Euphytica 109: 93106, 1999.

1999 Kluwer Academic Publishers. Printed in the Netherlands.

93
Phenotypic variation in a core collection of common bean (Phaseolus
vulgaris L.) in the Netherlands

A.C. Zeven
1
, J. Waninge
1
, Th. van Hintum
2
& S.P. Singh
3
1
Department of Plant Breeding (I.v.P.), Agricultural University, P.O. Box 396, 6700 AA Wageningen, the Nether-
lands;
2
Center for Plant Breeding and Reproduction Research (CPRO-DLO), Center for Genetic Resources, P.O.
Box 16, 6700 AA Wageningen, the Netherlands;
3
Kimberly Research and Extension Center, University of Idaho,
3793 North 3600 East, Kimberly, Idaho 83341, U.S.A. (formerly International Center for Tropical Agriculture
(CIAT), A.A. 6713, Cali, Colombia)
Received 15 December 1997; accepted 12 March 1999
Key words: common bean, core collection, dry-seed type, garden form, landrace, Phaseolus vulgaris, phenotypic
variation, the Netherlands
Summary
Forty accessions, forming a core collection of mainly bush type of the common bean (Phaseolus vulgaris L.)
germplasm in the Netherlands, were evaluated for 14 qualitative and quantitative traits at the Agricultural Uni-
versity, Wageningen (WAU), the Netherlands in 1992. These and an additional 117 Dutch accessions, mainly
collected in private home gardens, were also evaluated for phaseolin seed protein pattern, and morphological
and agronomic traits at the International Center for Tropical Agriculture (CIAT, Spanish acronym), Cali, Columbia
between 1987 and 1997. Multivariate and principal component analyses at both WAU and CIAT indicated existence
of one large group with no discernable patterns among Dutch common bean collections of landraces, garden forms
and cultivars. However, when phaseolin, an evolutionary, biochemical marker, was used as an initial classication
criterion followed by use of morphological markers, the two major gene pools; Andean and Middle American with
two races in each (Chile and Nueva Granada in Andean, and Durango and Mesoamerica in Middle American)
were identied. The Andean gene pool was predominant (136 of 157 accessions), especially the race Nueva
Granada (126 accessions) characterized by the bush determinate growth habit type I and T phaseolin. The new
core collection comprised 31 accessions. Bean races Chile, Durango, and Mesoamerica were represented by 10,
7, and 14 accessions, respectively. Of the 9 French or snap bean accessions six possessed characteristics of race
Mesoamerica and three belonged to Durango race. Occurrence of these and a large number of other recombinants
strongly suggested considerable hybridization and gene exchange between Andean and Middle American gene
pools, thus blurring the natural boundaries and forming a large single group of common bean germplasm in the
Netherlands. The inter-gene-pool recombinants of both dry and French beans should be of special interest to
breeders for use as bridging-parents for development of broad-based populations.
Introduction
Gepts & Bliss (1988) described dissemination path-
ways and introduction of common bean (Phaseolus
vulgaris L.) from its origin and domestication regions
in the Americas into Europe and Africa. Zeven (1997)

