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Crop Science 43:110-119 (2003)
© 2003 Crop Science Society of America

CROP BREEDING, GENETICS & CYTOLOGY

Low Soil Fertility Tolerance in Landraces and Improved Common Bean Genotypes

Shree P. Singh*,a, Henry Teránb, Carlos German Muñozc, Juan Manuel Osornoc, Juan Carlos Takegamib and Michael D. T. Thungd

a University of Idaho, 3793N 3600E, Kimberly, ID 83341-5076
b Centro Internacional de Agricultura Tropical (CIAT), A.A. 6713, Cali, Colombia
c Department of Agronomy & Soils, University of Puerto Rico, Mayaguez, Puerto Rico 00681
d EMBRAPA Arroz e Feijão, Caixa Postal 179, 75375-000, Goiania, Santo Antonio de Goias, Brazil

* Corresponding author (singh{at}kimberly.uidaho.edu)


   ABSTRACT
TOP
 NOTES
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Soil mineral deficiencies or toxicities adversely affect common bean (Phaseolus vulgaris L.) production worldwide. Cultivars tolerant to low soil fertility (LF) should support sustainable farming systems and reduce production costs and farmers' dependence on fertilizers. Our objective was to identify LF tolerant landraces and improved common bean genotypes. We systematically screened 5000 to 5500 landraces and improved genotypes for LF tolerance at Popayán and Quilichao, Colombia, between 1978 and 1998. Mean LF intensity index across locations for seed yield ranged from 0.35 to 0.68. Average seed yield reduction over five cropping seasons was 53%. Seed yield, biomass, and HI were positively associated in LF and high soil fertility (HF). LF tolerance was identified in eight landraces and 14 improved genotypes. All landraces were from Middle America (MA), belonging to common bean races Durango, Jalisco, and Mesoamerica. All improved genotypes except one (A 36) also possessed characteristics of and involved one or more LF tolerant MA landraces in their pedigree. There was considerable variation for seed, plant, and maturity characteristics among LF tolerant genotypes. In LF, mean seed yield for landraces ranged from 856 kg ha-1 for ‘Apetito’ to 332 kg ha-1 for G 19833. Among improved genotypes, A 774 had the highest (948 kg ha-1) and CAP 4 the lowest (651 kg ha-1) seed yield. Reduction in seed yield due to LF ranged from 31% for A 36 to 63% for CAP 4. All landraces and seven improved genotypes had either a below average or average LF susceptibility index. Use of these LF tolerant landraces and improved genotypes should be maximized in breeding and genetic studies to enhance sustainable farming systems.

Abbreviations: A, March to June growing season • B, September to December growing season • CIAT, Centro Internacional de Agricultura Tropical • HF, high soil fertility • HI, harvest index • LF, low soil fertility • LFII, low soil fertility intensity index • LFSI, low soil fertility susceptibility index • MA, Middle America • PR, percent reduction in seed yield due to LF


   INTRODUCTION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
DEFICIENCIES OR TOXICITIES of minerals in soils in common bean production regions occur throughout the world. For common bean, general symptoms of mineral deficiency or toxicity may include poor emergence; slow growth; seedling and adult plant stunting; leaf yellowing, chlorosis, and bronzing; early seedling death; reduced overall growth and dry matter production; delayed and prolonged flowering and maturity; excessive flower and pod abortion; low harvest index; reduced seed weight; deformed and discolored seeds; and up to 100% yield loss. Root growth may also be adversely affected (Cumming et al., 1992; Fawole et al., 1982a). These symptoms may vary with the type, severity, and duration of mineral stress.

To overcome mineral deficiencies and toxicities, common bean growers must use corrective soil amendments such as lime (Fageria et al., 1995; Westermann, 1992), manure or composted manure (Tarkalson et al., 1998), and fertilizers rich in macro- and micronutrients such as N, P, B, Fe and/or Zn (Edji et al., 1975; Henson and Bliss, 1991). Identification and use of cultivars tolerant to mineral deficiencies and/or toxicities are essential for reducing production costs and dependence of farmers on soil amendment inputs.

