Rachis Internode Research Articles

Streptomyces pratensis S10 inhibits spread of Fusarium graminearum invasive hyphae and toxisome formation in wheat plants.

Fusarium head blight (FHB) of wheat, mainly caused by Fusarium graminearum, leads to severe economic losses worldwide. Effective management measures for controlling FHB are not available, due to a lack of resistant cultivars. Currently, the utilization of biological control is a promising approach that can be used to help manage FHB. Previous studies have confirmed that Streptomyces pratensis S10 harbors excellent inhibitory effects on F. graminearum. However, there is no information regarding invasive hyphae of F. graminearum are inhibited by S10. Thus, we investigated the effects of S10 on F. graminearum strain PH-1 hyphae extension, toxisome formation, and TRI5 gene expression on wheat plants via microscopic observation. The results showed that S10 effectively inhibited spread of F. graminearum hyphae along the rachis, restricting the infection of neighboring florets via the phloem. In the presence of S10, the hyphal growth is impeded by formation of dense cell wall thickenings in the rachis internode surrounding the F. graminearum infection site, avoiding cell plasmolysis and collapse. We further demonstrated that S10 largely prevented cell-to-cell invasion of fungal hyphae inside wheat coleoptiles using a constitutively green fluorescence protein-expressing F. graminearum strain, PH-1. Importantly, S. pratensis S10 inhibited toxisome formation and TRI5 gene expression in wheat plants during infection. Collectively, these findings indicated that S. pratensis S10 prevents spread of F. graminearum invasive hyphae via the rachis.

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The Structural–Rhythmological Organization of Coelogyne (Orchidaceae Juss.) Inflorescences

The morphostructure of inflorescences in the genus Coelogyne Lindl. was studied for the first time using a structural–rhythmological approach. Three species of Coelogyne were used to describe one-season, intercalary, and all-season inflorescences. In C. monilirachis, a new type of all-season inflorescence was identified, characterized by a prolonged sympodial growth of the rachis, lack of a pronounced dormant period, and thickening of all rachis internodes except the first. This inflorescence has been determined to be a compound monochasial cyme, with each floral unit represented by a separate flower. C. ovalis has a one-season inflorescence, with the floral unit being a determinate bracteous spike (stachyoid), and C. prolifera developed intercalary inflorescences united in an indeterminate bracteous spike.

Mutations within the miR172 target site of wheat AP2 homoeologs regulate lodicule size and rachis internode length.

Closed fertilization in flowers, or cleistogamy, reduces the risk of fungal infection in Triticeae crops. In barley (Hordeum vulgare), cleistogamy is determined by a single recessive gene, cly1, which results from a single nucleotide polymorphism within the microRNA172 target site of the Apetala2 (AP2) transcription factor gene. The recessive cly1 allele negatively regulates the development of lodicules, keeping florets closed at anthesis. However, cleistogamy is not evident in hexaploid wheat (Triticum aestivum) cultivars. This study aimed at identifying mutations in wheat AP2 orthologs by ethyl methane sulfonate-induced mutagenesis and high-resolution melt analysis. Although flowers of AP2 mutants induced in the A and D genomes opened at anthesis, their lodicule size was significantly smaller, especially in the direction of depth, than that of wild-type plants. One of the mutants that carried a nucleotide replacement in AP2 from the D genome produced a compact spike caused by a substantial decrease in rachis internode length, analogous to the barley dense spike. Cleistogamous hexaploid wheat might be generated by combining effective mutant alleles of AP2-homoeologous genes.

Indole-3-butyric acid, an alternative to GA3 for bunch quality enhancing of table grape Vitis vinifera L. cv. Superior Seedless

Gibberellic acid (GA3) is the most widely used plant growth regulator to thin out bunches and increase berry size in seedless table grapes, but there is evidence of its negative effects, including loss of fertility and malformations of the rachis. This study analyses the effects of applying the auxin indole-3-butyric acid (IBA), as a novel alternative, on fruit yield, bunch structure, anatomy and postharvest quality of cv. Superior Seedless. The application of IBA and GA3 at different phenological stages (15 cm shoot length, full bloom, fruit set and pea-size berry) and doses (2, 20 and 50 ppm) were compared in a two-growing season experiment. Spraying IBA at full bloom or fruit set, improved bunch weight by increasing the size and weight of the berries, correlating with the promotion of rachis vascular tissues. Bunches treated with IBA retained a greater number of berries at harvest without generating compactness, since the elongation of the rachis internodes and lateral shoulders were also promoted. In addition, IBA augmented postharvest quality of bunches by reducing rachis browning and increasing berry firmness. These results suggest that the use of IBA is a beneficial technology to improve bunch structure and quality in seedless grapes. Highlights IBA effects applied at different dosages and phenological stages was evaluated. IBA at full bloom and fruit set improved bunch and berry weight and berry size. The number of berries per bunch increased with IBA at full bloom and fruit set. IBA increased lateral rachis internode and lateral rachis length via promotion of vascular tissues. IBA increased postharvest quality, reduced rachis browning and increased of berry firmness.

