Specific adaptation and breeding for marginal conditions

Summary Breeding has been very successful in generating cultivars that in favorable environments, and together with large use of fertilizer and chemical control of weeds, pest and diseases, have increased agricultural production several fold. Today the environmental impact of high input agriculture in more favorable environments causes growing concern. By contrast, the impact of breeding in marginal environments has been elusive. The paper discusses evidence showing that the use of breeding principles developed for, and successfully applied, in favorable environments may be the main reason for the lack of breeding progress in marginal environments. Very little breeding work has actually been done in marginal environments, although the theory of correlated responses to selection indicates that selection conducted in good environments or in well-managed experiment stations is not expected to be very efficient when genotype by environment interactions of a cross-over type exist. The assumptions that heritability is higher under good conditions and that there is a carry-over effect of high yield potential are not supported by experimental evidence. If the target environment is below the cross-over point, selection has to be conducted for specific adaptation to that environment. The concept of wide adaptation has more a geographical than an environmental meaning, and it reduces genetic diversity and increases genetic vulnerability. Eventually the issue of genetic heterogeneity versus genetic uniformity is discussed in relation to specific adaptation to marginal environments.     

Relationship between barley grain yield measured in low- and high-yielding environments

Summary The paper addresses the general question of identifying the optimum environment for selection in plant breeding programs for low input agricultural systems. After defining low-yielding and high-yielding environments based on the average grain yield of large numbers of barley genotypes in different cropping seasons, we examined: 1) the phenotypic relationships between the highest yielding genotypes in low- and high-yielding environments, and 2) the genetic correlation coefficients between grain yield in low- and high-yielding environments. The results indicate that the alleles controlling high grain yield in low-yielding conditions are at least partially different from those controlling high grain yield in high-yielding conditions. Therefore, selection in high-yielding environments is expected to produce a negative response or no response in low-yielding environments. This may explain why crop varieties bred under high-yielding conditions failed to have an impact in low-yielding agricultural systems. The results may be extrapolated to systems where environmental concern suggests a reduction of inputs by raising the question of whether crop breeding programs based on selection under high inputs are likely to generate the right type of germplasm for an environmentally friendly agriculture.     

Choice of selection strategy in breeding barley for stress environments

To determine the optimum selection environment for barley (Hordeum vulgare L.) targeted at low-input, stress environment, barley lines were selected for high yield under stress (YS), high yield under non-stress (YNS), or average yield in stress and non-stress conditions (YA) during three breeding cycles (cohorts) of three years each. The lines were then tested in a total of 21 year-location combinations with average grain yields ranging from 0.35 to 4.86 t ha^-1. Yield under stress of the YS lines was between 27% and 54% higher than that of the YNS lines, with the top YS lines yielding under stress between 16% and 30% more than the top YNS lines. Realized heritability was between 0.35 and 0.67 when selection was conducted under stress and was significant in all three cohorts. By contrast, selection under non-stress gave a significant response in only one cohort, and its efficiency in improving yield under stress was significantly lower than selection under stress. The best YNS line ranked only 19th for yield under stress. The highest-yielding lines under stress were not only selected under stress, but were also landraces collected in very dry areas (< 250 mm total annual rainfall). This confirms earlier findings and supports the idea that the most effective way to improve productivity of crops grown in less-favored areas is to use locally adapted germplasm and select in the target environment(s). This revised version was published online in July 2006 with corrections to the Cover Date.     

A methodological study on participatory barley breeding II. Response to selection

Farmer participation is increasingly seen as a key to develop technologies which are more relevant to farmers' communities. In plant breeding, farmer participation is seen as key to increase the probability of adoption of new varieties. This paper addresses the issue of selection efficiency in participatory plant breeding by testing the effect of selection environment and of who did the selection in one cropping season (1997) on the performance of the selected lines in the following cropping season (1998). Selection environment had a larger effect on response to selection than who did the selection, confirming the importance of decentralized selection. Selections made by the breeder and the farmers in 1997, differed in 1998 for a number of traits, but seldom for grain yield. When the difference for grain yield was significant, breeder's selection was more effective on station, while farmers' selection was more effective in farmers' fields. The results of this study indicate that it is possible to organize a plant breeding program with the objective of adapting crops to a multitude of both physical and socio-economic environments: such a breeding program will, at the same time, increase productivity and stability, enhance biodiversity and produce environmentally friendly cultivars. This revised version was published online in July 2006 with corrections to the Cover Date.     