Greenhouse, growth chamber, and/or field screening methods have been used to identify crop germplasm tolerant to mineral deficiency or toxicity (Duncan et al., 1983). Large genotypic differences among crops also have been reported (Dwivedi, 1996; Fageria et al., 1995). Within-species variation in common bean for P (Whiteaker et al., 1976) and Zn deficiency and response (Westermann and Singh, 2000) and Al tolerance (Foy et al., 1972; Noble et al., 1985) have been documented. At the Centro Internacional de Agricultura Tropical (CIAT), Cali, Colombia, extensive research was conducted on N2 fixation (Graham, 1981) and tolerance of P deficiency (Lynch and Beebe, 1995; Thung, 1990; Yan et al., 1995a, b; Youngdahl, 1990) and Al and Mn toxicity (Ortega and Thung, 1987). In each of these cases, large genotypic differences were found.

In real farming situations, deficiencies and toxicities of two or more mineral elements often occur simultaneously (Sanchez and Salinas, 1981; Wortmann et al., 1995). Furthermore, there can be strong interactions among different minerals (Bache and Crooke, 1981) and other abiotic and biotic factors. Therefore, a more holistic approach was adapted at CIAT to develop low-input, environmentally sensitive technologies for common bean and other species (Nickel, 1987). In regard to LF, multiple deficient or toxic mineral stresses were applied to screen common bean germplasm (Ortega and Thung, 1987; Singh et al., 1995) and conduct genetic (Urrea and Singh, 1989) and breeding studies (Singh et al., 1989a, b). It was believed that germplasm and cultivars thus developed would be better suited for poor farmers in the tropics and subtropics. Such LF tolerant cultivars with higher yield potential would also be valuable for environment-friendly, sustainable farming systems in other production regions and increase profit margins for growers. Our objective was to identify LF tolerance among landraces and improved common bean genotypes.


   MATERIALS AND METHODS
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS AND DISCUSSION
 REFERENCES
 
Common bean and other Phaseolus species germplasm have been systematically screened for abiotic and biotic stresses under field conditions at CIAT. For example, as many as 20000 germplasm accessions were screened for anthracnose [caused by Colletotrichum lindemuthianum (Sacc. & Magn.) Lams.-Scrib.] at Popayán (Pastor-Corrales et al., 1995; Schwartz et al., 1982) and for angular leaf spot [caused by Phaeoisariopsis griseola (Sacc.) Ferr.] (Pastor-Corrales et al., 1998), and common bacterial blight [caused by Xanthomonas campestris pv. phaseoli (Smith) Dye] (Singh and Muñoz, 1999) at Quilichao, Colombia. Both Popayán [with a fine loamy, mixed, isothermic, typic (Andic) Dystrandept (Inceptisol) soil, pH of 4.3; 18°C mean growing temperature; elevation 1750 m; and 1925 mm rainfall] and Quilichao (with a very fine kaolinitic, isohypothermic, plinthic Kandiudox soil, pH of 4.5; 24°C mean growing temperature; elevation 990 m; and 1750 mm rainfall) have high levels of exchangeable Al and Mn, thereby causing toxicity. Also, these soils are deficient in N, P, B, Ca, and Mg (Table 1). Because disease nurseries often were grown in residual soil fertility, in addition to identifying disease resistant genotypes, these field environments also permitted retention of genotypes that had a better overall plant performance. A similar evaluation scheme, including use of complementary nurseries for different abiotic and biotic stresses, was used each year for evaluation of improved genotypes (Singh, 1992). Thus, more than 5000 landraces and improved genotypes with potential for LF tolerance were assembled for screening for their response to LF.


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  		Table 1. Soil characteristics at Quilichao and Popayán, Colombia, between 1978 and 1999 used to evaluate common bean genotypes for low soil fertility tolerance.