Mycotoxin DON Accumulation in Wheat Grains Caused by Fusarium Head Blight Are Significantly Subjected to Inoculation Methods.

The disease severity and mycotoxin DON content in grains caused by fusarium head blight (FHB) have been two prioritized economical traits in wheat. Reliable phenotyping is a prerequisite for genetically improving wheat resistances to these two traits. In this study, three inoculation methods: upper bilateral floret injection (UBFI), basal bilateral floret injection (BBFI), and basal rachis internode injection (BRII), were applied in a panel of 22 near-isogenic lines (NILs) contrasting in Fhb1 alleles. The results showed that inoculation methods had significant influence on both disease severity and mycotoxin accumulation in grains, and the relationship between them. UBFI method caused chronic FHB symptom characterized as slow progress of the pathogen downward from the inoculation site, which minimized the difference in disease severity of the NILs, but, unexpectedly, maximized the difference in DON content between them. The BBFI method usually caused an acute FHB symptom in susceptible lines characterized as premature spike death (PSD), which maximized the difference in disease severity, but minimized the difference in DON content in grains between resistant and susceptible lines. The BRII method occasionally caused acute FHB symptoms for susceptible lines and had relatively balanced characteristics of disease severity and DON content in grains. Therefore, two or more inoculation methods are recommended for precise and reliable evaluation of the overall resistance to FHB, including resistances to both disease spread within a spike and DON accumulation in grains.

A 3D grape bunch reconstruction pipeline based on constraint-based optimisation and restricted reconstruction grammar

• Wine grape bunch architecture reconstruction from a 2D image. • Prediction of invisible bunch elements including their connectivity. • Element-wise comparison improves on existing methods. • Application on a larger scale would facilitate grapevine breeding trials. Phenotypic traits of grapevines are known to be closely related to grapevine yield, wine flavour and sensitivity to disease. Traditional phenotyping methods based on manual measurements face the bottleneck of intensely repetitive and time consuming measurement. As viticulturists turning their focus to the 3D automatic phenotyping domain, existing work on grape bunch phenotyping indicates deficiencies in incomplete reconstruction information, poor element coincidence with the ground truth and poor performance under field conditions. To this end, the proposed work introduces a novel reconstruction pipeline by dividing it into sub-problems of visible-berry-related reconstruction and invisible element prediction. By taking a 2D image of the target bunch as the only sensor input, visible berries are detected using image processing algorithms. With the detected berry information, their morphological positions are predicted, from which internodes that are associated with the detected berries are derived. Parameters of derived internodes are then estimated employing constraint-based optimisation theory, from which the visible-berry-related reconstruction is able to be achieved. Invisible element prediction is then conducted by filling elements according to a Restricted Reconstruction Grammar (RRG). A fully reconstructed bunch model is finally presented. Compared with existing work, the proposed grape bunch reconstruction pipeline achieved an improvement in quantity estimation of rachis internodes, tertiary internodes and pedicels, whose percentage errors were indicated as 21.2, 42.2 and 31.2% respectively. A better performance was also revealed in length estimations of secondary internodes and pedicels with percentage errors of 3.5 and 0.5%. This may largely facilitate related studies on disease control of grape bunches since internode numbers and lengths are closely related to bunch compactness which is the indicator of disease sensitivity. Especially, the proposed reconstruction pipeline shows an promising improvement in element coincidence with F 1 scores of 0.90, 0.77, 0.45, 0.43 for respective element types. Knowing that element coincidence may influence the inner space utilisation of a bunch, the proposed work provides a better option for 3D grape bunch phenotyping.

Basal Rachis Internode Injection: A Novel Inoculation Method to Evaluate Wheat Resistance to Fusarium Head Blight.

