Euphytica 135: 205–215, 2004.© 2004 Kluwer Academic Publishers. Printed in the Netherlands.

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Agronomical and quality traits of runner bean germplasm andimplications for breeding

M. Santalla∗, A.B. Monteagudo, A.M. Gonzalez & A.M. De RonLegumes Breeding Group, Mision Biologica de Galicia, CSIC, P.O. Box 28, 36080 Pontevedra, Spain; (∗authorfor correspondence, e-mail: msantalla@mbg.cesga.es)

Received 4 February 2003; accepted 1 November 2003

Key words: culinary quality, genetic resources, Phaseolus coccineus, phenotypic variation

Summary

White runner bean landraces are greatly appreciated in the North highlands of Spain due to their excellent culinaryseed quality. Runner bean cultivars are grown like pole beans. Diversity within a runner bean collection of 31accessions from the Iberian Peninsula (Spain and Portugal) was examined using morphological, agronomical andseed quality traits. Landraces showed significant differences for most of the agronomical and seed quality traitsstudied except for seeds per pod, water absorption, seed coat tenderness and floury texture. Runner bean landracesshowed sufficient variability to select inbred lines for future breeding. Genotype × environment interaction wassignificant for days to first flowering, days to first dry pod, seeds per pod and seed length. The majority of physicaland nutritional seed quality traits studied which are important to determine the commercial value of a varietywere not subject to environmental influences. Different selection pressures affecting to the runner bean geneticmaterial could have occurred in several regions of the Iberian Peninsula. Extra-large and high yielding runner beangermplasm was identified and represents a valuable source of genetic diversity that has potential for developmentof improved cultivars to be chosen for commercialisation.

Introduction

The scarlet runner bean (Phaseolus coccineus L.),probably the third most important Phaseolus specieseconomically, is a climbing perennial crop but it isoften grown as an annual for dry seeds and immaturegreen pods production. It is also grown as an orna-mental climber. It is of importance in some parts ofEurope, although of minor importance in the UnitedStates (Mullins et al., 1999). In the United King-dom, which seems to be the major consumer of runnerbeans, the unripe pods are usually sliced longitudin-ally or obliquely, then boiled. However, the dry beansare never used. The importance of runner bean in theUnited Kingdom is also reflected by the number ofregistered varieties as compared to those of commonbean (Plant Varieties & Seeds Gazette, 2002). In theNetherlands, young pods or dry seeds are consumed,but the crop is only grown in private gardens (Zevenet al., 1993). South Italy and Spain seem to have pre-

ferred runner bean cultivars, namely climbing cultivarsproducing white seeds (Campion & Servetti, 1991),which are grown commercially on a fairly small scale,while many people grow their own varieties on theirfarms. The cultivar ‘Judión de la Granja’ is a landraceof great commercial value cultivated in the highlandsof Spain. Its culinary qualities, especially the ten-derness of its seedcoat and its buttery texture, haveallowed it to withstand competition with common drybean varieties and ensured its high market value.

This species is native from Mexico, Guatemala andHonduras (Delgado, 1988), and likely wild forms arenot all ancestral to the cultivated form. The area(s)where the domestication took place is still unknown(Debouck & Smartt, 1995). The runner bean wasintroduced into Europe from Central America. Seedexchanges with Europe must have happened since thefirst visits of Europeans to the Americas, which musthave taken the attractively coloured seeds back hometo sow in their gardens on their return (Zeven et al.,

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1993). Spain is supposed to be the country of intro-duction into Europe which is evidenced for the Frenchname ‘Haricot d’Espagne’ referred to runner bean. Itsgaudy inflorescences may be the reason for its recentexpansion as an ornamental plant in Europe. Mix-tures of runner bean and common bean are commonlygrown in Eastern Europe (Poland, Czech and SlovacRepublics and Hungary and probably also in adjacentcountries) (Zeven et al., 1993) and Southern Europe(Spain) as noted by the authors. Natural hybridisationhas not been reported in these countries although nat-ural hybridisation between common bean and runnerbean occur within mixtures in Central America (Wall,1969).