 
Common bean could not be grown on newly cleared native pastureland at Popayán and Quilichao without added lime and nutrients. Therefore, in HF plots, up to 5000 kg dolomitic lime and 90 kg N, 39 kg P, 90 kg K, 10 kg Zn, 20 kg Mg, and 1 kg B ha-1 were applied at the beginning in 1978–1979. The LF plots received half that amount. Additional lime and fertilizers were regularly applied for the first 5 to 7 yr to homogenize the fields and ensure adequate crop growth. Dolomitic lime and fertilizer applications were gradually reduced such that the LF plots did not receive any fertilizer and lime between 1990 and 1998 at either location. Chemical fertilizers were applied in HF plots in each cropping season at the rate of 45 kg N, 20 Kg P and 45 kg K ha-1. Results from soil analyses showed that LF plots at Quilichao remained deficient in B, Ca, Mg, and P and had toxic levels of Al (Table 1). B, P, and Zn were deficient in both HF and LF plots at Popayán. Moreover, the average yield potentials of LF and HF plots were similar to that of bean production areas in tropical and sub-tropical Latin America. At both sites, trials were grown during the main cropping seasons (A = March to June, and B = September to December), following the bimodal rainfall distribution common throughout tropical Latin America. At Popayán trials were always treated as rain fed, and supplemental sprinkler irrigation was used at Quilichao whenever necessary.

More than 5000 promising germplasm accessions and improved genotypes of common bean were systematically evaluated in LF plots at Popayán and Quilichao between 1978A and 1994B (Singh et al., 1995). Each plot consisted of a single row, 3.0 to 5.0 m long without replication. The distance between rows at Popayán was 0.5 m and at Quilichao 0.6 m. Visual appraisal of the vegetative growth before flowering and overall performance (including pod load) at maturity were recorded on a 1-to-9 scale, where 1 = excellent and 9 = very poor. All genotypes receiving scores of 7 and higher were discarded. Selected genotypes (approximately 500) were again evaluated in LF at both locations in 1994B and 1995A. Each plot consisted of four rows without replication. The length of the rows and spacing between rows were similar to the previous experiment. Visual appraisal before flowering and at maturity and seed yield were used to select the 81 highest-yielding genotypes.

Thirty-five landraces and 46 improved genotypes were evaluated in LF at Popayán and Quilichao in 1995B. A 9-by-9 partially balanced lattice design with four replicates was used. Each plot consisted of four rows, each 3.4 m long. The 33 highest yielding landraces and 31 improved genotypes were selected for further evaluations in 1996A. The trial was planted at Popayán and Quilichao under both HF and LF. An 8-by-8 partially balanced lattice design with four replicates was used. Plot size and row spacing were similar to the 1995B experiment. Mean seed yield in HF and LF and percent reduction (PR) due to LF were used as selection criteria. The selected 17 landraces and 19 improved genotypes were again evaluated under both HF and LF at Popayán and Quilichao in 1996B by means of 6-by-6 partially balanced lattice design with four replicates. Plot size, row spacing, and agronomic management of the nursery were similar to the previous year. However, in addition to seed yield, data were also recorded for biomass yield and harvest index (HI). In 1997A and 1998A, 11 selected landraces and 14 improved genotypes were evaluated in similar conditions at both Popayán and Quilichao by means of a 5-by-5 partially balanced lattice design with four replicates. Only seed yield was recorded for all genotypes.

Seed and biomass yields were adjusted to 140 g kg-1 moisture by weight. Formulae from Fischer and Maurer (1978) were adopted to calculate low soil fertility intensity index (LFII) for each growing season (and location) as LFII = 1 - Xlf/Xhf, where Xlf and Xhf are the mean of all genotypes under LF and HF environments, respectively. LF susceptibility index (LFSI) for each genotype was calculated as follows: LFSI = (1 - Ylf/Yhf)/LFII, where Ylf and Yhf are mean yields of a given genotype under LF and HF environments, respectively. For further data analysis, years and replications were considered as random effects, and genotypes and fertility levels as fixed effects. Simple phenotypic correlation coefficients among seed and biomass yields and HI were determined for the trials conducted in 1996B. All data were analyzed with a SAS PROC GLM statistical package (SAS Institute, 1985).