Fusarium head blight (FHB) is one of the most destructive fungal diseases of wheat. However, difficulties in reliably phenotyping of this disease have greatly hindered the understanding of the mechanism of wheat-pathogen interaction and genetic improvement of FHB resistance. Here we report a novel inoculation method called basal rachis internode injection (BRII), in which inoculum is injected into the basal internode of a rachis instead of a floret, as is done in single floret inoculation (SFI). One of the prominent advantages of BRII over SFI and other traditional methods lies in its independence from the moisture-maintaining system that is necessary for all existing methods, making it insensitive to environmental humidity and hence cost-effective. Another unique feature of BRII is that this method produces nearly clear-cut reaction types, by which FHB resistance can be treated as a qualitative trait because generally no FHB symptoms appear on the spikelets of resistant genotypes. In addition, BRII outperformed SFI with a higher infection rate and better goodness of fit with known FHB resistance and quantitative trait locus components in a panel of 15 genotypes, as well as two populations of recombinant inbred lines segregating in Fhb1. Note that BRII and SFI methods are not mutually exclusive but rather complementary because each method has its own advantages in differentiating FHB resistance between genotypes. Combining these two methods would significantly improve the reliability and consistency of FHB phenotyping in wheat.

BELL1-like homeobox genes regulate inflorescence architecture and meristem maintenance in rice.

Inflorescence architecture is diverse in angiosperms, and is mainly determined by the arrangement of the branches and flowers, known as phyllotaxy. In rice (Oryza sativa), the main inflorescence axis, called the rachis, generates primary branches in a spiral phyllotaxy, and flowers (spikelets) are formed on these branches. Here, we have studied a classical mutant, named verticillate rachis (ri), which produces branches in a partially whorled phyllotaxy. Gene isolation revealed that RI encodes a BELL1-type homeodomain transcription factor, similar to Arabidopsis PENNYWISE/BELLRINGER/REPLUMLESS, and is expressed in the specific regions within the inflorescence and branch meristems where their descendant meristems would soon initiate. Genetic combination of an ri homozygote and a mutant allele of RI-LIKE1 (RIL1) (designated ri ril1/+ plant), a close paralog of RI, enhanced the ri inflorescence phenotype, including the abnormalities in branch phyllotaxy and rachis internode patterning. During early inflorescence development, the timing and arrangement of primary branch meristem (pBM) initiation were disturbed in both ri and ri ril1/+ plants. These findings suggest that RI and RIL1 were involved in regulating the phyllotactic pattern of the pBMs to form normal inflorescences. In addition, both RI and RIL1 seem to be involved in meristem maintenance, because the ri ril1 double-mutant failed to establish or maintain the shoot apical meristem during embryogenesis.

A Homolog of Blade-On-Petiole 1 and 2 (BOP1/2) Controls Internode Length and Homeotic Changes of the Barley Inflorescence.

Inflorescence architecture in small-grain cereals has a direct effect on yield and is an important selection target in breeding for yield improvement. We analyzed the recessive mutation laxatum-a (lax-a) in barley (Hordeum vulgare), which causes pleiotropic changes in spike development, resulting in (1) extended rachis internodes conferring a more relaxed inflorescence, (2) broadened base of the lemma awns, (3) thinner grains that are largely exposed due to reduced marginal growth of the palea and lemma, and (4) and homeotic conversion of lodicules into two stamenoid structures. Map-based cloning enforced by mapping-by-sequencing of the mutant lax-a locus enabled the identification of a homolog of BLADE-ON-PETIOLE1 (BOP1) and BOP2 as the causal gene. Interestingly, the recently identified barley uniculme4 gene also is a BOP1/2 homolog and has been shown to regulate tillering and leaf sheath development. While the Arabidopsis (Arabidopsis thaliana) BOP1 and BOP2 genes act redundantly, the barley genes contribute independent effects in specifying the developmental growth of vegetative and reproductive organs, respectively. Analysis of natural genetic diversity revealed strikingly different haplotype diversity for the two paralogous barley genes, likely affected by the respective genomic environments, since no indication for an active selection process was detected.

