Genetic variability and inheritance of aluminium tolerance as indicated by long root regrowth in lucerne (Medicago sativa L.)

The objective of this study was to quantify the genetic variability and determine the inheritance of Al tolerance in lucerne (or alfalfa) using a 4-parent diallel mating design. Regrowth root length (RRL), along with root length (RL) and total root length (TRL), was measured to indicate relative Al stress response using a hydroponic system in a 3 μM Al solution at pH 4.5. A diallel analysis indicated the significance of general combining ability (gca) variance for RRL but not the specific combining ability (sca) variance; the same result was obtained for TRL but not for RL. For both RRL and TRL, genetic variance appeared to be more important than the environmental variance. For RRL, a strong but non-significant correlation was indicated between parental performance and their gca effects; while mid-parent heterosis and/or over-dominance were detected, as associated with the expressed sca effects in several combinations. These results suggest the complex genetic nature and expression of Al tolerance in the 4-parent diallel crossing system tested. Al-tolerant parent, GAAT‘S’, was the most promising parent, conferring the highest gca effects for RRL as well as for other two characters. The existence of significant gca variance in RRL may also suggest the feasibility of improving Al tolerance through enhanced root regrowth using phenotypic recurrent mass selection to pyramid desirable Al-tolerant genes, focussing on parental lines and/or elite individual plants expressing long regrowth roots.     

Genetic components of pod shattering in soybean

Half diallel crosses among ten pure breeding lines of soybean were made in 1997 and 1998 to study the inheritance of pod shattering in soybean. Evaluation for pod shattering among F₂ segregating populations was carried out in an oven set at 80 ^°C for 12 hours. Diallel analysis was carried out to estimate genetic parameters and detect presence of non allelic interaction of genes affecting pod shattering. Hayman's diallel analysis indicated significant variation of Wr + Vr and Wr – Vr over arrays, suggesting epistatic gene action. Similarly results from a joint regression coefficient over replications were significantly (p < 0.05) different from unity and zero, suggesting presence of non allelic interaction of genes. The intercept was positive, suggesting partial dominance for the shattering trait. Both general combining ability (GCA) and specific combining ability (SCA) effects were significant (p < 0.05). This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic combining ability of glucoraphanin level and other horticultural traits of broccoli

Broccoli (Brassica oleracea L., Italica Group) is a source of glucosinolates and their respective isothiocyanate metabolites that are believed to have chemoprotective properties in humans. Glucoraphanin (4-methylsulfinyl-butyl glucosinolate) is a predominant glucosinolate of broccoli. Its cognate isothiocyanate, sulforaphane, has proven a potent inducer of phase II detoxification enzymes that protect cells against carcinogens and toxic electrophiles. Little is known about the genetic combining ability for glucosinolate levels or the types of genetic variation (i.e., additive vs. dominance) that influence those levels in broccoli. In this study, a diallel mating design was employed in two field experiments to estimate combining abilities for glucoraphanin content. The diallel population was developed by crossing nine doubled-haploid (inbred) parents in all possible combinations (36), excluding the reciprocals. Horticultural traits of all entries were assessed on a plot basis. In fall 2001, glucoraphanin concentration of broccoli heads ranged from 0.83 to 6.00 μmol/gdw, and in spring 2002, ranged from 0.26 to 7.82 μmol/gdw. In both years, significant general combining ability was observed for glucoraphanin concentration and total head content, days from transplant to harvest, head weight, and stem diameter. Conversely, no significant specific combining ability was observed for any trait in either year. Results indicate that a given inbred will combine with others to make hybrids with relatively predictable levels of head glucoraphanin as well as, other important horticultural traits. This should allow identification of inbreds that typically contribute high glucoraphanin levels when hybridized with others.Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the U.S. Department of Agriculture.     

A diallel crossing approach aimed on selection for ripening time and yield in breeding of new strawberry (Fragaria × ananassa Duch.) cultivars

The extension of the ripening season in open field production is of high economic interest for strawberry growers. Therefore, targeted breeding for extreme early or late ripening cultivars with high yield potential is of particular interest. Thirteen strawberry cultivars were crossed in a reciprocal way without selfing, and the 144 resulting F₁ populations were evaluated in a field trial over a period of two consecutive years. The data were analysed using a mixed‐model approach adapted for diallel crossing designs using SAS 9.3. The variability in the crossing approach is mainly based on the general combining ability (GCA) of the cultivars (additive effects). Specific and reciprocal combining abilities (non‐additive effects) appear less important. The highest GCAs for the trait Marketable Yield were found for the cultivars ‘Polka’ and ‘Yamaska’. The trait Earliness is bilateral with significantly low GCAs for early ripening in ‘Clery’ and ‘Daroyal’ and significantly high GCAs for late ripening in ‘Yamaska’ and ‘Florence’. Crosses with these cultivars are likely to deliver populations with both high yield and an extended ripening period.     