The cross-pollinated runner bean is a source ofvariability for several traits for the improvement ofcommon bean (Gepts, 1981; Singh, 2001; Vander-borght, 1983). This species contains several useful ag-ronomic and disease resistance attributes such as coldtolerance, lodging resistance due to thick stem bases,presence of a tuberous root system allowing a peren-nial cycle, long epicotyls and racemes, a high numberof pods per inflorescence (Vanderborght, 1983), res-istance to Ascochyta blight (Schmit & Baudoin, 1992)and resistance to Sclerotinia sclerotiorum (Gilmoreet al., 2002). The genetic improvement of commonbean through interspecific hybridization requires, asa preliminary step, the characterization of the wholegermplasm collections of the two donor species in or-der to identify the best populations. However, despiteof the potential of runner bean for breeding purposes,germplasm has not been sufficiently evaluated andused to the developing of interspecific breeding lines.Researchers have successfully introgressed from P.coccineus to common bean moderate levels of resist-ance for Xanthomonas (Miklas et al., 1994a; 1994b),for Fusarium root rot (Wallace & Wilkinson, 1965)and for white mold in dry bean (Miklas et al., 1998) aswell as in snap bean (Abawi et al., 1978; Lyons et al.,1987). Wilkinson (1983) suggested that this speciescould be a potential source of high yield for commonbean but practical achievements, in terms of release ofcommercial cultivars, have been few (Hucl & Scoles,1985; Singh, 1992).

A complete evaluation of runner bean landracesshould reveal the existence of potentially valuabletraits useful for improvement of the common beanbecause they are rare or non-existent in the commonbean germplasm. This study may show the most fa-vourable runner bean landraces to merit special preser-vation and selection for commercial use. Developing

cultivars that meet consumer, processor and farmer ex-pectations is a difficult challenge for the plant breeder.Hence, it is important to incorporate culinary qualityevaluations into breeding programs which must simu-late processing practices. The objectives of this studywere to: i) evaluate the available unimproved runnerbean germplasm diversity for morphological, agro-nomical and seed quality traits in different environ-ments, and ii) predict the potential of this germplasmas a source of genetic material for breeding programsand commercial use.

Materials and methods

Plant material

Thirty-one runner bean landraces, that have been col-lected in areas from the Iberian Peninsula (Figure 1)where traditional farming methods have encouragedthe presence of old varieties, were included in thisstudy. Spanish farmers traditionally used to main-tain the crop for several years by leaving the rootsin the field during the winter to produce new plantsagain in the spring although nowadays farmers growtheir crop each year from seeds saved by themselves.This genetic material is maintained in the germplasmcollection at the MBG-CSIC (Misión Biológica deGalicia, Spanish Council for Scientific Research) (Ronet al., 1997) and manual flower pollination is carriedout in net-houses to maintain the integrity and ge-netic variability of each accession and to produce asufficient seed quantity for further studies.

Experimental design

The field experiment was carried out at two locationsin Northwest Spain, Salcedo (42◦ 24’ N, 8◦ 38’ W, 40masl, 14 ◦C mean temperature, average annual rainfall1600 mm) and Soutomaior (43◦ 14’ N, 8◦ 16’ W, 20masl, 13 ◦C mean temperature, average annual rainfall1000 mm) in two consecutive years (2000 and 2001).The landraces were planted in 2-row plots, each 3.8 mlong, in a randomized complete-block design with 2replications. Distance between rows was 0.80 m andplants were spaced 0.25 m apart in the row. Seed washand-sown but over planted by 100% and thinned to 30plants per plot after emergence. Runner bean landraceshad trellis support (2.5 m height) because all of themhave a climbing growth habit. Plants were cultivatedin the absence of any disease or water stress. Fertilizer

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Figure 1. Regions where runner bean landraces were sampled. Accessions in the MBG-CSIC germplasm collection are identified by PHA plusa number.

was applied at planting and irrigation was suppliedwhen necessary.

