Diversity 2010, 2, 837-862; doi:10.3390/d2060837
OPEN ACCESS
diversity
ISSN 1424-2818
www.mdpi.com/journal/diversity
Review
Italian Common Bean Landraces: History, Genetic Diversity
and Seed Quality
Angela R. Piergiovanni * and Lucia Lioi
Cnr, Institute of Plant Genetics, via Amendola 165/A, 70126 Bari, Italy; E-Mail: lucia.lioi@igv.cnr.it
* Author to whom correspondence should be addressed; E-Mail: angelarosa.piergiovanni@igv.cnr.it;
Tel: +39-080 5583400; Fax: +39-080 5587566.
Received: 14 April 2010; in revised form: 6 May 2010 / Accepted: 12 May 2010 /
Published: 27 May 2010
Abstract: The long tradition of common bean cultivation in Italy has allowed the evolution
of many landraces adapted to restricted areas. Nowadays, in response to market demands,
old landraces are gradually being replaced by improved cultivars. However, landraces still
survive in marginal areas of several Italian regions. Most of them appear severely
endangered with risk of extinction due to the advanced age of the farmers and the
socio-cultural context where they are cultivated. The present contribution is an overview of
the state of the art about the knowledge of Italian common bean germplasm, describing the
most important and recent progresses made in its characterization, including genetic
diversity and nutritional aspects.
Keywords: genetic resources; germplasm; molecular markers; Phaseolus vulgaris;
phaseolin
1. Domestication and Dissemination Pathways of Common Bean
Over a time period of at least 7000 years, the common bean (Phaseolus vulgaris L.) has evolved
from wild growing into a major leguminous crop. Before domestication, wild P. vulgaris had already
diverged into two major gene pools, each with a proper geographical distribution. Domestication of
wild common beans occurred independently in Mesoamerica and Andean South America [1] and gave
rise to two major gene pools also within the cultivated forms [2]. Evidence supporting common bean
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organization into two major domesticated gene pools came from studies made on morphological,
agronomical, and adaptation traits [3-5], phaseolin type [6-8], isozymes [9,10], and molecular
markers [11]. Cultivars from Mesoamerica usually are small- or medium-seeded (<25 g or 25–40
g/100 seed weight, respectively) and have S phaseolin type. The South American counterparts have
larger seeds (>40 g/100 seed weight) with T, C, H, and A phaseolin patterns [4,6]. Some limited
germplasm exchange took place between the two gene pools in pre-Columbian age, while much more
extensive seed flow occurred after the 1500s. Despite their partial reproductive isolation, the two gene
pools still belong to the same biological species. Although viable and fertile progeny can be obtained
and gene transfer could be attempted, still today breeding programs have only partially comprised
systematic combinations of a wide range of genotypes from each gene pool [12,13].
The discovery of the Americas triggered a rapid exchange of crops between the Old and New
World. The pathways of beans dissemination in Europe is still unclear and currently under discussion,
since the initial input of common bean in Europe is largely unrecorded. It is likely that sailors and
traders brought the nicely colored and easily transportable bean seeds already from the first trips
towards the Americas in the 16th and 17th centuries. The initial common bean accessions were
introduced probably in Europe from Mesoamerica, since Columbus arrived in Central America in 1492
and Cortes reached Mexico in 1518, while Pizarro, exploring Peru in 1528, gave the chance to
introduce common bean from the Andes. The first European explorers certainly devoted great interest
towards this species. For example, Gonzalo Fernandez de Oviedo, who explored Panama and
Nicaragua in 1530, included in its travel reports detailed information on common bean cultivation
techniques used by the American natives [14].
There are strong evidences that common bean reached France already in 1508, probably without
value for human consumption at that time [15]. The first description of common bean in European
herbals was done by Fuchs (1542–1543), who reported that the common bean had climbing habit,
white or red flowers and red, white, yellow, skin-colored or liver-colored seeds with or without spots.
However, it cannot be excluded that Fuchs reported a combination of traits belonging to both
P. vulgaris and Phaseolus coccineus L. species. Further descriptions were done by Roesslin in 1550,
by Oellinger in 1553 and by Dodonaeus in 1554 [15]. Fuchs and Dodonaeus only referred information
about the climbing habit of the bean plant. A punctual selection of old manuscripts (1493–1774)
mentioning P. vulgaris or its synonyms was recently reported by Krell and Hammer [16]. Since the
Mesoamerican biotypes are not very frequent in Europe, McClean et al. [17] supposed that the
germplasm dispersed into Europe was predominantly of Andean origin. However, after the
introduction, a natural selection took place within bean germplasm for tolerance to long days, disease
and pest resistance, stress tolerance and ability to survive, combined with a man-drive selection for
plant habit, seed color, seed pattern type and also disease and pest resistance [15].
