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Showing 18 results for Heritability
Pariya Dorri, Saeed Khavari Khorasani, Mahmood Vali Zadeh, Parisa Taheri,
Volume 1, Issue 2 (2-2015)
Abstract
Selection of breeding methods for plant genetic improvement largely depends on the genetic basis of selected traits. This research was carried out in order to study the genetic parameters of phenological and morphological traits , yield and yield components of maize using generation mean analysis of inbred lines derived from crosses KE72012 (P1) × K1263 / 1 (P2) inbred lines. This study was conducted at Torough Station of Khorasan Razavi Agriculture and Natural Resources Research Center, Mashhad, Iran, during 2012. Six maize generations include BC2, BC1, F2, F1, P2 and P1 was evaluated in a randomized complete block design (RCBD) with three replications. Analysis of variance showed higher dominance gene effects than additive effects. Average dominance gene effects (H / D) ½ in all measured traits, showed complete and over dominance gene effects indicating the importance of dominance effect in studied traits. A simple additive-dominance model performed using weighted least square analysis. Results showed not it’s adequacy for traits, indicating the importance of epistatic effects in controlling genetic variations. Broad sense heritability for all measured traits in crosses KE72012 (P1) × KE1263 / 1 (P2) ranged between 0.10 to 0.88 and the range of narrow sense heritability in this cross was between 0.02 to 0.59. Estimation of number of genes controlling grain yield showed that the one to five genes are involved in KE72012 (P1) × K1263 / 1 (P2).
Mahdi Kakaei, Hojatollah Mazahery-Laghab,
Volume 2, Issue 1 (5-2015)
Abstract
In order to identify the correlation between different agronomical traits and the study of their relation using regression analysis, 46 alfalfa populations were studied in the research farm at Bu-Ali Sina University in 2013. Analysis of variance for different traits revealed a significant difference between populations for dry forage yield at the level of 1% probability. The genotypic ccorrelation coefficients of fresh forage yield with the amounts of leaf chlorophyll (SPAD) (0.952**), plant height at 10% flowering stage with fresh forage (0.987**), dry matter yield with fresh forage yield (-0.942**), dry forage yield with plant height at 10% flowering stage (0.676*) and dry matter percent (0.896*) showed significant correlations. Phenotypic correlation analysis showed a positive and significant correlation between damage percent and the number of larvae (0.767**), a negative and significant correlation between dry material percent and fresh forage yield (-0.450**). However, dry matter percent had a negative and significant correlation with dry forage yield (0.424**). The stepwise regression for fresh forage yield as dependent variable showed that dry forage yield, dry matter percent, plant height in 10% flowering stage and number of larvae were respectively entered to the model and with 89.29% of cumulative contrast coefficient confirmed the most variations of fresh forage yield. Furthermore, the maximum and minimum heritability percent was depended to the number of Larvae and dry forage yield respectively. Naragamet population (No. 30) with a high yield of dry forage and the number of larvae and also a relatively low level of damage percent among other populations could be suggested as a desirable and tolerant population
Reza Nikooseresh, Goodarz Najafian,
Volume 3, Issue 1 (9-2016)
Abstract
In order to study genetic diversity, heritability and relation of some important traits with grain yield in bread wheat, 20 bread wheat irrigated lines evaluated using a randomized complete block design with three replications during two cropping cycles/years. Based on combined analysis results, there were a significant difference between genotypes. Also, in all traits, coefficient of phenotypic variation was greater than coefficient of genetical variation, indicating the effect of environment on investigated traits. The significant correlation between grain yield and number of days to maturity, spikes per m2 and the grains per spike. number of days to heading and number of days to maturity had highest heritability among traits, with 87 and 69%, respectively. Results of this showed that selection of early heading or shorter no. of days to heading and early maturing or shorter number of days to maturity is recommended to breeders for selection materials favorable for terminal warm condition and water shortage, preventing grain yield reduction. Results of cluster analysis showed that genotypes are divided in six groups. Totally, six superior genotypes from the fourth cluster were selected as superior lines. Based on bi-plot analysis, genotypes 14 and 5 showed the lowest genotype × trait interaction, but genotypes 16 and 8 showed the highest genotype × trait interaction.
