than vernalization or photoperiodic response beheved to be growth temperature in differentially in fluencing growth and/or developmental rates between genotypes.     

Measurement of response to vernalization in Australian wheats with winter habit

Summary Development in wheat is strongly controlled by sensitivity to vernalization and photoperiod, and to a lesser degree by non-vernalizing temperature and intrinsic earliness. A method to measure effect of vernalization in wheats with winter habit is described. Twenty seven wheats with winter habit and eight with spring/facultative habit were studied, comprising breeding lines and cultivars with maturities suited to south-central New South Wales. Effect of vernalization on the development of these wheats was quantified by integrating responses to vernalizing treatments of differing duration. Intrinsic earliness was measured as time for vernalized seedlings to grow to ear emergece in an 18h photoperiod with day/night temperature of 21/16°C, and response to photoperiod as the difference in time to ear emergence between 9 and 18h daylengths. Integrated response to vernalization is sensitive to both cumulative and thresh-hold responses and is applicable to wheats of all habit type. Integrated response to vernalization and intrinsic earliness were positively associated within wheats with winter habit. Wheats were largely of restricted origin, so that there were few allelic differences at Vrn loci to disrupt this association, which suggests intrinsic earliness may modify response to vernalization. Though integrated response to vernalization was measured with artificial treatments, it was strongly associated with ear emergence for wheats with winter habit when grown at a site in New South Wales.     

Pre-anthesis development and number of fertile florets in wheat as affected by photoperiod sensitivity genes <Emphasis Type="Italic">Ppd-D1</Emphasis>and <Emphasis Type="Italic">Ppd-B1</Emphasis>

Lengthening the late reproductive phase (LRP) of stem elongation in wheat (Triticum aestivumL.), by changing its photoperiod sensitivity independently of the preceding phases, would improve the yield potential through increasing spike weight and the number of fertile florets at anthesis. This paper presents results of a two-year field experiment designed to determine the impact of Ppd-D1and Ppd-B1on (i) the duration of three pre-anthesis developmental phases, and (ii) spike weight and the number of fertile florets at anthesis under two photoperiods during the LRP (natural and an extension of six hours over that). Near isogenic lines of Mercia and single chromosome recombinant lines of Cappelle Desprez were used. Under natural photoperiod, Ppd-D1hastened time to anthesis ca. 500^∘C d in both backgrounds by reducing each of the three pre-anthesis phases. Ppd-B1hastened the time to anthesis under natural photoperiod by 178^∘C d, mainly by reducing the early reproductive phase. The response to photoperiod of the LRP under extended daylength depended on the Ppdlocus present: Ppd-D1was insensitive while Ppd-B1and the recessive controls were sensitive. For all lines, photoperiod treatments and years, the number of fertile florets was associated with spike dry weight at anthesis (R ²≅ 80%, p< 0.01) which, in turn, was positively related to the intercepted radiation accumulated during the LRP (R ² 45%, p< 0.05). Changing the duration of the LRP through extended photoperiod or through Ppd-D1produced similar results in both backgrounds and years. Thus, altering the duration of the LRP by manipulating photoperiod sensitivity may be an alternative to changing the fertile floret number in wheat. Nevertheless, as no particular allele was responsible for the photoperiod sensitivity only during the LRP, new alleles should be studied to identify the control of photoperiod sensitivity of individual phases to fine-tune the pre-anthesis wheat development.     

Responses to photoperiod before and after jointing in wheat substitution lines

It has been hypothesised that wheat yields may be increased by lengthening the duration of the stemelongation phase. This paper reports studies on the effects of chromosomes carrying major photoperiod genes (Ppd-A1, Ppd-B1, Ppd-D1) in different genetic backgrounds, on responses to photoperiod before and after jointing, when the onset of stem elongation occurs, and on number of grains per spike. A field experiment considered the effects of two photoperiods on Chinese Spring and 12 substitution lines, in which chromosomes 2A, 2B or 2D had been substituted by those from four contrasting cultivars. The phase from seedling emergence to jointing (EM-JO) was more responsive than that from jointing to anthesis (JO-ANT), but no relationship was found between the duration of these phases. EM-JO length affected leaf and spikelet number and consequently grains per spike, but this component was further influenced by JO-ANT duration. Our results confirmed that the phases are independent in sensitivity, supporting the hypothesis that genetic manipulation of phase duration could enhance yield, but no evidence was found of any particular Ppd allele being responsible for major responses to photoperiod during stem elongation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Production and genetic characterization of the co-isogenic lines of a common wheat Triticum aestivum CV. S-615 for ten major genes