Environment of selection and type of germplasm in barley breeding for low-yielding conditions

Summary Thee groups consisting of 332, 243 and 280 barley breeding lines (entries) of known selection history were evaluated in 10, 9 and 8 environments, respectively, to determine the relationship between grain yield in low yielding (LYE) or high yielding (HYE) environment, and selection history and type of germplasm. One cycle of selection in LYE produced on average five times more entries outyielding the best check in LYE than selection in HYE. A retrospective analysis indicated that the highest yielding lines in LYE were lower yielding (15%–28%) in HYE when compared with the best check, and by 20% and 38% compared with the best entries in HYE. In contrast, the highest yielding lines in HYE were lower yielding (4%–33%) in LYE when compared with the best check, and by 33% and 40% when compared with the best entries in LYE. The highest yielding lines in LYE did not differ consistently from the highest yielding lines in HYE for a number of morphological and developmental traits including days to heading. This suggests there are many paths to high yield in LYE and that analytical breeding based on individual traits may not be appropriate for variable environments. Only 0.07% of the highest yielding entries in LYE was selected for high yield in HYE conditions confirming previous results indicating that selection for high yield in HYE is an inefficient strategy for improving yield in low yielding conditions. This frequency is 28 times lower than the frequency of high yielding entries in LYE selected from landraces or crosses with landraces in low yielding conditions. The results imply that the most cost-effective strategy for barley breeding in low yielding conditions is to select repeatedly in low yielding conditions and to include adapted germplasm (landraces) in the breeding material.     

An alternative approach to breeding for high diastatic power in barley

Summary Several lines from the cross Akka x Feebar were observed to have -amylase activity considerably in excess of either parent. It is suggested that, from crosses between two-rowed and six-rowed varieties, two-rowed genotypes with enhanced levels of grain nitrogen and -amylase activity may be obtained. These can be successfully exploited in a breeding programme to produce barley varities with high diastic power.     

Farmer participation in barley breeding in Syria, Morocco and Tunisia

The paper describes experiments on farmer participation in plant breeding conducted in three countries (Morocco, Syria and Tunisia) on barley, which is the predominant annual rainfed crop in the most marginal areas of these countries. Trials with different types and number of breeding material were planted both on research stations and in farmers' fields. Selection was done by professional breeders and farmers and data were gathered on breeders' and farmers' selection criteria and selection efficiency. The trials reflected the situation of the crop in the three countries, with high yields on station, low yields in some of the most marginal farmers' fields, and poor correlations between research stations and farmers' fields, as well as between farmers' fields. Grain yield was by far the most commonly used selection criterion by the farmers. However, farmers also made a widespread use of selection criteria not normally used by breeders such as grain filling and straw yield, as well as other characteristics of the straw (color) and of the leaves because of the importance of the crop as source of animal feed. A major difference between the selection criteria used by breeders and farmers was disease resistance, almost entirely neglected by the latter. Farmer selection was effective in identifying some of the highest yielding lines in the farmers' own fields and also in those cases where they performed selection on station. The coincidence between entries selected by the breeder and the farmers was high in Morocco but very low in Syria and Tunisia. There were substantial differences between the lines selected by the breeders on station and those selected by farmers in their fields. In Syria, decentralized-participatory selection was significantly more efficient in identifying the highest yielding entries in farmers' fields than any other selection strategy. This work demonstrates that it is possible to organize a plant breeding program so that farmers become major actors in the selection of new cultivars. This revised version was published online in August 2006 with corrections to the Cover Date.     

Simultaneous selection of segregating two-row winter barley material in England and Italy

Summary Independent but simultaneous selection by two breeding teams of the same segregating two-row winter barley material at Cambridge, England and Fiorenzuola, Italy was carried out over the years 1978–1982 with the objective of comparing the individual selection of the two breeding teams and assessing the adaptation of the surviving selections. Visual selection in the generations F₃ to F₅ did not result in any clear cut compartmentalisation of the material and both teams were as likely to select from families previously selected only by the other team as from families they had themselves previously selected. Lines with specific adaptation to each site as well as lines well adapted to both sites were identified from yield trials carried out in F₅ and F₆. The former showed only a very small yield advantage over lines with general adaptation to both sites.     

Selection environment and environmental sensitivity in barley

Summary Groups of 10 barley genotypes were selected for high grain yield under either high yielding (two groups) or low yielding conditions (two groups). The genotypes had a similar average grain yield across a wide range of yielding conditions, but differed in their linear response over environments (environmental sensitivity). The genotypes selected for high grain yield under low yielding conditions were less sensitive to changing environments than genotypes selected for high grain yield under high yielding conditions. The higher stability of genotypes selected under low yielding conditions was shown by both the linear regression analysis and the comparison of coefficients of variation. The use of a safety-first index showed that the probability of a crop failure of genotypes selected for high grain yield under high yielding conditions was between 1.8 and 2.7 times higher than for genotypes selected for high grain yield under low yielding conditions. The development of new cultivars for areas where a large proportion of the crop is grown by subsistence farmers should therefore be based on selection under low yielding conditions.     

