小麦冬春轮回选择育种方法研究进展

江苏淮安地处中国南北气候分界线上(33°33′ N),冬季低于5℃的天数达112天,所有冬小麦都能通过春化,1月（最冷）平均气温在0.2℃以上,春小麦也可以安全越冬。经过多年研究与实践,利用中国独有的太谷核不育小麦、矮败小麦以及冬春性小麦品种在淮安地区开展冬春轮回选择育种,把轮回选择交配圃中生产的具有很高潜在育种价值的材料向全国更多地方的小麦育种单位提供,建立供种与优良材料部分返还制度,把冬春杂交、轮回选择和穿梭育种的优越性相结合,形成了一个投入小,效益高的小麦育种体系,并成功培育出了一系列小麦新品种,为提高小麦育种成效、丰富小麦育种手段开辟了新途径。总结了国内外轮回选择育种方法研究所取得的进展,并将以解决淮北地区小麦迟播、早熟、高产间矛盾;构建抗赤霉病轮回群体,创造出优异的赤霉病抗源;以轮回群体为材料诱导小麦单倍体育种;加强同国外科研单位的合作,引入国外优良种质资源,丰富群体的遗传变异;加强与矮败小麦育种体系单位的协作等方面的研究作为今后工作的重点,以期培育出适应性广、综合性状优良的小麦新品种。     

Maize participatory breeding in Portugal: Comparison of farmer's and breeder's on‐farm selection

“VASO” is a Portuguese participatory maize breeding project (1984), where several maize landraces such as “Pigarro” have been selected both by a farmer's (phenotypic recurrent selection) and a breeder's approach (S2 lines recurrent selection). The objectives of this study were to determine the phenotypic and genotypic responses to participatory selection using these two different approaches, to clarify to which extent both selection methods preserve genetic diversity, and conclude what is the preferred method to apply in sustainable farming systems. The results, obtained via ANOVA, regression analyses and molecular markers, indicate that for both selection methods, genetic diversity was not significantly reduced, even with the most intensive breeder's selection. Although there were some common outputs, such as the determinated versus indeterminated ears, cob and ear weight ratio per ear and rachis 2, specific phenotypic traits evolved in opposite directions between the two selection approaches. Yield increase was only detected during farmer selection, indicating its interest on PPB. Candidate genes were identified for a few of the traits under selection as potential functional markers in participatory plant breeding.     

显性核不育亚麻在育种上的应用研究初报

利用显性核不育亚麻的育性稳定、不育株标记性状明显等遗传特点,进行了轮回选择、品种改良、培育新品种等多种利用途径的探讨。选育出产量超对照（内亚二、三号）８．１％～１８．９％的优良品种“内亚四号”;改良了“内亚二号”不抗枯萎病的缺点;创建了两个轮选群体;选育出２０个品系和一大批优异的单株材料。     

Physiological traits and cereal germplasm for sustainable agricultural systems

Plant breeding is not a discipline that readily comes to mind when agricultural sustainability is being considered. Sustainability is normally associated with farming practices such as stubble retention, direct-drilling, or amelioration practices such as contour farming or liming, or rotation practices for nutrient management and disease control. The contribution of plant breeding will be in providing germplasm for these changed practices and devising new methods of selection. This paper reviews opportunities where plant breeding can contribute to improvements in sustainable farming practices. The emphasis is on rainfed cropping systems and cereal improvement. The main contribution for breeding is to (i) increase crop water and nutrient use so that less escapes from the root profile; and (ii) preserve the soil resource with conservation farming systems by developing cultivars specifically adapted to changed farming systems and competitive cultivars that reduce herbicide use. To achieve these outcomes identification of desirable traits, suitable selection methods and development of appropriate germplasm are discussed.     