   RESULTS AND DISCUSSION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
REFERENCES
 
Fields at Popayán and Quilichao possessed deficient or toxic levels of two or more minerals (Table 1). No fertilizer or other amendments were applied in LF plots, and only 45 kg N, 20 kg P, 45 kg K ha-1 were applied in HF plots in each growing season between 1996 and 1998. Growth and development of common bean, like other crops, remove mineral nutrients from soil (Thung, 1990). Hence, the composition of mineral deficiency and toxicity changed over years at the two locations (Table 1). Furthermore, B, Ca, Mg, and P deficiencies and Al toxicity in LF at Quilichao, and deficiencies of B, P, and Zn in both LF and HF at Popayán still persisted at the end of our experiments. Thus, for identifying superior common bean landraces and improved genotypes tolerant to a range of mineral deficiencies and toxicities sequential screening in nonreplicated trials between 1978 and 1995A (Singh et al., 1995) and in replicated trials from 1995B to 1998 (Tables 2– 8) were essential.


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  		Table 2. Number of genotypes (N), range, and mean seed yield for 81 common bean genotypes evaluated in low soil fertility at Popayán and Quilichao, Colombia, in September to December 1995.

 

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  		Table 8. Mean seed yield, percent reduction, and low soil fertility susceptibility index for 25 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, from 1995 to 1998.

 

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  		Table 3. Number of genotypes (N), range, and mean for seed yield for 64 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, in March to June 1996.

 

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  		Table 4. Origin, growth habit, seed color, 100-seed weight, maturity, seed yield, percent reduction, low soil fertility susceptibility index, biomass yield, and harvest index for 36 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, in September to December 1996.

 

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  		Table 5. Simple phenotypic correlation coefficients between seed yield, biomass yield, and harvest index for 36 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, in September to December 1996.

 

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  		Table 6. Analysis of variance for seed yield for 25 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, in March to June 1997 and 1998.

 

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  		Table 7. Means for seed yield, percent reduction, and low soil fertility susceptibility index for 25 common bean genotypes evaluated in high and low soil fertility at Popayán and Quilichao, Colombia, in March to June, 1997 and 1998.

 
Because the amount of fertilizers and other soil amendment inputs used in this study were considerably lower, the seed yields (especially in HF) were not as high as those reported by Singh et al. (1989a) and Yan et al. (1995b). The LF intensity index (LFII) ranged from 0.02 in 1996A (Table 3) to 0.81 in 1996B (Table 4). However, the LFII varied between 0.43 and 0.66, suggesting that a relatively more consistent and moderately high stress due to LF occurred at Popayán and Quilichao during the final trials in 1997 and 1998 (Table 7). Thus, the mean seed yields in HF and LF, and PR due to LF within and across locations were used as the principal selection criteria. Consequently, some of the highest yielding genotypes across both locations in the early stages such as ‘Porrillo Sintetico’ (1313 kg ha-1), MAM 13 (1310 kg ha-1), and Amarillo 154 (1152 kg ha–1) in 1995B got excluded from 1996B (Table 4) and subsequent trials (Tables 7 and 8). Nonetheless, in the final trials, significant interactions among genotype, soil fertility level, year, and location still occurred (Table 6). Consequently, significant changes in rankings among genotypes were observed for seed yield at both soil fertility levels and locations over the cropping seasons and years (Tables 4, 7, and 8). Yan et al. (1995b) also recorded significant interactions among genotypes, P levels applied, and locations. Because temperature fluctuations over the cropping seasons and years near the equator often are minimal, changes in the rankings of genotypes could largely be due to differences in soil fertility levels, rainfall, and their interactions with the common bean genotypes at the two sites. While supplemental sprinkler irrigation was applied at Quilichao, Popayán trials were always grown as rain-fed or dry-land crops. Thus, despite relatively higher rainfall at Popayán, occasional moisture stress, especially during the pod and seed development phases, could not be ruled out. Water stress is known to affect P uptake and utilization in common bean (Al-Karaki et al., 1995).