Contribution of Idaho Agricultural Experiment Station. Journal

Article No. 99701. University of Idaho, College of Agriculture,
Moscow, ID, U.S.A.
discussed the 16th century and later introductions of
the common bean into West Europe. Already around
1550, various types of common bean have been de-
scribed. In the Netherlands, common beans have been
grown as dry bean and as French bean (i.e., snap beans
for harvesting green pods for vegetables). Landraces
(i.e., unimproved cultigens grown as a eld crop) of
dry bean have disappeared, while some derived lines
from landraces, hereto referred as garden forms, are
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being maintained in home gardens and genebanks.
Many home gardeners still grow their old garden
forms of dry bean. But often unnamed derivatives of
old and new, improved, cultivars are grown, especially
for brown-seeded types. Some bean accessions are
grown both as French bean and as dry bean, the dual
purpose types. The variation existing around 1946
1948 has been described by Nijdam (1947), Anon.
(1949) and summarized by Zeven (1997). Accessions
were grouped as to their seed colour (whitish, brown-
ish, yellowish, black and variegated) types, most of
them being bush erect or semiclimbing beans.
Three major characteristics could be used by
gardeners and farmers as selection criteria. These are
(1) growth habit, (2) seed colour and seed colour
pattern, and (3) use (dry and/or French bean). Any pre-
ferred type would be selected, whereas deviating types
would be removed. Such types could, for instance,
originate from hybridization between dry bean garden
forms and French beans, as most home gardeners grow
both types.
In the period 19751985, a new collection of
common beans grown in home gardens in the Neth-
erlands was made. An early report was written by
Zeven (1979). This material was investigated for the
level of variation of various characters at the De-
partment of Plant Breeding, Agricultural University
(WAU), Wageningen, the Netherlands and at the In-
ternational Center for Tropical Agriculture (CIAT),
Cali, Colombia. The purpose was to determine the
within-accessions and between-accessions phenotypic
variation of unimproved landraces and garden forms,
and improved cultivars and to investigate the pres-
ence of any grouping of accessions, e.g., into landrace
groups (Zeven, 1986), gene pools (Gepts & Bliss,
1985; Khairallah et al., 1990) and/or races (Singh et
al., 1991a). Such a grouping could point to known and
unknown associations between these accessions. Con-
sequently, a new set of core collection was formed.
Material and methods
Research at the WAU
Preliminary study of 223 of 250 initially collected
garden forms and cultivars (see Zeven, 1997, Ap-
pendix 1 for accession numbers) was carried out for
several morphological, physiological, and biochem-
ical (isozymes) characters by Bazelmans & Smits
(1988), Buiteveld (1989), Cool (1989), Veldhuizen
(1991), and van Os (1991). Their results allowed se-
lection of 40 core accessions that were included in
this study. These 40 accessions were again grown
and observed for 14 morphological and agronomic
characters (Table 1) in 1992.
The characters were observed in two replications
each consisting of ve plants. Since the observations
in the two replications were highly correlated (Table 1)
the observations were standardized per replication by
subtracting from each observation the mean score of
the character in the replication and subsequently di-
viding by the standard deviation of the character in the
replication. The resulting standardized observations
were averaged over replications, resulting in scores
with the average 0.0 and standard deviation below
1.0. The lower the correlation between replicates, the
smaller the standard deviation.
For analysis of the phenotypic diversity the 14
quantitative characters (Table 1) as observed in Wa-
geningen in 1992 (data not presented here, they can
be obtained from the rst author) were used. A cluster
analysis was performed using city block distances and
a median sort clustering routine (Payne et al., 1993).
The averaged standardized scores of the 14 charac-
ters were also used to perform a principal component
analysis (PCA). All calculations were performed using
the statistical package Genstat 5 Release 3.1 running
on a VAX computer under the VMS operating system.
Research at the CIAT
Of 250 accessions sent to the CIAT in 1979, 157 acces-
sions were evaluated for biochemical, morphological
and agronomic characters between 1987 and 1997.
Reaction to anthracnose, caused by Colletotrichum
lindemuthianum (Sacc. & Magn.), was observed in
the screenhouse (using a mixture of isolates) at CIAT-
Popayan, and accessions with a resistant score were
again tested separately with Andean and with Meso-
American isolates in the greenhouse. Reaction to bean
common mosaic virus (BCMV) was determined, us-
ing the NL3 strain of BCMV, in the screenhouse at
CIAT-Palmira. Similarly, evaluations against leafhop-
per, Empoasca kraemeri (Ross & Moore), were made
in the elds at CIAT-Palmira, and that for angular leaf
spot caused by Phaeoisariopsis griseola (Sacc.) Ferr.,
in the elds at CIAT-Quilicaho, using mixtures of An-
dean and Middle American isolates of the pathogen
collected locally from the previous cropping seasons.
Each accession was analyzed for their phaseolin seed
protein patterns (Gepts et al., 1986). Data were also
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95
recorded for growth habit (Singh, 1982), shape of
the central trifoliolate leaf, shape of ower bracteole,
ower colour, seed colour, seed brilliance, seed shape,
100-seed weight, days to rst ower and days to ma-
turity. These data were used to obtain further informa-
tion on any classication and to forma core collection.
Multivariate and principal component analyses of the
phenotypic diversity were performed using quantitat-
ive traits, seed weight and phenological characters.
Subsequently, accessions were classied into gene
pools and races, using phaseolin as initial classica-
tion criterion (Singh et al., 1991b) and morphological
markers (Singh et al., 1991a). Finally, data on all 14
traits were used to form a new set of core collection.
Principal component and cluster analyses were
also performed using CIAT data. For cluster analysis
criteria used, listed here in order, were phaseolin type,
ower colour, growth habit, trifoliolate leaf shape,
and bracteole shape, using Wards minimum vari-
ance method. Thus, ve clusters were formed which
explained 86% variance. SAS PROC CLUSTER
(SAS,1988) program was used.
The three traits, namely, 100-seed weight, days to
ower, and days to maturity were used for the prin-
cipal component analysis. The graphical distribution
of common bean races was made using the rst two
principal components and SAS PROC PRINCOMP
program (SAS, 1988).
Results
Wageningen
Table 1 shows the range for the characters used for the
analysis, and the correlation coefcients between rep-
licates. Considerable variation among 40 accessions
was recorded for each of 14 characters. Also, positive
and highly signicant correlations existed between the
two replicates for each character.
Table 2 presents the correlation coefcients
between 14 characters used in the multivariate ana-
lysis. Growth habit was positively correlated with
number of pods per plant, pod height, and seeds per
pod. Flowering time was positively associated with
ripening time, pods per plant, and pod colour intensity.
Similarly, ripening time was positively correlated with
pod length, pod colour intensity, beak length, and seed
weight, length, and height. Positive associations were
recorded between pod length and pod height, seeds
per pod, seed weight, seed length, and seed height.
Table 1. The range and the correlation coefcients between
replicates for characters of 40 Dutch common bean collections
used for multivariate analyses of the phenotypic diversity at
Wageningen Agricultural University
Character Range Correlation
coefcient

Growth habit
a
1 4 0.84
Flowering time (days) 37 49 0.85
Ripening time (days) 55 118 0.89
Pods/plant (number) 11 225 0.79
Pod length (cm) 8 21 0.85
Pod height (mm) 7 13 0.83
Pod colour (02) 0 2 1.00
intensity
Beak curve
b
1 3 0.83
Beak length (mm) 5 20 0.77
Seeds per pod (number) 3 9 0.53
Seed weight (g per 20 71 0.86
100 seeds)
Seed length (mm) 9 19 0.96
Seed width (mm) 5 8 0.80
Seed height (mm) 6 9 0.87
a
1 = determinate, bush and 4 = indeterminate climbing (ac-
cording to Singh, 1982).
b
1 = straight and 3 = curved.

All values signicant at p <0.01.