Summary Co-isogenic lines of a common wheat, Triticum aestivum, cv. S-615 with each of the following ten major genes were produced by repeated backcrosses; The gene C on 2D chromosome for the compactum character, s on 3D for the sphaerococcum character, Hd on 4B, B1 on 5A, and B2 on 6B for awn suppression, Hg on 1A for glume hairiness, Hp on 4A for peduncle hairiness, Ne1 on 5B and Ne2 on 2B for hybrid necrosis, and v1 on 3B for virescence. Seven of them showed the typical mendelian fashion of inheritance, while three others (C, s and v1) were transmitted at lower frequencies than their corresponding normal alleles.The effects of those major genes on 24 characters of cv. S-615 were investigated, which are summarized as follows: C: Increased node diameter, number of spikelets per ear and spike density, but decreased lengths of all ear rachis, awn, anther, empty and outer glumes, and grain, and grain index. s: Increased culm diameter and thickness, and spike density, reduced lengths of all 1st and 2nd internodes, culm, flag leaf, rachis, awn, anther, empty and outer glumes, and grain, and grain index, and accelerated heading. Hd: Increased number of spikelets per ear and anther length, while decreased length of awn, and empty and outer glumes. B1: Increased 1st internode, rachis, and anther lengths, and grain index, but decreased spike density, awn length and grain thickness. B2: Increased rachis length, number of spikelets per ear, empty and outer glume lengths and grain index, but reduced awn length and grain thickness. Hg: No effects. Hp: Increased number of spikelets per ear. Ne1: Accelerated heading, and increased empty glume length. Ne2: Increased node and culm diameters. v1: No effects.Two species-specific genes, C and s, affected a large number of characters expressed in various developmental stages. These genes were found to have pleiotropic effects, namely, they influenced, at least, two groups of interdependent characters. Three awn suppressors influenced a limited number of characters, but no evidence of their pleiotropic effects was obtained.Contribution from the Laboratory of Genetics, Faculty of Agriculture, Kyoto University, Japan, No. 422. The work was supported in part by a Grant-in-Aid (No. 148,004) from the Ministry of Education, Japan.     

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.     

Cytological and microsatellite mapping of the genes determining liguleless phenotype in durum wheat

Liguleless phenotypes of wheat lack ligule and auricle structures on all leaves of the plant. Two recessive genes principally control the liguleless character in tetraploid wheat. The F₂ progenies of k17769 (liguleless mutant)/Triticum dicoccoides and k17769/T. dicoccum segregated in a 15:1 ratio, whereas the F₂ progenies of k17769/T. durum and k17769/T. turgidum segregated in a 3:1 ratio. A new gene, lg₃, was found on chromosome 2A. Segregation of F₂ progenies between k17769 and chromosome substitution lines for homoeologous group 2 chromosomes suggested that the liguleless genotype had occurred by mutation at the lg₃ locus on chromosome 2A, and then by mutation at the lg₁ locus on chromosome 2B, in the process of domestication of tetraploid wheat. The gene (lg₁) was linked to Tc₂ (11.9 cM), which determines phenol colour reaction of kernels, on the long arm of chromosome 2B. The distance of lg₁ to the centromere was found to be 60.4 cM, and microsatellite mapping established the gene order, centromere – Xgwm382 – Xgwm619 – Tc₂ – lg₁ on the long arm of chromosome 2B.     

Distribution of the photoperiod insensitive <Emphasis Type="Italic">Ppd-D1a</Emphasis> allele in Chinese wheat cultivars

Photoperiod response is of great importance for optimal adaptation of bread wheat cultivars to specific environments, and variation is commonly associated with allelic differences at the Ppd-D1 locus on chromosome 2D. A total of 926 Chinese wheat landraces and improved cultivars collected from nine wheat growing zones were tested for their genotypes at the Ppd-D1 locus using allele-specific markers. The average frequency of the photoperiod-insensitive Ppd-D1a allele was 66.0%, with the frequencies of 38.6 and 90.6% in landraces and improved cultivars, respectively. However, the Ppd-D1a allele was present in all improved cultivars released after 1970 except for spring wheats in high latitude northwestern China, and winter wheats in Gansu and Xinjiang. The presence of the Ppd-D1a allele in landraces and improved cultivars increased gradually from north to south, illustrating the relationship between photoperiod response and environment. Ppd-D1a in Chinese wheats is derived from three sources, Japanese landrace Akagomughi and Chinese landraces Mazhamai and Youzimai. The current information is important for understanding the broad adaptation of improved Chinese wheat cultivars. F. P. Yang and X. K. Zhang contributed equally to this work.     

