CIMMYT's approach to breeding for wide adaptation

Summary The wheat area in developing countries, including China, is around 100 million ha. To address the needs of these very diverse wheat growing areas, CIMMYT has defined 12 wheat mega-environments (ME). A ME is defined as broad, not necessarily continuous often transcontinental area with similar biotic and abiotic stresses, cropping systems and consumer preferences. The factors describing each ME are presented.CIMMYT's breeding methodology is centered around the development of widely adapted germplasm with high and stable yield across a wide range of environments. Segregating populations are alternating screened in two diverse environments in Mexico. One key requirement is that all germplasm is tested under near optimum conditions for its yield potential. The second one is multi-locational testing of advanced lines at sites that represent a given ME (key locations) and careful screening of germplasm for tolerance to abiotic and biotic stresses specific to that environment. This methodology has permitted the pyramiding of a large number of multiple resistance genes for use against a wide spectrum of diseases and tolerance to abiotic stresses within each ME. In addition, the widespread testing of lines allows the identification of traits which are beneficial in several environments. Data from international nurseries are used to further delineate environments within an ME. This approach has proven to be successful since around 70% of the spring wheat area in developing countries (excluding China) is planted to varieties derived directly or indirectly from CIMMYT germplasm. The performance of the bread wheat cultivar Pastor in international trials is given as an example for a wide adaptation.     

An evaluation of seed and seedling drought tolerance screening tests in wheat

Summary A series of experiments was performed in order to evaluate the significance of seed germination and seedling growth in osmotic media as screening methods for drought tolerance.Ten spring wheat (Triticum aestivum L. em Thell.) and one durum wheat (Triticum durum Desf.) were tested under controlled environments, using polyethylene glycol-6000 (PEG) solutions as the moisture stress inducing media.Tolerance in the rate of endosperm utilization, under stress, prior to the onset of germination varied among cultivars.Germination rate or injury to germination at various concentrations of PEG differed significantly among cultivars. Cultivar rating with respect to injury to germination changed with stress levels. Injury to germination did not correlate with endosperm utilization rate in PEG or in water.Germinating seedlings were tolerant to extreme desiccation up to the stage of emergence of the first leaf from the coleoptile.Growth of photosynthesizing seedlings was monitored as they were carried through an increasing concentration gradient of PEG solutions, ranging from –5.9 to –11.3 bars of water potential. Cultivars significantly differed in seedling growth tolerance to increasing levels of water stress. Seedling growth tolerance across cultivars was not correlated with their germination responses under srress.It is concluded that tolerance to water stress in growing seedlings can be screened for by using PEG-containing nutrient solutions. It can not be predicted from germination tests in osmotica.Work was done under a US-Israel Binational Science Foundation (BSF) Grant no. 1654/78.Contribution from Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel, No. 192-E, 1979 series.     

Assessment of genetic diversity in bread wheat genotypes for tolerance to drought using AFLPs and agronomic traits

Moisture stress greatly limits the productivity of wheat in many wheat-growing regions of the world. Knowledge of the degree of genetic diversity among parental materials for key selection traits will facilitate the development of high yielding, stress tolerant wheat cultivars. The objectives of this study were to: (i) use amplified fragment length polymorphisms (AFLPs) to assess genetic diversity among bread wheat lines and cultivars with different responses to drought stress in two distinct environments and, (ii) compare genetic diversity estimated by AFLPs with diversity evaluated on agronomic performance under drought stress. Twenty-eight genotypes, 14 from Iran and 14 developed or obtained by CIMMYT, were evaluated in the study. Phenotypic data on the 14 Iranian lines were obtained in Iran, and data on the 14 CIMMYT lines were collected in Mexico. Ten AFLP primer pairs detected 335 polymorphic bands among the 28 cultivars. At the 5th fusion level of the resulting dendrogram, 6 genotype clusters were identified. Thirteen of the 14 CIMMYT genotypes grouped into one cluster while 4 of the remaining groups were comprised only of Iranian genotypes. When the agronomic performance of the Iranian materials was compared with the AFLP diversity analysis, 5 of the 6 drought susceptible genotypes clustered together in the agronomic dendrogram, and were located in the same cluster in the AFLP dendrogram. However, the drought tolerant Iranian materials did not show the same degree of relationship. The CIMMYT materials did not demonstrate a significant association between agronomic performance and genetic diversity determined using AFLPs. Clearly these data show that there are genotypes with similar agronomic performance and different genetic constitutions in this study that can be combined in a breeding program to potentially improve tolerance to drought stress.     

