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Published in Agron J 99:1219-1225 (2007)
DOI: 10.2134/agronj2006.0301
© 2007 American Society of Agronomy
677 S. Segoe Rd., Madison, WI 53711 USA
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Drought

Drought Resistance in the Race Durango Dry Bean Landraces and Cultivars

Shree P. Singh*

Plant, Soil, and Entomological Sciences Dep., Univ. of Idaho, 3793 North 3600 East, Kimberly, ID 83341-5076

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

Received for publication October 30, 2006.

   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Increasing water deficit or drought is a problem for dry bean (Phaseolus vulgaris L.) production in the western USA. Identification of drought-resistant cultivars is crucial for sustainable production in the region. The objectives of this study were to (i) measure the effects of drought on seed yield, seed weight, and days to maturity; (ii) determine correlation among the three traits in drought-stressed (DS) and nonstressed (NS) environments; and (iii) identify drought-resistant dry bean landraces and cultivars. Three landraces and 17 cultivars of race Durango were evaluated in DS and NS environments. Drought intensity index ranged from 0.32 to 0.88. There was a mean reduction in yield of 60% due to drought stress. Also, seed weight was reduced by 14% and maturity was reduced by 4 d in DS. Yield, seed weight, and maturity in NS and DS were positively correlated, and so were seed weight and maturity in NS and DS. But, the DS yield was negatively correlated with seed weight in both DS and NS. The ability to detect yield differences among 20 genotypes decreased with increasing drought stress. Cultivars Viva, NW 63, UI 239, and Common Red Mexican landrace had high yield in both DS and NS and below average reduction in yield due to DS, thus, exhibiting drought resistance. Buster, UI 126, and UI 465 were most susceptible. The frequency and timing of irrigation for the four drought-resistant genotypes should be determined and breeding, genetics, and physiology research intensified to maximize yield and water use efficiency in the western USA, which is facing increasing water shortage.

Abbreviations: DII, drought intensity index • DSI, drought susceptibility index • DS, drought-stressed • NS, nonstressed • PR, percentage reduction due to drought stress


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
THE western USA and southwestern Canada are characterized by inadequate summer rainfall for agricultural production. The agricultural use of water in the western USA is >85% of total water use, moderate to severe intermittent or terminal drought is a common occurrence, and dry bean and most other crops cannot be grown without supplemental irrigation (Cook et al., 2004). Drought also limits adaptation of pulse crops in the northern Great Plains (Miller et al., 2002). Drought is endemic and complete crop failure is common in central and northern highlands of Mexico (Acosta et al., 1999) and northeastern Brazil (da Silveira et al., 1981), each with approximately 1.5 million ha of dry bean (Terán and Singh, 2002). Moderate drought toward the end of the growing season is also common in Central America (Frahm et al., 2004). Moreover, in many areas of the world such as Chile, coastal Peru, most of Europe, northwest Africa, the Middle East, and western Asia, dry bean cultivation is dependent on irrigation.

Consumptive or net water requirement for a 90- to 100-d dry bean crop ranges from 250 to 550 mm depending on soil type, agronomic management of crop, climatic factors, and cultivars (Allen et al., 2000). Drought during emergence and vegetative growth stages reduces plant population and biomass yield. However, in general, dry bean is more sensitive to drought during pre-flowering (10–12 d before anthesis) and flowering stages causing excessive flower, young pod, and seed abortion (Lizana et al., 2006; Nielsen and Nelson, 1998; Pimentel et al., 1999). Thus, the growth stages as affected by intensity and duration of drought determine the extent of losses in seed yield and quality. Drought affects water-use efficiency and plant and seed uptake and utilization of most major and minor nutrients (Muñoz-Perea et al., 2005, 2006, 2007). Drought reduces N partitioning and fixation (Ramos et al., 1999; Serraj and Sinclair, 1998). Drought reduces biomass and seed yield, harvest index, number of pods and seeds, seed weight, and days to maturity (Abebe and Brick, 2003; Muñoz-Perea et al., 2006; Padilla-Ramírez et al., 2005; Ramirez-Vallejo and Kelly, 1998). Moreover, drought increases cooking time and seed protein content on dry weight basis (Pérez Herrera et al., 1999), and charcoal root rot [caused by Macrophomina phaseolina (Tassi) Goid.] in the highlands of Mexico (Mayek-Pérez et al., 2002) and the lowland tropics (Frahm et al., 2004).