轮回选择在烤烟品种改良中的应用探讨

笔者在国外学者研究文献的基础上,归纳了轮回选择在烤烟品种改良中的应用价值,并结合烤烟作物自身特点,分析了烤烟育种应用轮回选择的优势,最后结合育种实践经验,指出创建烤烟轮回选择基础群体应注意亲本的选配、亲本的数目及基础群体的合成方式,提出多目标烤烟育种采用改良混合选择法结合S1选择法的轮回选择程序,同时建议采用开放式的轮回选择方案、适时引入适当遗传比率的优异种质或基因,根据基础群体所包含的种质状况、选择性状的遗传模型及遗传复杂性确定轮回选择后代种植群体规模,依据轮回选择群体的规模、轮回选择实际进程和目标性状遗传方式确定对后代的选择强度,采用约束指数对多个目标性状同时进行选择。     

Recurrent selection as a breeding procedure for rye and other cross-fertilized plants

Summary The inadequacy of the established breeding methods (mass selection, family selection) and the fact that a breeding procedure based on heterosis is not feasible for all crops, have given the incentive to search for more effective breeding methods, in particular for cross-fertilized plants.On the basis of experiments with rye, the writer comes to the conclusion that a breeding system founded on the polycross test is somewhat unsatisfactory, because it leads too soon to a narrowing of the material, so that the degeneration caused by inbreeding cannot be avoided.Better possibilities are offered by the methods of recurrent selection developed for maize, particularly by those schemes initiating from two entirely unrelated populations (reciprocal recurrent selection). The two populations are subjected to a test cross system whereby individuals from one population are outcrossed on representative groups of individuals from the other one, and conversely. Tests of the progenies obtained in this way reveal which individuals from the population K combine well with population L, and vice versa. By following the line described in the article, a narrowed population K (indicated as K₁) and a narrowed population L (indicated as L₁) are obtained which more or less complement each other. This complementary character is enhanced by repeating the selection process. Crossing in mass of these two narrowed populations may result in a valuable hybrid which is best referred to as a varietal hybrid. It seems that here the danger of inbreeding is reduced to a minimum.This method has already yielded favourable results at the Plant Breeding Station of the Central Bureau at Hoofddorp c.f. Koopman (7). From the current experiments at the Institute of Agricultural Plant Breeding at Wageningen it also appears that by the breeding system described in this paper many fallacies of the old procedures are eliminated.     

Number of crosses and population size for participatory and classical plant breeding

In the breeding of self-pollinating crops, crossing creates variation upon which selection is exerted. If the value of crosses cannot be predicted then this uncertainty means that many crosses need to be made. However, since there is a limit to the capacity of a breeding programme, more numerous crosses result in each cross having a small population size, fewer progenies in later generations and a lower probability of recovering good genotypes from each cross. Published theory on the optimum number of crosses in a plant breeding programme, for a predominantly self-pollinating crop, usually assumes that all crosses are equal value. This overestimates the number of crosses required. When the optimum size of a population in a favourable cross is considered, theory predicts that very large populations are desirable. The required population size is even larger if linkage of loci controlling different traits is also considered. Hence, in an inbreeding crop, one possible strategy is to select a small number of crosses that are considered favourable and produce large populations from them to increase the probability of recovering superior genotypes. In an out breeding crop, the analogy is a few composites with large population sizes. This low-cross-number strategy is ideally suited to the particular constraints and advantages of participatory plant breeding. Such an approach, although not essential, may still be advantageous in classical breeding. When a breeding programme is based on few crosses, which parents are chosen is crucial and farmer participatory methods are highly effective in narrowing the choice. Modified bulk population breeding methods, and recurrent selection are desirable strategies in the participatory plant breeding of self-pollinating crops when combined with a low-cross-number approach. This revised version was published online in August 2006 with corrections to the Cover Date.     

A new approach to the breeding of pigeon pea (cajanus cajan Millsp.): formation of composites

Summary Pigeon pea exhibits a high degree of potential outcrossing. It is suggested that this outcrossing potential should be utilized in the formation of random mating composites. These composites will serve as the dynamic reservoir of variability and can also be used as base populations for studies in natural selection, mass selection and recurrent selection. For developing countries, where breeding facilities are limited, a simpler breeding scheme based on the above composites has been proposed.     