Improved drought tolerance and root development as components of a scheme to breed blueberries for mineral soil adaptability

Blueberry breeding in North America started at the beginning of the 20th century and present cultivars are adapted to irrigated, high organic, acidic, sandy soils. However, sites meeting these requirements are limited, which has forced breeders to develop germplasm that is adapted to unamended mineral soils. Wild Vaccinium species can be found growing on a number of different soil types under a variety of climatic conditions, which indicates that genetic variability for mineral soil adaptability is present in Vaccinium germplasm. However, for a number of reasons progress in the development of mineral soil-adapted blueberry cultivars has been slow. In this paper a breeding scheme is presented that would allow the user to more efficiently access the variability that exists for mineral soil adaptability. The scheme is composed of two different seedling screening techniques (drought and seedling rooting screen) that are designed to eliminate the less vigorous individuals in a population before field testing. The field environment would be used primarily to evaluate individuals for fruit quality and yield characteristics. In blueberries, both general and specific combining ability can be fully exploited because they are vegetatively propagated which immediately fixes any unique combination of genes.     

Honeycomb selection for yield in early generations of spring wheat

Summary The effectiveness of the honeycomb selection method for yield in spring wheat (Triticum aestivum L.) was evaluated using progenies from two wheat crosses, Glenlea x NB131 and Glenlea x Era. Honeycomb selection was carried out in the F₂ and F₃ generations, grown at the University of Manitoba in the summers of 1980 and 1981, respectively. In both generations, divergent selection was made for both high and low yield. Plants selected in the F₃ generation were entered in an F₄ yield test in the summer of 1982. Results of the experiment showed that honeycomb selection for yield in the F₂ and F₃ generations was effective in identifying parents of high- and low-yielding lines. F₃ plants from highyielding F₂ selections gave higher yields than those from low-yielding F₂ selections by 11.5% and 13.0% for Glenlea x NB131 and Glenlea x Era crosses, respectively. The F₄ yield test showed that high yielding selections from both crosses significantly outyielded by 8.9% low yielding selections and by 14.4% the unselected composite lines. It is concluded that the honeycomb selection method can be used for early generation selection in spring wheat.     

Haploidy breeding and mutagenesis for drought tolerance in wheat

Several intraspecific crosses between known drought tolerant wheat varieties and stable high yielding recombinants were made with the objective to develop improved cultivars for the moisture stressed rainfed areas of Pakistan. Five of these crosses were selected for further creation of useful mutations through the application of low doses of gamma rays and development of doubled haploids through anther culture. Anther culture response of the selected irradiated F₁ generations was studied on liquid and solid induction media. The highest number of calli among almost all crosses was produced on Potato-2induction medium. All the crosses varied greatly in response to callus induction and maximum calli (75%) were obtained from Lyl-73/vee’s’ cross. Similarly, genotypic differences were found for green vs. albino regenerants. The highest number of green plantlets (12.1%) was recorded for Lu-26/3062. From the developed doubled haploid population 25 DH-mutants were initially selected and nine lines were finally included in multi-locational field tests. Two DH-mutants (i.e. DHML-50 and DHML-9) have potential for better grain yield, earliness, disease resistance and moisture stress tolerance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Major gene control of tolerance of bread wheat (Triticum aestivum L.) to high concentrations of soil boron

Summary The genetic control of tolerance of wheat to high concentrations of soil boron was studied for five genotypes. Each genotype represented one of five categories of response to high levels of boron, ranging from very sensitive to tolerant. Tolerance to boron was expressed as a partially dominant character, although the response of an F₁ hybrid, relative to the parents, varied with the level of boron applied. The F₁ hybrids responded similarly to the more tolerant parent at low B treatments and intermediate to the parents at higher treatments. Ratios consistent with monogenic segregation were observed for the F₂ and F₃ generations for the combinations (WI^*MMC) × Kenya Farmer, Warigal × (WI^*MMC) and Halberd × Warigal. The three genes, Bo1, Bo2 and Bo3, while transgressive segregation between two tolerant genotypes, G61450 and Halberd, suggested a fourth locus controlling tolerance to boron.     

Breeding for resistance to cereal cyst nematode in wheat

Summary The progress of a backcross breeding programme to introduce resistance against the cereal cyst nematode into wheat is described. Methods of resistance screening and criteria for selection are detailed and the results discussed with reference to alternative procedures for the introduction of new resistance genes into major breeding programmes.     