Seed weight was highly correlated with its three other
components.
Table 3 shows the percentages of variation for
the rst three PCs and the vector loadings per char-
acter and PC. The rst PC separates the accessions
mainly on seed (weight, height, and length) and pod
(height and length, colour intensity, beak curve and
length) characteristics whereas the second PC separ-
ates mainly on growth habit, pods per plant, and seed
length. The third PC separates mainly on owering
time, pods per plant, pod length, seeds per pod and
seed width.
Figure 1 presents the dendrogram of 40 bean ac-
cessions based on phenotypic diversity. Figure 2 gives
the distribution of the accessions on the rst two prin-
cipal components (PCs) based on phenotypic diversity.
Indicated are accession number and seed colour. As
it can be seen from these two Figures, despite differ-
ences in growth habit, pod, seed, and phenological
characteristics, Dutch common bean collection, on
the whole, forms one large group with no specic
discernable patterns.
euph4747.tex; 10/08/1999; 17:05; p.3
96
Table 2. Correlation coeecients among characters measured from 40 Dutch common bean collections and used for multivariate
analyses of the phenotypic diversity at Wageningen Agricultural University
Character Character
2 3 4 5 6 7 8 9 10 11 12 13 14
1 Growth habit 0.5 0.26 0.59

0.16 0.43

0.2 0.14 0.19 0.32

0.04 0.14 0.13 0.17

2 Flowering time 0.66

0.50

0.30 0.14 0.31

0.10 0.11 0.25 0.12 0.23 0.11 0.14

3 Ripening time 0.25 0.31

0.26 0.36

0.23 0.33

0.08 0.40

0.48

0.02 0.38

4 Pods/plant 0.9 0.29 0.00 0.14 0.03 0.25 0.06 0.5 0.09 0.13
5 Pod length 0.35

0.17 0.30 0.11 0.50

0.57

0.69

0.13 0.40

6 Pod height 0.22 0.47

0.31

0.11 0.53

0.23 0.49

0.71

7 Pod colour intensity 0.46

0.31

0.33

0.18 0.2 0.33

0.26
8 Beak curve 0.46

0.07 0.55

0.32

0.49

0.47

9 Beak length 0.04 0.45

0.23 0.50

0.37

10 Seeds per pod 0.7 0.09 0.03 0.03

11 Seed weight 0.81

0.48

0.88

12 Seed length 0.05 0.57

13 Seed width 0.62

14 Seed height

Signicant at p < 0.05 and p < 0.01, respectively.