Molecular and comparative mapping of genes governing spike compactness from wild emmer wheat

The development and morphology of the wheat spike is important because the spike is where reproduction occurs and it holds the grains until harvest. Therefore, genes that influence spike morphology are of interest from both theoretical and practical stand points. When substituted for the native chromosome 2A in the tetraploid Langdon (LDN) durum wheat background, the Triticum turgidum ssp. dicoccoides chromosome 2A from accession IsraelA confers a short, compact spike with fewer spikelets per spike compared to LDN. Molecular mapping and quantitative trait loci (QTL) analysis of these traits in a homozygous recombinant population derived from LDN×the chromosome 2A substitution line (LDNIsA-2A) indicated that the number of spikelets per spike and spike length were controlled by linked, but different, loci on the long arm of 2A. A QTL explaining most of the variation for spike compactness coincided with the QTL for spike length. Comparative mapping indicated that the QTL for number of spikelets per spike overlapped with a previously mapped QTL for Fusarium head blight susceptibility. The genes governing spike length and compactness were not orthologous to either sog or C, genes known to confer compact spikes in diploid and hexaploid wheat, respectively. Mapping and sequence analysis indicated that the gene governing spike length and compactness derived from wild emmer could be an ortholog of the barley Cly1/Zeo gene, which research indicates is an AP2-like gene pleiotropically affecting cleistogamy, flowering time, and rachis internode length. This work provides researchers with knowledge of new genetic loci and associated markers that may be useful for manipulating spike morphology in durum wheat.

Genetic mapping of the labile (lab) gene: a recessive locus causing irregular spikelet fertility in labile-barley (Hordeum vulgare convar. labile)

The recessive labile locus mapped on chromosome 5HL causes irregular spikelet fertility and controls floret development as well as row-type in barley. The labile-barley displays a variable number of fertile spikelets at each rachis internode (0-3 fertile spikelets/rachis internode) which is intermediate between that observed in two- or six-rowed types. Previous re-sequencing of Vrs1 in 219 labile-barley (Hordeum vulgare L. convar. labile) accessions showed that all carried a six-rowed specific allele. We therefore hypothesized that this seemingly random reduction in spikelet fertility is most likely caused by the labile (lab) locus, which we aimed to phenotypically and genetically define. Here, we report a detailed phenotypic analysis of spikelet fertility in labile-barleys in comparison to two- and six-rowed genotypes using scanning electron microscopy analysis. We found that the first visible morphological deviation occurred during the stamen primordium stage, when we regularly observed the appearance of arrested central floral primordia in labile but not in two- or six-rowed barleys. At late stamen and early awn primordium stages, lateral florets in two-rowed and only some in labile-barley showed retarded development and reduction in size compared with fully fertile lateral florets in six-rowed barley. We used two F2 mapping populations to generate whole genome genetic linkage maps and ultimately locate the lab locus as a recessive Mendelian trait to a 4.5-5.8 cM interval at approximately 80 cM on chromosome 5HL. Our results will help identifying the role of the lab gene in relation to other spikelet fertility factors in barley.

Six-rowed spike4 ( Vrs4 ) controls spikelet determinacy and row-type in barley

Inflorescence architecture of barley (Hordeum vulgare L.) is common among the Triticeae species, which bear one to three single-flowered spikelets at each rachis internode. Triple spikelet meristem is one of the unique features of barley spikes, in which three spikelets (one central and two lateral spikelets) are produced at each rachis internode. Fertility of the lateral spikelets at triple spikelet meristem gives row-type identity to barley spikes. Six-rowed spikes show fertile lateral spikelets and produce increased grain yield per spike, compared with two-rowed spikes with sterile lateral spikelets. Thus, far, two loci governing the row-type phenotype were isolated in barley that include Six-rowed spike1 (Vrs1) and Intermedium-C. In the present study, we isolated Six-rowed spike4 (Vrs4), a barley ortholog of the maize (Zea mays L.) inflorescence architecture gene RAMOSA2 (RA2). Eighteen coding mutations in barley RA2 (HvRA2) were specifically associated with lateral spikelet fertility and loss of spikelet determinacy. Expression analyses through mRNA in situ hybridization and microarray showed that Vrs4 (HvRA2) controls the row-type pathway through Vrs1 (HvHox1), a negative regulator of lateral spikelet fertility in barley. Moreover, Vrs4 may also regulate transcripts of barley SISTER OF RAMOSA3 (HvSRA), a putative trehalose-6-phosphate phosphatase involved in trehalose-6-phosphate homeostasis implicated to control spikelet determinacy. Our expression data illustrated that, although RA2 is conserved among different grass species, its down-stream target genes appear to be modified in barley and possibly other species of tribe Triticeae.