2. Common Bean Introduction in Italy
As for other European countries, common beans arrived in Italy mainly unregistered. Bean pictures
are present in festoons adorning the myth of Psyche decorating Villa Farnesina in Rome, painted by
Giovanni di Udine in 1515 [18]. Mentions in historical documents fixed 1532 as the year of common
bean introduction in Italy. At that time, the humanist and literate Pierio Valeriano received a bag of
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bean seeds like compensation for work at the court of Pope Clemente VII. This last greatly supported
the diffusion in the Italian peninsula of this new crop that he received from the Spanish Emperor
Charles V [14]. Valeriano sowed the precious seeds in its fields located in Belluno province (Veneto
region, Northeastern Italy). Subsequently, Valeriano described in a short poem in Latin entitled ‘De
Milacis Cultura’, the cultivation technique, the plant and seed morphology, and the supposed
therapeutic properties of bean seeds [14]. The diffusion of common bean in Veneto region occurred
quickly and still today the cultivation of this pulse has a great economic relevance in Belluno province.
The interest for common bean by literates living in Northeastern Italy may be perceived by the
presence of citations regarding this pulse in some books that were published in Venice at the end of
16th century. In 1565, the physician and botanist Pietro Andrea Mattioli included in a translation and
commentary of the works of the Greek botanist Dioscorides the description of common bean
(Figure 1), together with those of other plants brought from explorations of the new lands in progress
at that age [14].
Figure 1. Picture of common bean plant (called ‘Fagiuoli’) from Commentarii in Sex
Libros Pedacii dioscorides of Pier Andrea Mattioli (1501–1577).
The physician Baldassarre Pisanelli in his ‘Treatise of the nature of food and drinks’ published in
1583 estimated that common beans were “much worse than faves, but among them the red are the
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best”. Teofilo Folengo, a monk and poet of noble family, also reported some features of common bean
in a treatise dealing with food published in 1562 [14].
As we go through the list of popular 16th century New World foods in the Old World, it became
obvious why some crops throve and others struggled. Italians, as well as Europeans, did not recognize
common bean as a foreign species. They thought that this plant was just a new variety of a crop that
they had used to grow and eat since a very long time, the cowpea (Vigna unguiculata (L.) Walp.). The
only ones to really notice the differences of New World beans were botanists, but the confusion
between cowpea and common bean was usual in 16th and 17th centuries. It is noteworthy that in Italy
the substitution of cowpea with common bean took place gradually over the time so that certainly the
two species coexisted for a long period of time. Cultivation of cowpea landraces is yet practiced only
in restricted areas of Central and Southern Italy [19,20].
An important aspect regards the introduction of common beans in the Italian consumer diet. Elite
cookbooks published between the 16th and 17th centuries were reluctant to offer common bean
recipes, which were known to be a sign of poverty and rustic living, as mentioned by the Florentine
Luigi Alamanni in an ode to agriculture printed in 1546 [14]. Only in the early 18th century a recipe,
which used dry bean, was reported in a cookbook by the Jesuit priest Francesco Gaudenzio [18].
Although in the 17th century, chickpea and broad bean were the pulses predominant in the diet, at the
end of 18th century the main source of vegetable proteins for a large part of the population was
represented by common bean [21]. For this reason, it was frequently defined the meat of paupers.
3. Italian Landraces and Safeguard Actions
The selective pressure operated by Italian farmers over the time on the pool of common bean
accessions introduced in Italy produced a myriad of landraces. Adaptation to the soil type and climatic
conditions of the new environments, the geographical isolation of several growing areas, the peculiar
agro-technique (i.e., the consociation with maize), aesthetical and organoleptic preferences have
provided the prerequisites for the selection toward a large number of landraces. This process resulted
in the establishment of different groups of landraces in each Italian region, though it is not unusual that
some landraces are associated to sub-regional areas such as some close villages, a valley, or a small
island. Generally, because of particular thermal, humidity and edaphic requirements, each landrace or
group of landraces cannot be grown with success in areas different from that where they are
traditionally cultivated. At the beginning of 20th century, Comes [22] published a book describing 472
Italian common beans. Successive publications confirmed the high degree of diversity associated to the
most important traits both of plant and seed among Italian populations and attempted their
classification [23,24]. A very small collection representative of common bean diversity available in
Italy at the beginning of 20th century is presently stored at Vavilov Research Institute of Plant Industry
(VIR), the Russian gene bank. Vavilov collected a number of samples belonging to several species
during his trip in Italy (1924–1926), including 12 common bean varieties (Table 1).
Nowadays, common bean is widely cultivated in intensive agricultural systems and commercialized
mainly as green pod and snap seed. In this frame, new cultivars completely displaced the old
landraces. However, in the areas where traditional or low input agricultural systems are still present,
Italian farmers still grow autochthonous varieties not only for personal consumption, but also for sale
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as niche products or specialties in farmer markets. This means that despite the lack of coordinated
efforts, farmers have de facto practiced the on farm maintenance of this germplasm [19,25].
Table 1. List and brief description of Italian common beans collected by Vavilov and held
at Vavilov Research Institute of Plant Industry (VIR).