Table 3. Vector loadings and percentages of explained vari-
ation by the rst three principal components of 40 Dutch
common bean collections evaluated at Wageningen Agricul-
tural University
Principal components
1 2 3
Variation explained (%) 34.16 14.62 14.34
Growth habit 0.070 0.563 0.150
Flowering time 0.178 0.067 0.505
Ripening time 0.274 0.282 0.264
Pods/plant 0.120 0.376 0.373
Pod length 0.281 0.131 0.309
Pod height 0.325 0.273 0.067
Pod colour intensity 0.216 0.124 0.040
Beak curve 0.316 0.120 0.157
Beak length 0.250 0.042 0.217
Seeds per pod 0.064 0.249 0.316
Seed weight 0.407 0.151 0.151
Seed length 0.312 0.417 0.081
Seed width 0.237 0.276 0.423
Seed height 0.398 0.023 0.179
CIAT
Growth habit
The numbers of accessions divided over the four
growth habit classes (Singh, 1982) were 133 de-
terminate type I; 5 indeterminate upright type II; 18
indeterminate prostrate, semiclimbing-type III; and
2 indeterminate strong climbing type IV. The latter
group of two accessions were snap beans. Three of the
5 upright type II accessions possessed S phaseolin and
2 had T phaseolin and all were dry beans. Among type
III accessions 10 possessed T or H
1
phaseolin and 7
had S phaseolin.
Leaf shape
The ovate shape of the trifoliolate leaves was the most
predominant, represented by 125 of 157 accessions
studied. This was followed by the hastate types pos-
sessed by 28 accessions. The cordate leaf shape was
exhibited by only 4 accessions three of which pos-
sessed T phaseolin and were of determinate growth
habit type I, and one had S type phaseolin and growth
habit type III.
Bracteole shape
All four shapes of ower bracteole (Singh et al.,
1991a) were observed. The lanceolate and ovate
shapes were the most predominant types with 69 and
67 accessions possessing these bracteole shapes, re-
spectively. The triangular and cordate types were,
respectively, represented by 12 and 9 accessions. The
most interesting feature to note was that some of the
triangular (previously recorded only in race Chile) and
cordate types (more common in race Mesoamerica)
(Singh et al., 1991a) were found in accessions pos-
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97
Figure 1. Dendrogram of 40 Dutch common bean accessions on phenotypic diversity estimated from 14 characters evaluated at the Agricultural
University, Wageningen, the Netherlands, in 1992
sessing type I phaseolin and determinate growth habit,
characteristic of race Nueva Granada.
Flower colour
Three types of ower colour were recorded: 101
white, 41 pink, and 8 accessions with purple coloured
owers. In addition, 7 accessions were variable (i.e.,
possessed mixtures of either pink and purple, white
and purple, or white and pink). Flower standards in 11
accessions had a blotch or were spotted. Similarly, one
accession had stripes on the lower outer surface of the
ower standard, and the standard of one accession pos-
sessed small spots. All these accessions with blotch,
stripes, or spots on ower standard possessed either
T or H
1
type phaseolin, a characteristic of Andean
common bean gene pool. All white seeded acces-
sions investigated had white owers. However, white
owers may also occur in accessions with coloured
seeds.
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98
Figure 2. Distribution of 40 Dutch common bean accessions on the rst two principal components based on phenotypic diversity estimated
from 14 characters evaluated at the Agricultural University, Wageningen, the Netherlands, in 1992; indicated are accession number and seed
colour.
Seed colour and seed colour pattern
Although 15 seed colour and colour patterns were
found among 157 accessions the most predominant
were white, beige and cream mottled. In addition
some of the medium and large white seeded acces-
sions had sectorial spots or spots around the hilum.
About 10% accessions had grey seed colour. The seed
colour classied at CIAT may differ somewhat from
the classication made at Wageningen. For example,
the brown and yellow colours recorded at WAU may
correspond to the beige and sulfure colours of CIAT,
respectively.
Seed brilliance
Three classes of seed brilliance were recognized: 15
opaque, 43 shiny or brilliant, and 99 semi brilliant.
There was no association between seed shape or seed
weight and brilliance. Similarly, there was no associ-
ation between seed colour or seed colour pattern and
degree of seed brilliance.
Seed shape
The oval seed shape was the largest class represented
by 61 accessions. This was followed by the kidney
shape with 52 accessions and cylindrical types with
31 accessions. There were 14 accessions possess-
ing rhombohedric seed shape. All these possessed T
type phaseolin and thus could be classied as An-
dean and not to Middle American gene pool. Had
they possessed S, Sd or B phaseolin they would be
classied into Middle American gene pool, especially
race Durango, because it often is the distinguishing
characteristic of this group of landrace cultivars from
highlands of Mexico (Singh et al., 1991a).
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99
Phaseolin seed protein patterns
Of 157 accessions 132 possessed T, 19 S, and one
H
1
phaseolin. Five accessions could not be clearly
dened. All accessions possessing S phaseolin seed
protein pattern are classied into Middle American
gene pool. Similarly, all T and H
1
types were con-
sidered of Andean origin. Of the ve accessions
whose phaseolin types could not be determined by
one-dimensional electrophoresis three possessed mor-
phological marker traits of Andean germplasm, and
two those of Middle American gene pool.