Synflorescence analysis in South American species of <i>Andropogon</i> section <i>Leptopogon</i> (Andropogoneae, Poaceae): a tool to identify different ploidy levels

In southern South America, Andropogon sect. Leptopogon is represented by both diploid and hexaploid species. In order to compare the synflorescence structures and floral arrays in relation to ploidy levels in species of Andropogon , the section Leptopogon was used as model. In this study, the synflorescence structure was typologically characterized. The structural analysis was based on the typology system developed by Troll and Weberling, which has proved useful in describing inflorescences. A comparative analysis of the variations observed in the structure of the synflorescence and a morphometric analysis using principal component analysis were carried out. These results revealed that diploid and hexaploid species present differences mainly in the following parameters: synflorescence length, number of internodes of the enrichment zone main axis, number of internodes and length of paracladia of the trophotagma, number of long paracladia per unit of inflorescence, length of hairs on articulated rachis internodes, pedicellate spikelet length, pedicellate spikelet width, pedicel width, length of hairs on pedicel, sessile spikelet length, awn length and floral system arrangements. This study provides a useful tool to distinguish between diploid and hexaploid species of Andropogon sect. Leptopogon . We propose naming the group of diploid species from Central and South America as the Andropogon selloanus complex.

High resolution mapping of Dense spike-ar (dsp.ar) to the genetic centromere of barley chromosome 7H

Spike density in barley is under the control of several major genes, as documented previously by genetic analysis of a number of morphological mutants. One such class of mutants affects the rachis internode length leading to dense or compact spikes and the underlying genes were designated dense spike (dsp). We previously delimited two introgressed genomic segments on chromosome 3H (21 SNP loci, 35.5cM) and 7H (17 SNP loci, 20.34cM) in BW265, a BC(7)F(3) nearly isogenic line (NIL) of cv. Bowman as potentially containing the dense spike mutant locus dsp.ar, by genotyping 1,536 single nucleotide polymorphism (SNP) markers in both BW265 and its recurrent parent. Here, the gene was allocated by high-resolution bi-parental mapping to a 0.37cM interval between markers SC57808 (Hv_SPL14)-CAPSK06413 residing on the short and long arm at the genetic centromere of chromosome 7H, respectively. This region putatively contains more than 800 genes as deduced by comparison with the collinear regions of barley, rice, sorghum and Brachypodium, Classical map-based isolation of the gene dsp.ar thus will be complicated due to the infavorable relationship of genetic to physical distances at the target locus.

Harvesting practices and early Neolithic barley cultivation at el-Hemmeh, Jordan

Recent research strongly suggests polyphetic origins of multiple cultigens across Southwest Asia approximately 11,000 years ago during the Pre-Pottery Neolithic A period. The harvesting practices that contributed to the dedicated use of cultivation as a plant exploitation strategy remain largely unidentified. Archaeobotanical data from el-Hemmeh, a settlement site dating to ca. 10850 cal. b.p., provides an opportunity to examine in close detail the harvesting strategies that may have contributed to the development of domesticated forms. Initial analyses indicate a variety of wild plant foods including barley, lentils, vetch, Pistacia cf. atlantica and fig were exploited, while the presence of large predomesticated barley grains and potential weed species suggest cereal cultivation was also pursued at the site. Barley rachis internodes from el-Hemmeh typically possess a wild morphology, but 22% of specimens show evidence of a forced or “ripped” disarticulation. This suggests barley may have been harvested while ears were partially immature and required subsequent processing in order to disarticulate spikelets.

Quantitative trait loci (QTL) mapping for inflorescence length traits in Lablab purpureus (L.) sweet

Lablab purpureus (L.) sweet is an ancient legume species whose immature pods serve as a vegetable in south and south-east Asia. The objective of this study is to identify quantitative trait loci (QTLs) associated with quantitative traits such as inflorescence length, peduncle length from branch to axil, peduncle length from axil to lowermost flowering node, rachis length, node number of inflorescence, rachis internode length, node order of the first inflorescence and node order of lowest inflorescence, which are key characters affecting the output of vegetable cultivars of lablab. A molecular linkage map was constructed using a F 2 population derived from the cross (Meidou2012 × Nanhui 23). The map covers 1302.4 cm with 131 loci (122 RAPD and nine morphological markers) and consist 14 linkage groups. In the F 2 population and derived F 3 families, a total of 46 QTLs explained from 8.1 to 55.0% of phenotypic variance of the traits. Of them, 16 QTLs were detectable in the same linkage regions among different generation/season combinations, and 10 QTL clusters were mapped. It suggests that, genes which control inflorescence growth were pleiotropic or coincident involving more than one trait. Thus, these QTLs may be tagged for marker assisted selection to improve yield of lablab. Key words: Inflorescence length, lablab, linkage map, quantitative trait loci (QTLs), random amplification of polymorphic DNA (RAPD).