Bafrin Molaei, Mohammad Moghaddam, Seyed Siamak Alvaikia, Ali Bandeh-Hagh,
Volume 3, Issue 2 (3-2017)
Abstract
This experiment was conducted during 2013-2014 in Tabriz University research farm, Iran. In this research, the inheritance of some agronomic and physiological traits in different generations of Bam (tolerant to drought) and Artha (sensitive to drought) cross was studied in field conditions by generation mean analysis. The experiment was a split plot design using randomized complete blocks with two replications with the irrigation conditions in the main plots and generations in the sub-plots. The interaction between generations and irrigation conditions was significant only for grain yield. Based on the result of generation means analysis for the spike length and straw weight, the threeparameter model was found fitting the analysis. For the thousand-seed weight in the water stress condition and for the plant weight in the irrigation condition, a non-significant chi-square suggesting that the six-parameter model for these traits is suitable. For the other traits, the chi-square was significant in both conditions. For spike length and leaf area, the degree of dominance was greater than one (2.3 and 1.53, respectively) which showed the existence of over-dominance gene action in controlling these traits. Straw weight in normal condition, leaf area and thousand seed weight in the stress condition had a high broad-sense heritability. The narrow-sense heritability for all other traits was low, suggesting the need for exploiting dominance gene action in the breeding programs if hybrid varieties are produced in the bread wheat.
Reza Mir Drikvand,
Volume 3, Issue 2 (3-2017)
Abstract
Identification and application of genetic diversity are essential to breeding programs success. In this study, genetic diversity of 20 rainfed barley genotypes were assessed using morphological traits as well RAPD and intron-exon splice junction (ISJ), semi-random markers. Results of this study showed that there were significant differences among genotypes for all traits, indicating high genetic variation among them. The highest and lowest broad sense heritability was related to spike length and grain yield, respectively The estimates of genotypic coefficient of variation (GCV) and phenotypic coefficient of variation (PCV) were high for number of grain per spike, and low for 1000-kernel weight, respectively. Mean of polymorphic percentage in ISJ marker was higher than RAPD marker. Cluster analysis showed that the distinctions based on morphological traits did not correspond with the distinction based on molecular data.The results showed that RAPD and ISJ markers were able to distinct two and six-rowed and also hulless and hulled barley genotypes. Distinction of three clusters did not follow the same pattern.There was significant and negative correlation between similarity matrices of molecular data and morphological traits, but similarity matrices of two molecular markers was significantly and positively correlated.
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Volume 4, Issue 1 (9-2017)
Abstract
Drought is a worldwide risk seriously influencing production of agricultural plants such as coriander. In order to evaluate heritability and gene action for important quantitative traits in coriander in three irrigation regimes, including well water (WW), mild water stress (MWS) and severe water stress (SWS), a cross between two genotypes of TN-59-158 (Sensitive parent) and TN-59-230 (Tolerant parent) was done. Parents and progenies of triple test cross were evaluated in filed condition through a randomized complete block design with three replications. Days to 50% flowering (DTF), days to harvest (DTH), fruit yield (FY), thousand fruit weight (TFW), harvest index (HI), relative water content (RWC), essential oil yield (EOY) were measured. Results showed that total epistasis effect was observed for all traits except RWC in WW. Also, total epistasis effect was observed for all traits except DTF in MWS and SWS. Partitioning of total epistasis revealed that [i] type of epistasis was contributed to determining DTF and TFW in WW and MWS treatments, DTH, FY and RWC in MWS and SWS treatments, and HI and EOY in WW, MWS and SWS treatments.
Hossein Astaraki, Peyman Sharifi, Fatemeh Sheikh,
Volume 6, Issue 2 (3-2020)
Abstract
In order to estimation genotypic correlation and heritability of some faba bean traits, 26 faba bean genotypes were evaluated in a randomized complete block design with three replications during 2014-16 growing seasons in Agricultural Research Sation of Borujerd located in Lorestan province, Iran. The restricted maximum likelihood (REML) was used to estimate the genotypic and phenotypic correlations, broad sence heritability and genetic gain. Analysis of variance based on least squares and REML indicated significant effect of genotype on days to maturity, plant height, hundred seed weight, pod length and dry seed yield. Genotype ×year interactions were significant on all of the traits except of pod length. Borujerd cultivar (G26) and G20 had the highest dry seed yield in both of years. REML results indicated a significant positive genetic correlation between dry seed yield and biological yield and plant height. Also, there were a significant negative phenotypic correlation between dry seed yield and days to maturity, and significant positive phenotypic correlation between dry seed yield and plant height, biological yield and harvest index. So, the selection of early maturing genotypes with a higher yield is achievable and selection can be done to improve the performance of dry seed yield. Cluster analysis indicated variability among genotypes. According to values of broad sence heritability and genetic gain for plant height and high genetic correlation of this trait and dry seed yield; plant height can be used as a suitable trait for improving dry seed yield by selection.