There has been no continuous effort to breed dry bean for drought resistance as such in the western USA and southwestern Canada. Nonetheless, the USDA-ARS researchers at Prosser, WA, have used a "purgatory-plot" with general water, nutrient, and root rot stresses and alternate-year dry bean production at Roza, WA, for germplasm screening and selection for decades (Miller and Burke, 1983; P. Miklas, unpublished data, 2006). Consequently dry bean cultivars developed at Prosser such as NW 63 (Burke, 1982c) and Othello (Burke et al., 1995b) carry moderate to high levels of resistance to drought (Muñoz-Perea et al., 2006, 2007), Fusarium root rot (Burke and Miller, 1983), and low soil fertility (Westermann and Singh, 2000). Nleya et al. (2001) found that under a moderate intermittent drought stress in the growth room, indeterminate Othello and determinate ‘Agate’ had high seed yields. However, seed yield differences among six pinto cultivars were nonsignificant (P > 0.05) under severe drought stress.

To characterize and screen dry bean germplasm for drought resistance, researchers have used physiological, biomass, and seed yield traits in drought-stressed (DS) and nonstressed (NS) conditions (Frahm et al., 2004; Pimentel et al., 1999; White et al., 1994a). Early maturity may help escape terminal drought while late maturity, especially in indeterminate cultivars, may facilitate partial recuperation from a mild drought stress during flowering (Nleya et al., 2001). Similarly, the root genotype was more important than shoot genotype under drought stress (White and Castillo, 1989), and greater root growth increased dry bean cultivar efficiency of extracting soil moisture (Sponchiado et al., 1989; White et al., 1990) and nutrients. Cultivar differences in paraheliotropism (light avoiding leaf and leaflet movements, Bielenberg et al., 2003; Pastenes et al., 2004) and other traits associated with light reflectance such as leaf color, cuticular wax, and pubescence may be observed. Similarly, cultivars that tolerate higher temperature and low soil fertility and/or that remobilize carbohydrate and other root and shoot reserves to developing pods and seeds during drought stress should give higher yield and improve adaptation to the western USA and Canada. However, the most reliable integrated measure of cultivar response to drought is still seed yield measured in replicated trials across contrasting DS and NS environments (Abebe and Brick, 2003; Terán and Singh, 2002; White et al., 1994a).

In dry bean, the highest level of drought resistance is found in race Durango dry bean followed by races Mesoamerica and Jalisco (Padilla-Ramírez et al., 2005; Terán and Singh, 2002). Race Durango (synonymous with Gene Pool 5, Singh, 1989) dry bean originated in the semiarid central and northern highlands of Mexico (Singh et al., 1991). Cultivars of race Durango possessing indeterminate, prostrate Type III growth habit (Singh, 1982) also predominate in the western USA and southwestern Canada. The objectives of this study were to (i) measure the effects of drought on seed yield, seed weight, and days to maturity; (ii) determine correlation among the three traits in DS and NS environments; and (iii) identify drought-resistant dry bean landraces and cultivars.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 REFERENCES
 
Seventeen dry bean cultivars released between 1932 and 1999 and three landraces or selection thereof of great northern, pink, pinto, and red market classes were evaluated in DS and NS environments at the University of Idaho in Kimberly and Parma Research and Extension Centers in 2000 and 2001. A randomized complete block design with four replications was used each in NS and DS conditions. Each plot consisted of four rows of 7.62 m length, spaced 0.56 m apart. Furrow irrigation was used to apply five to seven irrigations (approximately 85 mm water irrigation–1) to NS and two to four irrigations to DS plots (Table 1). The groups of NS and DS plots were planted adjacent to each other in the same field separated by a band of eight rows of a drought-resistant dry bean in DS to reduce lateral movement of water from NS to DS plots.


