Yield QTL affected by heading date in Mediterranean grown barley

Knowledge regarding quantitative trait loci (QTL) has led to remarkable advances in breeding for a variety of traits, some of which have an effect on yield under particular environmental conditions. However, the same yield QTL are not usually found, even in the same population tested in different environments, as a result of large genotype-by-environment interactions. In this study, we aimed at identifying yield QTL in a series of experiments carried out in Spain. We used a barley doubled haploid population derived from a spring by winter cross. The relationship between heading date and yield, and between the principal heading date QTL and yield, changed depending on the environment. Allelic combinations causing early or late heading dates usually did not favour high yield, whereas intermediate heading dates were associated with not only higher variability but also higher yield potential. To identify QTL for grain yield independent of heading date, the population was divided into three classes: Early , Intermediate and Late . We found three new QTL affecting yield only at specific maturity groups. The implications of these findings in relation to barley breeding are discussed.     

A QTL for early heading in wheat cultivar Suwon 92

Heading date is an important trait that determines wheat adaptation to environments. A recombinant inbred line (RIL) population derived from CI 13227 × Suwon 92 was employed to tag the quantitative trait locus (QTL) for early heading in Suwon 92. This population was phenotyped for heading date in 1994, 1995, and 1997, and analyzed with AFLP and SSR markers. Two AFLP markers (XGCTG.CGCT118 and XGCTG.CGCT60) closely associated with heading date were identified. Across years, XGCTG.CGCT118 and XGCTG.CGCT60 explained 40.4% and 32.2% of the total phenotypic variances, respectively. Interval analysis revealed a major QTL for heading date, designated QHd.pser-2DS, between AFLP marker XGCTG.CGCT118 and SSR marker Xgwm261. Based on the linkage map, QHd.pser-2DS was about 41.2 cM proximal to the distal end of chromosome 2DS, and explained 40.5% of the phenotypic variance across three years. The identified markers associated with the early heading QTL have the potential to be used in wheat breeding programs.     

Genetic effects of Vrn genes on heading date and agronomic traits in bread wheat

Summary The Vrn1, Vrn2 and Vrn3 genes have different values of effects on heading date and related yield components. The genetic background and environment do not affect the ranking of Vrn genotypes according to earliness within near-isogenic line sets; however, they do influence the level of differences between heading dates of particular genotypes and between effect values, respectively. The frequencies of defined Vrn genotypes in the global set of spring bread wheat cultivars are associated with grain weight per plant predicted on the basis of Vrn gene effects averaged over backgrounds and over environments. Peculiarities of backgrounds and environments alter the grain yield ranges of Vrn genotypes. For early photoperiod-insensitive wheats, planted in stress conditions at grain filling, the highest yield was predicted for double dominant Vrn genotypes with Vrn3. This gene is rarely used by the breeders in middle latitudes and its wider adoption is encouraged.     

Genetic analysis of heading date and other agronomic characters in barley (Hordeum vulgare L.)

Genetic analyses of heading date, tiller number, plant height, grain yield, kernel weight, and plump and thin kernels were made in three six-rowed barley crosses (Hordeum vulgare L.) involving four cultivars. Six populations, P₁ , P₂ , F₁ , F₂ , BC₁ , and BC₂ , from each cross were grown and evaluated at Fargo and Prosper, North Dakota, 1982. Parental means within crosses generally were different except for tiller number. Comparison of generation means suggested that late heading was dominant to early, high kernel weight was dominant to low, and kernel plumpness was influenced by additive gene action. The relationship between yield and heading date was not consistent among crosses and positive r values were quite low. It should be possible to select early maturing, high yielding segregates with plump kernels. Heterosis over the mid-parent was quite similar among crosses for heading date, but there was no heterosis over the high parent. Inbreeding depression was fairly constant for heading date, but was less consistent for yield. The lack of uniformity for estimates of inbreeding depression can be related to environmental variation and to its influence on type of gene action. The ratio of additive to dominance variance was inconsistent among crosses for heading date and yield. These data suggest selection for these characters should be delayed past the F 2 generation. Broad sense heritabilities for heading date ranged from 42 to 86%. Values obtained for grain yield were more consistent among broad sense than narrow sense estimates. Genetic advance estimates were low due to lack of additive variance. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

QTL mapping for heading date,leaf area and chlorophyll content under cold and drought stress in two related recombinant inbred line populations (Japonica rice) and meta‐analysis