Genotypic and environmental influence upon the nutritional composition of barley grain

Summary The grain yield and contents of the quantitatively predominant nutritional constituents of barley grain were determined in nine adapted spring barley varieties each grown at seven European locations with three or four replications.The largest variation in nutritional composition was due to different environmental conditions, but genotypic effects were also present. Interactions between genotype and environment were small. The average protein content at different locations varied from 8.1 to 14.7 per cent of the grain dry matter, and was not simply related to the amount of fertilizer-N applied.The nutritional composition of the grain was influenced by the grain yield level. The percentage of dietary fiber and protein decreased with increasing grain yield, but some varietal differences which were independent of the grain yield level could be established.The protein quality depended upon the protein level, as the protein contained more prolamin relatively to non-prolamin protein at high than at low protein levels. A difference between two varieties in the prolamin/non-prolamin ratio was consistent over a wide range of variation in protein content.     

Adaptation of barley (Hordeum vulgare L.) to harsh Mediterranean environments

Summary Differences in development of the apex may be the reason for cultivar differences in adaptation of barley to terminal drought in Mediterranean environments. The present study was conducted to identify apical development patterns of barley adapted to terminal drought stressed Mediterranean environments and to determine plant characteristics which can be used as criteria to select for an adapted development. Thirty-five two-rowed barley (Hordeum vulgare L.) entries were grown at two sites in northern Syria (Tel Hadya and Breda) in 1988/89. Four apical development patterns were observed: a slow or fast vegetative development, depending on the vernalization requirement, combined with a slow or fast generative development, depending on the daylength response of the crop. Early heading was related to fast generative development. Leaf appearance rates on the main shoot were constant during a major part of the pre-anthesis period, but significant differences were observed among development patterns. Genotypic differences in main shoot tiller number were associated with differences in the onset of tiller appearance and not with differences in tiller appearance rate or final leaf number on the main shoot. Since vernalization requirements and daylength responses are largely independent of terminal drought stress, selection for an adapted phenology can be done in favourable environments. Morphological traits related to these responses (winter growth habit, cold tolerance, plant colour, growth vigour, heading date) can be used as criteria for selection for adaptation to low-rainfall Mediterranean environments.     

Genotype by environment interactions in barley (<Emphasis Type="Italic">Hordeum</Emphasis><Emphasis Type="Italic"> vulgare</Emphasis> L.): different responses of landraces,recombinant inbred lines and varieties to Mediterranean environment

Genotype-by-environment interactions (GEIs) can affect breeding progress because they often complicate the evaluation and selection of superior genotypes. This drawback can be reduced by gaining insights into GEI processes and genotype adaptation. Here, we have studied the GEIs over a set of 24 barley genotypes that were grown across six environments (location-by-year combinations) in Sardinia, Italy. Three groups of genotypes were analysed: barley landraces (LANs), recombinant inbred lines (RILs), and commercial varieties (VARs). The additive main effects and multiplicative interaction (AMMI) model was used for data analysis, and results evidenced no significant differences in grain yield averages for the 24 genotypes. However, there was a relevant GEI for yield mainly between two of the six environments (one characterised by warm pre-anthesis period and high spring rainfalls, and the other characterised by opposite features) and two groups of genotypes (VAR and LAN). Moreover, a negative trade-off between yield levels of genotypes was seen when the barley genotypes were grown in the contrasting environments. Overall, intermediate GEI levels were seen for RILs in comparison to LANs and VARs, and some of the RILs provided valuable yield levels (e.g. RILs 23 and 52). The results thus show the potential usefulness of LANs as a genetic resource for breeding, e.g. to obtain genotypes adapted to Mediterranean environments, such as the RILs analysed in this study. Most of the actual work was carried out when the first author was a PhD student in ‘Agro-meteorology and ecophysiology of agricultural and forest systems’ and she was affiliated to Dipartimento di Scienze Agronomiche e Genetica Vegetale Agraria, Università degli Studi di Sassari, Via E. de Nicola, Sassari 07100, Italy.     