Number of days to ower and maturity
Both number of days to ower (when the rst ower
opened in the accession) and maturity (harvest ma-
turity) were recorded. A great majority of accessions
owered in 32 or fewer days. Only seven accessions
owered in 34 days, and only one owered in 46 days.
The latter two groups were indeterminates of growth
habit types III or IV, all except one were from races
Durango and Mesoamerica. Fifty percent of the acces-
sions (79 out of 157) were early maturing (i.e., took
65 or fewer days to mature). Of these 28 matured in
60 days. All early maturing accessions, except six ac-
cessions fromrace Chile of growth habit type III, were
determinate growth habit type I.
Bean common mosaic virus
In 157 common bean germplasm collections, 12 ac-
cessions (7.6%) were resistant, and 145 susceptible
(92.4%), upon inoculation with the NL3 strain of the
bean common mosaic virus (BCMV). Of 12 resistant
accessions, seven were of Andean gene pool, because
they possessed T phaseolin. Also, they were of de-
terminate growth habit type I (Singh, 1982). All other
ve accessions resistant to BCMV were snap bean
cultivars; one of these possessed determinate growth
habit type I, and two each were of growth habit types
III and IV. All ve possessed S phaseolin and thus
were of Middle American gene pool.
Anthracnose
Of 157 accessions 4 could not be screened for an-
thracnose reaction. A comparatively large number of
the accessions (111 out of the 153) tested showed res-
istant or intermediate reaction to anthracnose in the
elds as well as in the screenhouse evaluations against
the mixture of Andean and Middle American isolates
of the pathogen populations found at CIAT-Popayan,
Colombia. Based on the phaseolin phenotype, propor-
Table 4. Vector loadings and percentages of explained
variation by the rst three principal components of 30
Dutch common bean core collection evaluated at CIAT
between 1987 and 1997
Principal components
1 2 3
Variation explained (%) 53.5 31.5 15.0
100 seed weight 0.33 0.93 0.18
Days to ower 0.69 0.11 0.72
Days to maturity 0.65 0.36 0.67
tionally higher number of accessions from the Middle
American gene pool were susceptible.
Angular leaf spot
Unlike anthracnose, there was only a small number
(i.e., 14 out of 157) which showed intermediate re-
action (scores of 4 to 6) to angular leaf spot, caused
by Phaeoisariopsis griseola (Sacc.) Ferr. No acces-
sions being resistant (scores 3) were found. The 22
accessions could not be evaluated, whereas 121 were
susceptible.
Leafhopper
Reaction to leafhopper (Empoasca kraemeri Ross and
Moore) was scored under eld conditions at CIAT-
Palmira. Thirteen accessions had an intermediate re-
action and 144 were susceptible. No resistant (i.e.,
receiving scores of 3 or lower) accessions were ob-
served. Among the accessions with an intermediate
reaction 10 were determinate growth habit type I, and
3 were of growth habit type III. All 10 accessions were
of the Andean gene pool.
Principal component analysis
The rst two principal components explained 84.9%
of the variance. Such high percentages could be ex-
pected since the PCA was based on three traits only.
The rst and third principal components separated the
30 Dutch core accessions of common bean mainly
on phenological traits, days to owering and maturity
(Table 4). The second PC separated them largely on
the basis of 100-seed weight.
Gene pool and race classication
From multivariate and principal component analyses
of quantitative traits (e.g., days to owering and ma-
turity, 100-seed weight) it was not possible to identify
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100
Figure 3. Distribution of 157 Dutch common bean accessions on the rst two principal components based on phenotypic diversity estimated
from 100-seed weight and days to owering and maturity recorded at CIAT-Palmira, Colombia, in 1997; indicated are common bean races
Chile (C), Durango (D), Mesoamerica (M), and Nueva Granada (N).
any groupings or patterns among 157 accessions (Fig-
ure 3) evaluated at CIAT. This supported the conclu-
sions drawn at WAU (Figures 1 and 2). Nonetheless,
when phaseolin seed protein, an evolutionary marker
(Gepts, 1988; Gepts & Bliss, 1985; Gepts et al., 1986)
was used as the initial classication criterion (Singh
et al., 1991b), 136 accessions were classied as of
Andean and 21 as of Middle American gene pools
(Table 4). Thus, the Andean gene pool was the largest
one, being more than six times that of the Middle
American gene pool. By using morphological mark-
ers such as shape of the central trifoliolate leaet and
ower bracteole, presence or absence of the stripes at
the outer surface of ower standard, and seed shape
and weight, it was possible to divide both Andean
and Middle American gene pools into races (Singh et
al., 1991a). For example, among the Andean germ-
plasm 10 were of the race Chile and 126 that of Nueva
Granada race. None belonged to race Peru. Of 21
germplasm accessions of Middle American gene pool
14 were from race Mesoamerica and seven of race
Durango. The race Jalisco was not represented.
The core collection
Since the 40 accessions forming the core collec-
tion initially identied at WAU were also included in
the CIAT evaluations, and phaseolin, morphological
markers and other agronomic data were obtained ad-
ditionally, attempts were made to identify redundancy
among 40 accessions and to forma newcore collection
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101
Table 5. Classication on biochemical (phaseolin seed protein) and morphological markers

