Genetic analysis of the components of winterhardiness in barley (Hordeum vulgare L.)

Winterhardiness in cereals is the consequence of a number of complex and interacting component characters: cold tolerance, vernalization requirement, and photoperiod sensitivity. An understanding of the genetic basis of these component traits should allow for more-effective selection. Genome map-based analyses hold considerable promise for dissecting complex phenotypes. A 74-point linkage map was developed from 100 doubled haploid lines derived from a winter x spring barley cross and used as the basis for quantitative trait locus (QTL) analyses to determine the chromosome location of genes controlling components of winterhardiness. Despite the greater genome coverage provided by the current map, a previously-reported interval on chromosome 7 remains the only region where significant QTL effects for winter survival were detected in this population. QTLs for growth habit and heading date, under 16 h and 24 h light, map to the same region. A QTL for heading date under these photoperiod regimes also maps to chromosome 2. Contrasting alleles at these loci interact in an epistatic fashion. A distinct set of QTLs mapping to chromosomes 1, 2, 3, and 5 determined heading date under 8 h of light. Under field conditions, all QTLs identified under controlled environment conditions were determinants of heading date. Patterns of differential QTL expression, coupled with additive and additive x additive QTL effects, underscore the complexity of winterhardiness. The presence of unique phenotype combinations in the mapping population suggests that coincident QTLs for heading date and winter survival represent the effects of linkage rather than pleiotropy.     

Zearalenone-Like Substance in Winter Plants and Its Relation to Vernalization

It was found that there is a zearalenone-like substance in the growing points of vernalized wheat seedlings and carrot plants. The content of this substance synchronously increased with the depth of vernalization. It was separated by thin layer chromatography. Its UV spectrum was similar to that of zearalenone. It is believed that this substance may be one of the important endogenous substances which control the vernalization of winter plants.     

Thermoinduction of genes encoding the enzymes of gibberellin biosynthesis and a putative negative regulator of gibberellin signal transduction in<Emphasis Type="Italic"> Eustoma grandiflorum</Emphasis>

Eustoma grandiflorum Shinn requires vernalization for the induction of stem elongation and flowering. To investigate the role of gibberellins (GAs) in vernalization, the expression levels of genes encoding enzymes of GA biosynthesis, copalyl diphosphate synthetase, GA 20-oxidase and GA 3-hydroxylase, were examined using two culitvars that show different responses to vernalization. The three genes were induced in a vernalization- and a cultivar-dependent manner. EgSPY, a putative negative regulator of GA signal transduction, was also induced during the vernalization period. The results suggest that the expression of the genes encoding GAs biosynthesis is regulated by vernalization. We postulate that EgSPY functions as a negative regulator of GA signal transduction during vernalization, inhibiting adventitious shoot elongation during vernalization.Communicated by K.K. Kamo     

The regulatory role of vernalization in the expression of low-temperature-induced genes in wheat and rye

Low temperature is one of the primary stresses limiting the growth and productivity of wheat (Triticum aestivum L.) and rye (Secale cereale L.). Winter cereals low-temperature-acclimate when exposed to temperatures colder than 10°C. However, they gradually lose their ability to tolerate below-freezing temperatures when they are maintained for long periods of time in the optimum range for low-temperature acclimation. The overwinter decline in low-temperature response has been attributed to an inability of cereals to maintain low-temperature-tolerance genes in an up-regulated state once vernalization saturation has been achieved. In the present study, the low-temperature-induced Wcs120 gene family was used to investigate the relationship between low-temperature gene expression and vernalization response at the molecular level in wheat and rye. The level and duration of gene expression determined the degree of low-temperature tolerance, and the vernalization genes were identified as the key factor responsible for the duration of expression of low-temperature-induced genes. Spring-habit cultivars that did not have a vernalization response were unable to maintain low-temperature-induced genes in an up-regulated condition when exposed to 4°C. Consequently, they were unable to achieve the same levels of low-temperature tolerance as winter-habit cultivars. A close association between the point of vernalization saturation and the start of a decline in the Wcs120 gene-family mRNA level and protein accumulation in plants maintained at 4°C indicated that vernalization genes have a regulatory influence over low-temperature gene expression in winter cereals.     