The type and number of minerals considered, level of stress applied, screening environment (field versus greenhouse or growth chamber), selection criteria, and diversity of germplasm used for screening drastically affect the outcome of experiments. For example, Yan et al. (1995a)(b), interested in genotypic differences and understanding the physiology of specific mineral uptake and utilization, screened six common bean genotypes each of Andean and Middle American evolutionary origins for P deficiency tolerance, P-use efficiency, and response. Popayán was one of the sites included in their experiment. They reported that Andean genotypes, including G 16140 and G 19833, were more tolerant to P deficiency and were more efficient in P use in both Andosol (Popayán) and Ultisol (Mondomo) soils (located at approximately 1300 m elevation between Popayán and Quilichao) compared with Middle American genotypes such as ‘Carioca’. The genotypes G 16140, G 19833, G 2333, Carioca, and ‘Rio Tibagi’, common to experiments by Yan et al. (1995a)(b) were also included in this study. Contrary to their findings, G 16140 and G 19833 were the lowest yielding, and all other genotypes had significantly higher seed yields in both LF and HF environments despite their comparatively low PR and LFSI values (Tables 4, 7, and 8). Thus, these two landraces should be classified as highly LF susceptible.

Over the last 25 yr, large-seeded Andean common beans consistently had significantly lower yield than their small- to medium-seeded Middle American counterparts in Colombia and elsewhere (Singh, 1991; White and Gonzáles, 1990; White et al., 1992). Similar yield differences between the two groups of germplasm have been recorded across dozens of locations in the Cooperative Dry Bean Nursery that is evaluated each year in the USA and Canada (Singh and Powers, 2000). Yan et al. (1995a)(b) used 2000 kg dolomite lime, 180 kg N, 203 kg K, 2.5 kg B, and 10 kg Zn ha-1 at Popayán irrespective of the P levels. In contrast, between 1990 and 1998 no lime and fertilizer were applied in LF, and only 45 kg N, 20 kg P, and 45 kg K ha-1 were applied in HF plots. Thus, differences in type and levels of mineral stresses imposed in the two studies could be largely responsible for these contrasting results.

Subsistence farmers in Latin America seldom apply fertilizer for common bean that is grown on residual soil fertility, as was the case for LF in this experiment. Moreover, farmers rarely apply >50 kg ha-1 each of N, P, and/or K, and they almost never use micronutrient fertilizers containing B, Fe, and/or Zn. Seed yields recorded in LF and HF environments in this study were similar to those reported for contrasting environments in the Americas and elsewhere in the world. For common bean germplasm screening and breeding for low-input sustainable farming systems in Latin America, the USA, and elsewhere, an integrated approach whereby combined stress to multiple mineral elements are simultaneously imposed should be preferred over stress for specific minerals at specific times. Of course, the latter might be essential for understanding the physiology and genetics of specific mineral uptake and utilization.

Considerable variability for LF tolerance exists among landraces and improved genotypes identified in this study. Among the landraces, Apetito, Carioca, Garbancillo Zarco, Garrapato, and J 117 were consistently higher yielding in both LF and HF environments (Tables 4, 7, and 8). While they had similar PR and LFSI values, they also had considerably higher biomass yields and harvest index, especially compared with large-seeded Andean landraces such as G 16140, G 19833, and G 20554 (Table 4). Among the improved genotypes, A 321, A 445, A 774, FEB 190, MAM 38, and MAM 46 often had slightly higher seed and biomass yields than the highest yielding landraces, especially in HF. The large-seeded A 36 had the lowest yields. Nonetheless, its yields in both LF and HF were significantly higher than G 16140, G 19833, and G 20554. Mean seed yield reduction due to LF was 53% (Table 8). On average, landraces had slightly lower PR than improved genotypes. All landraces including G 16140 and G 19833 and improved genotypes had an average to below average LF susceptibility index.