Rice is highly susceptible to drought and cold. The goal of this study was to identify the QTLs affecting the rice heading date (HD), leaf area (LA) and chlorophyll content (CC) under cold and drought stress. A total of twenty‐nine and thirty‐eight additive QTLs were detected from two crosses of ‘Dongnong422’/‘Kongyu131’ and ‘Xiaobaijingzi’/‘Kongyu131’, respectively. qHD1‐7‐1, qHD1‐7‐2, qHD1‐2‐1, qLA1‐7‐1, qLA1‐6‐3 and qCC1‐7‐1 show adaptive effects under cold treatment, while qHD2‐2‐3, qHD2‐2‐2, qLA2‐7‐3 and qCC2‐5‐1 were important for explaining the genetic mechanism during drought tolerance. Additionally, nine and five additive × environment interaction QTLs were detected for two RILs, respectively. RIL26 and RIL73 were two lines that are associated with cold and drought for HD. 339 QTLs related to HD, CC and LA of rice from database and our research were projected onto the genetic map. Nine cloned genes and nineteen homologous candidate genes related to HD, CC, cold tolerance and drought tolerance were mapped by meta‐analysis. These results lay the foundation for the fine mapping of QTLs related to HD, LA and CC for marker‐assisted selection.     

西藏青稞冬春性鉴定及抽穗期多样性与稳定性分析

为提高青稞种质资源的利用效率并筛选优异杂交亲本,分别在西藏林芝和拉萨两地的春播和秋播环境下对1605份青稞种质资源进行冬春性鉴定和抽穗期多样性分析。设拉萨春播2个播期,即正常春播I和晚10 d春播II,以2个播期的抽穗期变化作为冬春性区别依据。结果表明,96.2%的西藏青稞地方品种为春性,在西藏3个生态区均有分布,冬性品种仅有3.8%,且主要分布在以林芝为主的藏东南生态区;抽穗期的Shannon-Wiener's多样性分析表明,春播条件下抽穗期多样性高于秋播,拉萨春播条件下抽穗期多样性最高,林芝春播次之,林芝秋播最低;在相同环境下,地方品种的多样性高于育成品种;在环境稳定性分析中, 368份春性地方品种和21份育成品种在不同环境中抽穗期较为稳定,其中康青3号在两地所有参试品种中抽穗期稳定性最高。本研究为全面理解西藏青稞资源的冬春性、抽穗期多样性和环境稳定性提供了参考,为广适应性青稞品种培育筛选出环境稳定性佳的亲本材料。     

Effect of VRN‐1 and PPD‐D1 genes on heading time in European bread wheat cultivars

The variation of the vernalization (VRN‐1) and photoperiod (PPD‐1) genes offers opportunities to adjust heading time and to maximize yield in crop species. The effect of these genes on heading time was studied based on a set of 245 predominantly spring cultivars of bread wheat from the main eco‐geographical regions of Europe. The genotypes were screened using previously published diagnostic molecular markers for detecting the dominant or recessive alleles of the major VRN‐1 loci such as: VRN‐A1, VRN‐B1, VRN‐D1 as well as PPD‐D1. We found that 91% of spring wheat cultivars contain the photoperiod sensitive PPD‐D1b allele. Photoperiod insensitive PPD‐D1a allele has been found mainly in southern region of Europe. For this region the monogenic control of vernalization by VRN‐B1 or VRN‐D1 dominant alleles is common, whereas in the remaining part of Europe, the combination of photoperiod sensitive PPD‐D1b allele with dominant VRN‐A1, VRN‐B1 and recessive vrn‐D1 alleles represents the most frequent genotype. Also, we revealed a significantly later (5–8 days) heading of the monogenically dominant genotypes at VRN‐B1 as compared to the digenic VRN‐A1 VRN‐B1 genotypes.     

