Comparison of intra- and inter-pool crosses in faba beans (Vicia faba L.). II. Genetic effects estimated from generation means in Mediterranean and German environments

Genetic diversity between parents is important for hybrid breeding and for maximum usefulness of a cross in pure line breeding. However, wide crosses may suffer from poor adaptation in the target environments and recombination losses owing to disruption of favourable epistatic gene combinations. In this study we investigated the importance of various gene effects for different traits in the parents and generations F¹ to F⁵ derived from 99 intra- and inter-pool crosses among 20 lines of the Minor, Major, and Mediterranean germplasm pools of faba bean in Mediterranean and German environments. Yield performance of parental inbreds and progenies of crosses in different selfing generations was high in the region of adaptation of the respective germplasm pool. Heterosis for yield was greatest in interpool crosses of Minor × Mediterranean germplasm, amounting to 106% in Mediterranean and 72% in German environments. Interpool crosses exhibited significantly greater heterosis compared with intrapool crosses in both mega-environments. Generation means analysis revealed the importance of additive gene effects and dominance × dominance epistasis in Mediterranean and German environments. Dominance effects and additive × additive epistasis for yield were negligible in most crossing groups. Since intrapool crosses of Mediterranean germplasm in Mediterranean environments and crosses of Minor and Major germplasm in the German environments were most promising, we suggest crosses among adapted germplasm for hybrid breeding and for establishing new base material in line breeding in faba bean.     

Pattern of genetic inheritance of morphological and agronomic traits of sorghum associated with resistance to sorghum shoot fly, <Emphasis Type="Italic">Atherigona soccata</Emphasis>

Sorghum shoot fly, Atherigona soccata is an important pest of sorghum during the seedling stage, which influences both fodder and grain yield. To understand the nature of inheritance of shoot fly resistance in sorghum, we performed generation mean analysis using two crosses IS 18551 × Swarna and M 35-1 × ICSV 700 during the 2013–2014 cropping seasons. The F₁, F₂, BC₁ and BC₂ progenies, along with the parental lines were evaluated for agronomic and morphological traits associated with resistance/susceptibility to sorghum shoot fly, A. soccata. The cross IS 18551 × Swarna exhibited significant differences between the parents for shoot fly deadhearts (%) in the postrainy season. The progenies of this cross exhibited lower shoot fly damage, suggesting that at least one of the parents should have genes for resistance to develop shoot fly-resistant hybrids. Leaf glossiness, leafsheath pigmentation and plant vigor score during the seedling stage exhibited non-allelic gene interactions with dominant gene action, whereas 100 seed weight showed both additive and dominant gene interactions. Presence of awns showed recessive nature of the awned gene. Generation mean analysis suggested that both additive and dominance gene effects were important for most of the traits evaluated in this study, but dominance had a more pronounced effect.     

Genetic analysis of plant height using two immortalized populations of “CRI12 × J8891” in Gossypium hirsutum L.

Plant height is an important plant architecture trait that determines the canopy structure, photosynthetic capacity and lodging resistance of upland cotton populations. To understand the genetic basis of plant height for marker-assisted breeding, quantitative trait loci (QTL) analysis was conducted based on the genetic map of recombinant inbred lines (RILs) derived from the cross “CRI12 × J8891” (Gossypium hirsutum L.). Three methods, including composite interval mapping, multiple interval mapping and multi-marker joint analysis, were used to detect QTL across multiple environments in the RILs and in the immortalized F₂ population developed through intermating between RILs. A total of 19 QTL with genetic main effects and/or genetic × environment interaction effects were identified on 15 chromosomes or linkage groups, each explaining 5.8–14.3 % of the phenotypic variation. Five digenic epistatic QTL pairs, mainly involving additive × additive and/or dominance × dominance, were detected in different environments. Seven out of eight interacting loci were main-effect QTL, suggesting that these loci act as major genes as well as modifying genes in the expression of plant height. The results demonstrate that additive effects, dominance and epistasis are all important for the genetic constitution of plant height, with additive effects playing a more important role in reducing plant height. QTL showing stability across environments that were repeatedly detected by different methods can be used in marker-assisted breeding.     

Genetic Analysis of Resistance to Phytophthora Root Rot in Tomato (Lycopersicon esculentum Mill.)