Inheritance of early blight resistance in diploid potatoes

Early blight (Alternatia solani) is a fungal disease in hot and humid environments, which causes leaf, stem and tuber lesions. Early blight resistance should be incorporated into potato cultivars because the fungicide spraying is an expensive solution for developing countries. The diploid cultivated species Solanum tuberosum group Phureja and group Stenotomum are sources of resistance alleles. The elucidation of the inheritance for early blight resistance must help to decide what could be the best breeding procedure to improve this diploid germplasm and transfer the resistance to the tetraploid level. Three experiments were carried out under controlled and field conditions to determine the heritability of this trait using nested and diallel mating designs with haploid, species and haploid-species hybrids. The narrow-sense heritability estimates were relatively high (0.64–0.78). This means that additivity was the most important type of gene action for determining resistance to early blight at the diploid level. The results suggested that diploid parents showing highest levels of resistance, throughout the cycle of disease development, can be used in 4x×2x crosses to obtain resistant tetraploid progenies to this fungal disease.     

Correlation between late blight resistance and foliage maturity type in potato

The genetics of race-non-specific foliage resistance against Phytophthora infestans, of foliage maturity type, and of their association in potato (Solanum tuberosum) were studied. Six progenies were derived from a half-diallel set of crosses between diploid potato clones that represented a broad pool within the genus Solanum and were free of any of the 11 known R genes for late blight resistance. The progenies were evaluated for resistance to late blight and for foliage maturity type, and five of them showed a significant correlation between the two traits. The correlation did not account for all variation that was present for both traits, as reflected in the analysis in which the relative AUDPC values were adjusted for foliage maturity type. The present study adds to previous results: resistance against P. infestans always coincides with late foliage maturity. However, the results also indicate that some selection for late blight resistance without affecting the foliage maturity type should be possible.     

Inheritance of Corolla Size, Number of Stamens and Percentage of Plants with Petaloid Stamens in Evergreen Azaleas (Rhododendron Subsect. obtusa)

Research has been carried out on the inheritance of corolla size, number of stamens and percentage of plants with petaloid stamens (double flower) in evergreen azaleas (Rhododendron Subsect. obtusa). For this purpose two incomplete diallel crosses were used. The first one included seven species and one cultivar and the second diallel 13 cultivars. In both diallels for all three characters the GCA mean squares were high. Also, the positive correlation between the parental values and the GCA estimates for flower size and number of stamens indicates that the expression of these characters is mainly determined by additive gene action. A negative correlation was found between GCA estimates for die n number of stamens and the percentage of plants with petaloid stamens. When breeding azaleas with large double flowers, parents can be directly selected on the basis of these characters since additive genetic variance dominates. As no genetic correlations were found between corolla size and number of stamens (percentage of plants with petaloid stamens) the breeding will not be hampered by unfavourable linkage of genes for these characters.     

Origin,diversity and utilization of the Cuban germplasm of common bean (Phaseolus vulgaris L.)

Summary Wheat root characters which influence vital plant processes have scarcely been explored for their genetic control. This study was conducted to i) examine the diversity of root traits and associated shoot traits in spring wheat (Triticum aestivum L.) cultivars; ii) study the nature of genetic control of selected traits; and iii) examine associations among root and shoot traits. Three experiments were conducted in the greenhouse with plants grown in the vermiculite medium in clear plastic tubes. In the first experiment, 42 spring wheat cultivars were grown for three weeks and measurements were taken on root length, leaf length, root number, leaf number, root dry weight, and top dry weight. In the second study, 15F₁'s originating from a partial diallel mating of six cultivars along with the parents were evaluated for 4 weeks. The data on root length, leaf length, and root number were subjected to diallel analysis according to Griffing's method 4, fixed model. A third experiment consisted of studying 2 F₂ populations with 141 plants per population. In the first study the 42 cultivars exhibited a wide range of variation for all six traits. Diallel analysis revealed significant effects of both general combining ability (GCA) and specific combining ability (SCA) for root length, leaf length, and root number. Parents with high GCA estimates were identified. Significant positive correlation coefficients were found among root and shoot traits. Analysis of F₂'s for root length indicated quantitative nature of inheritance of root length.     