Data collection

Morphological and qualitative data were recordedwhen the plants reached maximum vegetative develop-ment of the main stem (IBPGR, 1983). The followingmorphological traits were determined on fifteen ran-domly chosen plants per plot and included: leafletlength and width (measured in millimetres on a ter-minal leaflet of a trifoliate leaf and the ratio indicatesthe shape of the leaflet), size of bracteole (measuredin millimetres on freshly opened flowers), both ex-

pressed as the mean of a sample of fifteen leaflets andflowers, respectively. Pod dimensions (suture string,length, width and height expressed in millimetres, andthe ratio height/width which indicates the shape of thepod) and pod weight, determined in grams as the meanof a sample of fifteen pods, were measured when podsreached the optimal maturity stage for fresh consump-tion, that is when the pods have the minimum fibrecontent. In order to obtain a better description of run-ner bean landraces, the qualitative data such as colourand pattern of seed coat, and colour of flower werealso considered.

Agronomical data were measured on the wholeplot (Schachl & De la Rosa, 2001) and included: days

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to first flowering (determined from sowing until oneplant had at least one open flower), days to beginningof flowering (50% plants had at least one open flower),end of flowering (50% plants had flower abscission),period of flowering (beginning until 50% plants hadflower abscission), days to first dry pod maturity (de-termined from sowing until one plant had at least onedry pod), number of pods per plant, number of seedsper pod and seed yield (expressed in grams per plant).

Seed quality data were measured on dried, soakedand cooked bean seeds (IBPGR, 1983; Schachl &De la Rosa, 2001). These included: seed dimensions(length, width and height expressed in millimetresand the ratios length/width and height/width whichare related with the shape of the seed, and they weremeasured on 10 random seeds per plot after drying for72 h at 80 ◦C), dry seed weight (determined in gramson 100 dry seeds per plot), proportion of seed coat(defined as the relation in weight between seed coatand cotyledon plus seed coat, after removing the seedcoat from the cotyledon and keeping them for 24 h at105 ◦C), water absorption (measured as the amount ofwater that dried seeds absorbed during soaking), andnutritional seed traits such as crude protein, starch andtotal sugars, determined on dried material using theNear Infrared Transmittance (NIT) method.

Culinary seed evaluations were performed on a ho-mogeneous sample of each runner bean landrace. Thesamples were soaked for 15 h at 25 ◦C and cookedunder pressure for 5 min. Twelve independent observ-ers evaluated samples of the runner bean landraces onthe following culinary characteristics which conformthe global culinary value: visual appearance (rangedfrom cracked to whole seeds), coat perception (rangedfrom smooth to wrinkled seeds), coat and albumen ten-derness (ranged from tender to tough seeds), creamytexture (ranged from non-creamy to creamy seeds),granulated texture (ranged from non-granulated togranulated seeds) and floury texture (ranged from non-floury to floury seeds) of the cooked beans. Sampleswere also assessed for taste (ranged from non-flavourto flavoured seeds), veining appearance (ranged fromveined to non-veined seeds) and other traits related toglobal culinary value. The rating of bean germplasmfor culinary quality was conducted on 5-point scaleaccording to Sanz & Atienza (2001).

Data analyses

Initially, agronomic and seed quality data for eachenvironment were analysed separately. Then, homo-

geneity of error variances was tested (Bartlett, 1947),followed by combined analysis of variance by a ran-dom model, whereby landraces and environments(locations and years) were considered random effects.Because error variances were heterogeneous for someagronomic traits, data were transformed before tothe combined analysis. Data were analysed with theSAS (2000) statistical package. The least signific-ant difference (LSD) method (p <0.05) was used toevaluate differences between landrace means for thesequantitative traits. Standard errors and coefficients ofvariation were determined for the quantitative traitsstudied. Morphological and qualitative traits were de-termined for each runner bean landrace by using fre-quencies in order to have a good description of thematerial studied. A good agreement was found among12 observers for culinary seed quality characters takenon each sample of each landrace. LSD multiple com-parison test was used on both the original and thestandardized culinary data and standardization did notchange the ranking of the means. Hence, analyses ofvariance were performed on mean culinary values ofthe determination made on each sample. Agronom-ical and dry seed quality characters were analysedby principal component analysis using the NTSYS-pcpackage (Rohlf, 2000).