Local name
Donor
Code
Collection
year
VIR 2096
1924
VIR 2279
1925
Fagiolo Burro
Rampicante
Raparino Nano
VIR 2284
1925
Raparino Gigante
VIR 3559
1926
Fagiolo Metis
VIR 3560
1926
Ingegnoli, Rome
VIR 3562
1926
Fagiolo Cento per
uno
Regino Nano
VIR 3563
1926
Fagiolo
Orvieto, Umbria
VIR 3564
1926
No name
Orvieto, Umbria
VIR 3565
1926
No name
Orvieto, Umbria
VIR 5009
1926
Fagioli scritti
Lucca, Tuscany
VIR 5010
1926
Fagioli rossi
scritti
Lucca, Tuscany
VIR 5105
1926
Fagiolo Bianco
comune
unknown
Ingegnoli, Milan
Instituto Superiore
Abburo, Bologna
Instituto Superiore
Abburo Bologna
Ingegnoli, Rome
Ingegnoli, Rome
Plant and seed description
climbing bean; very late maturity; sphaericus x
ellipticus; seeds large, dark-violet to black
bush bean; very late maturity; green pod;
ellipticus; seeds pale reddish-brown with red
stippling
semi-climbing bean; late maturity; ellipticus;
seeds pale reddish-brown with red stippling
bush bean; middle maturity; flowers white; seeds
bicolored: half is white and half black
bush bean; middle maturity; pods flat; seeds
small, pale brown
late maturity; plant with curly top; pods green
with striped purple; seeds pale reddish-brown
with red stippling
bush bean; late maturity; flowers pale pink;
green pod; seeds buff
semi-climbing bean; late maturity; flowers white;
seeds small, white
semi-climbing bean; late maturity; flowers white;
green pods; seeds slightly flattened, white
bush bean; late maturity; flowers pale pink; pods
green and flat; seeds pale reddish-brown with
pink stippling; productivity medium
bush bean; late maturity; flowers pink; green
pod; seeds reddish-brown with purple stippling;
low productivity
semi-climbing bean; very late maturity; flowers
pink; seeds flat, white
Historical information dealing with cultivation area, description of agro-technique, traditional
recipes, etc., are available only for some landraces. For example, it is proved that already in the 18th
century, the garden pea cultivation was completely replaced by the common bean in the neighborhood
fields of Lamon village, Belluno province (Figure 2). Bellati [26] reported how common bean had a
relevant economic role for farmers of this province in the 19th century due to a commercialization of a
large portion of annual harvest in neighboring regions.
Nowadays, common bean cultivation represents a lucrative activity for Lamon farmers and four
appreciated landraces, generically named ‘common bean from Lamon’ are cultivated. It should be
interesting to underline that the common names of three of these varieties probably relate with the
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beginning of common bean cultivation at Lamon. In fact, the names ‘Spagnol’, ‘Spagnolet’ and
‘Spagnolon’ are alterations of the Italian word ‘spagnolo’ (Spanish) and this circumstance recalls
Valeriano’s work, as previously mentioned. The high appreciation by consumers of these borlotto
types (striped red seeds and striped pods), was recognized by the attribution of the Protected
Geographical Indication (PGI) (GUCE 163/96 2 July 1996), one of the European Union marks
attributable to traditional foods (E.C. reg. n. 2081/92 and 2082/92).
Figure 2. Map of Italy with provenance of some common bean landraces. (a) Piedmont;
(b) Veneto; (c) Tuscany; (d) Marche; (e) Lazio; (f) Campania; (g) Basilicata; (h) Sardinia;
(i) Sicily. Arrows indicate the landrace cultivation area.
Another landrace that has certainly been cultivated for a very long time is the common bean from
Gradoli, (Figure 2) also named ‘Fagiolo del Purgatorio’, literally bean of Purgatory. According to
historical sources, this white small-seeded common bean has been cultivated near the Lake of Bolsena
since 17th century [27]. At that times, the brotherhood named ‘Confraternita del Purgatorio’ involved
in the assistance of paupers, began the tradition to organize a lunch every year during Lent reserved for
poor persons, consisting of dishes prepared with local common bean. The economic value attributed in
the past centuries to this common bean is ascertained by the practice to reward the clergymen who
celebrated religious rituals with bean seeds. Although the Lenten tradition is steadfastly maintained
every Ash Wednesday, the ‘Fagiolo del Purgatorio’ is scarcely requested by consumers and
consequently fated to extinction in a very short time.
Conversely, ‘Billò’ common bean is an appreciated landrace with a relatively short tradition of
cultivation. Oral information collected by interview of elderly farmers indicated that the cultivation of
this bean in Cuneo province, Piedmont region (Figure 2), started about 60 years ago. At that time,
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843
seeds from Lamon were introduced by some farmers in the area presently devoted to Billò cultivation.
A recent study based on molecular markers has evidenced that several decades of cultivation in two
different environments without exchange of material is producing a differentiation between the
original material grown at Lamon and that collected in Cuneo province [28].
A detailed analysis of the recent scientific literature shown that on farm survival of local common
bean varieties is a relevant phenomenon in Italy [28-34]. These findings assume a particular
importance in the areas located along the Apennine ridge and in several small islands where
agricultural systems still retain traditional forms. Unfortunately, the cultivation of these old varieties is
generally practiced by elder farmers without any action of sustain by governmental or local
institutions. These circumstances cannot assure the survival of this precious germplasm, so that a large
fraction of common bean landraces is severely endangered with the risk of extinction. Recent
European Community regulations introducing the possibility to attribute marks of origin and quality,
PGI and PDO (Protected Designation of Origin), to local typical products, could give an important
support to on farm survival of élite common bean landraces, but at same time, the abandonment of the
less appreciated ones could be encouraged [35]. This discrepancy can be overcome through the
application of both on farm and ex situ conservation to the same germplasm. Presently, only three PGI
marks have been assigned to Italian common bean; they are ‘Fagioli di Lamon’, ’Fagioli di Sarconi’
and ‘Sorana’ landraces.
