Combining ability of low phytic acid (<Emphasis Type="Italic">lpa1</Emphasis>-<Emphasis Type="Italic">1</Emphasis>) and quality protein maize (QPM) lines for seed germination and vigour under stress and non-stress conditions

Grain yield is more likely to be compromised by poor seed germination and vigour in low phytic acid (LPA) and quality protein maize (QPM) than normal maize (Nm), especially when grown under stressful tropical environmental conditions. The objectives of this study were to determine the effect of stress conditions on seed germination and vigour traits (percentage germination, seedling dry weight, average root and shoot length and vigour index) and to determine the GCA effects of the parental lines and SCA effects of the crosses. A ten parent half diallel (LPA, Nm, QPM lines) was subjected to the standard germination (non-stress) and accelerated aging (AA) (stress) tests. All seed lots were at the same physiological age and produced under the same season and conditions. Genotypic and group differences were investigated. General combining ability (GCA) and specific combining ability (SCA) effects were significant (P ≤ 0.001) for all traits under both stress and non-stress conditions indicating that both additive and non-additive gene effects were significant. Generally SCA effects were superior for all traits. The LPA lines displayed 61 % reduction in germination and 23–52 % reduction in vigour under stress conditions thereby underscoring challenges that are expected in deploying LPA maize in stress conditions. However, LPA line CM 31 exhibited large positive and mostly significant GCA effects, while two LPA × LPA crosses showed significant positive SCA effects. Results indicate breeding is required to improve both germination and vigour of the LPA lines to adapt them to tropical conditions that are generally stress-prone.     

食用向日葵产量与籽粒主要性状配合力及遗传分析

选用4份分枝型恢复系和6份雄性不育系材料作亲本,按照不完全双列杂交（NCII）设计配制24份向日葵杂交组合,对组合的小区产量、单株产量、单盘实粒数、百粒重、籽仁率、籽粒长度和籽粒宽度7个性状的亲本配合力、遗传参数及竞争优势进行分析。结果表明,24份组合的7个性状存在真实性的遗传差异。在亲本不育系一般配合力（GCA）选择的基础上,应注意组合特殊配合力（SCA）的选择,GCA效应值高的亲本育种潜力较大,组配出优势组合的概率大;籽粒长度和籽粒宽度的狭义遗传力较低,分别为7.91%和14.89%,不宜进行早代选择;GCA较高的不育系为亲本A和亲本D,其中亲本A在各性状上均为正向GCA效应值,且在供试不育系中,其GCA效应值最大,其次为亲本D;组合1×A小区产量的SCA效应值及竞争优势最高,为本试验中的最优组合,其他籽粒性状与对照相比也均有提升。     

Combining ability and heterotic patterns of extra-early maturing white maize inbreds with genes from <Emphasis Type="Italic">Zea diploperennis</Emphasis> under multiple environments

Low soil nitrogen (Low N), Striga hermonthica and recurrent drought are major constraints to maize production and productivity in sub-Saharan Africa (SSA). Only a few extra-early maturing hybrids with combined tolerance to drought, Striga and low N have been commercialized in SSA. The objectives of the study were to determine the general combining ability (GCA) and specific combining ability (SCA) effects of grain yield and other traits, classify the inbreds into heterotic groups using the SCA effects of grain yield, and the heterotic group’s SCA and GCA of grain yield (HSGCA) methods, and examine the performance of hybrids under contrasting environments. Sixty-three extra-early white maize inbred lines containing genes from Zea diploperennis were crossed to four elite testers to obtain 252 single-cross hybrids and evaluated together with four checks at four locations for 2 years under drought, Striga-infested, low N and optimal environments in Nigeria. The GCA and SCA effects were significant (P ≤ 0.01) with preponderance of GCA over SCA effects for all measured traits indicating that additive genetic effects were predominant in the lines under all the contrasting environments. The HSGCA was more efficient than the SCA method in the classification of the inbreds into heterotic groups. The hybrids TZdEEI 74 × TZEEI 13 and TZdEEI 74 × TZEEI 29 were high yielding and most stable across research environments. These hybrids should be further evaluated in on-farm trials to confirm the consistency of performance for commercialization in SSA.     