In addition to LF tolerance, landraces possess many other useful traits. For example, landrace cultivars Apetito, Flor de Mayo IV, and Garbancillo Zarco also have high levels of resistance for drought stress (Terán and Singh, 2002). De Celaya and J 117 have high levels of resistance to bean pod weevil (Apion godmani Wagner) (Garza et al., 1996, 2001); Carioca is highly resistant to root-knot nematode (Meloidogyne chitwoodi Golden et al.) (S. Hafez and P. Sundararaj, personal communication, 2001); Garrapato has a high level of resistance to Bean golden yellow mosaic virus (Morales and Niessen, 1988; Urrea et al., 1996); Colorado de Teopisca has three independent dominant genes for resistance to many races of C. lindemuthianum (Pastor-Corrales et al., 1994; Young et al., 1998); and Compuesto Chimaltenango 2 has a broad-based resistance to rust [caused by Uromyces appendiculatus (Pers.) Ung.] (Stavely and Grafton, 1989). Moreover, Garbancillo Zarco has been a popular cultivar, grown on hundreds of thousands of hectares in association with corn (Zea mays L.) in moderately infertile soils in the state of Jalisco, Mexico. Similarly, Carioca has been the most popular cultivar for decades occupying more than two million hectares in infertile soils in Brazil. Owing to its high and stable yield, Carioca's cultivation has also been extended to Argentina, Bolivia, and Africa. Because these landraces were domesticated under subsistence farming systems in the absence of chemical fertilizers, herbicides, pesticides, and irrigation, they may possess resistance to many abiotic and biotic stresses. These useful intrinsic landrace characteristics therefore should be introgressed in cultivars destined for low-input sustainable farming systems.

No yield gains in LF and HF environments would be expected from intraracial common bean populations lacking genetic variation (Singh et al., 1989b). A 752 was specifically selected in the LF environment at Popayán (Singh et al., 1989a). It also exhibited good levels of tolerance to LF in this study. LF tolerant Carioca and ‘Flor de Mayo’ were among the parents used in the interracial population from which A 752 was derived. These two cultivars also are in the parentage of MAM 38. Like MAM 38, all other improved genotypes except APN 115, CAP 4, DICTA 11, and DICTA 17 were bred at Popayán and Quilichao under moderate biotic and abiotic stresses, and exhibited comparatively high LF tolerance. Moreover, LF tolerant landraces identified in this study or similar genotypes were used in broad-based interracial populations that were often subjected to the mass-pedigree method of selection (Singh et al., 1989a) with (Singh et al., 1990) or without early generation yield tests across locations (Singh et al., 1991b, 1992, 1993). For pedigree of improved genotypes refer to Rodríguez et al. (1995). A 321 exhibited LF tolerance in Africa (Wortmann et al., 1995). A 774 has out yielded most genotypes, including Carioca, in Brazilian national trials since it was introduced in the country in 1989 (Thung et al., 1993). MAM 38 was the highest yielding genotype in its market class across environments in the Mexican highlands (Acosta-Gallegos et al., 1995).

Because rigorous selection was applied over a period of several years for the identification of most promising LF tolerant landraces and improved genotypes, researchers interested in a much broader range of LF tolerant germplasm may wish to evaluate an additional portion or all 81 genotypes included in the 1995 and 1996 trials. In addition, root growth in P deficient conditions (Fawole et al., 1982a), P-use efficiency (Fawole et al., 1982b; Lindgren et al., 1977), K-use efficiency (Shea et al., 1967), resistance to chlorosis induced by Fe (Coyne et al., 1982; Zaiter et al., 1987a, b) and Zn (Singh and Westermann, 2002) deficiencies, seed Zn accumulation (Forster et al., 2002), and tolerance to LF as measured by seed yield (Urrea and Singh, 1989) are heritable traits in common bean and other crops (Clark and Duncan, 1991). Thus, much larger gains and higher levels of LF tolerance should be expected from the broad-based interracial populations involving landraces and improved genotypes identified in this study (Singh et al., 1989a). Moreover, owing to the fact that for N2 fixation considerably higher levels of P are required (Graham and Rosas, 1979), LF tolerant genotypes identified in this study might be useful for N2 fixation at low P levels.