Distribution of photoperiod-insensitive allele Ppd-A1a and its effect on heading time in Japanese wheat cultivars

The Ppd-A1 genotype of 240 Japanese wheat cultivars and 40 foreign cultivars was determined using a PCR-based method. Among Japanese cultivars, only 12 cultivars, all of which were Hokkaido winter wheat, carried the Ppd-A1a allele, while this allele was not found in Hokkaido spring wheat cultivars or Tohoku-Kyushu cultivars. Cultivars with a photoperiod-insensitive allele headed 6.9–9.8 days earlier in Kanto and 2.5 days earlier in Hokkaido than photoperiod-sensitive cultivars. The lower effect of photoperiod-insensitive alleles observed in Hokkaido could be due to the longer day-length at the spike formation stage compared with that in Kanto. Pedigree analysis showed that ‘Purple Straw’ and ‘Tohoku 118’ were donors of Ppd-A1a and Ppd-D1a in Hokkaido wheat cultivars, respectively. Wheat cultivars recently developed in Hokkaido carry photoperiod-insensitive alleles at a high frequency. For efficient utilization of Ppd-1 alleles in the Hokkaido wheat-breeding program, the effect of Ppd-1 on growth pattern and grain yield should be investigated. Ppd-A1a may be useful as a unique gene source for fine tuning the heading time in the Tohoku-Kyushu region since the effect of Ppd-A1a on photoperiod insensitivity appears to differ from the effect of Ppd-B1a and Ppd-D1a.     

Segregation for heading date and stem enlargement in kohlrabi x broccoli crosses

Summary F₁ and F₂ progenies from crosses of an early broccoli (Brassica oleracea var. italica L.) line with kohlrabi (B. oleracea var. gongyloides) indicated that days to flower bud maturity in annual segregates is quantitative and additive in inheritance. It appeared that the biennial kohlrabi parent strongly contributed genetic factors for late maturity. F₁ plants of both crosses, 98% of F₂ plants from early broccoli × kohlrabi, and 81% of the F₂ plants from late broccoli × kohlrabi were annual in 1980. In 1986, 96% of early broccoli × kohlrabi F₂ and 87% of late broccoli × kohlrabi F₂ plants were annual. Kohlrabi stem enlargement showed a continuous range of expression in the F₂ with some dominance of broccoli type. Few F₂ plants were close to the kohlrabi parent in degree of stem enlargement.     

Photoperiod sensitivity gene controlling heading date in rice cultivars in the northernmost region of Japan

Rice is grown in diverse environments at latitudes ranging from 53^°N to 40^°S. In Japan, Hokkaido is the northernmost rice cultivation region(42–45^°N latitude). Only extremely early maturing (heading) varieties that have extremely low photoperiod sensitivity are adapted to this area. Heading date is the most important trait in adaptation to this particular environment. Also, the efficient manipulation of heading date is a crucial component of rice improvement. To determine the genetic basis for heading date among cultivars grown in Hokkaido, the heading behaviour was analyzed. Clear segregations were observed. To estimate the photoperiod sensitivity of the genes involved, the cultivars and F₁ plants from crosses between the cultivars were grown under different daylength conditions. The results indicated that the genes controlling heading date are photoperiod sensitive, suggesting they play important roles in the northernmost rice cultivation regions in Japan, to which only cultivars with extremely low photoperiod sensitivity are adapted. This revised version was published online in August 2006 with corrections to the Cover Date.     

Development and evaluation of pyramiding lines carrying early or late heading QTLs in the indica rice cultivar ‘IR64’

The heading date is an important trait for determining regional and climatic adaptability in rice. To expand the adaptability of the indica rice cultivar ‘IR64’, we pyramided multiple early or late heading quantitative trait locus (QTLs) in the ‘IR64’ genetic background by crossing previously developed near-isogenic lines (NILs) with a single QTL for early or late heading. The effects of pyramiding QTLs were observed in three different climatic zones of the Philippines, Madagascar, and Japan. The early heading pyramiding lines (PYLs) headed 6.2 to 12.8 days earlier than ‘IR64’ while the late heading PYLs headed 18.8 to 27.1 days later than ‘IR64’. The PYLs tended to produce low grain yield compared to ‘IR64’. The low yield was not improved by combining SPIKE, which is a QTL that increases the number of spikelets per panicle. Conversely, ‘IR64-PYL(7+10)’ carrying Hd5 and Hd1 headed earlier, produced more tillers, and more panicles per m² than ‘IR64’, and mitigated the yield decrease in early heading. These results suggest that the effects of pyramided QTLs on heading date were consistent across various environments and PYLs could be used to enhance the adaptation of ‘IR64’ in other rice growing environments.     

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.     