Inheritance of threshing percentage in pearl millet

Summary The threshing percentage (TH%) has been suggested as a selection criterion to identify the pearl millet (Pennisetum glaucum (L.) R.Br.) lines with improved ability to fill and set grains under water limiting conditions. In this study, eight genetically diverse pearl millet inbreds and their 28 crosses produced by half diallel crossing design were used to examine range in TH%, to evaluate general combining ability (GCA) of parents and specific combining ability (SCA) effects of crosses. The results showed significant variation among parental lines for TH%. The inbreds differed for their GCA effects and crosses for their SCA effects. Parents with high TH% and positive GCA effects were identified. The results of the study revealed that both additive and dominance components, with the preponderance of later, were important in the inheritance of TH%. Heritability in narrow sense was moderate (55%) indicating that selection for high TH% might be effective.     

A comparison of progeny from diallel crosses of diploid potato with regard to the frequencies of 2n-pollen grains

Summary Selection of diploid potato genotypes that produce high frequencies of 2n-pollen grains in different environments is an important requirement for analytic breeding as well as for true potato seed production. In this context, quantitative estimates of 2n-pollen formation were made in two type of diploid potato populations. One population set consisted of the progeny from diallel crosses of five well-known diploid potato clones (A=US-W 5293.3, B=US-W 5295.7, C=US-W 5337.3, D=US-W 7589.2 and E=772102-37). Another population set was obtained by pollinating three genotypes of Solanum phureja, IvP35, IvP48 and IvP101 with clones A, C and E. It was observed that the behaviour of the parental clones with respect to 2n-pollen grains formation in different environments, such as in Europe and China, was stable. The correlation coefficient between means of 2n-pollen grains frequency of combinations and percentage of genotypes with more than 5% 2n-pollen grains in a given combination were estimated and that between mean and maximum frequency of 2n-pollen grains formation in each combination. The results showed that the former coefficients were significant wheareas the latter were not. The combining ability analysis revealed variation in 2n-pollen grains formation between families. Specific combining ability (SCA) effects contributed much more than general combining ability (GCA) to the variation in 2n-pollen grains formation. The broad-sense heritability of 2n-pollen grains formation was estimated to vary between 0.45 and 0.53 and the narrow-sense heritability was calculated to be 0.15 and 0.20 in the two types of populations. This means that additive variance was relatively low. Two to four genes were estimated to control the inheritance of 2n-pollen grains formation.     

Combining ability of seed vigor and seed yield in soybean

Studies have shown no consensus in relationships between seed yield and vigor in soybean [Glycine max (L.) Merrill]. The lack of information regarding the inheritance of seed vigor prompted this study to determine the types of gene action and combining ability estimates for seed vigor and its related traits. Five high and six low seed vigor soybean genotypes were crossed in a diallel, and selfed to produce 55 F₂ progenies, which were examined, along with the parents, for seed vigor, yield, and seed weight. Significant genotype and environment effects were found for seed vigor and yield. General combining ability (GCA) effects for seed vigor and seed yield were significant (p≤ 0.01) and larger than specific combining ability (SCA) effects. Significant GCA and SCA effects were found for seed weight, indicating that both additive and non additive genetic effects were involved in conditioning seed weight. The ratios of mean square, 2GCA / (2GCA+SCA), were 0.96 for seed vigor and 0.93 for seed yield. These ratios indicated that additive gene effects were more important than non additive gene effects for seed vigor and seed yield in these crosses. Mean seed vigor(83.8%), as determined by accelerated aging germination, and mean seed yield (2,155 kg ha^-1)in high vigor × high vigor crosses were higher than the high vigor × low vigor and low vigor × low vigor crosses. Mean percent accelerated aging germination rates in F₂ populations from diallel crosses were significantly related to mid-parent seed vigor(r² = 0.52^**) and midparent seed size (r² = 0.31^**). These results indicated that levels of seed vigor can be improved through breeding, while maintaining high yields because of the predominance of GCA effects in both seed vigor and seed yield. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distribution of the cis-sabinene hydrate acetate-chemotype in accessions of marjoram (Origanum majorana L.)