As concerns the ex situ conservation of Italian germplasm, collections of about 1,500 accessions are
held in the CNR, Institute of Plant Genetics (IGV, Italy) and the Leibniz Institute of Plant Genetics and
Crop Plant Research (IPK, Germany), while several small collections, consisting of at least 100
accessions, are maintained by some Italian Universities such as those of Turin, Perugia, Ancona,
Basilicata, Viterbo, and Sassari. Moreover, the increasing attention towards the safeguard of
biodiversity has encouraged some regions to create their own facilities for the ex situ conservation
such as those created by the Tuscany and Friuli regions [36]. Nowadays, a very small fraction of
Italian landraces is held in the regional gene banks. The knowledge about the levels of diversity
present in these collections needs a robust scientific work with the aim of cataloguing and
characterizing the stored material. These efforts constitute an important goal in prospecting adequate
strategies for ex situ conservation management as well as for the utilization of this precious germplasm
in breeding programs. At the same time, for on farm conservation, which can be seen as one of the
most effective strategies applied to the safeguard of crop genetic resources, knowledge about the level
of diversity within each landrace is fundamental in order to plan the most appropriate program for their
survival.
Still today no systematic cataloguing of all common bean landraces cultivated in the Italian
peninsula has been attempted. In the absence of appropriate financial resources and coordinate
initiatives, only sporadic efforts have been tried. For example, some regional agricultural development
agencies have published catalogues describing the common bean landraces grown in the own region.
This is the case of the Agricultural Research Service of the Regions: Basilicata (ALSIA), Abruzzo
(ARSSA), and Calabria (ARSSA) [37-39]. The description of some local common beans has been
included in a number of publications on crop biodiversity at regional level edited by Friuli region [40]
and Park of Dolomiti in Veneto region [41]. Web catalogues have been created within the frame of
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regional projects devoted to rural area development by the Department of Environmental
Biotechnologies, University of Perugia and Agricultural Research Service of Lazio region (ARSIAL).
4. Characterization and Evaluation of Italian Common Bean Landraces
4.1. Phenotypic Variation
In spite of increasing use of DNA-based markers in investigating genetic diversity within
germplasm collections, the evaluation of phenotypic variation is still crucial in determining adaptation,
agronomic potential and breeding value of landraces.
A considerable diversity among the genotypes grown by Italian farmers has been observed in seed
morphological traits: shape, size, coat color, and type and color of pattern. The first description of seed
shape variation within Italian germplasm was that attempted by Comes [22] who analyzed 472
populations and grouped them into four classes: (1) kidney; (2) cylindrical; (3) elliptic; (4) round.
Moreover, Comes [22] described a wide range of coat color from white to violet, in addition to a lot of
bicolored, spotted and striped types using a visual inspection. It is surprising to observe that these
classifications greatly agreed with the seed descriptors included in the official common bean
descriptors released by IBPGR in 1982 [42]. More recently, a computer-aided image analysis system
has been specifically designed for the characterization and discrimination of Italian common bean
landraces by phenotypic traits [43]. The automatic acquisition of parameters such as size, shape, color,
and texture of seeds gives scientists a reliable and quick tool for the identification of landraces (see
also an extensive review by Dell’Aquila [44]). A survey of the scientific literature dealing with Italian
landraces has evidenced that a large part of phenotypic variation described by Comes [22] has survived
until now [30,34,37,45-51]. However, as a consequence of different aesthetical preferences of local
farmers, the coat color variation is not homogeneously distributed in the Italian peninsula.
As shown in Table 2, white coat seed populations and borlotto types are present in all regions,
while unusual coat types are associated to restricted areas indicating the prominent role played by
aesthetical preferences of local farmers in the choice of the own set of local populations. For instance,
black coat types are cultivated only in Central Italy and are named ‘Marconi al palo’, ‘Stortino di
Lucca’ and ‘Mangiatutto nano’ grown in Lazio, Tuscany and Marche region, respectively (Figure 2).
Two groups of landraces with yellow coat are described: the bush types named ‘Zolfino’ and
‘Solfarino’ cultivated in Tuscany and Lazio region, respectively [52,53], and the climbing ones named
‘Gialet’, and ‘Cesarin’ and ‘Centut’ grown in Veneto and Friuli region, respectively [54-56]. Presently,
a study is in progress to ascertain if ‘Gialet’ and ‘Cesarin’ can be originated from the same genetic
stock (F. Miceli Pers. Comm.). In contrast with the high phenotypic variation of seed traits among the
populations, the literature reports a very low phenotypic variation at intra-population level. This trend
is attributable to the custom of farmers to preserve only a few plants for each population to obtain
seeds
for
the
next season.
In regard to plant habit, a wide variation from bushy to aggressive climbing ability has been
observed within Italian common bean landraces, though climbing types are widely predominant. It is
known that the predominance of one growth habit type is related to ecological adaptation as well as to
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the cropping system. The practice, now abandoned, to grow common bean mixed with maize or
alternatively, the presence of woods that assured abundant and cheap raw material to make the stakes,
could have favored the predominance of climbing habit.