我国西南地区玉米育种面临的挑战及相应对策探讨

在剖析西南地区发展玉米产业面临挑战的基础上,反思了西南地区玉米杂交育种存在的主要差距,并根据国内外玉米杂交育种发展趋势和我们的实践,从育种层面提出了应采取的对策措施。育种目标上,重点是抗病与抗虫、耐旱与耐瘠、适合全程机械化生产和耕作制度改革,以及拓展利用营养体优势新型“饲草玉米”。种质创新上,应明确杂优类群、简化杂优模式,系统开展基础种质评价与分析,合成与改良育种用群体,创制育种特异新材料。突破性自交系选育上,应注重特异新基因的发掘与利用,把产量GCA作为重要选择标准,把自交衰退慢的株系作为主要选择对象,自交、姊妹交或混粉交替进行,提高优良基因型频率;加大选系的鉴定力度,并测定产量。育种方法上,应增加种植密度,加大选择压力,实施南北穿梭育种,加快我国南方玉米种子生产基地建设,同时加快高新技术在玉米育种上的应用。     

Repetitive cycling and simple recurrent selection in traditional barley breeding programs

Summary A critical examination of three representative barley development programs from three barley producing areas of the world reveals that the evolution of each can be explained by the concepts of simple or phenotypic recurrent selection. Typically, each has utilized only 13 to 16 separate genetic sources in their program. Most of the sources were introduced over 50 years ago and have been progressively recombined through a simple recurrent selection process. The time period required for selection, testing, and reincorporation of improved genetic recombinants into the breeding program, or the completion of a cycle of recurrent selection, ranged from 6.5 to 10.5 years. If the major operative force in barley breeding is recurrent selection, then additional attention should be given to increasing the genetic base and reducing the time required to complete cycles of selection.     

Genetic diversity among populations and breeding lines from recurrent selection in Brassica napus as revealed by RAPD markers

Recurrent selection facilitated by dominant male sterility has been conducted to broaden the genetic basis for cultivar development in Brassica napus. This study aimed to evaluate the genetic variation in four base populations (C0‐C3) and breeding lines from two of the populations produced during recurrent selection by random amplified polymorphic DNA (Rapd) markers. Genetic variation in four populations declined gradually with the advance of selection cycles as measured by expected genetic heterozygosity (from 0.2058 in C0 to 0.1536 in C3) but the decline was not statistically significant. When compared with the average genetic distances for 21 germplasm collections with wide geographical and genetic origins (0.4712) and seven breeding lines from pedigree selection (0.2059), seven breeding lines selected from the C1 population and 11 from the C3 population had a larger average genetic distance (0.5339 and 0.5486, respectively). Clustering analysis indicated that the lines from recurrent selection had a much lower genetic similarity than lines from pedigree selection. Our results suggest that base populations derived from recurrent selection could provide a wider genetic variation for selection of breeding lines with more broad genetic bases.     

Efficiency of Recurrent Selection Methods to Improve the Line Value of a Population

Five breedings methods have been studied for their efficiency in improving the line value of a population, i.e. the value of all lines that can be derived from the population. These methods are: half-sib family recurrent selection (HSF-RS), full-sib family recurrent selection (FSF-RS), half-sib progeny recurrent selection (HSP-RS), recurrent selection (S₁-RS) and the single-doubled-haploid descent recurrent selection (SDH-RS). All breeding methods are compared on the basis of the genetic advance per unit of time, with the constraint of the same effective size after the first cycle of selection. Two modalities of SDH-RS are considered for an annual plant without an off-season generation: one with a cycle in four years (SDH₁-RS) and the other with a cycle in five years (SDH₂-RS). For all methods the possibility of one off-season generation for intercrossing is considered. Fields of efficiency of the best methods are determined considering five main parameters, the heritability, the importance of additive variance, the correlation QL between the additive effect for line value and the additive effect for the per se value, the length of the cycle for SDH-RS and the selection intensities. SDH-RS is the best method when the heritability is low, with a low proportion of additive variance and with a relatively low correlation QL between additive effects for per se and line values. Without off-season generation, S₁-RS is the best method among the remaining realistic situations; with one off-season generation for intercrossing it is FSF-RS. The use of an off-season generation reduces the domain of efficiency of SDH-RS while an increase in the selection intensity increases this domain. With a selection intensity for SDH-RS equal to that for S₁-RS, SDH-RS is the best in a large range of realistic situations except for SDH₂ with one offseason generation and medium to high correlation QL.     