Farshad Fallah, Danial Kahrizi, Abbas Rezaeizad, Alireza Zebarzadi, Lila Zarei,
Volume 6, Issue 2 (3-2020)
Abstract
After cereals, oilseeds are the second-largest food reserves in the world. According to available statistics, more than 95 percent of Iran's oil needs are imported. Given the growing need for edible oils in Iran, it is important to identify fatty acids in the oilseed crops. Camelina sativa L. is an oil-medicinal plant and belongs to the Brassicaceae family that requires very little water and fertilizers. It is known as a low input plant. In this study, to analyze the fatty acid profile for breeding programs and specific industries, 137 doubled haploid camelina lines were evaluated in terms of fatty acid composition and variability of fatty acids trait, to estimate phenotypic coefficient of variation (PCV), genotypic coefficient of variation (GCV), heritability, and expected genetic advance. The determination of fatty acid by gas chromatography showed that 18 types of fatty acids were detectable in camelina seed oil. It is shown that the two fatty acids (C14:0 and C16:1) have the highest PCV and GCV. The highest heritability for C20:2, C20:3 and C20:0 fatty acids was estimated 98.92, 98.59 and 96.49 percent, respectively. In this study, two lines with linoleic acid of 35.81-36.67% and four lines with values ranged from 22.08-23.00% were introduced. The ratio of omega-6 to omega-3 (0.479-0.759) was obtained in the studied lines.
Saman Valizadeh, Ahmad Ismaili, Hadi Ahmadi, Omid Ali Akbarpour, Bijan Bajalan, Ashkboos Amini,
Volume 6, Issue 2 (3-2020)
Abstract
Wheat is mostly cultivated at rainfed condition in Iran, so, water deficit stress has much effect on yield reduction. Hence, breeding activities are necessary for introduction of wheat tolerant genotypes to water deficit stress. In order to estimate the heritability and genetic correlation between traits of 36 wheat genotypes, an experiment was conducted in two separate conditions (water stress and non-stress) based on a randomized complete blocks design with three replications. Studied traits in wheat genotypes under water stress and normal condition showed significant differences for environment, genotype and genotype× environment interaction at 1 and 5% level of probability. The results of the factor analysis showed that the 6 first factor in normal condition explained 81.13% of total variance, and the 5 first factor in stress condition explained 74.96% of total variance. Estimation of genetic correlations based on REML approach revealed that biological yield, harvest index and number of grains per spike had the highest correlation with grain yield and these characteristics are of important for selecting the varieties with high yield under non-stress and stress conditions. Estimation of heritability based on REML approach showed that number of days to heading had the highest amount of heritability in both normal and stress conditions.
Maryam Tahmasbali, Reza Darvishzadeh, Amir Fayaz Moghaddam,
Volume 7, Issue 1 (9-2020)
Abstract
In a breeding program, it is important to find out information about the genes action, because knowledge in this field could help the researchers in their crossing programs and realizing effective selection. In this study, breeding values of different agronomic traits in oriental tobacco were predicted using the best linear unbiased prediction (BLUP) procedure. For this purpose, 89 tobacco genotypes were evaluated in a randomized complete block design with three replications under normal (without broomrape) and stress (with broomrape) conditions at Urmia Tobacco Research Centre, during two successive years. Broomrape stress was applied by mixing 0.06 gr broomrape seed with soil in pots. C.H.T.209.12e × F.K.40-1 genotype had high yield under both normal and broomrape stress conditions and was one of the desirable genotypes in terms of yield tolerance and stability index. The Rustica genotype was the best genotype in terms of the breeding value of most of studied traits in both normal and broomrape stress conditions. The result from cluster analysis based on the breeding values of the studied traits showed that, tobacco genotypes were divided into 6 and 5 groups in normal and broomrape stress conditions, respectively; but the distribution of genotypes within the groups was different depending on the conditions. The highest heritability was observed for root fresh weight under normal condition and for leaf fresh weight under broomrape stress conditionss. The results showed that a genotype with good phenotypic performance may have low breeding value. Therefore, considering breeding value information along with phenotypic mean of traits can increase the efficiency of breeding programs.