Results and discussion

The 31 runner bean landraces are classified accordingto the qualitative traits studied as white flowered typesnamed P. coccineus var. albiflorus and red floweredtypes which are named P. coccineus var. coccineus(Zeven et al., 1993). The white flowered landracesproduced white seeds while the red flowered landraceshad either red-purple seeds with few black stripes orpurple seeds with many black spots and stripes. Bassetet al. (1990) demonstrated that purple to black seedcoats and pure red flowers in P. coccineus can only beachieved by having Vwf allele accompany the gene(s)for red flower colour. If those materials were P. vul-garis, they would carry V allele at the V locus and havethe corresponding bluish flower colour but non purered flowers. Thus, runner bean landraces have main-tained the presumed adaptive value of the dark seedcoat colours without sacrificing the presumed adaptivevalues of the pure red flower colours (Basset, 1997).The P. coccineus var. bicolor type was not found in theIberian Peninsula where most of the landraces belongto the P. coccineus var. albiflorus (71%) compared to

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Table 1. Mean squares, mean, standard error (S.E.), coefficient (%) and range of variation for agronomical characters of runner bean landracesgrown at northwestern Spain

Sources of Mean squares

variation d.f.a First End of Period of First dry Pods/plant Seeds/pod Seed yield

flowering flowering flowering pod

days g/plant

Year (Y) 2 2579.1∗∗ 2701.4 6346.9 178.5 18610.4 5.06∗∗ 114501.7

Location (L) 2 1703.6∗∗ 20215.5 6899.1 33096.5 14232.3 0.93∗∗ 110838.5

Year∗Location 1 0.6 16015.2∗∗ 9611.3∗∗ 7820.9∗∗ 3618.9 0.01 16414.7

Replication(Y∗L) 4 26.3∗ 71.8 55.1 314.7∗∗ 1418.6∗ 0.62 10472.3∗Landraces 30 73.3∗ 154.1∗ 219.6∗ 120.8∗ 1207.2∗ 0.78 7314.9∗∗Landraces∗Y 30 13.1 77.4 87.7 77.9 1209.7 0.69 8498.5

Landraces∗L 30 28.7∗ 31.1 52.4 72.6 862.7 0.48 5686.1

Landraces∗Y∗L 30 13.9 44.8 41.8 59.3∗∗ 1155.8 0.92∗∗ 7609.2

Error 120 9.1 36.3 39.1 24.9 722.8 0.38 5035.9

Mean ± S.E. 46.9±1.06 134.1±2.13 83.1±2.21 114.4±1.76 40.3±9.49 3.19±0.219 108±25.1

White group 47.3 134.4 83.2 115.2 41.7 3.13 108

Red-purple group 45.4 131.8 82.9 113.8 35.5 4.12 117

Mixture group 45.7 133.6 84.0 112.5 37.0 3.24 105

LSD (0.05) ns ns ns ns ns ns ns

C.V.b 6.48 4.49 7.50 4.37 66.78 19.41 66.01

Range of variation 39.3–61.7 125.7–166.0 64.0–113.3 109.1–132.3 24.5–83.9 2.67–4.12 29–212

a Degrees of freedom.b Coefficient of variation.c ∗,∗∗ = significant at p ≤ 0.05, p ≤ 0.01, respectively.d ns = not significant at p ≤ 0.05.

the P. coccineus var.coccineus (3%). The P. coccineusvar. bicolor was documented in The Netherlands andin Great Britain (Zeven et al., 1993). The pattern ofvariation for this qualitative trait showed that in somerunner bean landraces (26%) a whole combination offlower colours can be observed and plants with whiteand red corollas occur together, which are maintainedby farmers as one single crop. Runner bean landracesare vigorous perennial vines while bushy varietieswere not found. Leaflets are round and from inter-mediate to long in length, while ovate or lanceolateleaflets were not observed. Bracteoles are intermediatein shape.