All LF tolerant landraces originated in tropical and subtropical Latin America and improved genotypes were developed at locations close to the equator. It is therefore likely that many genotypes identified in this study are sensitive to long summer days in the temperate environments of North America (White and Laing, 1989). Thus, it would be essential first to test these genotypes for photoperiod response and general adaptation before their use in breeding programs in the USA. Sensitive genotypes would need to be grown in short-day (<=12 h light) conditions for hybridization with adapted elite parental germplasm and cultivars. Furthermore, because most genotypes possess undesirable sprawling or climbing, indeterminate growth habit Type III or IV (Singh, 1982), and have noncommercial seed types in the USA, some form of backcrossing or recurrent selection may need to be used for introgression of LF tolerance and other useful traits into North American cultivars.

Association between seed and biomass yields often are positive, and both are negatively correlated with harvest index (White et al., 1992). However, in this study, all three traits were positively correlated among themselves in both LF and HF environments (Table 5). This would suggest that the three traits were interdependent and that similar mechanisms were largely involved in their expression in both LF and HF environments. Despite these findings, the high expenses associated with conducting yield trials, and the significant interactions existent among genotypes, fertility levels, locations, and years (Table 6 and Yan et al., 1995b), the use of biomass yield and vegetative organs at any growth stage and/or HI as indirect selection criteria for LF tolerance in common bean are not proposed. Similarly, the use of LFSI or PR alone as selection criteria for LF tolerance is not advocated. Exceptions occurred such that genotypes with high biomass yields did not always have the highest seed yield and/or HI (A 750 and CAP 2) and genotypes with high HI did not always have the highest seed yield (Ojo de Cabra 24 MU and SEA 12). Moreover, these are highly selected genotypes, and even in the HF environments moderate stress due to low soil fertility existed. Thus, a positive correlation among the three traits might be expected. The proposed exclusive reliance on seed yield as the selection criterion for LF tolerance is contrary to the earlier proposal by Lynch and Beebe (1995) who advocated selection for mechanisms of tolerance or their linked markers. The latter may be of some use in the early stages of a germplasm-screening program when researchers do not have access to dependable field screening facilities. Nevertheless, seed yield testing across contrasting environments must be an integral part of any successful LF tolerant breeding program, should seed remain the principal harvestable product. After all, seed yield is the final product resulting from integration of all physiological processes controlling growth and development during the entire crop cycle (Wallace, 1985).

A positive correlation between seed yields in LF and HF environments would not justify a separate breeding program for each environment (Atlin and Frey, 1989). Singh et al. (1989a) found that the amount and type of genetic variation among common bean populations were more important than selection in high- versus low-input environments. The highest seed yield gains were realized in interracial populations among Middle American common bean races Durango, Jalisco, and Mesoamerica (Singh et al., 1991a). The Andean x Middle American intergene pool population resulted in the lowest yielding genotypes regardless of selection environment (Singh, 1995; Singh et al., 1989a). Yield testing of early generation broad-based populations involving high yielding parents with positive general combining ability in both LF and HF environments, development of advanced generation lines in HF only from promising populations that do well in both environments, followed by yield testing in both LF and HF environments might be a worthwhile breeding strategy for development of high yielding common bean cultivars for sustainable farming systems.


   ACKNOWLEDGMENTS
 
We humbly dedicate this article to Dr. John L. Nickel, Director General of CIAT, Cali, Colombia, from 1974 to 1990 for his vision, insight, interest, leadership, patience, and unconditional support for research projects designed to promote low-input sustainable common bean-based cropping systems in the tropics and subtropics.


   NOTES
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 NOTES
ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Published as Idaho Agric. Exp. Stn. J. Article No. 02714. University of Idaho, College of Agriculture & Life Sciences, Moscow, ID 83844.

Received for publication December 31, 2001.


   REFERENCES
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 ABSTRACT
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 MATERIALS AND METHODS
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The SCI Journals Agronomy Journal Vadose Zone Journal
Journal of Plant Registrations Soil Science Society of America Journal
Journal of Natural Resources
and Life Sciences Education		 Journal of
Environmental Quality