A new semi-dwarfing gene identified by molecular mapping of quantitative trait loci in barley

Semi-dwarfing genes have been widely used in spring barley (Hordeum vulgare L.) breeding programs in many parts of the world, but the success in developing barley cultivars with semi-dwarfing genes has been limited in North America. Exploiting new semi-dwarfing genes may help in solving this dilemma. A recombinant inbred line population was developed by crossing ZAU 7, a semi-dwarf cultivar from China, to ND16092, a tall breeding line from North Dakota. To identify quantitative trait loci (QTL) controlling plant height, a linkage map comprised of 111 molecular markers was constructed. Simple interval mapping was performed for each of the eight environments. A consistent QTL for plant height was found on chromosome 7HL. This QTL is not associated with maturity and rachis internode length. We suggest the provisional name Qph-7H for this QTL. Qph-7H from ZAU 7 reduced plant height to about 3/4 of normal; thus, Qph-7H is considered a semi-dwarfing gene. Other QTLs for plant height were found, but their expression was variable across the eight environments tested.

Flt-2L, a locus in barley controlling flowering time, spike density, and plant height

Flowering time represents an important adaptive trait for temperate cereal crops and may also impact on frost damage in cereal reproductive tissues by enabling escape or by influencing accumulation of genuine tolerance. The Flowering time-2L (Flt-2L) quantitative trait locus (QTL) on the distal end of barley chromosome arm 2HL overlaps with QTL for rachis internode length and reproductive frost damage. Flt-2L was also found to be associated with plant height. By combining marker analysis with phenotyping in progeny families of selected Amagi Nijo x WI2585 F(6) recombinants, we were able to map quantitative flowering time, rachis internode length, and plant height effects on 2HL as discrete Mendelian traits. The three developmental characters showed codominant modes of expression and perfectly cosegregated with one another in a 1.3-cM marker interval, indicating control by the same gene or closely linked genes. Twelve genes were identified in the related intervals in the rice and Brachypodium distachyon genomes. The HvAP2 gene cosegregated with Flt-2L and represents a plausible candidate for Flt-2L, since it is highly similar to the wheat domestication gene Q which has similar developmental effects. These data will contribute to isolation of the Flt-2L gene(s) and help establish the basis of the frost damage QTL.

Genes and traits associated with chromosome 2H and 5H regions controlling sensitivity of reproductive tissues to frost in barley

Frost at flowering can cause significant damage to cereal crops. QTL for low temperature tolerance in reproductive tissues (LTR tolerance) were previously described on barley 2HL and 5HL chromosome arms. With the aim of identifying potential LTR tolerance mechanisms, barley Amagi Nijo x WI2585 and Haruna Nijo x Galleon populations were examined for flowering time and spike morphology traits associated with the LTR tolerance loci. In spring-type progeny of both crosses, winter alleles at the Vrn-H1 vernalization response locus on 5H were linked in coupling with LTR tolerance and were unexpectedly associated with earlier flowering. In contrast, tolerance on 2HL was coupled with late flowering alleles at a locus we named Flt-2L. Both chromosome regions influenced chasmogamy/cleistogamy (open/closed florets), although tolerance was associated with cleistogamy at the 2HL locus and chasmogamy at the 5HL locus. LTR tolerance controlled by both loci was accompanied by shorter spikes, which were due to fewer florets per spike on 5HL, but shorter rachis internodes on 2HL. The Eps-2S locus also segregated in both crosses and influenced spike length and flowering time but not LTR tolerance. Thus, none of the traits was consistently correlated with LTR tolerance, suggesting that the tolerance may be due to some other visible trait or an intrinsic (biochemical) property. Winter alleles at the Vrn-H1 locus and short rachis internodes may be of potential use in barley breeding, as markers for selection of LTR tolerance at 5HL and 2HL loci, respectively.