Table 2. Distribution of the seed coat types (according to IBPGR) within Italian common
bean landraces as described in the literature.
Region
N.
pop
Coat without pattern
White
Brown pale
Yellow
Black
Coat with pattern
Other
bicolor
to dark
Abruzzo
10
3
1
Basilicata
65
18
11
Calabria
1
1
Campania
3
2
Pattern around
type
hilum
4
2
[28,47,49]
16
4
[30,37,117]
1
4
Lazio
19
6
4
Liguria
2
1
1
Marche
23
8
4
Piedmont
5
1
1
Sicily
4
Tuscany
24
Veneto
8
168
10
Borlotto
[118,119]
Friuli
Total
6
2
1
1
1
1
51
[67,102]
2
[56]
5
[28,49,53,119]
[119]
1
3
7
[120]
3
3
11
Ref.
2
1
1
1
25
5
1
1
2
1
3
9
4
[119]
1
[100,119]
2
[59,121]
5
19
46
[34,54,122,123]
10
Currently, about 90 % of landraces cultivated in Basilicata region have climbing habit [30,45,46]. A
similar frequency has been estimated in Cuneo province, Piedmont region [51]. Determinate and
indeterminate growth type (40 vs. 60 %, respectively) were detected in Campania region [57] as well
as within borlotto-like landraces of Marche region [58]. Variation of habit within one landrace has
been also reported. As an example, the common bean named ‘Piattella Pisana’ or ‘San Michele’,
cultivated in Tuscany exhibits a high variation of several morphological and physiological plant traits.
Baldanzi and Pardini [59], analyzing 33 populations belonging to this landrace, observed that the
climbing ability is highly variable. Similar results were reported for the common bean named
‘Decimino’ or ‘Turco della Garfagnana’, another landrace cultivated in Tuscany. The surveyor of
material grown by farmers evidenced differences in habit, flower color, and seed shape within
‘Decimino’ [59]. Two distinct growth habits, determinate and indeterminate, that differ in plant height,
number of nodes and pods, and yield, were detected also within the ‘Fagiolo del Purgatorio’ [61]. The
presence of a different habit within these three landraces could due to farmer preferences since
Tuscany and Lazio regions are geographically close (Figure 2). An aggressive climbing ability
characterizes the common bean named ‘Fagiolo a pisello’ cultivated in Colle di Tora on the steep
hillsides of Turano Lake. Plants of this common bean can been more than 3 m tall but flowers appear
only in the upper part [62].
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4.2. Seed Storage Proteins as Biochemical Markers
Unlike in other legumes, storage proteins in common bean are mainly constituted by vicilin, in this
species called phaseolin. It accounts for 50% of the total proteins in mature seeds, consisting of a
number of polypeptides in the Mr 54–44 kDa, and is considered a biochemical marker. Phaseolin
electrophoretic analyses of wild-growing and domesticated materials supported the hypothesis of the
presence of two major gene pools within common bean germplasm, the Mesoamerican and Andean
ones [6]. Multiple domestication events are thought to be the cause of parallel geographic phaseolin
patterns variation between wild and cultivated forms. The Mesoamerican wild forms showed S as well
as a number of M phaseolin types, while S patterns predominated within the domesticated counterpart.
Similarly, T, C, H, and A phaseolin types were found among wild and cultivated materials from the
Andes [8]. It has been shown that phaseolin is a useful marker to follow the dispersal pathway of
common bean from domestication areas into Europe. A higher frequency of Andean phaseolin types
with respect to Mesoamerican ones was recorded in European germplasm first by Gepts and Bliss [7].
A more recent study on 544 European accessions confirmed that Andean phaseolin types T (45.6%)
and C (30.7%) prevailed over the Mesoamerican S type (23.7%) [63]. Lioi [64], who analyzed a large
Italian collection, showed the predominance of Andean genotypes (73%) over Mesoamerican ones,
and that phaseolin C types had a higher frequency than T ones. The prevalence of the Andean gene
pool within the Italian common bean germplasm stored in some international gene banks has been
recently confirmed by Logozzo et al. [63], who reported 88% frequency of Andean types with a
predominance of C type (51.5%), and only 12% of S types. Tiranti et al. [65] studied 159 common
bean landraces collected from farmers and local markets in 10 Italian regions (six from Piedmont, 14
from Liguria, 11 from Friuli, 25 from Tuscany, four from Marche, 52 from Umbria, 25 from Lazio, 17
from Abruzzo, three from Basilicata, two from Sicily). The presence of the three major phaseolin types
(C, T and S) was observed in seven regions, suggesting a homogeneous distribution of phaseolin types
in the sampled regions. The observed frequencies were 40, 28, and 32% for Andean (C, T) and
Mesoamerican (S) types, respectively. These results agree largely with Lioi’s data [64], while the
frequency of resulting S types was more than double the 12 % reported by Logozzo et al. [63].
Results of several studies carried out at regional level are summarized in Table 3. Despite a large
variation in the number of analyzed populations and sampling strategies, among these investigations a
prevalence of the Andean phaseolin types was observed.
At least for the more sampled regions such as Basilicata, Sicily, Campania and Sardinia (Figure 2),
the frequency of each phaseolin type at regional level can be depicted by these studies. The clear
predominance of C type has been observed for populations from Basilicata region [45,46].