The resistant accession, LA1312, and the susceptible cultivar ‘Peto 343′, were crossed to develop F₁, F₂ and BC₁ populations for genetic analysis of resistance in tomatoes to Phytophthora parasitica Dastur, the causal agent of Phytophthora root rot. There was no maternal effect on resistance. Generation means analysis indicated that tolerance to Phytophthora root rot was under genetic control with both simple (additive and dominance) and digenic interaction (additive × additive and additive × dominance) effects contributing to the total genetic variation among generation means. Weighted least square regression analysis indicated that the majority (ca. 96 %) of the genetic variation could be explained by simple additive effects alone. Narrow sense heritability was estimated as 0.22. Based on effective factor formulae, at least five effective factors controlled the resistance. Implications for breeding procedures are discussed.     

Involvement of higher order interactions addressing complex polygenetically controlled inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot] resistance in pearl millet [Pennisetum glaucum (L.) R.Br.]

Inheritance of downy mildew [Sclerospora graminicola (Sacc.) Schrot]resistance was studied using generation mean analysis in pearl millet [Pennisetum glaucum (L.) R.Br.]. Eleven basic generations, namely, P₁, P₂, F₁, F₂, B₁, B₂, B₁F₂, B₂F₂, L₁, L₂ and L₃ of three crosses involving six diverse lines for downy mildew incidence were evaluated under artificial epiphytotic conditions over two environments. The downy mildew incidence was best fitting for digenic, trigenic and tetragenic ratios when fitted into classical Mendelian ratios demonstrating involvement of two or more genes. Digenic and trigenic interaction models were adequate in the case of crosses I and III respectively, to account for the total variability in generation means. Unlike severity, comparative estimates of gene effects over two environments were mostly consistent in all crosses for prevalence. Most of the epistatic and major gene effects were found significant in all crosses for both the disease traits. Non-allelic interactions particularly at three-gene loci viz., w (additive × additive × additive) and y (additive × dominance × dominance) in cross II and all trigenic interactions in cross III were predominant. Duplicate dominance (cross I) and complementary epistasis (crosses II and III) were observed for both the traits revealing inconsistency of gene effects over crosses. The gd₁ (interaction of additive gene effect with e₁) and gh₁(interaction of dominant gene effect with e₁) were significant in crosses I and II, indicating interaction of additive and dominance gene effects with environments. Thus a breeding method that can mop up the resistant genes to form superior gene constellations interacting in a favorable manner against pathotype I would be more suitable to accelerate the pace of resistance improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Estimation of additive, dominance and digenic epistatic interaction effects for certain yield characters in Vigna sesquipedalis Fruw

Inheritance of yield and yield contributing characters were investigated using generation mean analysis, utilising the means of six basic populations viz., P₁, P₂, F₁, F₂, BC₁P₁ and BC₁P₂ in four crosses of Vigna sesquipedalis. The analysis reiterated that the importance of dominance (h) gene effects for pod yield/plant and pods/plant as compared to additive (d) gene effects. However, significant and positive additive effects were noticed for pod yield/plant, pods/plant, pod weight and seed weight in different crosses. The three types of gene interactions (additive, dominance and epistasis) were significantly involved for pods/plant in cross KU 7 ×KU 8. Among the digenic epistatic interactions, both additive ×additive (i) and dominance × dominance (l) contributed more for pod yield/plant and pods/plant, however, it varied among the crosses. Populations having earliness can be developed as indicated by reducing dominance effects. Pedigree selection and heterosis breeding is suggested to exploit the fixable and non fixable components of variation respectively in Vigna sesquipedalis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of drought resistance related traits in two crosses of groundnut (<Emphasis Type="Italic">Arachis hypogaea</Emphasis> L.)

Groundnut (Arachis hypogaea L.) is an important oilseed crop grown in more than 100 countries across wide range of environments. Frequent occurrence of drought is one of the limiting factors adversely affecting groundnut productivity, especially in rainfed areas, and hence genotypes having high water use efficiency (WUE) under limited available water need to be developed. In groundnut, WUE is correlated with SPAD chlorophyll meter reading (SCMR) and specific leaf area (SLA). These two traits, SCMR and SLA, can be used as surrogate traits for selecting for WUE. In order to improve SCMR and SLA, and in turn WUE in groundnut, a good knowledge of the genetic system controlling the expressions of these traits is essential for the selection of the most appropriate and efficient breeding procedure. The present investigation was conducted to determine the gene action controlling the inheritance of SCMR and SLA in two crosses, ICG 7243 × ICG 9418 and ICG 6766 × Chico, and their reciprocals. Six generations of each cross (P₁, P₂, F₁, F₂, BC₁P₁, and BC₁P₂) were evaluated for SCMR and SLA at two stages of the crop growth viz., 60 and 80 days after sowing (DAS). For SCMR at 80 DAS, additive effects were important in both the crosses whereas predominance of dominance effects with duplicate epistasis was observed for SCMR at 60 DAS and SLA at both stages in both the crosses. Predominance of additive effect for SCMR at 80 DAS suggested effective selection could be practiced even in early generations whereas for SCMR at 60 DAS and SLA at both stages in both crosses, it would be better to defer selection to later generations. Further, recording of SCMR and SLA should be done between 60 and 80 DAS for screening the germplasm lines for drought tolerance.     

