Vertical differentiation in root placement is one of the potential mechanisms of plant niche differentiation. It can be due to the remarkable plasticity of roots in response to nutrients and neighbours, but most data on it come from pot or garden experiments. The roles of vertical differentiation and of plasticity in it in the field are thus not well known. We examined species‐specific root vertical distribution in a montane grassland using quantitative real‐time PCR. We asked whether individual species differ in their rooting depths, whether such differences are associated with above‐ground functional traits (such as height or specific leaf area), and whether they respond to the presence of a competitor. This response was assessed by comparison of species‐specific vertical profiles between control plots and plots where the dominant species, Festuca rubra, had been removed. Vertical profiles of individual species varied considerably, from species with most root biomass concentrated in the uppermost (<2 cm) soil layer, through species with uniform vertical distribution, to a species with roots predominantly below 8 cm (Nardus stricta). Species at the fast end of the plant economy spectrum were more likely to place their roots in the uppermost layers. Grassland species, thus, exploit different parts of the below‐ground resources in spite of their short stature, minor differences in height above‐ground and shallow soil. While below‐ground and above‐ground biomasses of most species were higher in the removal plots, species rooting patterns did not change in response to the removal. The interspecific differences in vertical profiles were thus due to species' innate differences, not to plastic responses to the presence of the dominant species. Synthesis. The findings imply that vertical root differentiation in the field is strong and can contribute to niche differentiation. However, the role of root plasticity in natural systems may be considerably weaker than in artificial systems with few species and strong nutrient gradients. This absence of the plastic response in the field is likely to be due to a fairly homogeneous distribution of nutrients in the soil and to the predominantly symmetric nature of below‐ground competition.
Therefore, a doubled haploid (DH) mapping population (n = 122) was created by crossing SusPtrit with Golden Promise to develop a 'Golden SusPtrit', i.e., a barley line combining SusPtrit's high susceptibility to non-adapted rust fungi with the high amenability of Golden Promise for transformation. We identified nine genomic regions occupied by resistance quantitative trait loci (QTls) against four non-adapted rust fungi and P. hordei isolate 1.2.1 (Ph.1.2.1). Four DHs were selected for an Agrobacterium-mediated transformation efficiency test. They were among the 12 DH lines most susceptible to the tested nonadapted rust fungi. The most efficiently transformed DH line was SG062n (11-17 transformants per 100 immature embryos). The level of non-adapted rust infection on SG062n is either similar to or higher than the level of infection on SusPtrit. Against Ph.1.2.1, the latency period conferred by SG062n is as short as that conferred by SusPtrit. SG062n, designated 'Golden SusPtrit', will be a valuable experimental line that could replace SusPtrit in nonhost and partial resistance studies, especially for stable transformation using candidate genes that may be involved in rustresistance mechanisms.
Key messageRphq2, a minor gene for partial resistance toPuccinia hordei, was physically mapped in a 188 kbp introgression with suppressed recombination between haplotypes ofrphq2andRphq2barley cultivars.AbstractPartial and non-host resistances to rust fungi in barley (Hordeum vulgare) may be based on pathogen-associated molecular pattern (PAMP)-triggered immunity. Understanding partial resistance may help to understand non-host resistance, and vice versa. We constructed two non-gridded BAC libraries from cultivar Vada and line SusPtrit. Vada is immune to non-adapted Puccinia rust fungi, and partially resistant to P. hordei. SusPtrit is susceptible to several non-adapted rust fungi, and has been used for mapping QTLs for non-host and partial resistance. The BAC libraries help to identify genes determining the natural variation for partial and non-host resistances of barley to rust fungi. A major-effect QTL, Rphq2, for partial resistance to P. hordei was mapped in a complete Vada and an incomplete SusPtrit contig. The physical distance between the markers flanking Rphq2 was 195 Kbp in Vada and at least 226 Kbp in SusPtrit. This marker interval was predicted to contain 12 genes in either accession, of which only five genes were in common. The haplotypes represented by Vada and SusPtrit were found in 57 and 43 %, respectively, of a 194 barley accessions panel. The lack of homology between the two haplotypes probably explains the suppression of recombination in the Rphq2 area and limit further genetic resolution in fine mapping. The possible candidate genes for Rphq2 encode peroxidases, kinases and a member of seven-in-absentia protein family. This result suggests that Rphq2 does not belong to the NB-LRR gene family and does not resemble any of the partial resistance genes cloned previously.Electronic supplementary materialThe online version of this article (doi:10.1007/s00122-015-2627-5) contains supplementary material, which is available to authorized users.
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