Significant differences among the runner beanlandraces were noted for most of the agronomicalcharacters studied (Table 1) except for seeds per pod.Table 1 shows that when the landraces were brokendown into subgroups according to their seed andflower colour (white, red and mixture groups), over-all seeds per pod and seed yield was higher in thered-purple subgroup than in the white subgroup incontrast to pods per plant although the differenceswere not significant. Other authors (Zeven et al.,1993) found in landraces from Hungary that all seed

colour groups produced almost the same number ofpods per plant and seed yield per plant. The desir-able landrace means observed for the measured traitswould lend themselves to directional selection for fur-ther agronomical breeding. In addition, some of theagronomical traits considered were subjected to envir-onmental influences. Of the seven traits studied, oneshowed a significant landrace × location interaction(days to first flowering) and two showed a significantlandrace × location × year interaction (days to firstdry pod maturity and seeds per pod). This means thatthe data related to agronomic aspects can only providea frame of reference, and comparisons are only accept-able in the case of data collected from a common setof experiments.

Significant differences were observed amonglandraces for most of the physical and nutritionalseed quality properties except for water absorption(Table 2). A non significant variation was observed forseed size for subgroups of landraces, and it is assumedthat there is no association between flower colour andseed weight in agreement with Zeven et al. (1993).Average seed weights found in Spanish landraces areslightly higher than values reported in materials from

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Table 2. Mean squares, mean, standard error (S.E.), coefficient (%) and range of variation for dry seed characters of runner bean landracesgrown at northwestern Spain

Sources of d.f.a Mean squares

variation Length Width Height 100 seeds- Water Coat Crude Starch Total

dry weight absorption proportion protein sugars

mm %

Year (Y) 2 84.39 14.54 0.69∗ 20493.85 274.4 0.19 0.68 1.672 1.339

Location (L) 2 3.99 0.49 2.89∗∗ 227.38 86.2 4.87 27.58 0.256 0.872

Year∗Location 1 2.25 1.74 0.01 218.01 1260.3 8.03 10.35 30.472∗ 0.975

Replication(Y∗ L) 4 0.83 0.89∗ 0.33 426.13 476.4∗∗ 3.63∗ 7.95∗∗ 1.591 0.814∗∗Landraces 30 28.18∗∗ 8.81∗∗ 2.56∗∗ 8889.05∗∗ 134.3 6.62∗ 2.11∗ 4.636∗∗ 0.338∗∗Landraces∗Y 30 0.99 0.36 0.19 264.14 106.8 1.77 1.82 1.908 0.152

Landraces∗L 30 1.01∗ 0.42 0.10 279.29 64.3 0.91 1.11 2.694 0.145

Landraces∗Y∗L 30 0.53 0.47 0.18 263.39 107.4 1.65 1.83 2.881 0.201

Error 120 0.63 0.33 0.14 203.43 91.2 1.43 1.27 2.035 0.131

Mean ± S.E. 19.78±0.279 12.12±0.202 8.19±0.131 136.9±5.04 109.9±3.43 9.39±0.421 24.29±0.399 46.95±0.502 4.49±0.131

0.279 0.202 0.131 5.04 3.43 0.421 0.399 0.502 0.1316

White group 19.63 12.17 8.12 138.3 110.7 8.99 24.11 46.62 4.58

Red-purple group 20.46 10.78 7.97 123.5 101.4 12.03 25.55 48.56 4.36

Mixture group 20.09 12.16 8.39 135.1 109.3 10.07 24.66 47.65 4.28

LSD (0.05) ns 0.74 0.40 ns 6.7 0.53 ns ns ns

C.V.b 3.99 4.73 4.52 10.42 8.68 12.72 4.64 3.04 8.05

Range of variation 17.38–26.11 10.49–15.88 7.21–9.76 96.0–266.7 97.1–119.2 7.82–12.03 23.38–25.55 46.10–48.03 3.81–5.10

a Degrees of freedom.b Coefficient of variation.c ∗,∗∗ = significant at p ≤ 0.05, p ≤ 0.01, respectively.d ns = not significant at p ≤ 0.05.