Combining ability under drought stress relative to SSR diversity in common wheat

Eight-parental diallel cross and SSR molecular markers were used to determine the combining ability of common wheat lines grown under well-watered (WW) and water-stress (WS) conditions. Analysis of variance of yield indicated highly significant differences among the progenies. General combining ability (GCA) determined most of the differences among the crosses. Specific combing ability (SCA) was also significant but less important. The estimates of GCA effects indicated that one line was the best general combiner for grain yield under drought. Neis genetic distance, measured using SSR markers, differed from 0.20 to 0.48 among the eight genotypes. The correlation of Neis genetic distance with SCA for grain yield and heterosis ranged from 0.4 to 0.5. These results indicate that the level of SCA and heterosis depends on the level of genetic diversity between the wheat genotypes examined. Microsatellite markers were effective in predicting the mean and the variance of SCA in various cultivars combinations. However, selection of crosses solely on microsatellite data would miss superior combinations.     

Genetic analysis of drought-adaptive traits at seedling stage in early-maturing maize inbred lines and field performance under stress conditions

Maize hybrids that are tolerant to drought at the seedling stage are needed to boost productivity in the rainforest agro-ecology of West Africa. Genetics of tolerance of maize seedling to drought stress is not well understood and is poorly documented. The objectives of this study were to screen early-maturing maize lines for seedling drought tolerance, determine the inheritance and the combining ability of selected inbred lines, and evaluate the performance of seedling drought-tolerant hybrids under field conditions. Forty-nine early maize lines were screened for drought tolerance at the seedling stage. Ten drought-tolerant and two susceptible inbred lines were selected and used in diallel crosses to generate 66 hybrids. The twelve inbred lines and their hybrids were evaluated under induced drought at seedling stage in the screen house and under marginal growing conditions on the field for two seasons. Data collected were subjected to analysis of variance using the DIALLEL-SAS program. Mean squares for both GCA and SCA were significant for most traits in all research environments, indicating that additive and non-additive gene actions are controlling seedling traits under stress conditions. However, for most traits, SCA was preponderant over GCA in all environments, indicating overdominating effect of non-additive gene action. Which in turn implied that the best improvement method for the traits is hybridization. Inbred TZEI 7 had the best GCA effect for seedling traits under screenhouse conditions and for grain yield and other agronomic traits under drought conditions in the field. Hybrids TZEI 357?×?TZEI 411 and TZEI 380?×?TZEI 410 showed superior SCA effects under screen house conditions. In conclusion, the study established wide genetic variability for drought tolerance at seedling stage among tropical early-maturing maize germplasm however, the non-additive gene action was more important for most seedling traits.     

印水型杂交粳稻产量及产量构成因素的配合力分析

为确定新育成印水型杂交粳稻亲本的潜力,并为印水型杂交粳稻育种提供理论基础,本研究以4个不育系和4 个恢复系为亲本进行不完全双列杂交,研究产量及其构成因素的配合力。结果表明：产量性状一般配合力(GCA)方差均达到极显著水平,除结实率外,父本的一般配合力方差大于母本,单株产量、有效穗和结实率的特殊配合力(SCA)方差达到显著或极显著水平。139A和晚轮422 的GCA效应较高,产量性状以Ⅱ类为主,139A×晚轮422 和辽粳10A×晚轮422 的单株产量水平及组合SCA效应较高。父本GCA效应和组合SCA效应对产量性状的影响较大,母本GCA效应对产量性状的影响相对较小。本研究中,父本的遗传差异大于母本,对产量性状的影响也大于母本,139A和晚轮422 有较大的应用潜力。     

Combining ability,heterosis and genetic diversity in tropical maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) under stress and non-stress conditions