Angioi et al. [66] recorded a high frequency of the C phaseolin among Sardinian common bean
landraces (Table 3). Moreover, Mesoamerican types appear to be less represented in Sardinia; only one
genotype with S phaseolin type was found only among 73 genotypes analyzed. A similar result has
been recorded for the Campania region (Table 3), where only 9.2% of tested populations belonged to
the Mesoamerican gene pool [57,67].
The predominance of C types has been observed also within Spanish common bean landraces by
Rodiño et al. [68]. This similarity between Italian and Spanish germplasm strongly suggests that
common bean may have been introduced in Italy mostly from Spain more than directly from America.
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This is not surprising because several Italian regions were under Spanish rule in the 16th and
17th centuries.
Table 3. Distribution of the phaseolin pattern frequencies within Italian common bean
landraces as described in the literature.
Region
N pop
Phaseolin type
T
C
Abruzzo
9
1
1
Basilicata
66
18
38
Calabria
4
Campania
98
Lazio
17
4
1
Piedmont
5
1
3
Sardinia
73
21
51
Sicily
28
3
17
Tuscany
1
Veneto
12
4
5
Total
313
49
119
T+C
A
H
1
Mix
Andea
n
Ref.
S
3
8
3
89
Mix AndeanMesoamerica
n
3
[49]
2
[45,46,104]
1
[118]
9
1
4
3
2
[57,67]
2
1
1
89
2
4
8
[34,49]
[34,50]
1
[66]
7
*
1
[104]
3
[34,54,104,122]
31
8
*Piergiovanni unpublished.
The presence of two or more phaseolin patterns within some populations from Abruzzo, Lazio,
Piedmont and Sicily regions (Table 3) is indicative of the informal selection from which these
landraces have originated. As shown in Table 3, the populations from Central Italy (Lazio and
Abruzzo regions) showed the highest polymorphism of phaseolin protein fraction. In addition to the
three main types (T, C, and S), within these populations phaseolin patterns uncommon in Europe, such
as the Andean types A and H [47,49] were observed. This suggests that Central Italy retains higher
germplasm variability than the other Italian regions, as was recently reported for the Italian lentil
germplasm [69].
The existence of no small-seeded populations (100 seed weight > 40g) showing S phaseolin type
has been evidenced by some authors [45,46,63]. These kinds of biotypes are uncommon within the
European germplasm [70] and absent in the Mesoamerican gene pool. It has been hypothesized that
they could be the result of a deliberate selection towards large-seeded genotypes within the
Mesoamerican biotypes carried out by European farmers over time, being large-seeded types generally
more accepted from consumers. Alternatively, when Mesoamerican and Andean genotypes began to be
Diversity 2010, 2
848
cultivated in proximity in the European environments, recombination resulted in increasing
large-seeded types showing S phaseolin pattern [71].
Phytohemagglutin (PHA) is the second most abundant protein in common bean seeds. PHA exhibits
a relatively limited heterogeneity but it may be used in addition to phaseolin for the characterization of
landraces [72]. Although gene families coding for phaseolin and PHA are not linked, in cultivated
materials there is a narrow association between specific phaseolin and PHA patterns [73]. As
suggested by Gepts [74], these findings could be due to the establishment of two distinct gene pools,
where specific alleles of different genes may be associated with one another more often than random
assortments would predict. An extensive screening for PHA variation in Italian material was carried
out by Lioi [75], who analyzed about 350 accessions. Of 10 different patterns detected, two (TG2 and
SG2) were predominant and were generally co-present with Andean and Mesoamerican phaseolin
patterns, respectively.
Figure 3. Electrophoretic patterns of phaseolin (PHAS) and phytohemagglitinin (PHA) in
seeds of ‘Fagiolo del Purgatorio’ from Gradoli populations.
A more recent study also demonstrated that PHA could be utilized for the characterization of
landraces. Lioi et al. [27] found an interesting variation of the PHA electrophoretic pattern within the
populations of ‘Fagiolo del Purgatorio’. The PHA patterns, MG2 and SG2 (Figure 3), together with
several different traits such as vegetative cycle, yield, and pest resistance, suggested that more than
one constitutive nucleus has contributed to the genetic background of this landrace [76]. Therefore, an
adequate number of populations should be preserved to ensure the retention of the diversity present in
this landrace.
4.3. Molecular Diversity
Although phaseolin has been a valid tool in identifying geographical patterns of domestications and
dispersion, its scarce polymorphism does not permit discrimination among closely related landraces.
The advent of molecular techniques has greatly improved the ability to understand the genetic
structure of common bean landraces. Studies of the level and structure of genetic diversity in common
bean landraces are of fundamental importance in tracing reasonable patterns of dispersion and
diversification of this species, and in understanding relationships among landraces, genetic resources
Diversity 2010, 2
849
management and plan safeguard actions for their preservation, improvement, and promotion. Although
in the last decade the attention devoted by researchers to Italian common bean landraces has greatly
increased, studies on this topic are far from being exhaustive.