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  		Table 1. Number of irrigation and amount of water applied to three dry bean landraces and 17 cultivars in nonstressed and drought-stressed environments, rainfall, and mean minimum and maximum temperatures from June to September at Kimberly in 2000 and Kimberly and Parma, ID, in 2001.

 
Growth habit was recorded according to Singh (1982) during flowering and verified at the end of pod maturation. Days to maturity was recorded when 90% of the pods changed from green to yellow or beige color. The two central rows were cut at maturity, threshed 8 to 10 d later, cleaned, dried, and seed yield recorded (kg ha–1) at 12% moisture by weight. Weight of 100 seed (g) was recorded. A mixed model (McIntosh, 1983) was used for data analyses where environments (i.e., trial at a location in any year) and replications were considered random and water stress treatments (DS vs. NS) and genotypes (landraces and cultivars) were fixed effects. Data for each environment were analyzed separately and the homogeneity of variances was tested for the combined analyses (Bartlett, 1947). Drought intensity index (DII) for each trial and mean drought susceptibility index (DSI) and percentage reduction (PR) due to drought stress were calculated for each landrace and cultivar according to Fischer and Maurer (1978): DII = [1 – (YDS/YNS)], where YDS and YNS are the mean seed yield of all 20 genotypes under DS and NS environments, respectively. PR = [1 – (YDSg/YNSg)] x 100, where YDSg and YNSg are the mean seed yields in DS and NS environments, respectively, for each genotype. DSI = PR/DII. Also, simple phenotypic correlation coefficients among traits were determined using the mean values in NS and DS for the three environments. All data were analyzed using the SAS (v 9.1.3) PROC GLM statistical package (SAS Institute, 2004). Subsequently, the ANOVA, summary of results tables, and a figure showing classification of cultivars and landraces based on their mean seed yield in DS and NS environments were prepared.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 REFERENCES
 
Mean squares for environment (except for seed weight), treatment, genotype, and their all first-order interactions were highly significant (P < 0.01) for seed yield, seed weight, and days to maturity (Table 2). The mean reduction in seed yield due to drought stress ranged from 32% at Kimberly to 88% at Parma in 2001 (Table 3). The corresponding reduction in mean seed yield at Kimberly was 62% in 2000. Thus, mean seed yield for all genotypes in NS and DS were the lowest at Parma in 2001 compared with the 2 yr at Kimberly. Furthermore, even though one additional irrigation was applied in both the NS and DS plots (Table 1) the drought stress in 2001 was nearly three times more severe at Parma (DII = 0.88) than at Kimberly (DII = 0.32) (Table 3).


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  		Table 2. Mean squares from a combined analysis of variance over three drought-stressed and nonstressed environments for three dry bean landraces and 17 cultivars evaluated at the University of Idaho, Kimberly and Parma Research and Extension Centers, Idaho in 2000 and 2001.

 

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  		Table 3. The year of release, growth habit, seed yield, percent reduction due to drought stress, drought susceptibility index, 100 seed-weight, and days to maturity for three dry bean landraces and 17 cultivars evaluated in three nonstressed and drought-stressed environments at the University of Idaho, Kimberly and Parma Research and Extension Centers, Idaho, in 2000 and 2001.