Allelic relationships of genes controlling number of flowers per axis in chickpea

Genetic variation for number of flowers per axis in chickpea (Cicer arietinum L.) includes single-flower, double-flower, triple-flower and multi-flower traits. A double-flowered (DF) line ICC 4929, a triple-flowered (TF) line IPC 99-18 and a multi-flowered (MF) line JGM 7 were intercrossed in all possible combinations and flowering behavior of parents, F₁s and F₂s was studied to establish allelic relationships, penetrance and expressivity of genes controlling number of flowers per axis in chickpea. The F₁ from ICC 4929 (DF) × IPC 99-18 (TF) cross were double-flowered, whereas F₁ from ICC 4929 (DF) × JGM 7 (MF) and IPC 99-18 (TF) × JGM 7 (MF) crosses were single-flowered. The F₂ from ICC 4929 (DF) × IPC 99-18 (TF) cross gave a good fit to a 3:1 ratio for double-flowered and triple-flowered plants. The F₂ from ICC 4929 (DF) × JGM 7 (MF) cross segregated in a ratio of 9:3:3:1 for single-flowered, double-flowered, multi-flowered and double-multi-flowered plants. The F₂ from IPC 99-18 (TF) × JGM 7 (MF) cross segregated in a ratio of 9:3:4 for single-flowered, triple-flowered and multi-flowered plants. The results clearly established that two loci control number of flowers per axis in chickpea. The double-flower and triple-flower traits are controlled by a single-locus (Sfl) and the allele for double-flowered trait (sfl ^d ) is dominant over the allele for triple-flower trait (sfl ^t ). The three alleles at the Sfl locus has the dominance relationship Sfl > sfl ^d > sfl ^t . The multi-flower trait is controlled by a different gene (cym). Single-flowered plants have dominant alleles at both the loci (Sfl_ Cym_). The double-flower, the triple-flower and the multi-flower traits showed complete penetrance, but variable expressivity. The expressivity was 96.3% for double-flower and 76.4% for double-pod in ICC 4929, 81.2% for triple-flower and 0.0% for triple-pod in IPC 99-18, and 51.3% for multi-flower and 24.7% for multi-pod in JGM 7. Average number of flowers per axis and average number of pods per axis were higher in JGM 7 than double-flowered line ICC 4929 and triple-flowered line IPC 99-18. The results of this study will help in development of breeding strategies for exploitation of these flowering and podding traits in chickpea improvement.     

Transfer of resistance to race 2 of Plasmodiophora brassicae from Brassica napus to cabbage (B. oleracea var. capitata). II. Meiosis in the interspecific hybrids between B. napus and 2x and 4x cabbage

Summary Meiosis in 14 interspecific F₁ hybrids with three chromosomal levels (triploid, tetraploid, hexaploid; 2n=28, 37 and 55) between Brassica napus L. and 2x and 4x cabbage (B. oleracea var. capitata L.) was studied. The oleracea genome from B. napus maintained close homology with the c genome of cabbage while the campestris genome of B. napus showed partial homology with the c genome contained in the hybrids. Genotypic influence on chromosome pairing was indicated. Structural chromosome differences and spontaneous chromosome breakage and reunion were suggested as causes for the abnormalities which related to the unbalance of the genotypes. The divergence of the genomes of B. napus and B. oleracea and the need for the qualification of the term secondary association were discussed.Contribution No. J. 673, Research Station, Agriculture Canada, St. Jean, Québec.     

Relationship between cold resistance,heading traits and ear primordia development of wheat cultivars

Summary For breeding early heading wheat cultivars with resistance to frost damage which are well adapted to dry areas of West Asia and North Africa, the relationships between winter hardiness, ear primordia development and heading traits, i.e. veernalization requirement, photoperiodic response and narrow-sense earliness, were assessed using a total of 30 genotypes of wheat (Triticum aestivum L.) grown in an experiment in Syria. The results of artificial freezing tests indicated that cultivars with good winter hardiness were to be found only in the winter wheat cultivars which required 50 or more days of vernalization treatment. These winter wheat cultivars did not initiate internode elongation without vernalization even at 95 days after planting. Thus their ear primordia were still underground and were protected from frost injury at this stage. Photoperiodic response and narrow-sense earliness were not associated with winter hardiness and earliness of internode elongation, but were related to the number of days to heading after planting. This indicated the possibility for breeding early heading cultivars with winter hardiness and tiller frost avoidance by combining high vernalization requirement, short narrow-sense earliness and neutral response to photoperiod.     