Genetic effects and heterosis of within-boll yield components in upland cotton (Gossypium hirsutum L.)

Within-boll yield components are the most basic contributors to lint and seed yield of cotton (Gossypium hirsutum L.), which is a major source of natural fiber and edible oil throughout the world. Little information is available on genetic effects and heterosis of these traits in cotton. Three cotton cultivars and six breeding lines differing in within-boll yield components were used for this study. Parents and their F₁ progeny with reciprocals from a 3×6 factorial mating design were grown at Jiangxi Agricultural University experimental farm in 2008 and 2009. Seven within-boll yield components and two boll bur characters were analyzed under an additive-dominance genetic model with genotype and environment interaction. Additive effects were significant for all traits and dominance effects were significant for all traits except seed mass per seed. Genetic variances for lint mass per seed, SM/S and boll bur weight were primarily additive variances ranging from 39.6 to 58.9 %. Lint mass per boll and seed number per boll variances were primarily due to dominance genetic effects ranging from 36.4 to 48.8 %. Dominance and additive effects were equally important for boll weight, seed mass per boll and boll bur percentage. Additive and additive × environment effects were more important than other effects for lint percentage. A802-1 had the best mean performance and additive effects increasing BW, SM/B, S/B and SM/S, but reduced LP and LM/S. A9-1 and Lu40534 had additive effects associated with increasing LP. The two crosses A9-1×Lu40534 and Tezsh×33B were detected with favorable heterozygous dominant effects and heterosis over best parent for BW, LP, LM/B, SM/B and S/B. Favorable genotypic and phenotypic correlations were identified between within-boll lint yield components (LM/B, LM/S) and within-boll seed yield components (SM/B, S/B, SM/S). These results indicate that simultaneous genetic improvement of multiple within-boll yield components can be expected in breeding populations derived from these cotton cultivars and breeding lines.     

Genetics of resistance to early blight (Alternaria solani Sorauer) in tomato (Lycopersicon esculentum L.)

The genetic nature of early blight resistance in tomato was studied in three crosses at seedling and adult plant stages. A six generation mean analysis of the cross Arka Saurabh (susceptible) × IHR1939 (resistance) and its reciprocal cross revealed that the resistance to early blight was conferred by recessive polygenes at both seedling and adult plant stages. This polygenic early blight resistance revealed the importance of additive and additive × additive gene effects at seedling stage and higher magnitude of dominance and dominance× dominance gene effects at adult plant stage. Evaluation of parents, F₁, F₂ and backcross generations of IHR1816 (resistance) × IHR1939 (resistance) revealed that the early blight resistance genes in IHR1816 (Lycopersicon esculentum NCEBR-1) and IHR1939 (Lycopersicon pimpinellifolium L4394) are independent. This revised version was published online in July 2006 with corrections to the Cover Date.     

Quantitative resistance to Cercospora leaf spot disease caused by Pseudocercospora cruenta in cowpea

Six populations (Parent 1, Parent 2, F1, F2, BC1 and BC2) generated from each of four crosses involving four resistant and two susceptible varieties of cowpea (Vigna unguiculata L. Walp) were evaluated for resistance to Cercospora leaf spot (CLS) disease caused by Pseudocercospora cruenta under induced epiphytotic conditions, in four separate field experiments. Climatic conditions determined the onset of CLS disease in the susceptible cultivar and varied in the four experiments from 35 to 48 days after planting (DAP). Genetic analysis revealed that the mode of inheritance of resistance to P. cruenta can be oligogenic or polygenic depending upon the cross. This is the first report of polygenic inheritance of CLS resistance. Number of nodes infected fitted a simple additive dominance model with predominance of additive effects based on generation mean analysis. Oligogenic resistance was observed for the other three crosses, with the most plausible models being: a single gene model with incomplete dominance in CB27 × IT87D-939-1; a single gene model with complete dominance in CB27 × VRB-10; and a triger model in Los Banos Bush Sitao × IT86D-792, based on segregation analysis of symptomatic : non-symptomatic plants. The role of minor genes was also indicated in the above crosses. Suggested approaches to breeding for resistance to CLS are discussed.     