Six years of maize selection in ‘Kitale synthetic II’, ‘Ecuador 573’, and ‘kitale composite a’ using methods of the comprehensive breeding system

Summary Evaluation of the results of six years of selection in the Kitale maize breeding methods study proved reciprocal recurrent selection to be an effective interpopulation improvement method. Ear-to-row selection was effective in improving Kitale Composite A (KCA). Data from a diallel of the cycle-6 ear-to-row substrains showed that where less than 10 lines were selected, inbreeding appeared to depress gain, but differences among the various ear-to-row experiments were not significant. Intererossing the substrains was predicted to reduce the effect of inbreeding. Yield gains in reciprocal recurrent selection and ear-to-row selection were associated with increases in ears per 100 plants. The reciprocal recurrent selection variety cross yield gain was estimated at 3.5% per year (7%/cycle) and that of the best ear-to-row procedure at 2.6% per year. The variety cross syn-2 from reciprocal recurrent selection was predicted to improve at the same rate as the best ear-to-row procedure (E7) with one long growing season per year. Under a cropping system with two similar seasons per year, however, ear-to-row selection should be more effective than reciprocal recurrent selection or its derived syn-2. Mass selection at 10% selection intensity produced significant improvement, but not at 2% selection intensity.S₁ and three half-sib selection methods were ineffective in KCA. Inbreeding and the lack of linkage equilibrium in KCA were discussed as contributing. Theoretical substrain compositing required 30 to 40 lines selected to equal the effectiveness of ear-to-row selection with 10 lines. Full-sib selection was effective in improving yield at 1.2% per year. All the recurrent selection methods showed a significant increase in ears per 100 plants. Population size was shown to be important in reducing losses due to inbreeding.If a hybrid or variety cross is potentially useable, reciprocal recurrent selection provides the best population approach. Initially, the variety cross syn-2 could be released with no additional breeding effort. Transition to variety crosses or traditional hybrids from extracted lines could be done as farmer husbandry improves and seed production facilities develop. Otherwise, open-pollinated varieties superior to existing maizes can be effectively developed using ear-to-row selection. The other methods may be effective in breeding populations when larger effective population sizes are used, but they were not effective in KCA when only 10 lines were recombined each cycle.Contribution from the East African Agriculture and Forestry Research Organization of the East African Community, the U.S. Agency for International Development (Project 618-11-110-657, East African Food Crop Research), the Kenya Ministry of Agriculture, and the Agricultural Research Service of the U.S. Department of Agriculture.     

On some aspects of reciprocal recurrent selection

Comments are given on several critical points regarding the merits of reciprocal recurrent selection as a method in hybrid breeding. A review of definitions reveals that the method utilizes general combining ability only. In simplified terms some theory of population genetics is presented and used in comparing the effectiveness of reciprocal recurrent selection with that of recurrent selection for specific combining ability to a single cross. It is found that the latter method need not be inferior if at loci exhibiting overdominance multiple alleles mutually interacting in this way may be assumed to exist. Further comments concern the use of a constant tester, the expected interactions of testcrosses with years and places, and the possibilities of an immediate utilization of testcrosses in commercial hybrids.It is concluded that reciprocal recurrent selection should not be regarded as a shortcut but as a way to enhance the genetic diversity of breeding stocks for the future. Other methods may be preferable if one has the twofold need for an early output of better hybrids and a long-sighted improvement of the same material.Paper presented at the First Meeting of the Maize Section of Eucarpia, Rome, February 23–26, 1960.     

水稻恢复系选育的轮回选择法及其应用效果研究

利用野败型细胞质雄性不育性作为遗传工具,进行水稻恢复系的轮回选择。试验证明：（1）由于构建基础群体时,加入了一定比例具有保持系性质的常规品种,以及在后续世代中,不断将经过后裔测验,证实在主要恢复基因位点上均为杂合型的中选可育株种子返回群体,给不育株授粉,因而保证了每轮次轮回选择群体内不育株比例基本稳     