Jamal Rahimi Darabad, Varahram Rashidi, Hossein Shahbazi, Mohammad Moghaddam Vahed, Ebrahim Khalilvand,
Volume 7, Issue 2 (3-2021)
Abstract
In order to determine the heritability and genetic parameters of some agronomic traits in barely (Hordeum vulgare L.) cultivars, a seven-parent half diallel (F1 crosses + parents) was conducted in the non-stress and salt stress (8 and 12 ds m-1) conditions in a randomized complete block design with three replications. Genetic analysis was performed by Hayman’s method and Griffing’s fixed model, method 2. The slope of linear regression of Wr on Vr were significantly higher than 0 and had not significant difference with 1 indicating the additive-dominant model was satisfied in all cases. The narrow-sense heritability of traits was medium to high (0.4-0.8) but their broad-sense heritability was estimated relatively high (0.7-0.9). Results of regression graphs showed that Afzal parent had the most dominant allele. The significance of “a” component in most of the studied traits indicated the presence of the additive effects in controlling of traits. The significance of “b” component in most of the studied traits indicated the presence of the dominance effects in controlling of traits. The proportion of positive and negative genes was lower than 0.25 in all of the traits (except for grain weight per spike in 12 ds m-1 salinity), indicating the presence of asymmetry in the distribution of the positive and negative alleles in the parents. Based on general combining ability effects, it was concluded that under salinity, cultivar “Kavir” had favorable alleles in plant height, grain weight per spike and 100 grain weight traits and can be used as a general parent in breeding programs. Estimates of high broad-sense heritability and narrow-sense heritability in most traits indicated that these genetic materials were promising for breeding under normal and salinity stress conditions.
Ali Dowlatshah, Ahmad Ismaili, Hadi Ahmadi, Karim Khademi, Daryoush Goudarzi,
Volume 7, Issue 2 (3-2021)
Abstract
Plant breeding researches is based on genetic diversity and evaluation of genetic diversity is also one of the most important steps in introduction of new cultivars. In this study, genetic diversity of 25 grass pea genotypes was studied based on randomized complete block design with three replicates in Khorramabad (Iran). Analysis of variance showed significant differences among genotypes for most of traits. Mean comparison showed that genotype IF1312 with the highest grain yield and genotypes IF1332 and IF471 with the highest dry and fresh forage yield had the best yield. Principal component analysis showed that the first 3 factors explained 62.64% of total variance. Based on cluster analysis, genotypes IF1307, IF1872 and IF471 with the highest grain and forage yield are belonged to one cluster. REML method was used to estimate genetic correlation and heritability of different traits. The highest amount of heritability (0.87) was estimated for number of immature grains and the least heritability (0.10) was estimated for total dry weight. Grain yield had a high and positive genetic correlation with forage yield, and biomass, percentage of leaf and dry forage yield also had a high and positive genetic correlation with fresh forage yield. Totally, genotype IF1307 had the best performance for most of traits compared to the other genotypes and had an acceptable forage yield among genotypes.
Reza Amiri, Sohbat Bahraminejad, Kianoosh Cheghamirza,
Volume 8, Issue 1 (8-2021)
Abstract
The study of the genetic structure controlling agronomic traits is one of the preconditions for selecting the appropriate breeding method. In order to analyze genetic of some agronomic traits of bread wheat using generation mean analysis, parents and different generations resulting from the cross of Marvdasht × MV-17 wheat cultivars were assessed in a randomized complete block design with three replicates under normal and terminal drought stress conditions in Research Farm of Razi University (Iran) during 2015-2016 cropping season. Based on the results of weighted ANOVA, a significant difference was observed between different generations for most of the studied traits under both conditions. Generations mean analysis revealed that in addition to the additive and dominance gene effects, a variety of epistatic effects also played a role in the inheritance of most traits, and therefore cannot hope for the success of selection in first generations. The role of additive gene effect was greater than the dominance one for plant height, peduncle length and awn length, indicating usefulness of using recurrent selection to aggregate these genes followed by selecting lines with favorable agronomic characteristics. The model of genetic control for most of the traits was similar under both conditions in terms of the presence or absence of non-allelic interactions and it has not been widely affected by drought stress. The broad-sense heritability for plant height, peduncle length and awn length was estimated to be high under both conditions. The broad-sense heritability for kernel yield was moderate estimated under both conditions, but the narrow-sense heritability was very low. In general, considering the greater role of the non-additive gene effect for most of the traits, the selection is suggested in advanced generations and after access to a high level of gene fixation.