Drought and low soil fertility are considered the most important abiotic stresses limiting maize production in sub-Saharan Africa. Knowledge of the combining ability and diversity of inbred lines with tolerance to the two stresses and for those used as testers would be beneficial in setting breeding strategies for stress and nonstress environments. We used 15 tropical maize inbred lines to (i) evaluate the combining ability for grain yield (GY), (ii) assess the genetic diversity of this set of inbred lines using RFLP, SSR, and AFLP markers, (iii) estimate heterosis and assess the relationship between F₁ hybrid performance, genetic diversity and heterosis, and (iv) assess genotype × environment interaction of inbred lines and their hybrids. The F₁ diallel hybrids and parental inbreds were evaluated under drought stress, low N stress, and well-watered conditions at six locations in three countries. General combining ability (GCA) effects were highly significant (P < 0.01) for GY across stresses and well-watered environments. Inbred lines CML258, CML339, CML341, and CML343 had the best GCA effects for GY across environments. Additive genetic effects were more important for GY under drought stress and well-watered conditions but not under low N stress, suggesting different gene action in control of GY. Clustering based on genetic distance (GD) calculated using combined marker data grouped lines according to pedigree. Positive correlation was found between midparent heterosis (MPH) and specific combining ability (SCA), GD and GY. Hybrid breeding program targeting stress environments would benefit from the accumulation of favorable alleles for drought tolerance in both parental lines.     

Mode of inheritance of common bean (Phaseolus vulgaris L.) traits for tolerance to low soil phosphorus (P)

Low soil phosphorus (P) has been singled out as a major constraint leading to perpetually low bean (Phaseolus vulgaris L.) grain yield far below the expected yield potential. In developing countries beans are mainly produced by small-scale farmers who have little capacity to use inorganic fertilizers to replenish their soils. Yet bean production contributes significantly to their income and provides a cheap source of protein to rural and urban populations. The genetics of inheritance of the traits conferring low soil P tolerance is not well understood. The identification and understanding of the mode of inheritance of the traits for P-efficiency in bean will go along way in boosting bean yields through development of varieties adapted to low soil P. The objective of this study was to determine the inheritance of the traits conferring adaptation to low soil P, for the important large seeded red mottled bean market class. Three parents with known tolerance to low soil P were crossed with five adapted but non-tolerant genotypes in an 8 × 8 half diallel mating scheme. The resulting F₁ seeds were evaluated under medium and low soil P conditions at two locations. Both general combining ability (GCA) and specific combining ability (SCA) variances were highly significant (P ≤ 0.01) for all five characters studied except SCA variance for root dry weight at one location. The magnitude of GCA variance was up to twelve times higher than the SCA variance. The GCA:SCA ratio varied from 0.62 to 0.96 for the characters studied. The additive genetic variance was more important than the dominance variance for tolerance to low soil P.     

Estimating gene action,combining ability and heterosis for grain yield and agronomic traits in extra-early maturing yellow maize single-crosses under three agro-ecologies of Ghana

Maize (Zea mays L.) is the most important cereal crop produced in Ghana. However, yield of the crop is generally low, producing just about 1.7 t/ha. The low yield is attributed to continuous use of local/unimproved varieties. Generally, hybrid varieties have proven to out-yield the local/unimproved varieties due to improved vigour. Development of hybrid varieties depend on good understanding of combining ability and inheritance of important quantitative traits such as grain yield (GY). 45 half-diallel crosses generated from 10 extra-early maturing yellow inbred lines were evaluated in 2015 under rain-fed conditions. The objectives were to determine the genetic control, breeding value and estimate heritability for GY and agronomic traits of the inbred lines under contrasting growing environments in Ghana. General combining ability (GCA) and specific combining ability (SCA) were important in the inheritance of GY and agronomic traits of the inbred lines. However, GCA was more important than SCA across environments to suggest that additive gene action was more important than non-additive gene action in the inheritance of GY and agronomic traits in the inbred lines. High broad-sense heritability, for GY and other agronomic traits indicated preponderance of additive gene action in trait expression, thus, selection based on phenotypic expression could be feasible. Inbred lines P1, P4 and P8 were good combiners for high GY. The genotype, P4 × P8, was identified as the ideal and most yielding single-cross hybrid across research environments and should be further tested on-farm before commercialization.     