高产多抗中强筋小麦陕512的选育研究（Ⅰ）

陕512小麦是西北农林科技大学农学院以陕麦150太谷核不育系为母本,以陕354为父本,1997年田间杂交,轮回选择结合系谱法育成,2001年出圃,2004年9月提前通过审定。因长势好、产量高,籽粒大、外观商品性好、中强筋、适口性优,抗病、抗干热风、抗穗发芽,被多家种子公司列为统供品种,迅速推广开来。该品种选育推广之快说明轮回选择是提高产量因子之间及产量、品质、抗性之间在更高水平上和谐的有效途径。     

Three-Way and Four-Way Recurrent Selection in Plant Breeding

Three or four-way recurrent selection is proposed for the development of three or four-way crosses. In term of genetic effects, it could be justified if there are interactions between more than three genes, whether alleles or not, if autopolyploids are also considered. For autotetraploids, in particular with tetragenic interaction, four-way recurrent selection could be more effective than reciprocal recurrent selection (two-way selection). According to the types of progenies used to develop the tests, different selection procedures are considered, the two extremes being called method 1 and method 3. With method 1 (selection for three or four-way general combining ability) a plant from a population is used to produce three-way or four-way progenies with the other populations used as testers. With method 3 (selection for three or four-way specific combining ability) three-way or four-way crosses are developed with one plant from each population. To study the genetic advance, the theory is developed to consider either diploidy or autotetraploidy. It is shown that method 3 will allow a greater genetic advance than method 1 for low heritability and low selection intensity. The way towards hybrid development with three and four-way pedigree selection is discussed: again selection can be for general or specific three or four-way combining ability. It is emphasized that an advantage of recurrent selection for three or four-way specific combining ability is that it allows direct varietal development with pedigree selection on three or four-way specific combining ability. Some other practical aspects are also considered.     

太谷显性核不育基因用于抗逆性育种的效应

利用太谷核不育小麦，采用改良半姐妹和混合轮选及隔代回交轮选三种方法进行抗逆性育种。结果表明，改良半姐妹和混合轮选对群体主穗粒数、单株粒重、百粒重等主要产量性状和抗干热风能力均有显著改良效果。改良半姐妹比混合轮选改进幅度大，且对降低群体株高有极显著效果；而混合轮选对降低株高效果不明显；两群体各主要经济性状有向选择目标集中，变异度下降的趋势。用隔代回交轮选法改造丰抗２号效果显著，升选系抗干热风，抗寒性     

利用小麦Tal基因进行株粒重、株穗数和株高群体改良的研究

利用太谷核显性核不育基因，开展小麦株粒重、株穗数和株高三个性状的轮回选择，研究群体改良的效果。经四轮选择看出：（1）株粒重增加8.25g，平均每轮增加2.06g；株穗数增加5.25个，平均每轮增加1.31个；株高降低8.82cm，平均每轮降低2.21cm。三个性状态参数的论间差异显著。（2）随轮选次数的递增，群体平均值朝着附合育种     

Development of a recurrent apple breeding programme in New Zealand: a synthesis of results,and a proposed revised breeding strategy

This paper describes an apple breeding population that has been developed in New Zealand since 1990 to broaden the genetic base for cultivar breeding. The population is now in its third generation, with recurrent selection for general combining ability being the underlying approach. During the 20 years of the programme, nearly 1,000 trees have been progeny tested. An early focus on selection within open-pollinated tests has been replaced by selection within control-pollinated tests. Genetic evaluation methods along with estimates of genetic parameters (e.g. heritability, genetic correlations, and genotype-by-environment interactions), genetic gain, and genetic diversity are summarised in this paper. The third-generation crossing programme involving nearly 350 parents is almost completed and seedlings from these crosses are in the nursery. In the revised breeding strategy, three hierarchal populations are envisioned to better integrate this broad-based population with the more commercially focused cultivar breeding programme: main breeding population (MBP), elite breeding populations (EBP) and cultivar development populations (CDP). A two-subline structure and a minimum of two crosses per parent will be implemented in the MBP. EBP will focus on developing parents for breeding goals of the CDP. Forward-selected (based on estimated breeding values) candidates from the main- and elite populations will be used as parents of the CDP. Cloning of seedlings in elite populations and also in the first-stage of cultivar development, will be investigated as a tool for enhancing predictability and shortening the cycle for delivering genetic gain.