at CIAT, Colombia, of 157

Dutch common bean collections into gene pools and races. The number of accessions forming each gene pool or race are
given in parenthesis
Gene Race Phaseolin Growth Leaf Bracteole Seed Seed Bean
pool type habit shape shape shape size type
Andean Nueva T I Ovate Ovate Cylindrical Large Dry
(136) Granada II Hastate Lanceolate Kidney Medium French
(126) III Cordate Triangular Oval Small
Cordate Rhombohedric
Chile H
1
I Ovate Ovate Oval Large Dry
(10) T III Hastate Lanceolate Medium French
Cordate Triangular Small
Cordate
Middle Durango S III Ovate Ovate Rhombohedric Large Dry
American (7) Hastate Lanceolate Kidney Medium French
(21) Cordate Triangular Oval Small
Cordate
Mesoamerica S I Ovate Ovate Kidney Large Dry
(14) II Hastate Lanceolate Rhombohedric Medium French
III Cordate Triangular Oval Small
IV Cordate Cylindrical

There is no linkage among characters within rows across columns. Together they refer to the characteristics of the common
bean race.
(Table 5). This newset of core accession comprised 22
accessions that were common to the group of 40 core
collection identied initially, and 8 new accessions.
Hopefully, the 30 accessions included in this core col-
lection should represent the genetic variation present
among 250 accessions collected between 19751978
in the Netherlands. Since most of the genebank ac-
cessions prior to 1940 were lost during the World
War II, and there has been considerable changes in
farming systems and eld crops grown since then, it
will never be known whether or not the 250 acces-
sions initially collected or the core collections listed
in Table 4 are the true representation of the genetic
variation that might have existed in common bean in
the Netherlands. However, many accessions collected
in home gardens were inherited from parents and
grandparents.
Discussion
In both the results of the cluster analysis (Figure 1)
and principal component analysis (Figure 2), no clear
groups among the 40 core accessions evaluated at
WAU or in 157 accessions evaluated at CIAT (Fig-
ure 3) could be distinguished. As the 40 core ac-
cessions represent groups obtained by Bazelmans
& Smits (1988), Cool (1989), Buiteveld (1989),
Veldhuizen (1990) and van Os (1991), and from
over 50% of total Dutch collections evaluated in
CIAT, we conclude that all dry-bean accessions of
the 19751985 collection, including bush and climb-
ing landraces, garden forms and, may be, cultivars,
adapted to the Dutch environments form one large
group. Maybe, after the, probably, many introductions
of landraces and garden forms into the Netherlands
(and elsewhere in Europe) they hybridized with each
other exchanging genes and blurring genetic boundar-
ies. But growers wishing to describe their material use
characters of dry-bean types such as plant habit, seed
colour, seed colour pattern and use.
Both the Kogelboon accessions (accs. 2.08,
2.09), Transvaalse Bonte met draad (acc. 3.13) and
the Citroenboon (acc. 4.05) are identied as Middle
American accessions. However, Gepts & Bliss (1988)
scored both the Kogelboon accessions as having the
T phaseolin phenotype, which points to an Andean
origin. In any case, both Kogelboon accessions re-
semble each other and may be closely related. In
the upper part of Figure 2 the two Walcherse Witte
euph4747.tex; 10/08/1999; 17:05; p.9
102
accessions occur. Walcherse Witte (acc. 1.03) is de-
rived from the landrace of that name, formerly grown
in the province of Zeeland, whereas Walcherse Witte
Blanka (acc. 1.04) was collected in a part of the
province of Drenthe, where inhabitants from Zeeland
had settled some 150 years ago (Zeven, 1997). Both
accessions belong to the Middle American gene pool;
Walcherse Witte (acc. 1.03) is scored as belong-
ing to the Mesoamerica race, and Walcherse Witte
Blanka (acc. 1.04) to the Durango race. The latter
was scored by Gepts & Bliss (1988) as possessing
the S phaseolin phenotype, which also indicates that
this accession originally came from Middle America.
Their near-location in Figure 2 and the fact that they
both have a Middle American origin supports the fact
that Walcherse Witte and Walcherse Witte Blanka
are related. So, the latter accession very likely derives
frommaterial introduced fromthe province of Zeeland
by the Zeeland settlers. Furthermore, at that time, the
landrace Walcherse Witte must have included plants
of the Mesoamerica and the Durango races. Buiteveld
(1989) also found a near-resemblance of the above two
accessions of Walcherse Witte and a third accession
(acc. 1.20) of this landrace.
The accessions aff. Berna (accs. 2.07, 2.16, 2.17)
are named as such as they arrived in our collection
without a name, but as the seeds resemble those of
the cultivar Berna we named them aff(inity) Berna.
Unfortunately, Berna was not among the 40 repres-
entative accessions. The three accessions are closely
situated to each other in Figure 2. Accessions 2.07
and 2.16 belong to the Andean gene pool, whereas
acc. 2.18 belong to the Andean race Nueva Granada.
Berna is a BC2 of the recurrent parent Beka, which
is selected from the landrace Noord-Hollandse Bru-
ine. Beka belongs to the Andean Nueva Granada
race. This points to an Andean origin of Noord-
Hollandse Bruine.
The two (climbing) Stokkievitsboon accessions
(accs. 3.17, 3.18) both come from the same region.
These two accessions with four other Kievitsboon-
accessions (accs. 3.07, 3.14, 3.15, 3.46) are located
at the left side of Figure 2. This is caused by their
heavy seed weight. One of the six accessions belongs
to the Andean gene pool and two to the Nueva Granada
race, whereas Gepts & Bliss (1988) scored acc. 3.07 to
possess T phaseolin. The (small-seeded) Kleinzadige
Kievitsboon is situated more to the center of Figure 2.
It belongs to the Nueva Granada race. This inform-
ation points to an Andean origin of the (bush and
climbing) (large- and small-seeded) Kievitsboon ac-
cessions. Transvaalse bonte met draad (with string,
acc. 3.13) belongs to the Mesoamerica race and pos-
sesses the Middle-American T phaseolin. Its coun-
terpart Transvaalse bonte zonder draad (without
string, acc. 3.13) belongs to the Andean Nueva
Granada race. This points to a Middle-American and
an Andean origin respectively of these accessions. If
this is the truth their similar appearance and name are
in contradiction with their separate provenance.
Roodbonte Krobbe (acc. 3.23) also has an An-
dean origin, whereas the race of Kleine Krobbe
(acc. 3.54) has not been identied. The fact that iden-
tication was difcult and its position in Figure 2
could indicate a hybrid origin of an Andean and a
Middle-American accession.
The six Strogele accessions (accs. 4.06, 4.07,
4.12, 4.13, 4.30, 4.31) are grouped together. Gele
boon (acc. 4.27) also belongs to this group. This may
mean that their variation for the characters investigated
is low. All belong to the Andean gene pool. This points
to an Andean origin.
The yellow-seeded beans Gele Krobbe (acc.
4.26) and Citroenboon (acc. 4.05) are clearly sep-
arated from the Strogele accessions. Gele Krobbe
belongs to the Andean gene pool, and Citroenboon
to the Middle American gene pool. Each must have
had its own breeding history.
The three Soldatenboon- and two black-seeded
accessions are scattered about Figures 1 and 2. This
points to a separate breeding history of each of them.