Variation for resistance to Puccinia hordei in Ethiopian barley landraces

Summary Up to 100 single plant derived lines of 18 Ethiopian barley landraces were evaluated for infection type in the seedling and adult plant stage, and for latent period in the adult plant stage only. A low infection type indicates the presence of race-specific resistance genes of the hypersensitive type, while the latent period is the major component of the polygenic, partial resistance.In the seedling stage 1721 of these single plant derived landrace lines were assessed for infection type against two barley leaf rust races. In the adult plant stage 1227 from these 1721 lines were evaluated for infection type against one race. In the seedling stage 2 (against race 1-2-1) and 29 against race A) out of the 1721 lines showed an infection type lower than 6–7 on the 0 to 9 scale. In the adult plant stage none of the 1227 lines had an infection type lower than 6–7 against race 1-2-1.The variation between and within the landraces for latent period in the adult plant stage was large. Some landraces such as landrace 212845 showed a highly significant and longer mean latent period than most other landraces. Virtually all plants in all landraces carry at least some partial resistance.The near-absence of race-specific, major, resistance genes and the high frequency of moderate levels of partial resistance indicates that the durability of leaf rust resistance in Ethiopian barley landraces is due to the latter type of resistance, and that the multiline principle does not operate.     

The use of reversed-phase high performance liquid chromatography as an aid to the identification of European barley cultivars

Summary A reversed-phase high performance liquid chromatography (RP-HPLC) system was used to separate the storage proteins (hordeins) extracted from European barley cultivars. From a total of 38 barleys tested, 26 types of hordein patterns could be distinguished after RP-HPLC. This appears to be a marked improvement in resolution over that achieved in a similar survey of European barley cultivars using SDS polyacrylamide electrophoresis (32 hordein patterns resolved by SDS PAGE from a total of 160 spring and winter barleys tested).Different hordein patterns were resolved by RP-HPLC within each of two groups of barley previously classified by SDS PAGE as indistinguishable within groups (three distinct patterns identified in a total of five cultivars tested from group 1A and five patterns observed among eight cultivars from group 3B). Thus RP-HPLC achieves a higher resolution than undirectional electrophoresis and promises to be a valuable aid in the identification of European barley cultivars.     

Breeding for yield stability in unpredictable environments: single traits,interaction between traits,and architecture of genotypes

Summary Attempts to identify individual traits to use as an indirect measure for grain yield have shown some degree of success in environments where crop yields are affected by no stress or only by predictable stresses. However, analytical breeding has been largely unsuccessful in the case of a) stressful environments characterized by low yields due to high variability in the frequency, timing, duration and severity of a number of climatic stresses, and b) breeding programs where the major objective is greater yield stability defined as a reduction in the frequency of crop failures. Experimental evidence suggests that, when environmental variability is high due to unpredictable differences in frequency, timing and severity of various climatic stresses, each time different combinations of several traits are likely to confer drought resistance. Interaction among traits in determining overall crop response to variable stresses is expected to occasionally enhance the importance of a specific trait in a specific stress situation. In this context it becomes difficult to consider drought resistance as a character with its own identity in terms of inheritance.Assessment and verification of traits based on the use of isogenic lines tends to oversimplify the interactions between traits, as the approach provides information on the effect of a specific trait only in a specific genetic background.At the population level of organisation, the dilemma between selection for individual traits and specific combinations of traits can be translated into the dilemma between selection for individual genotypes and specific combinations of genotypes. The evidence that natural selection under stress conditions has not been able to identify either a single trait or a single genotype with a given adapted architecture of traits, is discussed in relation to germplasm development philosophies to stabilize yield in unpredictably stressed environments.     

Attempts to overcome partial incompatibility between Hordeum vulgare L. and H. bulbosum L.

Summary Partial incompatibility has already been reported between Hordeum vulgare and H. bulbosum during the course of doubled haploid production, and in this paper attempts to overcome the breeding barrier are described. The methods which seem to offer most success are those of environmental adjustments and the adoption of new genotypes of H. bulbosum. Further cultivars of H. vulgare exhibiting this phenomenon are also noted.     

The grain yield of uniculm barley (Hordeum vulgare L.) in two contrasting environments

Summary Four six-rowed uniculm lines of spring barley were grown in two yield trials at different sowing densities. One trial was grown on light sandy soil in 1990. The second trial was grown on clay soil in 1991 and included the two-rowed, tillering cultivar Golf. In the trial on sandy soil, the grain yield of the uniculm lines approached that of Golf grown in an adjacent trial, but in 1991 when the growing conditions were more favourable, Golf yielded significantly more than the uniculm lines. Uniculm lines apparently perform relatively better under marginal growing conditions than in high yielding environments.The uniculm lines do not conform to the ideotype proposed for wheat by Donald (1968) but the results show that a drastic change in plant type need not to imply a large drop in grain yield.     