Diallel analysis of acid soil tolerant and susceptible maize inbred lines for grain yield under acid and non-acid soil conditions

Maize is not inherently tolerant to soil acidity but due to the ever increasing demand for the crop in the developing world, production of maize on acid soils continues to expand. Breeding for maize acid soil tolerance is the best strategy to improve yield under these conditions. Therefore, the current study was done to determine the general combining ability (GCA) of eight acid-soil tolerant and susceptible inbred lines and the specific combining ability (SCA) of cross combinations of these lines for grain yield under acid and non-acid soils. The eight lines were crossed using a diallel mating design to produce 28 single cross hybrids for evaluation under acid and non-acid soils at four sites for two seasons. Line C2 was the best general combiner under both soil environments while A2/C1 and A1/C2 had the highest desirable SCA effects under optimal conditions. Loss in grain yield and sensitivity to low pH stress was higher among genotypes in light textured soils than heavy soils. Non-additive gene action was more important than additive gene action in conditioning grain yield under both environments. Results revealed that it was feasible to improve grain yield under low pH and optimum soils from the set of genotypes used in the current study.     

Genetic distance and hybrid value in tropical maize under P stress and non stress conditions in acid soils

An emphasis in maize breeding for areas with acid soils is the development of varieties with tolerance to P-deficiency plus high yield potential in acidic as well as normal soils. This study was carried out to assess the (i) genetic diversity within a set of tropical inbred lines developed from acid soil-tolerant populations; (ii) F1 yield performance, mid-parent heterosis (MPH), high-parent heterosis (HPH), and specific combining ability (SCA) in a diallel set of crosses under P stress (low P) and non-stress (high P) conditions; and (iii) the effect of P stress on the relationship between genetic distance (GD) and hybrid performance. Using field evaluation and molecular marker studies, the results show that these germplasm from the South American maize breeding program of CIMMYT for improving tolerance to acid soils had only a moderate level of genetic diversity. The utility of GD as a predictor of hybrid value is best up to a certain threshold, as correlations with GD became inconsistent when the inbred parents were greatly divergent. There was no correlation between GD and F1 grain yield, MPH, HPH and SCA when the GD was >0.77. The high correlation of GD with F1 grain yield and with SCA in specific subsets of crosses having a narrower range of GD shows that GD can be put to practical use in predicting hybrid performance. The highest correlation between GD and SCA, seen in the subset of crosses between lines within a cluster, was reasonably stable even when the environment had a severe effect on yield.     

Combining ability and heritability of soybean resistance to groundnut leaf miner

Groundnut leaf miner (GLM) (Aproaerema modicella) (Deventer) is one of the most destructive pests of soybean and groundnuts. In this study, the mode of inheritance, general combining ability (GCA), specific combining ability (SCA) effects, maternal effects of resistance to GLM and grain yield ha^?1 were determined. Thirteen soybean parental genotypes and 81 F₂ populations were evaluated for resistance to GLM in a 5?×?19 alpha lattice diallel design with two replications under natural GLM infestation in northern (Arua) and eastern (Iki-iki) Uganda during September to December 2016 rainy season. Highly significant differences were observed among parental genotypes and F₂ populations for GLM incidence, severity, and grain yield. The estimates of GCA effects were significant for GLM incidence and severity scores but not for the number of larvae per plant and grain yield ha^?1. SCA effects were non-significant for all the studied traits, suggesting that GCA effects were the major component responsible for soybean resistance to GLM with additive gene effects being more important for these traits. Baker’s ratio ranged from 0.44-1.0 for most of resistant traits except number of larvae per plant and grain yield ha^?1. The results indicated also that cultivars Maksoy1 N, PI615437, PI578457A and NIIGC4.1-2 were good combiners against GLM incidence and severity. Parent PI615437 was a good combiner for grain yield and Maksoy1 N?×?PI615437 was a superior cross for grain yield and against GLM incidence. There were no maternal effects for the inheritance of resistance to GLM. The study provides a basis for understanding patterns of inheritance of soybean resistance to groundnut leaf miner for an efficient breeding program.     