Thirteen accessions could not be identied at the
CIAT. The cause could be that they are hybrids
between accessions of both races. So, identical to the
origin of the above-mentioned Kleine Krobbe. The
place and time of hybridization is unknown. A re-
cent hybridization event lead to the presence of the
segregating accession Gevlekte Moddermannen.
The cause can be as stated in the paragraph on
growth habit that during composing the collection
bush types and hence, unconsciously, a certain gene
pool was selected for.
At CIAT, phaseolin seed protein pattern (Gepts,
1988; Gepts & Bliss, 1985, 1988; Gepts et al., 1986),
shape of the central trifoliolate leaet and ower brac-
teole, presence or absence of stripes at the outer base
of ower standard, and seed shape, were used to clas-
sify the Dutch common bean germplasm into Andean
versus Middle American gene pools, and subsequently
into races (Singh et al., 1991a). Furthermore, the pres-
ence or absence of bers in the pod wall and sutures
helped us identify the French, snap, or green beans
euph4747.tex; 10/08/1999; 17:05; p.10
103
from dry beans. Of 157 accessions thus characterized
at CIAT, 136 were of Andean origin or gene pool and
21 belonged to Middle American gene pool. Among
the Andean types, bush determines of Nueva Granada
race possessing T phaseolin were most predominant,
only 10 were fromrace Chile and none fromrace Peru.
It could be suggested that with time more Andean
accessions reached the Netherlands, than Middle-
American ones, that Andean accessions are better
adapted to the Dutch environment, or that people se-
lected for some reasons for Andean types or against
Middle American types. Or there were more contacts
with South America (e.g., Dutch Brazil and Surinam),
than with Middle America. The same suggestions
could not hold to explain the distribution of accessions
within a gene pool and for the absence of acces-
sions belonging to the Andean Peru race and Middle
American Jalisco race, which are highly photoperiod
sensitive and are characterized by a strong climbing
habit and adaptation to over 2000 m elevation in the
tropics and subtropics of Latin America.
As it was intended to set up a collection of bush
dry bean accessions, it is not surprising that most
of the accessions (80.9%) have the bush determin-
ate growth habit type I. Furthermore, accessions such
as Walcherse Witte which were classied in growth
habit type III were considered by the farmers as bush
types. The cause was that plants of this landrace were
formerly grown on poor soil where they developed
sub-optimally, i.e. they did not reach growth habit type
III, but stopped growth after reaching a growth habit
equivalent to types I or II (H. Lamberts, pers. comm.,
1975).
From close examination of seed weight it was
evident that in Andean germplasm there had been se-
lection for smaller seed size after initial introduction
into the Netherlands of common bean from the Amer-
icas. In the center of their domestication, in Andean
South America, the 100-seed-weight of the landraces
and garden forms from both races Chile and Nueva
Granada, are often higher than 40 g. Moreover, occur-
rence of dozens of accessions possessing T phaseolin
but rhombohedric seed shape (a characteristic of race
Durango of Middle America), less than 25 g/100
seed weight (often characteristic of race Mesoamer-
ica), cordate trifoliolate leaet shape, and/or cordate
shape of ower bracteole provided strong evidence
for recombination and introgression of Middle Amer-
ican alleles into Andean germplasm. Since formation
and development of large seeds would require com-
paratively longer period, especially in environments
with cool summer nights such as occurring at higher
latitudes (including the Netherlands) and large seeds
are undesirable for French beans, any selection for
earliness and/or for dual purpose harvests would have
resulted in gradual reduction in seed weight.
The nine strong climbers (growth habit IV) are
Witte Stokboon (acc. 1.38), Beige Gele Boon (acc.
4.25 ex Germany), Wulleboon (acc. 1.55), Veense
Snijboon (accs. 1.58, 1.71), Snijboon (acc. 1.66),
and Franse Prinses (acc. 1.78). Most of them are of-
ten grown as French bean. Two variable accessions are
Bruine Kogelboon (acc. 2.30) and Witte Sabelboon
(acc. 1.56).
Based on research at Wageningen it was concluded
on other grounds that both Kogelboon accessions
may have come from the transition zone between the
Andean or Middle-American areas of origin. This
conclusion is supported by the above mentioned clas-
sication. Many (Stro)gele accessions also scored as
either of Andean or Middle-American origin. Maybe
both the Kogelboon and (Stro)gele accessions ori-
ginate from this transition zone. The other accessions
could originate from the same transition zone, or are
interracial hybrids.
Only one accession viz. Witte boon (acc. 1.79)
is classied as Middle/American group/Andean pod
beak type. Maybe it also came from the above trans-
ition zone or it is a hybrid of an Andean and a
Middle-American accession, originated in the New or
Old world.
Although no efforts have been made to systemat-
ically characterize snap or French bean germplasm,
and trace back their origins, most of the popular cul-
tivars of French bean are believed to be of Andean
origin. Thus, occurrence of six French bean acces-
sions possessing characteristics of race Mesoamerica
and three French beans possessing characteristics of
race Durango including S phaseolin protein pattern
is the rst report of this kind, to the best of our
knowledge. While it can not be known with any de-
gree of certainty whether these French beans were
originally introduced from the Americas or else de-
veloped locally from hybridization between Andean
and Middle American gene pools, they should be of
immense value as a diverse source of photoperiod in-
sensitive germplasm for future breeding programs. If
these and over four dozen other accessions of dry bean
types marked as recombinants are proven to be of
hybrid origin, derived from Andean Middle Amer-
ican inter-gene-pool crosses, these may be of special
interest and may serve as bridging parents in future
euph4747.tex; 10/08/1999; 17:05; p.11
104
inter-gene-pool crosses because they may have re-
duced problems of incompatibility often associated
with such crosses (Singh & Gutirrez, 1984; Singh &
Molina, 1996; Gepts & Bliss, 1985; Kornegay et al.,
1992).
A great majority of accessions possessed shiny or
semishiny seeds, very few were opaque. It could be
imagined that this character attracted the special atten-
tion of the farmers, gardeners and breeders, during the
seed picking. Hence, one could assume that brilliant
seed attracted the attention of the seed picker. No clear
separation for seed colour was found.
Most of the BCMV resistant accessions are either
French bean (princesseboon) or come from a few
provenances. Those from Rockanje and Venray come
from small-scale breeders of French beans, who ap-
parently selected for BCMV-resistance. This may
also be the cause of the varying accessions 1.71
(Veense Snijboon) and 1.72 (Soepboon) from Ven-
ray. Most of tolerant accessions are either white- or
black-seeded.
One wonders whether the occurrence of higher
number of accessions of Andean gene pool possess-
ing resistance to anthracnose was an indicative of the
fact that at CIAT-Popayan, Colombia, freely occurring