The effects of homoeologous group 3 chromosomes on grain colour dependent seed dormancy and brittle rachis in tetraploid wheat

`Langdon' (LDN), a durum wheat (Triticum turgidum L. var. durum) cultivar, and a set of chromosome substitution lines of Langdon, where A or B genome chromosome were replaced with a homologous chromosome of wild emmer wheat, T. turgidum ssp. dicoccoides (DIC), were used to assess the effect of the specific chromosome on seed dormancy in tetraploid wheat. The LDN(DIC 3A) and LDN (DIC 313) lines showed significantly lower seed germination than Langdon. It appears that LDN(DIC 3A) and LDN(DIC 3B) have red grain whose allele were designated as R-A1b and R-B1b, respectively and the rachises of LDN(DIC 3A) and LDN(DIC 3B) were fragile. The alleles for brittle rachis were designated as Br ₂ for LDN(DIC 3A) and Br ₃ for LDN(DIC 3B). From the F₂ of the crosses, Langdon/LDN(DIC 3A) and Langdon/LDN(DIC 3B), Br ₂ was located approximately 44.2 cM from the R-A1b locus and Br ₃ approximately 47.0 cM from the R-B1b locus, respectively. Recombinant inbred chromosomal lines for 3A and 3B were used to assess (1) the linkage relationship between grain colour and fragile rachis, and (2) the effect of grain colour on germination. Estimated distance between R-B1b – Br ₂ was 39.6 cM. For the 3A population, germination percentage of both colour groups was 12.4% for the red grain group and 68.6% for the amber group, respectively. For the 3B population, germination percentage of the red group was 7.3% and that of the amber group was 82.1%. For both populations, differences were statistical significant by t-tests. We considered that seed dormancy of T. turgidum ssp. dicoccoides was dependent on grain colour. It raised the possibility that brittle rachis is due to a paralogous gene set on homoeologous group 3 chromosomes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Factors influencing the development of bread wheat plant types to be grown in the ‘transitional zone’ in northern Syria

Summary This study sought to identify factors that influence wheat development in the transitional wheat growing zone of northern Syria. Three development factors were studied, intrinsic earliness, and responses to vernalization and to photoperiod. Two sets of wheat were studied, each composed of lines with differing combinations of development factors. Set 1 comprised 20 parental and breeding lines utilized by the CIMMYT/ICARDA facultative and winter wheat breeding program based at Tel Hadya. Set 2 comprised 19 parental and breeding lines utilized by an Australian winter wheat breeding program based at Temora. Field development was recorded in greatest detail at one site. Tel Hadya, using the state of differentiation of the apex of the main tiller of sampled plants. To extend findings, development was also recorded as the time from sowing to ear emergence for later sowings of wheat at Tel Hadya, and in sowings at four other regional sites.The significance of each development factor was tested in multiple regressions that predicted either stage of apical development at Tel Hadya, or time to ear emergence in all trials. It was found that intrinsic earliness was the major factor associated with development, in both sets of wheat. Response to photoperiod had a much smaller and less consistent effect. Response to vernalization had least effect on development, possibly because low temperature in winter delayed development for a longer period than was required to fully vernalize winter wheats. Our results suggested it may not be directly relevant whether spring or winter wheats are grown in the transitional zone of northern Syria. The desired phenotype for the region, of slow development prior to double ridge, then fast development to ear emergence, cannot be simply achieved from combinations of the three development factors. Selection for improved adaptation to the region must continue to rely on direct field observations.     

Mapping the <Emphasis Type="Italic">compactum</Emphasis> locus in wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) and its relationship to other spike morphology genes of the Triticeae

The spikes of club wheat are significantly more compact than spikes of common wheat due to the action of the dominant allele of the compactum (C) locus. Little is known about the location of C on chromosome 2D and the relationship between C and to other spike-compacting genes. Thus, a study was undertaken to place C on linkage maps and a chromosome deletion bin, and to assess its relatedness to the spike compacting genes zeocriton (Zeo) from barley and soft glume (Sog) from T. monococcum. Genetic mapping was based on recombinant inbred lines (RILs) from a cross between the cultivars Coda (club) and Brundage (common) and F₂ progeny from a cross between the club wheat Corrigin and a chromosome 2D substitution line [Chinese Spring (Ae. tauschii 2D)]. The C locus was flanked by Xwmc144 and Xwmc18 in the RIL population and it was completely linked to Xcfd116, Xgwm358 and Xcfd17 in the F₂ population. C could not be unambiguously placed to a chromosome bin because markers that were completely linked to C or flanked this locus were localized to chromosome bins on either side of the centromere (C-2DS1 and C-2DL3). Since C has been cytogenetically mapped to the long arm of chromosome 2D, we suspect C is located in bin C-2DL3. Comparative mapping suggested that C and Sog were present in homoeologous regions on chromosomes 2D and 2A^m, respectively. On the other hand, C and Zeo, on chromosome 2H, did not appear to be orthologous.