 
There were large differences for each trait within and across the three environments and between the DS and NS treatments within each environment (Table 3). As shown in Table 3 the mean seed yield in NS over the three environments ranged from 4432 kg ha–1 for ‘UI 239’ (Myers et al., 1997) to 3236 kg ha–1 for ‘Kodiak’ (Kelly et al., 1999a). The corresponding range in DS was 1920 kg ha–1 for NW 63 (Burke, 1982c) and 1112 for ‘UI 126’ (see Dean, 2000). In both NS and DS in the three environments, mean seed yield of cultivars of pink and red market classes tended to be higher than great northern and pinto market classes (Table 3). The great northern ‘UI 425’ had the highest mean seed yield in the NS, but in the DS, ‘Matterhorn’ (Kelly et al., 1999b) had the highest average seed yield. Among pink, ‘Viva’ (Burke, 1982a) had significantly higher seed yield than ‘Harold’ (Burke et al., 1995a) in NS, but differences among the three cultivars were nonsignificant (P > 0.05) in DS. In pinto, ‘NW 590’ (Burke, 1982b) followed by ‘Bill Z’ (Wood et al., 1989) had the highest yield in both NS and DS. Kodiak in NS and UI 126, ‘Buster’, Kodiak, ‘UI 114’, and Common Pinto in DS had the lowest seed yield. The three red cultivars had the highest seed yield in NS and DS among all genotypes. Buster and UI 126 had the highest mean reduction (70%) in seed yield due to drought stress. The cultivars with the least reduction in seed yield included Viva (53%), Harold (54%), and NW 63 (54%). These also had mean DSI values <1.0.

When the mean seed yield in DS was plotted against the NS yield, the three landraces and 17 cultivars fell in four quadrants (Fig. 1 ). Common Red Mexican landrace and six cultivars that yielded well in both DS and NS fell in the top-right quadrant. However, the seven genotypes in the top-right quadrant fell in three sub-groups: UI 239 and Common Red Mexican had the highest mean seed yield in both NS and DS followed by NW 63 and Viva. The cultivars in the third group included NW 590 (relatively higher yield in NS) and ‘UI 537’ (relatively higher yield in DS) at the two extremes and ‘US 1140’ falling between the two. Bill Z, UI 425, and ‘Montrose’ (Brick et al., 2001), although situated in the top-left quadrant, were closer to the latter group of cultivars. Harold, Matterhorn, and Beryl performed well in DS, but did not respond as well as all the above cultivars in NS. The remaining six lower yielding cultivars and Common Pinto landrace in the lower-left quadrant formed three groups: Buster, UI 126, and ‘UI 465’ had near average yield in NS but relatively low yield in DS. In contrast, Kodiak and ‘UI 59’ had low yield in NS but near average yield in DS. Cultivar UI 114 and Common Pinto landrace fell between the two groups.


Figure 1
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  		Fig. 1. Classification of three dry bean landraces and 17 cultivars based on mean seed yield in three nonstressed and drought-stressed environments at the University of Idaho, Kimberly and Parma Research and Extension Centers, Idaho in 2000 and 2001.

 
Pink cultivar Viva and Common Red Mexican had the lowest (28 g) and pinto Kodiak had the highest (41 g) mean 100 seed weight in NS (Table 3). Viva followed by Common Pinto had the lowest and pinto Buster and Kodiak had the highest 100 seed weight in DS. Mean 100 seed weight of pinto market class was slightly higher than that of others in both NS and DS. Mean reduction in 100 seed weight due to drought stress ranged from 0% for Common Red Mexican to 22% for Bill Z. Other cultivars with relatively higher reduction in 100 seed weight included NW 590, UI 114, UI 126, UI 425, and UI 465. In general, most drought-resistant cultivars and landraces had a smaller seed weight reduction due to drought stress.

The mean maturity ranged from 90 d for Common Pinto landrace to 96 d for great northern cultivar UI 425 in NS (Table 3). Common Pinto (82 d) followed by US 1140 (84 d) was the earliest and Buster (93 d) followed by Kodiak and Common Red Mexican (both maturing in 92 d) were the latest maturing dry bean cultivars in DS. The mean days to maturity of all 17 cultivars and three landraces was reduced by 4 d in DS compared with NS.