普通玉米3个籽粒品质性状的配合力分析

以10个普通玉米自交系及其90个杂交组合在北京和山西2个地点的试验数据,分析了蛋白质、淀粉和油分含量的配合力效应。结果表明：(1) 各性状的一般配合力(GCA和特殊配合力(SCA) 均方极显著,且GCA均方大于SCA均方,说明它们在遗传上主要受加性基因效应的影响。(2) 杂交种的3个品质性状与亲本GCA平均值和SCA极显著正相关,但与亲本GCA均值间的相关性相对较强,其相关系数(r)分别为0.708**(蛋白质)、0.756**(淀粉)和0.772**(油分),表明提高亲本GCA对改良杂交种品质具有更大的作用。(3) 自交系BT1和E28的蛋白质含量GCA较高,掖478、昌7-2、掖107和48-2的淀粉含量GCA较高,E28、黄C和P138的油分含量GCA较高。     

Combining ability of maize inbred lines containing genes from Zea diploperennis for resistance to Striga hermonthica (Del.) Benth

Maize ( Zea mays ) is a staple food crop in sub-Saharan Africa, but its production is threatened by Striga hermonthica (Del.) Benth. Transfer of resistance genes from wild relatives may increase resistance to S. hermonthica in tropical maize. The objective of this study was to determine the combining ability of resistance to S. hermonthica among lines containing Zea diploperennis and tropical germplasm. Forty-five diallel crosses of 10 inbred lines were evaluated in an alpha-lattice design with and without artificial Striga infestation at two locations each in the Republic of Benin and Nigeria for 3 years. Results of analyses showed that only general combining ability (GCA) mean square was significant (P = 0.01) for number of emerged Striga plants (NESPP), while both GCA and specific combining ability (SCA) mean squares were significant for host damage score (HDS) and grain yield under Striga infestation. The ratio of GCA to SCA mean squares for the three traits varied from 3.5 to 57.5. Although GCA × environment interaction was significant for the three traits, two inbred lines containing Z. diploperennis (ZD 551) and tropical (TZL TC 87) germplasm had negative and significant GCA effects for NESPP and HDS and positive GCA effects for grain yield under Striga infestation in the two countries. Correlation between NESPP and HDS was strong and significant ( r  = 0.87, P = 0.01). Our results highlight the importance of harnessing useful genes from wild relatives to improve resistance to S. hermonthica in adapted maize germplasm.     

食葵主要经济性状的配合力及杂种优势分析

选用农家种改良过的6个食葵细胞质雄性不育系和4个优良的分枝型恢复系为亲本,按NCⅡ遗传交配设计,配成24个F₁杂种,研究了9个主要经济性状,即株高、茎粗、生育期、盘径、单盘总粒数、单盘实粒数、结实率、百粒重、单株产量的配合力及杂种优势表现。结果显示：（1）不育系中有8个性状,恢复系中有7个性状的一般配合力差异达显著或极显著水平;有4个性状的特殊配合力差异达到显著或极显著水平;（2）株高、茎粗、生育期、盘径、百粒重等5个性状主要受基因的加性效应控制;单盘总粒数、单盘实粒数、结实率、及单株产量等4个性状则受基因的加性效应和非加性效应共同影响;（3）从GCA分析看,参试的不育系以A6最为理想,A2次之;恢复系则以C3最为理想。从SCA分析看,组合A2C1单株产量效应值最高;（4）杂种优势分析表明,大部分经济性状存在杂种优势,甚至相当一部分存在超亲优势,并且这些优势都是可利用的,F₁多倾向于早熟、粗壮、高杆、大盘、多粒亲本。     

Inheritance of evapotranspiration and transpiration efficiencies in diallel F1 hybrids of durum wheat (Triticum turgidum L. var. durum)