The bicyclic monoterpenes cis-sabinene hydrate and cis-sabinene hydrate acetate are considered to be responsible for the special flavor of marjoram (Origanum majorana L.). Analysis of single plants of different marjoram accessions revealed the presence of a chemotype that divided the samples into two groups, with cis-sabinene hydrate acetate present either at high amounts of above 19% or at low amounts below 2%. Additionally, an arrow negative correlation of the concentration of cis-sabinene hydrate to its acetate was observed (R² = 0.92).Since no trans-sabinene hydrate acetate could be found in the analyzed samples, the formation of cis-sabinene hydrate acetate seems to be a result of a specific enzymatic conversion in marjoram. Among 20 accessions ofmarjoram studied, 17 were heterogeneous for the cis-sabinene hydrate acetate chemotype with frequencies of the low cis-sabinene hydrate acetate type between10% and 50%. If further sensorial studies can give leads as to which of the two compounds contributes to the preferred flavor, the variation observed in this study among natural populations could be used in breeding towards an optimized quality of marjoram. The analytical technique applied in this study can be used to screen high sample numbers with a reasonable effort and time. This revised version was published online in August 2006 with corrections to the Cover Date.     

Geographical differentiation and diallel analysis of seed dormancy in barley

Seed dormancy is one of the most important parameters affecting the malting process and pre-harvest sprouting in barley (Hordeum vulgare L.). Variation of seed dormancy in 4365 cultivated and 177 wild barley (ssp. spontaneum) accessions derived from different regions of the world was investigated in Okayama University, Kurashiki, Japan. Seed dormancy of each accession was estimated from their germination percentages at 0, 5, 10 and 15 weeks post-harvest after-ripening periods. All of the wild barley accessions showed less than 10% germination at 0 week after-ripening period. Level of seed dormancy in 4365 cultivated barley accessions showed a clear geographical differentiation. Seventy seven percent of Ethiopian accessions showed high germination percentages, while 86% of Japanese, Turkish and North African accessions showed low germination percentages at 0 week after-ripening period. A half diallel cross using eleven barley accessions with different level of dormancy revealed that seed dormancy was predominately controlled by additive gene effects. These results suggest that large genetic diversity for seed dormancy in barley is explained as different levels of additive accumulation of genetic factors. Barley varieties showing appropriate dormancy could be developed by crossing among barley germplasm accessions used in the present study.     

Heterosis and genetic analysis of iron concentration in grains and leaves of maize

Iron (Fe) is a major micronutrient affecting animal and plant health. Maize (Zea mays L.) is an important crop for both food and fodders. The concentration of Fe in grains and vegetative organs, therefore, may exert a direct or indirect influence on human nutrition. Improving Fe nutrition by breeding is a promising way, but little is known about the genetics and heterosis of Fe nutrition in maize. In the present study, nine inbred lines were used to analyse the heterosis, combining ability and genetic ability of Fe concentration in grain and in the leaves supporting the cobs (ear‐leaf) of maize by a diallel mating design. It was showed that heterosis of Fe concentration in grains was very low, indicating that traditional hybrid breeding may not be efficient for improving this trait. Iron concentration of the ear‐leaf was controlled by additive gene effects and heterosis was high, indicating that hybrid breeding can be used to improve Fe nutrition of the vegetative organs of maize.     

Influence of plant age,disease incubation time,and host plant resistance on Verticillium wilt symptom expression in lucerne (Medicago sativa)

Summary The most economical means to control Verticillium wilt in lucerne, caused by Verticillium albo-atrum, is through resistant cultivars. Most Verticillium wilt resistance evaluations use some form of root-cut-soak inoculation procedure. In Trial 1, three lucerne cultivars were inoculated at eight plant ages and plant symptoms were scored at ten successive weekly intervals. Trial 2 consisted of two cultivars inoculated at ten plant ages and symptoms were evaluated at eight weekly intervals. In an experiment to study the genetics of resistance to this pathogen, four plants showing no visual symptoms were selected from each of four germplasms. The selected populations were crossed in diallel fashion. The population crosses plus the parents were evaluated for resistance to V. albo-atrum. Results from Trial 1 and Trial 2 showed disease severity increased with increasing inoculation age and with increasing scoring age. Cultivar symptom expression interacted with inoculation age, but not with scoring age. The diallel analysis supported resistance to Verticillium wilt being conferred by additive gene action, as heterosis was not detected and general combining ability mean square was 3.75 times larger than the specific combining ability mean square.     

Induced polygenic variation and genetic architecture in Indian mustard

Summary The F₁ seed obtained from an eight-parent diallel (excluding reciprocals) in Indian mustard was treated with 100 Kr of gamma rays, grown and selfed to obtain F₂M₂. The mean was decreased in the majority of the crosses in an irradiated set whereas the coefficient of variation increased. Combining ability analysis revealed the change in relative ranking of the parents and crosses and suggested the genotype-specific responses to irradiations. Components of combining ability and heterosis were also changed considerably. In some cases irradiation resulted into an increase in additive genetic variance.     