Ahmad Kaab Omeyr, Payam Pourmohammadi, Abdolali Gilani, Khalil Alami-Saeid, Mohammad Fakhari,
Volume 8, Issue 2 (3-2022)
Abstract
Various strategies have been developed to reduce water consumption in rice cultivation, including rice cultivation in water-saturated soil, drying and wetting of the soil (periodic irrigation), and aerobic rice cultivation. In this study, in order to identify and grouping rice cultivars suitable for aerobic cultivation, 34 cultivars of native and improved rice cultivars were selected and evaluated. Factor analysis based on principal components analysis based on all traits showed that 6 factors were extracted with special values of more than one. These 6 factors, after the Varimax rotation, justify the 6 main and independent factors, justifying 78.08% of the total variation. In the first harvest, cluster analysis using WARD method, cultivars were grouped into 4 groups, which cluster 3 being the largest cluster. Discriminant analysis of first harvest, showed that all cultivars were grouped correctly. In canonical discriminant the first two canonical functions with eigenvalues greater than one explained a total of 88.9% of the variance. In cluster analysis based on the average of the data obtained from Raton product, genotypes were divided into two different groups. The genotypes of the first group were higher than the genotypes of the second group in terms of yield and yield components and were introduced as superior genotypes. Heritability with genetic improvement for leaf area duration, number of seedling (grown plants), number of spikes per unit area, number of full seeds and degree of maturity showed that these traits can be transferred to hybrid offspring and selections based on these traits are effective.
Mohammad Zabet, Fahime Barazandeh, Alireza Samadzadeh,
Volume 10, Issue 1 (9-2023)
Abstract
To investigate the genetic structure of different sesame traits, 7 genotypes, including Ardestan, Sirjan, Fars, Sabzevar, Jiroft, Oltan and TS-3 were studied in the form of a 7 x 7 one-way diallel design in the research farm of the Faculty of Agriculture at University of Birjand during 2015-2018.The traits measured included plant height, height to the first capsule, number of auxiliary branches, number of leaves, leaf length, days to 50 % flowering, days to 90 % flowering, days to physiological maturity, number of capsules per plant, biological yield, grain yield, harvest index, number of seeds per capsule, capsule weight, capsule length, capsule width, chlorophyll a, b and total chlorophyll, oil and protein percentage.Hayman′s variance analysis showed that there were significant components a, b and b3 in all traits and components b1 and b2 in most traits. Parameters D, H1, and H2 were significant and the parameters F, h2 and E were not significant for most traits. The average degree of dominance (√H1/D) showed incomplete dominance and over dominance for all traits. The H2/(4H1) parameter was less than 0.25 in all traits except the day to 50 % flowering, suggesting that increasing and decreasing genes did not have symmetrical distribution among the parents. The parameter (√(4DH1)+F)/√(4DH1)-F ) showed symmetrical and asymmetrical distributions for the studied traits. In most of the traits, there was a dominant gene block controlling the traits. Broad sense and narrow sense heritabilities ranged 0.47-0.99 and 0.17-0.98, respectively. In general, all traits were controlled by genes with additive and dominance effects, so it is possible to select and produce hybrids in sesame. Considering that Fars, Oltan and TS-3 genotypes had the most dominant alleles, it is recommended that in future studies, attention should be paid to these three genotypes.