Genetic effects and genotype × environment interactions for cooking quality traits in Indica-japonica crosses of rice (Oryza sativa L.)

Indica-japonica hybridization is an important approach for developing superior performing hybrids in rice (Oryza sativa L.). In view of the scanty information available on cooking quality characters in indica-japonica crosses, an investigation was undertaken to estimate genetic and genotype × environment variance and covariance components of amylose content, gel consistency and alkali digestion value, and to determine the relative importance of direct genetic effects, maternal genetic effects and cytoplasmic effects in the genetic variations of the three quality characters. Two indica photo-sensitive genic male sterile (PGMS) lines and four japonica varieties were used as parents to make crosses. Genetic model with genotype × environment interactions for triploid endosperm was used for genetic studies of the three cooking quality characters. Variance component analysis revealed that genetic variations of the three characters were mainly attributable to direct additive and maternal additive effects, and the three traits had significant direct and maternal heritabilities. Genotype × environment interactions were mainly dominance × environment (including direct dominance × environment and maternal dominance × environment) and cytoplasm × environment interactions. Environment factors could only affect the expression extent of dominant genes, without changing their directions. Predicted values of genetic effects indicated that the parental lines, ‘VI-70’ and ‘H9304-1’, appeared to be best for amylose content, ‘T 1950’ and ‘Suxuan’ appeared to be best for gel consistency and alkali digestion value. This revised version was published online in August 2006 with corrections to the Cover Date.     

The inheritance of quantitative traits in Brassica napus ssp. rapifera (swedes): Augmented triple test cross analyses of production characters

Two F₂ triple test crosses, augmented with F₃s, produced from crosses between different inbred lines of swedes (Brassica napus ssp.rapifera L.) were assessed in field trials at Dundee in 1988 and 1989,respectively. This paper reports the analyses of resistance to powdery mildew, neck length, growth cracks, sugar content and hardness; analyses of yield have been published previously. Additive genetical variation was found for all traits while non-additive variation was less important, the highest degree of dominance being 0.44 for hardness. There was evidence of additive × dominance and dominance × dominance epistasis for mildew and additive × additive epistasis for neck length and hardness. Significant,consistent reciprocal differences were found and these were particularly large for neck length and growth cracks. Sugar determination was carried out on the basic generations of the second cross, the parental lines of which showed large differences in concentration of glucose, fructose and sucrose. Directional dominance was found for high glucose but not for fructose or sucrose. The implications of these results for swede breeding are discussed and it is proposed that inbred cultivars would be a more practical option than F₁ hybrids. This revised version was published online in August 2006 with corrections to the Cover Date.     

Generation Mean Analysis of Inheritance of Resistance to Pyrenophora teres in Barley

F1, F2, Fl × parentl (BC1), and Fl × parent2 (BC2) generations resulting from four crosses among seven cultivars of barley used in national and international breeding programmes were tested at the seedling stage for their resistance to a mixture of five isolates of Pyrenophora teres. Four methods were used to assess disease resistance: infection type (IT), average lesion size (ALS), number of lesions per unit leaf area (NL) and percent leaf area infected (PLAI). Gene actions were estimated by generation mean analysis on each of the four crosses and on each of the evaluation methods. Significant additive and additive × additive epistatic effects were found. Infection type and percent leaf area infected were found to be highly correlated in all four crosses. These results suggest that barley breeders could improve the level of resistance to P. teres by making appropriate crosses between highly susceptible barley cultivars.     

Genetic analysis on oil content in rapeseed (Brassica napus L.)

High oil content is one of the most important characteristics of rapeseed (Brassica napus L.) breeding. In order to understand the genetic basis of seed oil content, a series of reciprocal crosses between rapeseed parents with high oil content (53110, 61616 and 6F313), medium-oil content (Zhongshuang 9) and low oil content (51070 and 93275) were conducted. It was found that the oil content of F₁ hybrid seeds in rapeseed was mainly controlled by the maternal genotype. The maternal effect value of oil content was estimated to be 0.86. The pollen parent had a xenia effect on oil content, estimated to be 0.14 which changed the mean value by 1.86 percent. The inheritance of oil content was studied in a set of 8 × 8 diallel crosses of different varieties. The results indicated that the inheritance of oil content could be explained by an additive-dominant-epistasis model. Although the dominant and additive effects played major roles and accounted for more than 70% of the total variance, there was also a small epistatic effect. The broad and narrow sense heritability of oil content was 83.88 and 36.94%, respectively. Based on the oil content differences between the reciprocal crosses in the same offspring generation (F₁ and F₂) in rapeseed, it could be concluded that there were significant cytoplasmic effects on oil content. In this study, two lines with significantly cytoplasmic effects, either positive or negative, were selected.     