Identification of triploid Citrus by isozyme analysis

Summary Seedlessness is a desirable horticultural attribute in Citrus and is positively associated with triploidy. The conventional cytological method for triploid identification is a laborious technique involving the preparation of root tips for chromosomal analysis. Digital densitometry of isozymes, however, offers the possibility of distinguishing triploid Citrus from large populations of seedlings both quickly and cheaply. Where there are no gene dosage regulation effects, greater band density should be evident in the allozyme contributed by the diploid gamete for a heterozygous locus. The isozymes of 4 enzymes; malate dehydrogenase, 6-phosphogluconate dehydrogenase, shikimate dehydrogenase, and phosphoglucose isomerase, were investigated with polyacrylamide gel electrophoresis. Band densities of these isozymes for triploid Citrus, their diploid siblings and diploid progenitors were measured using a digital densitometer. Of the 4 enzymes investigated only allozymes for shikimate dehydrogenase exhibited consistent differences over a wide range of Citrus cultivars. Greater band density was evident in the allozyme contributed by the diploid gamete. The band density ratio between allozymes for triploid Citrus was close to 0.5, while for diploid Citrus band density ratios were close to 1.0. This effect is due to the extra protein coded by the additional gene dose and was not observed in diploids. Shikimate dehydrogenase proved to be an accurate molecular marker for distinguishing between diploid and triploid Citrus for heterozygous progeny.Abbreviations PAGE Polyacrylamide gel electrophoresis - MDH malate dehydrogenase - 6PGD 6-phosphogluconate dehydrogenase - SkDH shikimate dehydrogenase - PGI phosphoglucose isomerase     

Genetic analysis of resistance to Karnal bunt (Tilletia indica (Mitra)) in bread wheat

Summary An analyis of an F1-based incomplete diallel was conducted involving 11 parents with different levels of resistance to Karnal bunt (Tilletia indica (Mitra)). It demonstrated that general combining ability (GCA) and thus additive or additive × additive gene effects were very important in the inheritance of resistance, accounting for 86.9% of the variation. Further analysis concentrated on F3 lines derived from individual random F2 plants from crosses with resistant varieties having the highest negative GCA effects. It was shown that the varieties Weaver and W499 have single dominant genes of resistance, which are different from each other, and which differ from a single allelic gene in varieties K342 and Cruz Alta. The majority of the crosses did not demonstrate a relationship between Karnal bunt infection and the number of days to heading. Resistant F3 lines varied in the number of days to heading from 80 to 100.     

Combining abilities of in vitro traits in wheat (Triticum aestivum L.) immature embryo cultures

Summary Combining ability for six in vitro culture traits in wheat were studied in a 8×8 diallel cross (excluding reciprocals). Specific combining ability effects (sca) were significant for all six traits derived from immature embryos on two media protocols, whereas general combining ability (gca) variances were significant only for five of them. Furthermore, based on ratios obtained by comparing the ratio of K² gca to K² sca, sca was more important than gca for all six traits. Genetic correlations between shoot formation and other in vitro traits, except callus weight and root formation, were higher in magnitude than the corresponding phenotypic correlations estimates, indicating the importance of genetic effects.     

Combining ability analysis of resistance to head blight caused by <Emphasis Type="Italic">Fusarium graminearum</Emphasis> in spring wheat

Fusarium head blight (FHB) caused by Fusarium spp. is one of the most important fungal diseases of wheat (Triticum aestivum L.) in regions with wet climatic conditions. Improvement of the FHB resistance by developing new varieties requires sound knowledge on the inheritance of resistance. An 8 × 8 diallel analysis was performed to estimate general (GCA) and specific (SCA) combining ability of resistance to FHB. The F₁s and parental lines were evaluated under artificial inoculation at the experimental field of IFA-Tulln, Austria during 2001 and 2002. Disease severity was evaluated by repeated scoring of the percentage of infected spikelets and calculating an area under the disease progress curve (AUDPC). The analysis of combining ability across two years showed highly significant GCA and non-significant SCA effects indicating the importance of additive genetic components in controlling FHB resistance. The significant GCA-by-year interaction presented the role of environmental factors in influencing the FHB reaction of wheat lines. The comparison of the crosses with low FHB infection and GCA effects of their parents showed that such crosses involved at least one parent with high or average negative GCA effect. The results revealed that it is feasible to use highly or moderately resistant genotypes and conventional breeding methods to achieve genetic improvement of FHB resistance in spring wheat.