Mostafa Khodadadi, Behzad Sorkhilalehloo, Seyed Mohammad Mahdi Mortazavian, Jahangir Abbasi Kohpalekani, Mahmoud Bagheri, Milad Karbasi,
Volume 10, Issue 1 (9-2023)
Abstract
Eggplant is a highly nutritious vegetable that is widely consumed. The aim of this study was to evaluate the genetic diversity between eggplant accessions from the National Plant Gene-Bank of Iran. In the first year, a preliminary evaluation was conducted using 168 accessions. Based on preliminary evaluation results, 40 accessions were selected for complementary evaluation in the second year. The evaluation was based on 23 quantitative and qualitative traits. The results of the preliminary evaluation showed statistically significant (P<0.01) differences between accessions for all traits. Fruit shape frequencies were rounded (35.89 percent), elongated (32.18 percent), oval (13.67 percent), Semi-elongated (13.15 percent), and mace-shaped (5.11 percent). In the complementary evaluation, there were significant differences between accessions for all traits. Qualitative traits such as flower color (1.56) and fruit shape (1.53) exhibited the highest genetic variation, while fruit color (0.5) showed the lowest. Cluster analysis analysis results revealed four groups for accessions and the highest (22.34) and least (0.12) genetic distances between 1 and 2 and between 7 and 21accessions, respectively. Factor analysis showed that the first three factors explained 68.06 percent of total variation in data. The first and second factors were related to yield and yield components, respectively. Also, fruit yield traits showed high heritability and there was significant genetic correlation between these traits. Therefore, high heritable and high-scoring traits in these factors should be considered when selecting progenies in segregating populations for improvement in terms of fruit yield and shape.
Mozhgan Shirinpour, Ehsan Atazadeh, Ahmad Bybordi, Saeid Aharizad, Ali Asghari, Ashkboos Amini,
Volume 10, Issue 1 (9-2023)
Abstract
Considering the importance of maize production and the impact of water deficit stress on reducing the yield of maize, estimating the genetic components and heritability of traits for determine the breeding method under water deficit stress is essential in breeding programs. The generations drived from a cross between two inbred lines of maize including B73 (maternal line) and MO17 (paternal line), SC704 (F1) as well as F2, BC1, BC2 and F3 generations in order to estimate the genetic effects and heritability of yield, yield components and morphological traits were studied. Seven maize generations using the generations mean analysis under the full irrigation, mild and severe water deficit conditions were evaluated. The experiment was conducted in the form of randomized complete block design with 20 replications per experimental unit during two cropping seasons (2018-2019) at the Agricultural Research Station of University of Tabriz. The results of two-year combined analysis of variance and mean comparisons under three different irrigation regimes showed that water deficit stress significantly reduced all of the studied traits (except root/shoot ratio). The generations mean analysis showed the high contribution of non-additive gene effects for the genetic control of grain yield, ear diameter, number of kernel row, ear weight (in full irrigation conditions), 100 grain weight, plant height, fresh shoot weight and biological yield traits. According to these results, selection in the advanced generations and the breeding method based on hybridization can be effective to improve these traits. Also, the significant contribution of additive gene effects in controlling the inheritance of ear length, ear weight (in both stress conditions) and root/shoot ratio traits indicated that selection in early segregating generations and inbred parents can be effective for breeding of these traits and taking advantage of additive variance. Hybrid SC704 and inbred MO17 compared with the inbred B73 showed the lowest variation percentage under the water deficit stress conditions, which indicated their high yield potential and stability in the stress conditions.
Mohammad Nader Ebrahimi, Hadi Ahmadi, Mostafa Darvishnia, Daryoush Ghoudarzi,
Volume 10, Issue 2 (2-2024)
Abstract
Wheat fusarium head blight (FHB) is one of the most important diseases of wheat in humid and hot regions of the world,which causes a decrease in yield and grain quality. Fusarium head blight is caused by Fusarium graminearum. In order to investigate the resistance of some wheat traits against FHB, 27 varieties and lines of bread wheat were sown in autumn of 2021 at the farm of Faculty of Agriculture, Lorestan University. The spikes were infected at the first flowering stage by spray inoculation. The results showed that there was a great diversity among the 27 varieties of bread wheat tested in terms of growth traits, yield components, and grain yield. The results of simple correlation and also stepwise regression analysis showed that the traits of 1000 seed weight, plant height, number of seeds per spike, number of spikes, spike length and plant height are very important traits for grain yield. According to values of broad sense heritability and genetic gain for plant height and high correlation of this trait and seed yield, plant height can be used as a suitable trait for improving seed yield by selection. The overall results showed that Shiraz, Pishtaz, Quds and Morwarid-2 cultivars are suitable for transferring resistance to wheat spike blight disease in breeding programs.
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