A variety of PCR-based molecular markers are useful tools for the study of genetic diversity. In
particular, SSRs (Simple Sequence Repeats) or microsatellites, are short (mostly 2–4 bp) tandem
repeats of DNA sequence; their polymorphism originates from a different number of repetitive core
motifs present at one locus [77]. They are useful genetic tools although their development is time
consuming and requires the identification of SSRs. The first strategy used to search for microsatellite
motifs was the screening of publically available DNA sequence databases developed first by
Yu et al. [78,79] and then by Guerra-Sanz [80]. Another strategy used to enlarge the number of
microsatellite loci in P. vulgaris was the sequencing of size-fractionated BAC clones [81]. A number
of polymorphic microsatellite markers have been isolated from both cDNA and genomic
libraries [82-85]. The number of gene-based microsatellites was recently increased through a cDNA
library screening approach by a large scale library developed from multiple source tissues of an
Andean gene pool genotype [86]. SSRs were used by several authors to construct genetic maps
[79,87,88], to evaluate genetic diversity in commercial variety or lines or genotypes of common bean
[89-91], or to separate more effectively wild and cultivated accessions as well as Andean and
Mesoamerican gene pools [92,93].
Figure 4. UPGMA dendrogram of 14 ‘Fagiolo del Purgatorio’ from Gradoli populations
based on SSR molecular markers.
The first approach that applied SSRs to characterize Italian germplasm was performed by
Masi et al. [94] on three common bean landraces from the Basilicata region. This study showed that
the frequency of polymorphic SSRs was fairly high, with some examined loci being multi-allelic,
confirming the usefulness of these markers to the study of genetic diversity in bean collections. Some
studies on common bean landraces from different Italian regions analyzed by SSR markers revealed
the presence of both gene pools with predominance of landraces of Andean origin [34,58]. A study
conducted on 33 local populations belonging to seven Italian common bean landraces revealed that
Diversity 2010, 2
850
they retain a considerable level of heterogeneity. SSR markers used were highly variable depending on
the locus, with detection of 53 alleles at 14 loci examined. Populations belonging to the same landrace
were grouped in clearly distinguishable sub-clusters with only one exception, suggesting that different
farmers maintain distinct materials. In particular, the ‘Fagiolo del Purgatorio’ populations were
grouped in two sub-clusters (Figure 4) confirming biochemical and agronomic data and suggesting that
more than one constitutive nucleus has contributed to the genetic background of this landrace [34].
A low level of diversity was detected in a Sardinian and a Ligurian common bean collection,
characterized using SSR and RAPD markers [66,95], probably due to a strong founder effect. Despite
a relatively low level of diversity, the landraces were clearly distinct from cultivated materials. A
detailed study about the organization of the ‘Fagiolo a pisello’ diversity was reported by Tiranti and
Negri [96]. The aim of this study was to define an appropriate on farm conservation strategy, which
could be used as a model for other populations. Results showed a high level of genetic similarity
suggesting that populations were closely related to each other, probably as consequence of a single
introduction event. Nevertheless, all the populations were morphologically and genetically clearly
differentiated, with a considerable within-population genetic variation.
Similar studies conducted at local level, and in particular on some landraces from Marche region,
revealed the presence of genotypes intermediate between Andean and Mesoamerican gene pools,
suggesting probable hybridization events which might generate additional variation [58]. Moreover,
molecular investigations revealed that some landraces from northeastern Italy could have played a role
in the genealogy of commercial varieties [56].
Another class of molecular markers, AFLPs (Amplified Fragment Length Polymorphisms), due to
their capability to detect polymorphism at a great number of loci, were used to assess diversity among
wild material, landraces and common bean cultivars [97,98]. From three to seven AFLP primer
combinations were used to estimate genetic variation among common bean landraces maintained on
farm from different Italian regions. They revealed a quite high percentage of polymorphism, with each
accession showing a unique pattern. The branching patterns showed two main clusters, regrouping
accessions belonging to Mesoamerican and Andean gene pools, respectively. Within each cluster,
accessions were well separated from each other [34].
Detailed studies effectuated on some Central Italy landraces (Umbria, Lazio and Abruzzo regions)
showed that genetic differences among landraces could be ascribed to individual farmer’s preference
and selection as well as to different introductions occurring in each location [31-33].
In the last years, some studies were carried out devoted to the fingerprinting of specific landraces
using different molecular markers such as AFLP, RAPD, semi-random, ISSR, and SSR. Some
landraces of great interest such as: ‘Fagiolo Badda’ (Sicily) [99,100], ‘Occhio nero di Oliveto Citra’
and ‘Fagiolo di Controne’ (Campania) [101-103], ‘Fagioli di Lamon’ (Veneto), ‘Fagioli di Sarconi’
and ‘Fagioli di Rotonda’ (Basilicata), ‘Zolfino’ (Tuscany) [104], ‘Fagiolo a pisello’, and ‘Fagiolo del
Purgatorio’ (Lazio) [29,32,105,106] were analyzed. Through the data obtained, relationships among
landraces regarding problems related to the homonymy and distribution of the diversity within
landraces have been clarified. They represent preliminary actions necessary for drawing up
disciplinary rules for conservation consortia, as well as for the attribution of European PGI marks.