Flooding tolerance of spelt (Triticum spelta L.) compared to wheat (Triticum aestivum L.) – A physiological and genetic approach

In marginal, agroclimatic zones, yield is often affected by flooding, but the effect is much less for winter spelt (Triticum spelta L.) than for winter wheat (Triticum aestivum L.). This study evaluates the reaction of a wheat x spelt population (F₅ RILs of Forno x Oberkulmer) to flooding stress in the early phase of germination. Lines with greater tolerance to 48 h flooding just after imbibition showed less electrolyte leakage (r = -0.79) indicating greater membrane integrity and better survival. Five QTL explaining 40.6% of the phenotypic variance for survival to flooding were found, and localized on the chromosomes 2B, 3B,5A, and 7S. The tolerance to 48 h flooding four days after sowing was best correlated with the mean germination time (r = 0.8), indicating that the plants with a fast coleoptile growth during flooding are less susceptible to flooding. Ten QTL were found for seedling growth index after flooding explaining 35.5% of the phenotypic variance. They were localized on chromosomes 2A, 2B, 2D, 3A, 4B, 5A, 5B, 6A, and 7S. Standard varieties of spelt and wheat showed the same tolerance characteristics. The possibility to use marker assisted selection for flooding tolerance is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

The drought response of landraces of wheat from the northern Negev Desert in Israel

Summary Diverse landraces of wheat, collected from the semi-arid (150 to 250 mm of total annual rainfall) Northern Negev desert in Israel were considered as a potential genetic resource of drought resistance for wheat breeding. These materials were therefore evaluated for their reponses to drought stress in agronomical and physiological terms. Up to 68 landraces, comprising of Triticum durum, T. aestivum, and T. compactum were tested in two field drought environments, in one favourable field environment, under post-anthesis chemical plant desiccation which revealed the capacity for grain filling from mobilized stem reserves, under a controlled drought stress in a rainout shelter and in the growth chamber under polyethylene glycol (PEG)-induced water stress. Biomass, grain yield and its components, harvest index, plant phenology, canopy temperatures, kernel weight loss by chemical plant desiccation, growth reduction by PEG-induced drought stress and osmotic adjustment were evaluated in the various experiments.Landraces varied significantly for all parameters of drought response as measured in the different experiments, which was in accordance to their documented large morphological diversity. Variation in grain yield among landraces under an increasing drought stress after tillering was largely affected by spike number per unit area. Kernel weight contributed very little to yield variation among landraces under stress, probably because these tall (average of 131 cm) landraces generally excelled in their capacity to support kernel growth by stem reserve mobilization under stress. Yield under stress was reduced with a longer growth duration of landraces only under early planting but not under late planting. Landraces were generally late flowering but they were still considered well adapted phenologically to their native region where they were always planted late.Landraces differed significantly in canopy temperature under drought stress. Canopy temperature under stress in the rainout shelter was negatively correlated across landraces with grain yield (r=0.67^**) and biomass (r=0.64^**) under stress. Canopy temperature under stress in the rainout shelter was also positively correlated across landraces (r=0.50^**) with canopy temperature in one stress field environment. Osmotic adjustment in PEG-stressed plants was negatively correlated (r=–0.60^**) with percent growth reduction by PEG-induced water stress. It was not correlated with yield under stress in any of the experiments. In terms of yield under stress, canopy temperatures and stem reserve utilization for grain filling, the most drought resistant landrace was the Juljuli population of T.durum.     

Tolerance to paraquat is correlated with the traits associated with water stress tolerance in segregating F2 populations of barley and wheat

To identify scorable marker traits that can be used in cereal breeding programs for selecting drought tolerant individuals, we investigated the correlation among the drought-associated traits in two F₂ populations derived from the crosses made between drought tolerant and sensitive barley and wheat parental genotypes. The parental genotypes of these crosses also differed by at least three other traits – paraquat tolerance, leaf size, and the relative water content. These three traits were scored in two F₂populations of 80 individuals for each barley and wheat cross. Analysis of results indicated that the enhanced tolerance to paraquat was correlated with reduced leaf size and increased relative water content, two traits associated with water stress phenotypes of the drought tolerant barley and wheat parents. Our results suggested that the selection based on paraquat tolerance istechnically less demanding and thus useful for rapid screening of individuals for enhanced drought tolerance in segregating populations. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genotypic differences in wheat Al tolerance