Generation mean analysis of resistance to downy mildew in adult muskmelon plants

Downy mildew (Pseudoperonospora cubensis) is the most devastating disease in muskmelon (Cucumis melo). A generation mean analysis study was designed to determine the types of gene action and estimate the heritability for resistance to downy mildew in four selected crosses of muskmelon. Generation mean analysis revealed that genetic dominance may be of greater importance for expression of resistance to downy mildew in both greenhouse and field experiments and in all the crosses. The F₁ mean was significantly lesser than the mid-parent value and skewed towards resistant parent in all the crosses. Negative sign associated with gene effects indicated, in those crosses, that disease level could be decreased in relation to midparent. All the crosses expressed significant and positive additive (d) gene effects. Dominance (h) and dominance × dominance (l) gene effects had opposite sign in all crosses and both experiments, which implied duplicate type of gene action. High mid-parent heterosis in all the crosses indicated strong dominance effects (as combination of parental alleles) for resistance to downy mildew. In all the crosses, both resistant and susceptible parent contributed one or more dominant/partially dominant factors for resistance. Estimates of broad-sense heritability were high and relatively consistent in both experiments. The two different screening experiments showed that fixable gene effects (d + i) were lower than the non-fixable gene effects (h + l) in all the crosses indicating greater role of non-additive effects in the inheritance of resistance to downy mildew. Resistance to downy appeared to be controlled mainly by dominance effects, therefore the inbred lines IIHR 121 and IIHR 122 could be used strategically to exploit heterotic effects.     

Diallel analysis of Fusarium head blight resistance in genetically diverse winter wheat germplasm

New sources of partial resistance to Fusarium head blight (FHB) in wheat have been identified over the past decade; however, little is known of their breeding value. A 20 parent partial diallel that included resistant genotypes from the U.S., Europe, China and South America was used to evaluate the potential of these sources of resistance as parents in wheat breeding programs. Eight plants replication⁻¹ of each of 190 crosses and 20 parents were point-inoculated with Fusarium graminearum under greenhouse conditions in two replicated experiments. Both general (GCA) and specific combining ability (SCA) were significant. Most of the variance for FHB severity was associated with additive genes; however, estimates for SCA ranged from highly negative to highly positive in both resistant × resistant and resistant × susceptible crosses which suggest that improving FHB resistance through gene pyramiding strategies based on additive genetic variation may be complicated by interaction effects that condition FHB resistance.     

Epistasis and heritability of resistance to <Emphasis Type="Italic">Phytophthora nicotianae</Emphasis> in pepper (<Emphasis Type="Italic">Capsicum annuum</Emphasis> L.)

Gene effects of resistance to two isolates of Phytophthora nicotianae in two crosses of pepper were investigated using separate generation means analysis. Additive-dominance models were inadequate in all cases. Digenic parameter models were adequate in three cases and the probability of goodness of fit of models was negatively correlated with the aggressiveness of the pathogen. None of these models explained variation among generation means in the combined cross Beldi × CM334 with P. nicotianae isolate Pn_2. Additive × additive, dominance × dominance and dominance × additive effects were significant in most cases. Additive and dominance effects (of negative sign) contribute more to resistance than to susceptibility. Additive variance was greater than environmental and dominance variance and ranged from 0.038 to 0.224. Narrow-sense heritabilities were dependent upon the cross and inoculate and ranged from 86 to 92%. The results of this study indicate that selection with more aggressive isolates of the pathogen will be useful for enhancing resistance in pepper.     