other origins (Zeven et al., 1993), specially for whiteseeded landraces. The largest seed weights observedin some white landraces could be due to farmer selec-tion according to consumer preferences in the Southof Europe, which are for large white seeds, flat orvery flat and slightly ovate in shape. Differences forwater absorption were associated with a more rapiduptake of water and a lower seed coat proportion inthe white subgroup compared to red-purple subgroup.Beninger & Hosfield (1998) observed marked differ-ences for seed physical characteristics among commonbean genotypes with contrasting seed colours. Theseauthors found the lowest ratios of seedcoat and thelargest weights of whole dry bean in the lightest col-oured seed coat genotypes. In addition, seed proteincontent in runner bean landraces was lower in whitelandraces as compared to red-purple landraces. Thecontrary was observed for total sugars, although thedifferences were not significant. Ortega et al. (1978)reported a higher average protein content in colouredcommon bean genotypes than in white ones. The av-erage protein content estimated in the runner bean

landraces (24.29%) was higher than the average valueof 16.33% detected in another reduced set of Spanishrunner bean varieties (Alvarez et al., 1998) but it wassimilar to values reported in materials from other ori-gins (Ortega et al., 1978). Average protein and sugarcontent values are also similar to the values of 22.6%and 4.29% respectively, found for Spanish commonbean varieties (Escribano et al., 1997). In addition, theaverage starch content estimated in the runner beanlandraces was slightly higher than values reported forcommon bean varieties from Spain (Escribano et al.,1997). The high contents of protein and total sug-ars and large seed weights observed in some of therunner bean landraces studied were considered to bedesirable, as would be low levels of starch contentand seed coat. Variability was reported in the runnerbean landraces studied for the physical and nutritionaltraits determined which would permit further qualityimprovement. Furthermore, the majority of physicaland nutritional seed traits considered determinant ofcommercial value were not subject to environmentalinfluences. Of the nine traits studied, one had a sig-

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nificant location effect (seed length). This is relevantto breeding programs as it makes selection in oneenvironment desirable and simple to organise.

No significant differences (Table 3) in coat tender-ness and floury texture were found among the runnerbean landraces studied, although significant differ-ences were observed for taste, veining appearance andthe other traits affecting the culinary global quality.These results suggest that there is variability in theculinary perception of the genetic material studied.Overall results on global culinary quality are similarfor most of the landraces studied except for landracesPHA-0456, PHA-1021 and PHA-1022, which wereconsidered to be inferior to the rest. The inferior globalquality of these landraces was due to a tough albumenperception coupled to non-creamy and granulated seedculinary-perceptual aspects.

A principal component analysis was carried outamong runner bean landraces for agronomical andseed quality traits. Up to 73% of the total variationwas explained by the first three axes, which accoun-ted for 32%, 27% and 14% of the observed variation,respectively. Discrimination along the first principalcomponent was accounted by variation for seed size,along the second principal component by variationfor seed quality related to characteristics such as seedcoat proportion, crude protein, crude starch, total sug-ars and global quality and through the third principalcomponent by variation for seed yield per plant. Thelargest seed size landraces plotted on the right side ofthe graph while the best seed quality landraces plottedon the bottom of the graph (Figure 2A). The best culin-ary perception of the runner bean landraces is due to alow level of seed coat and to the proportions of somechemical components. Thus, the most highly regardedrunner beans are those that contain a higher proportionof total sugars, which makes them sweeter, and due totheir lower content of protein and starch. Runner beanlandraces with a higher starch content had a floury tex-ture. The graph showed extra-large seed germplasmwith a very good quality (landraces PHA-1031 andPHA-1028 with a seed weight of 257 and 267 g/100seeds, respectively), which represent germplasm fromthe highlands, and small seed germplasm (landracePHA-1015 with a seed weight of 96 g/100 seeds),which is cultivated on the Northern Iberian Peninsula.Germplasm with a poor seed quality is also pointedout (landraces PHA-1016, PHA-0456, PHA-1021 andPHA-1022) and these landraces could correspond torunner beans used in food dishes as young pods due totheir long pods.