Effects of environment, disease progress, plant height and heading date on the detection of QTLs for resistance to Fusarium head blight in an European winter wheat cross

In order to characterise quantitative trait loci (QTLs) for Type I and Type II resistance against Fusarium head blight (FHB) in wheat, a population of recombinant inbred lines derived from the cross Cansas (moderately resistant)/Ritmo (susceptible) was evaluated in spray-inoculated field trials over three seasons. Map-based QTL analysis across environments revealed seven QTLs on chromosomes 1BS, 1DS, 3B, 3DL, 5BL, 7BS and 7AL (QFhs.whs-1B, QFhs.whs-1D, QFhs.whs-3B, QFhs.whs-3D, QFhs.whs-5B, QFhs.whs-7A, QFhs.whs-7B) associated with FHB resistance. They accounted for 56% of the phenotypic variance. QFhs.whs-1D primarily appeared to be involved in resistance to fungal penetration, whereas the other QTLs mainly contributed to resistance to fungal spread. FHB resistance was significantly correlated with plant height (PH) and heading date (HD). Including all single environments, corresponding overlaps of QTLs for FHB resistance and QTLs for PH/HD occurred at six loci, among them two consistently detected QTLs, QFhs.whs-5B and QFhs.whs-7A. When significant effects of PH and HD on FHB resistance were eliminated by covariance analysis, a second QTL analysis revealed possible escape mechanisms for the majority of the coincidental loci.     

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.     

Identification of a locus for asynchronous heading in rice, <Emphasis Type="Italic">Oryza sativa</Emphasis> L.

We have identified a locus for asynchronous heading in rice. The length of heading period in a plant was measured as the indicator of asynchronous heading in 98 backcross inbred lines (BILs). These lines were derived using a single-seed descent method from a backcross of (japonica Nipponbare/indica Kasalath)/Nipponbare, and we used this mapping population in quantitative trait locus (QTL) analysis. We located a single QTL related to asynchronous heading (qah7) in the region of R2401–C39 on chromosome 7, and its contribution to the total variation was 0.14 (r²), with the positive effect due to the Kasalath allele. A near isogenic line (NILah7) that carries a Kasalath chromosomal segment corresponding to qah7 in the Nipponbare genetic background was selected and analyzed to identify a locus for asynchronous heading. The length of heading period in NILah7 was significantly longer (P < 0.001) than that in Nipponbare.     

Locating supplementary RFLP markers on barley chromosome 7 and synteny with homoeologous wheat group 5

In order to develop QTL applications, eight new loci were mapped on barley chromosome 7 using 124 doubled haploid lines of the North American Barley Genome Mapping Project (NABGMP) progeny (‘Steptoe’בMorex’)- These loci involve six genomic DNA restriction fragment length polymorphisms (RFLPs) and two cDNA-RFLPs including a puroindoline gene. The distribution of these markers on barley chromosome 7 was compared with that of homoeologous wheat counterparts, i.e. wheat group 5. One locus on chromosome 7 was associated with a QTL for β-glucanase activity measured in green and finished barley malt.     

Distribution of photoperiod-insensitive alleles Ppd-B1a and Ppd-D1a and their effect on heading time in Japanese wheat cultivars

The genotypes of photoperiod response genes Ppd-B1 and Ppd-D1 in Japanese wheat cultivars were determined by a PCR-based method, and heading times were compared among genotypes. Most of the Japanese wheat cultivars, except those from the Hokkaido region, carried the photoperiod-insensitive allele Ppd-D1a, and heading was accelerated 10.3 days compared with the Ppd-D1b genotype. Early cultivars with Ppd-D1a may have been selected to avoid damage from preharvest rain. In the Hokkaido region, Ppd-D1a frequency was lower and heading date was late regardless of Ppd-D1 genotype, suggesting another genetic mechanism for late heading in Hokkaido cultivars. In this study, only 11 cultivars proved to carry Ppd-B1a, and all of them carried another photoperiod-insensitive allele, Ppd-D1a. The Ppd-B1a/Ppd-D1a genotype headed 6.7 days earlier than the Ppd-B1b/Ppd-D1a genotype, indicating a significant effect of Ppd-B1a in the genetic background with Ppd-D1a. Early-maturity breeding in Japan is believed to be accelerated by the introduction of the Ppd-B1a allele into medium-heading cultivars carrying Ppd-D1a. Pedigree analysis showed that Ppd-B1a in three extra-early commercial cultivars was inherited from ‘Shiroboro 21’ by early-heading Chugoku lines bred at the Chugoku Agriculture Experimental Station.