isolates of the anthracnose pathogen are largely that of
the Middle-American types or that pathogen popula-
tions of South American types are less virulent. Partial
support for the former could be obtained from the fact
that proportionally more Middle American germplasm
were susceptible to anthracnose.
Leafhopper resistance is of no importance in the
Netherlands. Genes conditioning intermediate resist-
ance may still be present because of the lack of any
negative selection pressure for these genes. From re-
peated evaluations of several thousand accessions of
common bean at CIAT it is known that moderate levels
or intermediate resistance to leafhoppers mainly oc-
cur in a few small-seeded accessions belonging to
the Middle American gene pool, i.e. the Mesoamer-
ica race. Thus, this identication of some Andean
germplasm with intermediate resistance against leaf-
hopper should be of special value to breeders in the
tropics and subtropics of Latin America and Africa.
A general lack of resistance to angular leaf spot
might be a function of lack of the occurrence of the
pathogen in the Netherlands. But one would have
expected much higher frequency of accessions with
resistance to BCMV as it was the case with an-
thracnose. It is believed that both anthracnose and
BCMV were severe problems endemic on common
bean in Western Europe including the Netherlands
(Hubbeling, 1957, 1969), France and Spain. Since
anthracnose affected pods and seeds have an ugly ap-
pearance and low market value there would have been
strong selection against susceptible genotypes by the
home gardeners. Moreover, since considerable breed-
ing for anthracnose was carried out in the country
(Hubbeling, 1957; Kruger et al., 1977; Mastenbroek,
1960) it is likely that the comparatively higher fre-
quency of anthracnose resistant accessions is probably
reection of that work. And owing to relatively more
recent research on BCMV that was done about the
same time when germplasm was collected (Drijfhout,
1978; Drijfhout & Bos, 1977; Drijfhout et al., 1978;
Hubbeling, 1969), its signicant impact, therefore,
could not be observed at the production level.
As most accessions are pure, a small sample (12%
of original collection of 250 accessions) of 30 core ac-
cessions should represent the genetic variation found
among Dutch common bean germplasm for most bio-
chemical, morphological, and agronomic traits. These
may be of special interest for those researchers who
for some reason could not study all 250 accessions.
Also, the initial screening of these core accessions
could provide a clue as to in which group of landraces
and garden forms to search for more useful variation.
Thus, comparatively larger quantities of seed of these
core accessions may need to be kept for distribution to
frequent users.
Tolerant accessions for BCMV are Witte boon
(acc. 1.45, 1.47, 1.55), Kookboon (acc. 1.64), Bru-
ine Gevlekte Moddermannen (acc. 1.50), Soepsnij-
boon (acc. 1.67). All three Walcherse Witte acces-
sions (accs. 1.04, 1.20, 1.41) are sensitive to BCMV.
This supports the assumption that this landrace was
wiped out on the Island of Walcheren after the large
scale introduction of the propagation cultivation of
gladiolus (Zeven, 1997). Among the Kievitsboon
accessions several accessions with intermediate res-
istance are found. The Grootzadige Kievitsboon-
accessions fromWageningen are probably related, and
the same may be true for the accessions from Nieuwe-
Wateringen. Among the white-, brown-, (straw-)
yellow- and black-seeded accessions, accessions with
intermediate resistance to leafhopper was rare.
Due to the large numbers of resistant and suscept-
ible accessions there is no clear relationship with seed
colour. However, 15 of the 20 observed (Stro)gele
accessions (originally coming from South America,
see above) are resistant to both the Andean and
Mesoamerican isolates. The same is true for ve of the
euph4747.tex; 10/08/1999; 17:05; p.12
105
9 observed Eiboon accessions. The remaining 4 ac-
cessions possessed intermediate resistant with scores
close to the resistance score.
Most brown-seeded accessions, including all ac-
cessions of the cultivar(s) Berna and aff. Berna
were susceptible to one or both groups of isolates.
All Kievitsboon, Krombek and Citroenboon ac-
cessions were intermediate or susceptible. The high
incidence of resistance in various bean types may
point to a close relationship within the type. This
may point either to little variation in the region of
provenance or to a founder effect.
Only three of the other identied cultivars were
resistant to both groups of isolates. These cultivars
were the brown-seeded Beka and Ceka and all three
accessions of the small-seeded Kievitsboon Renka.
Within-accession variation was very low. Origin-
ally, the within-variation of landraces may have been
higher, but due to their method of maintenance in
gardens (seeds of only one or a few plants are used
for the next generation) this variation was probably
lost (Zeven, 1997). So Walcherse Witte must have
originated from hybridization of at least two Middle
American races. Although the Strogele accessions
came from various areas in the Netherlands they are
quite similar.
Conclusions
The multivariate and principal component analyses of
quantitative traits at WAU and CIAT showed that the
landraces, garden forms and many cultivars of com-
mon bean in the Netherlands belong to one large group
when phaseolin seed protein pattern, the seed col-
our and seed colour pattern, and other morphological
markers are not taken into consideration. The cultiva-
tion of various dry-bean types close to each other and
close to French bean garden forms and the hybridiz-
ation between these types during the past 500 years
must have removed genetic boundaries, and resulting
in the development of one large gene pool. Farm-
ers and gardeners selected among these hybrids for
plant habit, seed colour and seed colour pattern, and
for use as to dry and/or French bean. Seed exchange
over short distances may have taken place and regional
preference for certain types may have caused selec-
tion and maintenance of regional types. Nonetheless,
using phaseolin seed protein pattern, an evolutionary
marker (Gepts, 1988; Gepts et al., 1986) and other
morphological markers (Singh et al., 1991a) it was
possible to divide Dutch common bean germplasm
into Andean and Middle American gene pools and into
four races, and identify recombinants between the two
gene pools. Most of the landraces and garden forms
were derived from South America and were of dry
determinate bush bean type.
Only, the landrace Walcherse Witte and a Cit-
roenboon accession came from Middle America. The
origin of the two Kogelboon accessions is not clear.
They, maybe, come from the transition zone between
the Middle American bean and Andean bean acces-
sions.
The accessions with the name Berna, a cultivar,
and aff. Berna are close to each other. This may in-
dicate that aff. Berna is indeed Berna and that its
recent origin has not yet lead to diversity.
Acknowledgements
We thank Cesar Cardona for leafhopper evaluation,
Daniel Debouck for phaseolin determination, Fran-
cisco Morales for bean common mosaic evaluations,
Henry Teran for data recording and analyses, and to
Isabel Cristina Giraldo for typing of the manuscript.
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