The differences among landraces by the first andthird principal components (Figure 2B) identified oneextra-large and high yielding landrace plotted onthe right-top side of the corresponded graph whilemedium-seed and high yielding landraces plotted onthe left-top side of the graph. This observation is im-portant because attempts by bean breeders to recom-bine traits such as the large seed size of the Andeangene pool with the yield potential of the Mesoamer-ican gene pool have generally failed (Welsh et al.,1995). Thus, incorporation of runner bean germplasmin interspecific breeding programs may facilitate pro-duction of high yielding common bean lines (Wilkin-son, 1983). A non linear relationship between seedyield and seed size was found, although some outliersemanate on the graph. These outliers correspond tothe landraces PHA-1031, which showed an extra-largeseed size (267 g/100 seeds) associated with a high seedyield (157 g/plant), and the landraces PHA-0352 andPHA-0311, which presented a small-medium seed size(105 and 119 g/100 seeds, respectively) but a very highseed yield (212 and 199 g/plant, respectively). The av-erage yield per plant found in this study was 0.108 kgat the density of 50000 pl/ha. However, the yield perplant in those landraces was quite high as regard toyields published for Mexico. Thus, Reyes & Kohashi(1978) estimated a yield of 1000 kg/ha for Chapingo,and Arias (1980) 3125 kg/ha for Chiapas. In spite ofthe low density used in those studies (6770 pl/ha), theaverage yield per plant was of 0.46 kg. In addition,studies carried out in Budapest (Zeven et al., 1993)revealed an average yield per plant of 0.245 kg, whichwas 3828 kg/ha at the density used (15625 pl/ha).

The study of the morphological, agronomical andseed quality aspects, shows there is variability amongthe runner bean landraces studied. Thus, different se-lective pressures seem to have occurred in severalregions of the Iberian Peninsula. Although, someSpanish germplasm look uniform on the basis of seedcharacteristics, those that farmers use to recognizeand designate landraces and varieties, variation in ag-ronomical and seed quality characteristics has beenobserved in these varieties. This result could be due tooutcrosses among landraces which usually grow closeto each other and to seed interchanges among farmersaccording to seed characteristics which allow for theirrecognition. Studies carried out on Mexican landraces(Escalante et al., 1994) indicated that apparently thedomestication process had not eroded the level of ge-netic variation of P. coccineus and that the similarlevels of genetic variation among wild and cultivated

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Figure 2. Relationship among runner bean landraces from the Iberian Peninsula. A) Principal component plot of seed size and quality diversityof runner bean landraces. B) Principal component plot of seed size and yield diversity of runner bean landraces.

Figure 2. Continued.

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Table 3. Means of the culinary analysis and assessment of degree of taste and appearance of cooked runner bean landraces grown atnorthwestern Spain

Landrace Taste Veining Visual Coat Coat Albumen Creamy Granulated Floury Globala

appearance appearance perception tenderness tenderness texture texture texture quality