Inheritance of water use and transpiration efficiency was studied in diallel F₁ hybrids from six cultivars of T. durum, selected for their different responses to drought stress. Plants were grown in 10 l pots at optimal (control) and low soil moisture levels in the glasshouse. GCA and SCA effects were significant at both soil moisture levels. Results demonstrated that evapotranspiration (ETE) and transpiration efficiency (TE) were under additive and dominant gene control. Interactions of GCA and SCA with moisture levels were also highly significant. GCA effects were the major components of the genetic variance of the biological and generative measures of ETE and TE. Narrow sense heritabilities for both ETE and TE based on grain yield were higher at the soil stress moisture level. Heritability estimates of ETE and TE based on the total biological yield, however, declined under the lower soil moisture level. Measures of ETE and TE showed significantly high and positive genotypic and phenotypic correlations among them as well as with grain yield and harvest index. In most cases, genotypic correlations were higher than phenotypic correlations. The results suggest that selection for high ETE and TE in terms of grain yield should be undertaken under both stress and optimal conditions. Selection in early segregating generations may lead to effective identification of desirable recombinants. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Development Traits Affecting Time to Low Ear Moisture in Maize

A diallel cross of six inbred lines of maize (Zea mays L.) from the Corn Belt of the USA and one inbred line (NZ1A) with characteristics of maize from the highlands of Mexico was grown in a cool, temperate environment. The objective was to investigate interrelationships and combining abilities of plant development traits likely to influence the time required to reach low ear moistures in the field. The traits were days to silking, days to physiological maturity, ear moisture at physiological maturity, grain filling duration and ear drying rate after physiological maturity. Grain yield and grain moisture at harvest were also considered. Differences among the hybrids of this diallel cross for ear and grain moisture at the usual harvest time were largely determined by differences in ear drying rate after physiological maturity, and not by differences in days to silking, days to physiological maturity, or in ear moisture at physiological maturity. General combining ability (GCA) effects were much larger than specific combining ability (SCA) effects for all traits. NZ1A had positive GCA effects for grain filling duration and negative GCA effects for ear moisture at physiological maturity which were outside the range of the Corn Belt lines suggesting that this line, or similar Mexican germplasm, could be useful for developing hybrids with high grain yields and low ear moistures at harvest in cool, temperate environments.     

Genetic analysis of drought tolerance in adapted × exotic crosses of maize inbred lines under managed stress conditions

Introduced maize (Zea mays L.) germplasm can serve as sources of favorable alleles to enhance performance in new maize varieties and hybrids under drought stress conditions. In the present study, the combining abilities of 12 exotic maize inbred lines from CIMMYT and 12 adapted maize inbred lines from IITA were studied for grain yield and other traits under controlled drought stress. The inbred lines from each institution were separated into groups using SSR-based genetic diversity and were intercrossed using a factorial mating scheme to generate 96 hybrids. These hybrids were evaluated under both controlled drought stress and well-watered conditions at Ikenne in Nigeria in 2010 and 2011. Average mean yields of hybrids under drought stress represented 23 % of the average yield of hybrids under full irrigation. General combining ability (GCA) effects accounted for 49–85 % of the observed variation for several traits recorded under both well-watered and drought stress conditions. Specific combining ability effects for grain yield, though positive in most hybrids, were not significant under drought stress conditions. All the twelve exotic and nine adapted lines had positive GCA effects (female, male, or both) for grain yield under either drought stress or full irrigation, or both environments. EXL03 and EXL15 that had positive and significant female and male GCA effects for grain yield under both environments can be used to improve their adapted counterparts for grain yield and drought tolerance. Normalized difference vegetation index had weak but significant correlation with grain yield.     

Ga糯玉米自交系的配合力分析与评价

摘要：利用不完全双列杂交NCⅡ设计,以3个经遗传改良的Ga糯玉米自交系为父本,与8个糯玉米自交系进行杂交。对24个杂交组合进行农艺性状鉴定,分析Ga糯玉米自交系主要农艺性状的配合力效应。结果表明,父、母本各性状的一般配合力和特殊配合力效应均达极显著差异。Ga糯玉米自交系具有良好的配合力,其中,CM327的一般配合力效应比较高,其组配的杂交种在产量、穗长、行粒数等性状上具有明显优势,具有很好的杂优利用潜力。Ga糯玉米自交系的选育利用,有助于糯玉米种质的扩增与杂种优势的利用。