Comparison of European with West Asian and North African winter barleys in tolerance to boron toxicity

Three plastic-house experiments were conducted to compare the tolerance of European with West Asian and North African (WANA) winter barleys to boron (B) toxicity. Experiment I screened 24 winter barley entries with diverse origins. Experiment III tested 420 random accessions from seven European and seven WANA countries. Plants were screened in a soil mixed with boric acid (50 mg B/kg) and foliar B-toxicity symptom scores were recorded. Lower scores indicated higher B-toxicity tolerance. In Experiment II, five lines/varieties from each of the European and WANA groups were grown in pots with two soil B levels (0 and 25 mg B/kg). The West Asian landrace barleys had a lower mean B-toxicity symptom score than the European ones. The Syrian landrace variety normally grown in drier areas had a lower score than the Syrian landrace variety grown in wetter areas. Dry weights of the European and WANA groups were not different without adding B, but dry weight under 25 mg B/kg was lower for the European group than the WANA group. European accessions had a higher mean B-toxicity symptom score than the WANA accessions. Iranian and Afghan accessions had the lowest mean scores among countries. These results support the hypothesis that European winter barley varieties and accessions are less tolerant to B toxicity than those WANA accessions and varieties developed from local landraces. The lower B-toxicity tolerance could be a factor adversely affecting the performance of European winter barley varieties in the highlands of WANA. This revised version was published online in July 2006 with corrections to the Cover Date.     

Yield stability and adaptation of Nordic barleys

Summary Grain yield was studied in a collection of 220 Nordic barley lines at diverse locations in the Nordic countries. Two-row (2r) and six-row (6r) lines differed very significantly in reaction to the growing conditions within and between the two locations, Svalöv (in southern Sweden) and Højbakkegård (in Denmark). This difference was also highly significant at Viikki (in Finland), but not at As (in Norway) or between Viikki and As. Genotype × location (GL) and genotype × year (GY) variance components were used to estimate phenotypic yield stability by Shukla's stability variance (% MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaaiaacaqabeaadaqaaqaaaOqaaiaabo8adaahaa% WcbeqaaiaabkdaaaGcdaWgaaqcbaCaaiaabMgaaSqabaaaaa!3B73!\[{\text{\sigma }}^{\text{2}} _{\text{i}} \]). Only 7 lines did not contribute significantly to GL- and GY-interactions, and their yield levels were 7–27% lower than that of the highest yielding line (5057 kg/ha). Estimates of GL- and GY-stability parameters were not significantly correlated. Neither responsiveness, measured by the regression coefficient (b _i ), nor phenotypic yield stability, measured by the deviations from regression (Tai's _i ) were correlated with yield. Pedigree studies showed that both b _i and % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaaiaacaqabeaadaqaaqaaaOqaaiaabo8adaahaa% WcbeqaaiaabkdaaaGcdaWgaaqcbaCaaiaabMgaaSqabaaaaa!3B73!\[{\text{\sigma }}^{\text{2}} _{\text{i}} \] can be changed by recombination and/or induced mutations. Mixing of near isogenic lines with different resistance genes, and selection within a landrace, also resulted in changes in responsiveness. Recently released 2r-cultivars were more unstable than older 2r-cultivars revealed by positive correlation between the year of release and _i . Cultivars originating from southern Scandinavia were higher yielding than cultivars originating from the central or the northern regions of Scandinavia.     

Resistance to powdery mildew in selections from Moroccan barley landraces

Seventy-five barley landraces from Morocco were tested for resistance to powdery mildew and a number of different resistance genes were detected. Thirty-five isolates of Blumeria graminis f. sp. hordei and the Pallas isoline differential set were used. Isolates used in the experiment had virulences corresponding to all major resistance genes used in Europe. Forty-four of the tested landraces showed resistant reactions. From each of these landraces, one to five resistant plants were selected and 92 single plant lines were created. Six lines selected from 3landraces were assumed to carry the mlo gene but they were discarded after microscopic investigation. Seventeen lines were tested in the seedling stage with 17isolates and another 69 lines were tested with 23 differential isolates. These lines showed 71 reaction spectra to isolates of powdery mildew. Eight lines (9%), 255-3-3, 282-3-4, 286-1-1, 294-2-3,294-2-4, 295-1-2, 308-1-2 and 327-2-1, selected from 7 landraces showed resistance to all isolates. Seventy-eight lines (90%) showed a resistant infection type 2with more than 50% of the isolates used. In most of the selected lines (86%) unknown genes, alone or in combination with known specific resistance genes, were detected. Four different resistance alleles (Mlat, Mla6, Mla14 and Mla1) were postulated to be present in the tested lines. The most common was Mlat, which was postulated in 35 (41%) lines. The use of newly identified sources of powdery mildew resistance in barley breeding is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.