Seed yield, seed weight, and days to maturity in NS were positively correlated with their respective values in DS (Table 4). The DS yield was negatively correlated with seed weight in both NS and DS. Seed weight in DS was also positively associated with days to maturity in NS and DS. All other phenotypic correlation coefficient values were nonsignificant (P > 0.05).


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  		Table 4. Simple phenotypic correlation coefficients among mean seed yield, days to maturity, and seed weight for three dry bean landraces and 17 cultivars evaluated in three nonstressed and drought-stressed environments at the University of Idaho, Kimberly and Parma Research and Extension Centers, Idaho in 2000 and 2001.

 

   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
REFERENCES
 
The interaction of dry bean cultivars and landraces with treatments (DS and NS) and environments (location/year) suggested that in general, the seed yield of 17 cultivars and three landraces changed significantly from one growing condition to the other with or without changing their rank order. For example, in great northern market class, while UI 59 consistently had the lowest yield in all three NS environments, UI 425 had the highest yield at Kimberly in 2000, but Matterhorn yielded the highest at Kimberly and UI 465 at Parma in 2001. Similarly, UI 425 had the highest seed yield in DS at Kimberly in 2000, but its yield was the lowest at Parma in 2001. On average, Parma has higher mean and maximum temperatures than Kimberly during the summer months because it is at a lower elevation (703 m Parma vs. 1195 m Kimberly). Consequently, mean daily humidity is often lower and evapotranspiration values are higher at Parma than Kimberly. These climatic differences between Kimberly over the 2 yr and Parma in 2001 may have been primarily responsible for large differences in DII values in the three environments (Muñoz-Perea et al., 2006, 2007). Frahm et al. (2004), Schneider et al. (1997), and Terán and Singh (2002) also reported a wide range (from 0.02 to 0.90) in DII values.

The desired level of drought stress or DII value for dry bean germplasm screening and selection for drought resistance will depend on the production region, its climatic conditions, and diversity of germplasm used. Environments with DII values usually lower than 0.50 and hence a milder drought stress, such as Kimberly in 2001, may permit identification of cultivars and landraces with an intermediate level of drought resistance. But, environments with higher DII values are required for identification of the highest levels of drought resistance for unpredictable fluctuating drought conditions in nonirrigation assisted and dryland production regions such as those occurring in the northeastern Brazil and semi-arid highlands of Mexico. Thus, considering the mean seed yield in NS and DS, PR, and DSI values UI 239, Common Red Mexican, NW 63, and Viva would be selected for cultivation in both NS and relatively severe DS conditions. However, caution must be exercised while imposing drought stress because under very severe terminal drought stress the genetic differences among germplasm being screened may not be significant (Nleya et al., 2001). Furthermore, for production regions with irregular rainfall and uncertain irrigation water supply cultivars such as the seven genotypes in the lower-left quadrant in Fig. 1 with relatively lower mean yield in NS and DS, an average or above average reduction due to drought stress (PR value), and relatively higher DSI values would be the least desirable.

It is worth noting that cultivars UI 239 and NW 63 have the Common Red Mexican landrace in their pedigree (Burke, 1982c; Miklas, 2000; Myers et al., 1997). Common Red Mexican was used extensively as a source of adaptation to semiarid environments and resistance to Beet curly top virus (a leafhopper-vectored curtovirus) in the northwestern USA dry bean breeding programs. Furthermore, of 17 cultivars and three landraces evaluated, only Common Red Mexican did not have any reduction in seed weight due to drought stress and it had the smallest change (1 d) in the mean days to maturity between the DS and NS conditions. Common Red Mexican has typical characteristics of race Durango (Singh, 1989; Singh et al., 1991). Its leaves are small, relatively dark, and it often does not produce any guide. Also, the lower internodes of Common Red Mexican are shorter and leaves stay green for a longer period covering the soil surface and conserving moisture around the roots. These traits may reduce evapotranspiration, increase tissue water retention capacity, and facilitate seed filling for longer duration, compared with drought-susceptible cultivars. However, it is not known if these characteristics are linked with or have pleiotropic effects on genes and quantitative trait loci (QTL) determining drought resistance as such in Common Red Mexican.