Genetic analysis of resistance to <Emphasis Type="Italic">Fusarium</Emphasis> root rot in common bean

Fusarium root rot (FRR) is a major disease of common bean worldwide. Knowledge of the inheritance of resistance to FRR would be important in devising strategies to breed resistant varieties. Therefore, a 12 × 12 full diallel mating scheme with reciprocal crosses was performed to generate 132 F₁ progenies, which were then advanced to the F₃. The progenies were evaluated for resistance to FRR under green house conditions in Uganda. General combining ability (GCA) effects were highly significant (P ≤ 0.01) for disease scores. Specific combining ability effects were not significant (P > 0.05) in the F₁, but were highly significant (P < 0.01) in the F₃ generation. These results indicate that resistance to FRR was governed by genes with additive effects in combination with genes with non-additive effects. Reciprocal differences were also significant (P = 0.01) at F₁ and F₃, primarily reflecting a large influence of maternal effects in both these generations. In fact, susceptible parents did not differ significantly (P > 0.05) for disease scores when used as paternal parents in the F₃, but differed strongly as maternal parents (P = 0.0002). Generally, the progenies were distinctly more resistant when the resistant parent was used as the female in crosses, especially as observed in the F₃. The maternal effects were strong in the F₃ generation, suggesting a complex form of cytoplasmic–genetic interaction. The non-maternal reciprocal effects in the F₃ were significant (P < 0.05) in both the resistant × resistant diallel, and in the resistant × susceptible crosses. Mid-parent heterosis (MPH) occurred in most crosses, with average heterosis approximately equal in each of the three generations, indicating that epistasis was probably more influential than dominance of individual genes. Gene-number formulas indicated that several genes were involved in resistant × susceptible crosses. Among resistant × resistant crosses, many produced continuous distributions of F₁ progeny scores, suggesting polygenic inheritance, while bi-modal distributions were characteristic of the F₃ distributions, and fit expected ratios for two or three loci segregating in each cross. Dominant forms of epistasis favoring resistance were strongly indicated. Parent–offspring heritability estimates were moderate. Overall, the results indicate that resistant parents contain a number of different resistance genes that can be combined with the expectation of producing strong and durable resistance. The lines MLB-49-89A, MLB-48-89, RWR719 and Vuninkingi, with large and negative GCA effects, contributed high levels of resistance in crosses and would be recommended for use in breeding programs.     

Analysis of decision-making coefficients of three main fiber quality traits for upland cotton (Gossypium hirsutum L.)

The analysis of genetic correlations between fiber length (Len), strength (Str), micronaire, and 12 other traits was conducted using the additive (A)-dominance (D) genetic model, which considers genotype × environment interaction effects, in intraspecific upland cotton (Gossypium hirsutum L.) hybrids to effectively improve the quality of cotton cultivars in high planting density cases. Decision-making coefficients were computed based on the genetic correlation and path analysis of three fiber quality traits. The decision-making coefficient analysis of three fiber traits in cross breeding was beneficial for the improvement of Len by increasing the additive effects of Str and length of boll (LB) and decreasing lint percentage (LP), boll number of the top three fruit-bearing branches. The analysis was also beneficial for the improvement of Str and fiber fineness by increasing or decreasing the additive effects of other traits. Utilizing heterosis in hybrids was beneficial to the heterosis of Len by selecting the high dominance effects of number of nodes of the 1st fruit-bearing branch and LB and decreasing the dominance effects of diameter of boll (DB) and LP and for improving Str by increasing the dominance effects of DB and decreasing the dominance effects of number of fruit-bearing branches and number of nodes of the main stem (NNMS). Utilizing heterosis was also beneficial for improving fineness by increasing the dominance effects of LB, Str, and lint yield and decreasing the dominance effects of NNMS and Len.     

The biological characters and polyploidy of progenies in hybridization in 4×–2× crosses in <Emphasis Type="Italic">Dianthus caryophyllus</Emphasis>

Crosses were carried out between tetraploid Dianthus caryophyllus cv ‘Butterfly’ (2n = 4×= 60) and seven diploid cultivars (2n = 2×= 30). Fewer seeds were obtained and the low seed germination was found which suggested the post-fertilization barrier in 4×–2× crosses. 12 progeny were obtained from 5 crossing combinations. Chromosome analysis revealed that they consisted of 5 triploid hybrid plants and 7 tetraploid hybrid plants, suggesting that unreduced male gamete maybe be involved in polyploid formation. Various flower shapes and colours were observed in the polyploid progenies, showing that sexual polyploidization results in greater variability and fitness. The hybrids obtained by 4×–2× crosses showed the flower-size intermediate between the parents or larger than the parents. Some favourable characters of parents such as flower shape, flower colour and resistance to Fusarium oxysporum, were successfully transmitted to the hybrids. Since polyploid hybrids have some of the profitable characters of the parents, they are expected to be used for future breeding in carnation.