Summary Aluminium tolerance of 83 genotypes from Croatian and Yugoslav Triticum aestivum germplasm was evaluated in nutrient solutions having Al³⁺ activities of 0, 12.5 and 25 M. Relative root length (25 M Al³⁺/0 Al) of various genotypes ranged from 2 to 97% (from very sensitive to tolerant to Al). No genotype with Al tolerance close to that of very tolerant cultivar Atlas-66 was found. Soil, climatic, fertilization, and liming effects that wheat plants giving seeds for the nutrient solution Al-tolerance screening had been subjected to during their growth cycle did not influence the Al-tolerance ranking. Significant correlation was found between screening wheat for Al tolerance in nutrient solutions and in acid Pseudogley soil amended with five rates of limestone in a greenhouse experiment. Seed protein concentration was significantly related to the Al-tolerance ranking (r² = 0.962). Such a significant correlation was not obtained in a case of rheological and other quality characteristics of seeds. Al-tolerant wheat genotypes identified in this study will be used in breeding for improved Al tolerance.Abbreviations HSD Tukey's Honestly Significant Difference - RRL-2 relative root length, in % (12.5 M Al³⁺/0 Al) - RRL-4 relative root length, in % (25 M Al³/0 Al)     

Heat tolerance for yield and its components in different wheat cultivars

Summary Twenty one diverse, standard and experimental cultivars of common spring wheat (Triticum aestivum L.) were tested for the effect of heat stress on phenology, yield and its components by growing the materials for 2 years under full irrigation during the hot summer (offseason), and the cool winter (normal) conditions. Heat tolerance was estimated for each variable by the heat susceptibility index (S) which scales the reduction in cultivar performance from cool to hot conditions relative to the respective mean reduction over all cultivars.Genotypes differed significantly in S for yield and its components. The ranking of cultivars in S over the 2 years was consistent for yield, kernels per spike and kernel weight, but not for spike number. Of the three yield components, the greatest genotypic variation in S was expressed for kernels per spike. However, S for yield could not be simply attributed to S in a unique component across all cultivars. On the other hand, a general linear model regression of summer yield on its components revealed that the most important yield component affecting yield variation among cultivars under heat stress was kernel number per spike. Kernel number per spike was positively associated across cultivars with longer duration and greater stabilty of thermal time requirement from emergence to double ridge. It is therefore concluded that kernel number per spike under heat stress is a reasonable estimate of heat tolerance in yield of wheat and that this tolerance is operative already during the first 2 to 3 weeks of growth.     

Breeding for resistance to cereal cyst nematode in wheat II. Use of test tube cultures in selection

Summary The use of soil. naturally infested with Heterodera avenae, to select resistant heterozygotes in backcross progenies of wheat, was tested for reliability. Selfed progenies from plants selected as resistant were cultured monoxenically in test tubes with nematodes hatched from single cysts, while backcross progenies from the same parent plants were grown in pots of naturally infested soil. Cyst counts were made after two months' growth. The results showed that over 50% of the backcross lines, screened in previous generations with naturally infested soil, had been erroneously selected as resistant. The test tube cultures clearly differentiated lines carrying resistance from those which were susceptible and corroborated results from pot tests.     

Genetic changes in resistance to environmental stresses by U.S. Great Plains wheat cultivars

Increased grain yield potential of newcultivars of wheat (Triticum aestivumL.) is attributed to morphological traits,but actual yield is determined mostly byenvironmental conditions. Our objectivewas to ascertain the contribution ofresistance to freezing, high temperature,drought, and defoliation to advances inyield of landmark cultivars of winter wheatin the U.S. Great Plains. Eight cultivarsthat represented significant improvementbetween 1874 and 1994 were compared bystandard electroconductivity measurementsof stability of seedling cell membranes tofreezing, high temperature, and desiccationand by grain yield and its components inplants subjected to freezing during theseedling stage and to high temperature,drought, and defoliation during maturation. Genetic changes relative to `Turkey'(introduced 1874) in stability of cellmembranes to freezing, high temperature,and desiccation were small andinconsistent. Advances in grain yieldunder control conditions were similar togains in field studies. Most cultivars hadlittle genetic change in yield afterfreezing, drought, or defoliation, probablybecause high levels of resistance areincompatible with high yield potential andthe stresses are episodic. Genetic advancein grain yield under high temperature wasphenotypically correlated with change inyield under control conditions, suggestingthat the trait is essential forproductivity because of the ubiquitousoccurrence of the stress in the region. Weconcluded that changes in resistances tofreezing, drought, and defoliationcontributed little to advances inproductivity of winter wheat in the GreatPlains, but that resistance to hightemperature was important for new cultivars.     