PHA-0127 2.60 4.50 3.31 1.02 2.05 1.04 3.14 1.51 2.70 22.5

PHA-0163 2.69 4.17 3.72 1.18 2.11 1.15 3.18 1.27 2.51 23.2

PHA-0166 2.79 2.54 3.36 0.82 2.48 0.93 3.00 1.52 2.70 23.4

PHA-0175 2.34 3.62 3.13 0.83 2.93 1.10 2.80 140 2.60 24.1

PHA-0282 3.10 3.49 3.13 0.88 2.63 1.00 3.35 1.43 2.48 21.8

PHA-0311 2.61 2.36 3.63 1.23 2.30 1.20 3.06 1.60 2.61 25.1

PHA-0322 2.91 3.90 3.15 1.17 2.34 0.97 3.48 1.37 2.70 21.9

PHA-0344 2.73 2.12 3.98 0.96 2.93 0.97 3.02 1.47 2.90 26.8

PHA-0352 2.50 2.66 4.00 1.00 1.30 1.70 3.30 1.50 2.30 22.8

PHA-0406 2.87 3.79 3.28 1.20 2.52 1.22 3.32 1.58 2.36 23.4

PHA-0409 2.80 2.90 2.60 1.25 2.45 0.55 3.20 1.65 2.40 20.4

PHA-0456 2.18 3.77 3.00 2.25 2.92 2.20 2.57 2.65 2.45 33.8

PHA-0469 2.63 4.10 4.07 1.40 2.40 0.87 3.62 1.57 2.12 23.3

PHA-0659 2.56 3.61 3.17 1.37 2.63 0.93 2.97 1.43 2.91 24.9

PHA-0664 2.69 3.25 2.97 1.02 2.54 0.80 2.98 1.70 2.78 23.3

PHA-0669 2.74 1.78 3.93 1.33 3.15 1.20 3.21 1.36 2.48 27.0

PHA-1005 2.37 2.29 3.61 1.22 2.85 1.00 3.02 1.73 2.81 26.8

PHA-1015 3.00 5.00 3.30 1.70 2.00 1.70 3.70 1.30 1.30 20.5

PHA-1016 1.96 4.71 2.02 1.58 2.95 0.85 2.58 2.00 3.10 25.7

PHA-1018 2.66 2.88 3.04 0.88 2.88 1.10 2.97 1.55 3.04 25.2

PHA-1019 2.45 3.17 2.67 1.41 2.23 1.33 3.27 1.57 2.60 22.5

PHA-1020 2.68 4.60 2.92 0.87 2.76 1.00 3.43 1.18 2.58 21.0

PHA-1021 2.06 2.61 3.87 1.52 3.13 1.67 2.62 2.25 2.98 33.6

PHA-1022 2.42 4.25 4.20 1.52 2.75 1.36 2.77 1.76 2.61 29.9

PHA-1023 2.31 4.02 3.51 1.31 2.95 1.26 2.36 1.41 2.78 28.4

PHA-1024 3.60 4.60 3.52 1.17 2.42 0.92 3.90 1.32 2.70 21.7

PHA-1025 2.74 3.32 3.76 1.40 3.06 1.00 3.23 1.56 2.78 27.2

PHA-1027 2.58 3.75 3.05 1.25 2.85 1.55 3.15 1.75 2.50 25.9

PHA-1028 2.68 1.97 2.91 1.32 2.67 1.10 3.17 1.34 2.78 23.5

PHA-1029 2.77 2.70 3.11 1.06 2.50 0.95 3.11 1.83 2.50 23.4

PHA-1031 3.26 3.16 3.57 1.12 2.85 1.30 3.40 1.47 2.37 24.5

LSD (0.05) 0.70 1.18 0.94 0.62 ns 0.60 0.69 0.71 ns 0.73

a Scores above 40 = bad quality; 39–30 = acceptable quality; 29–20 = very good quality; scores lower than 19 = excellent quality.b ns = not significant at p ≤ 0.05.

materials was mainly due to the outcrossing rates inthis species. Data on Spanish varieties (Alvarez etal., 1998) concluded that the genetic variation main-tained after its introduction into Spain could be highalthough the data are too scarce. The germplasm stud-ied represents a valuable source of genetic diversitythat probably would be highly useful for future breed-ing programs. Thus selection within local germplasmwould offer possibilities for the long-term generationof other materials.

The apparent correspondence between superiorcommercial quality and high seed yielding in runner

bean germplasm is noteworthy. In light of the results,the landrace PHA-1031 was considered to have themost favourable characteristics to merit special protec-tion and promotion for commercial use as a large whiteseeded and high yielding runner bean. The landracesPHA-0311 and PHA-0352 were also remarkable fortheir very high yield. There may be a potential marketfor large white seeded runner bean varieties as a sub-stitute for white kidney butter beans, and the range ofnew cultivars for food consumption could be increasedand diversified.

214

Acknowledgements

Research was supported by the projects AGF97-0324 and PGIDT99AGR29102 from the Spanish andGalician Governments, respectively. The authors aregrateful to the Centro de Recursos Fitogenéticos (Min-istry of Science and Technology, Alcalá de Henares,Spain) for supplying some of the bean landraces stud-ied, to the Diputación de Pontevedra and P. Rodiñofor on-farm facilities and to Mercedes Taboada fortechnical assistance.

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