Padilla-Ramírez et al. (2005), Terán and Singh (2002), and White et al. (1994a, 1994b) identified drought-resistant dry bean landraces Apetito (synonymous with G 13637) and others belonging to common bean races Durango and Jalisco from Mexican highlands. Similarly, San Cristobal 83, a landrace from the Dominican Republic belonging to Mesoamerica race, was also drought-resistant (Terán and Singh, 2002; White et al., 1994a, 1994b). It is not known for sure if these drought-resistant landraces of different evolutionary origins and belonging to different races possess different complementary mechanisms, genes, and QTL for drought resistance. But, White et al. (1994b) found positive general combining ability for seed yield under drought stress between race Mesoamerica and the two Mexican highland races, Durango and Jalisco. Furthermore, Singh (1995) and Terán and Singh (2002) selected high-yielding drought-resistant breeding lines from such interracial populations within the Middle American gene pool. Thus, the use of drought-resistant Common Red Mexican landrace (and others noted above) and cultivars such as Viva, UI 239, and NW 63, and breeding lines such as B98311, L88-63, SEA 5, SEA 13, V 8025, and others (Abebe and Brick, 2003; Frahm et al., 2004; Muñoz-Perea et al., 2006, 2007; Singh et al., 2001; White et al., 1994a) should be maximized. These should be used for anatomical, biochemical, genetical, morphological, and physiological studies to understand the basis of drought resistance and adaptation to semi-arid and arid environments, and to identify and map favorable genes and QTL present in drought-resistant genotypes. The judicious use of these germplasm may also be pivotal for the future development of high-yielding, broadly adapted, drought-resistant cultivars for dryland and irrigation-assisted sustainable production systems, and for determining irrigation schedule and amount of water to be applied to each drought-resistant cultivar to maximize water usage in the western USA and southwestern Canada.

In dry bean, seed weight, color, and shape are important components that determine the recovery percentage (i.e., percentage of good quality marketable seed in the harvested lot) and commercial value of edible bean. Drought stress reduced the mean 100 seed weight by 5 g (i.e., 14%), supporting the findings of Muñoz-Perea et al. (2006), Singh (1995), and Terán and Singh (2002). Thus, cultivars such as Common Red Mexican that did not exhibit any reduction in seed weight in DS should be preferred over those with marked reduction due to drought stress. It may also be worthwhile to determine the genetic basis of seed weight in Common Red Mexican that set it apart from all other cultivars and landraces. Two drought susceptible cultivars, namely Buster and Kodiak that also exhibited little reduction in seed weight, may respond to drought stress by partial seed abortion and repartitioning photosynthate to the remaining seeds within the developing pods.

The mean days to maturity was reduced by 4 d in DS compared with NS. Under a terminal drought stress, similar to the one imposed in this study, days to maturity was also reduced (Terán and Singh, 2002). Furthermore, days to maturity was positively correlated with seed yield in NS and early maturity helped escape terminal drought (White and Singh, 1991). In this study, there was no association between seed yield and days to maturity in either NS or DS. However, days to maturity was positively correlated with seed weight in both NS and DS, suggesting that later maturing cultivars and landraces, on average, had higher seed weight and early maturity reduced seed weight under both NS and DS conditions in race Durango dry bean cultivars and landraces.


   ACKNOWLEDGMENTS
 
The author extends thanks to Marie Dennis, Richard Hayes, and Craig Robinson for assisting with the field trials, and to Henry Terán and Margarita Lema for data analysis and preparation of the tables and the figure.


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 




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