Heading date QTL in a spring × winter barley cross evaluated in Mediterranean environments

Heading date is a key trait for the adaptation of barley to Mediterranean environments. We studied the genetic control of flowering time under Northern Spanish (Mediterranean) conditions using a new population derived from the spring/winter cross Beka/Mogador. A set of 120 doubled haploid lines was evaluated in the field, and under controlled temperature and photoperiod conditions. Genotyping was carried out with 215 markers (RFLP, STS, RAPD, AFLP, SSR), including markers for vernalization candidate genes, HvBM5 (Vrn-H1), HvZCCT (Vrn-H2), and HvT SNP22 (Ppd-H1). Four major QTL, and the interactions between them, accounted for most of the variation in both field (71–92%) and greenhouse trials (55–86%). These were coincident with the location of the major genes for response to vernalization and short photoperiod (Ppd-H2 on chromosome 1H). A major QTL, near the centromere of chromosome 2H was the most important under autumn sowing conditions. Although it is detected under all conditions, its action seems not independent from environmental cues. An epistatic interaction involving the two vernalization genes was detected when the plants were grown without vernalization and under long photoperiod. The simultaneous presence of the winter Mogador allele at the two loci produced a marked delay in heading date, beyond a mere additive effect. This interaction, combined with the effect of the gene responsive to short photoperiod, Ppd-H2, was found responsible of the phenomenon known as short-day vernalization, present in some of the lines of the population. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Regions associated with repression of the barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis>) <Emphasis Type="Italic">VERNALIZATION1</Emphasis> gene are not required for cold induction

Activity of the VERNALIZATION1 (VRN1) gene is required for flowering in temperate cereals such as wheat and barley. In varieties that require prolonged exposure to cold to flower (vernalization), VRN1 is expressed at low levels and is induced by vernalization to trigger flowering. In other varieties, deletions or insertions in the first intron of the VRN1 gene are associated with increased VRN1 expression in the absence of cold treatment, reducing or eliminating the requirement for vernalization. To characterize natural variation in VRN1, the first intron of the barley (Hordeum vulgare) VRN1 gene (HvVRN1) was assayed for deletions or insertions in a collection of 1,000 barleys from diverse geographical regions. Ten alleles of HvVRN1 containing deletions or insertions in the first intron were identified, including three alleles that have not been described previously. Different HvVRN1 alleles were associated with differing levels of HvVRN1 expression in non-vernalized plants and with different flowering behaviour. Using overlapping deletions, we delineated regions in the HvVRN1 first intron that are associated with low levels of HvVRN1 expression in non-vernalized plants. Deletion of these intronic regions does not prevent induction of HvVRN1 by cold or the maintenance of increased HvVRN1 expression following cold treatment. We suggest that regions within the first intron of HvVRN1 are required to maintain low levels of HvVRN1 expression prior to winter but act independently of the regulatory mechanisms that mediate induction of HvVRN1 by cold during winter. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nucleotide sequence data reported are available in the DDBJ/EMBL/GenBank databases under the accession numbers 1179825, 1179833, 1179836, 1179858.     

Quantitative-genetic analysis of reciprocal crosses between a winter and a spring cultivar of common wheat (Triticum aestivum L.)

The winter wheat cultivar Sakhalin (parent A) and the spring wheat cultivar Khush-hal (parent B), assumed to be both nuclear-genetically and plasmotypically different, were reciprocally crossed and the F₁ generation of the reciprocals backcrossed to either parent. The populations (AxB)F₁, (AxB)F₂, [(AxB)F₁xA]F₁ and [(AxB)F₁xB]F₁, and their reciprocals (BxA)F₁, (BxA)F₂, [(BxA)F₁xA]F₁ and [(BxA)F₁xB]F₁, assumed to have the plasmotypes of the parents A and B respectively, were spring sown in the same field from unvernalized (experiment #1) and vernalized (experiment #2) seeds. The results of the analyses of variance of the data recorded in the two experiments are fairly similar. In both the experiments the plasmotypically A populations have produced more heads and higher grain yields per plant than their plasmotypically B reciprocals. The components of the family means and of half the differences between the reciprocals within the families show that vernalization has not affected the efficiency of the nuclear genes of parent A but has reduced the efficiency of the nuclear genes of parent B in both the homozygous and the heterozygous states. This in turn has affected the components of family mean squares and those of the family x reciprocals interaction mean squares in the analysis of variance tables.     

Flowering requirements of Polymnia canadensis (Asteraceae) and their influence on its life history variation

The purpose of this study is to determine the flowering requirements of Polymnia canadensis and how they correspond to the occurrence of winter annuals, biennials, and short-lived monocarpic perennials in this species. Polymnia canadensis has a vernalization requirement for flowering, and even very small plants (i.e., those with one pair of leaves) can be vernalized. Vernalized plants can flower under both long- and short days. However, to flower plants must attain a minimum postvernalization size. Plants of this primarily monocarpic species that do not die after they flower once require another period of vernalization to flower a second time (i.e., to be dicarpic). Vernalized plants exposed to high temperatures can be devernalized; these must be re-vernalized in order to flower.