Molecular mapping of an aluminum tolerance locus on chromosome 4D of Chinese Spring wheat

Summary The tolerance of aluminum (Al) of disomic substitution lines having the chromosomes of the D genome of Triticum aestivum L. cv. Chinese Spring individually substituted for their homoeologues in T. turgidum L. cv. Langdon was investigated by the hematoxylin method. The disomic substitution lines involving chromosome 4D were more Al tolerant than Langdon. The tolerance was found to be controlled by a single dominant gene, designated Alt2, that is in the proximal region of the long arm of chromosome 4D. The locus was mapped relative to molecular markers utilizing a population of recombinant chromosomes from homoeologous recombination between Chinese Spring chromosome 4D and T. turgidum chromosome 4B. Comparison of the location of Alt2 in this map with a consensus map of chromosomes 4B and 4D based on homologous recombination indicated that Alt2 is in a vicinity of a 4 cM interval delineated by markers Xpsr914 and Xpsr1051. The Alt2 locus is distal to marker Xpsr39 and proximal to XksuC2. The Altw locus is also proximal to the Knal locus on chromosome 4D that controls K⁺/Na⁺ selectivity and salt tolerance. In two lines, Alt 2 and Knal were transferred on a single 4D segment into the long arm of T. turgidum chromosome 4B.     

Polyacrylamide gel electrophoresis of wheat gliadins: the use of a moving boundary for improved resolution

Summary The process of the separation of gliadin proteins of wheat, using polyacrylamide gel electrophoresis, was studied in detail. It was shown that electrophoresis is a dynamic process during which the pH of the gel changes together with the potassium ion concentration. The most positive effect on the separation of the gliadins was generated by a moving front, which is a boundary between regions with a low concentration of K⁺ ions and a low pH, and region with a high concentration of K⁺ ions and a high pH, after optimization of the concentrations of the cations and anions in the electrode solutions. The finding was exploited for the development of an extremely simple electrophoresis system, in which buffers were not needed for obtaining a high resolution. The system was further improved by applying a stacking gel. The advantages of this system are discussed. This new approach may be useful for improving electrophoresis systems for other applications.     

Endosperm peroxidase electrophoresis patterns to distinguish tetraploid from hexaploid wheats

Summary A simple method is proposed to distinguish hexaploid (Triticum aestivum L.) from tetraploid (Triticum turgidum L., durum wheat) cultivated wheats on the basis of peroxidase isozymes coded by genome D. It can also be used as a first step to detect possible contamination by tetraploid genotype mixtures. The peroxidase patterns of endosperm and of embryo plus scutellum found among 349 entries of a durum wheat world basis collection are shown.     

Response to selection for grain yield and harvest index in F2, F3 and F4 derived lines of two wheat crosses

Summary Aspects of selection for yield and harvest index were investigated by simulating selection using data from random pedigreed F₂, F₃, F₄ and F₅ derived lines from two crosses grown in plots at two sites over two years.Improvement in yield through selection was obtained when the response was measured at the same site and in the same year as the selection. Selecting the best 10 per cent of F₂ to F₄ derived lines gave F₅ derived lines that outyielded random selections by 19 to 53 per cent for one cross and 5 to 23 per cent for the second cross. These lines were 41 to 50 per cent better than the mid-parent in one cross, but were less than the mid-parent in the other cross.However, the response to selection when measured in a different year was little better than random selection. The effect of different sites also reduced the effectiveness of selection.Selection of harvest index in early generations for improvement of yield was ineffective when response was measured at the same site in the same year, or in different years.Contrary to some theoretical proposals, the same improvement in yield was obtained by selecting in early or late generations. While high yielding genotypes may be lost by delaying selection, this is counteracted by the better predictive value of late generations due to their greater homozygosity and homogeneity.     

Transmission of somaclonal variation in wheat

Summary Transmission of somaclonal variation was studied in callus derived (SC1) plants of wheat, Triticum aestivum L. Second (SC2), third (SC3) and fourth (SC4) generations were compared with those obtained from embryo culture (E1-E4). SC2 generation had significantly lower grain yield (–37.3%), reduced thousand kernel weight (–7.8%) and shorter height (–2.1%) than that of E2, and included 5.7% seed-sterile and 15.0% partially seed-sterile plants. SC3 generation showed reduced yield (–12.6%), lower thousand kernel weight (–4.9%) and shorter plant height (–1.9%) than E3 generation. SC4 progeny of a short height, partially sterile SC1 variant included aberrant plants: a haploid, few aneuploids, mixoploids, some showing multiploidy, spindle abnormalities, multivalents, bridges and fragments, and tillers having supernumerary spikes with branched rachis. Somaclonal variation resulted from genome instability in callus, and likely involved transposable elements.