4.4. Seed Quality
Diversity 2010, 2
851
The common bean is the world’s second most important pulse after soybean. It is well known that
dry seeds represent an affordable and inexpensive source of proteins despite being deficient in the
sulfur amino acids. In addition, common bean seeds are rich in complex carbohydrates, dietary fiber,
starch, minerals and vitamins. Like other legumes, common bean seeds contain a number of bioactive
compounds including enzyme inhibitors, lectins, phytates, oligosaccharides and phenolic compounds,
which play metabolic functions in humans or animals. These features may be regarded at the same
time as disadvantageous (reduction of protein digestibility or mineral bioavailability, flatulence) or
favorable (antioxidant effect or pre-biotic activity). As discussed in previous sections of this review, a
wide range of genetic variation exists within the Italian local common bean varieties, so differences in
compositional and physic-chemical traits may be predictable. Unfortunately, there are relatively scarce
studies on the nutritional and nutraceutical value of the Italian landraces as well as on their comparison
with the modern cultivars.
Table 4. Protein content relative to some Italian landraces derived from the literature. A
range of values is reported when several populations belonging to the same landrace were
analyzed.
Local name
Northern Italy
Billò
Bianco di Bagnasco
Spagnolet nano
Gialet
Bala rossa
Bonei
Lamon
Central Italy
Zolfino
Fagiolo del Purgatorio
Fagiolo romanesco
Fagiolo regina
Fagiolina arsolana
Cioncone
Pallini
A pisello peligni
Pane aquilano
Pane peligni
Sourthern Italy
Bianco di Rotonda
Ciuoto
Verdolino
Bianco del Pollino
Badda bianco, Badda nero
and Monaco Mussu Niuru
Cultivation region
Protein (% dm)
Ref.
Piedmont
Piedmont
Veneto
Veneto
Veneto
Veneto
Veneto
27.4
23.2
21.8–23.8
26.1–27.8
24.5
24.2
28.0
[50]
[27]
[27]
[123]
[54]
[54]
[108]
Tuscany
Lazio
Lazio
Lazio
Lazio
Lazio
Lazio
Abruzzo
Abruzzo
Abruzzo
21.5
21.4–27.6
26.8
27.7
23.6–27.3
24.4–25.9
23.4
[124]
[27]
[125]
[125]
[125]
[125]
[125]
[125]
[125]
[125]
Basilicata
Basilicata
Basilicata
Calabria
Sicily
26.6
20.8
22.2
23.6–27.2
26.8–27.1
[34]
[34]
[34]
[118]
[108,126]
Protein content is generally considered a very important trait to estimate the nutritional quality of
common bean seeds. The evaluation of 21 local populations cultivated in Basilicata region for three
Diversity 2010, 2
852
consecutive growing seasons evidenced a broad variation of protein content (22.0–28.2% dm) within
this pool of samples. The same study showed that seven tested landraces had an average protein
content falling in the range 25.0–28.8% dm, the same recorded for six cultivars used as references
[48]. A similar protein range (21.8–29.2% dm) was recorded by Perazzini et al. [107], who evaluated
eight populations cultivated in Central Italy. Other landraces have been sporadically investigated, the
information available in the literature is shown in Table 4. These studies suggest that a large fraction
of Italian landraces have a good nutritional quality, at least in terms of total protein content.
More recently, the evaluation of minor seed components (starch, fat, fiber, macro and
microelements, etc.) started, but this field is far to be sufficiently examined [48,108,109]. Data
collected up to now suggest that a narrow range of variation among the landraces exists for these seed
components, and that differences in soil type, adopted cropping technique and environmental
conditions strongly affect these grain traits.
It is well known that common bean grains are relatively rich in trypsin inhibitors (TI), a class of low
molecular weight proteins able to inhibit trypsin. In addition to this action, these inhibitors act as
defensive agents against pests, regulators of germination, and reserve of sulfur-amino acids [110]. The
range of TI variation within Italian landraces has been investigated, together with the year-to-year
variation recorded only for a very narrow number of genotypes [111]. This study revealed not only a
three-fold variation of TI values among the set of tested landraces (14.3–37.4 TIU mg-1 dm) but also a
large effect of year-to-year climatic changes on these anti-nutritional compounds. The increase of TI
expression might possibly be related to the drought stress suffered by plants during the vegetative
growth stage.
Generally, researchers recognize that the pigments responsible for seed coat color in common bean
are flavonoids and that these compounds could have positive health benefits as antioxidants. At the
present, soybean is considered the most relevant source of flavonoids with beneficial health effects
among legume crops. However, soybean is not a traditional crop in Italy, while common bean is
usually consumed by a large part of the population. The scarce knowledge about the flavonoid content
requires additional research work on all legume crops of the Mediterranean area, including Italian
common bean landraces.
It is known that in the Phaseolus genus flavonols and other phenolic compounds are stored in seed
coat because they have anti-pathogen and anti-feeding activities, assuring a good protection of seeds
from external attacks. The flavonoid composition of three phenotypes of ‘Zolfino’ landrace was
described by Romani et al. [112]. Dinelli et al. [113] characterized and quantified the flavonoid
present in 23 Italian landraces traditionally grown in Basilicata, Tuscany and Veneto. Kaempferol and
related conjugated forms were found in all the ‘Zolfino’ accessions and in the ‘Verdolino’ landrace
from Sarconi, while they were absent in 'Fagioli di Lamon' as well as in the other landraces from
Sarconi. These data reflect the different coat pigmentation of tested samples since ‘Zolfino’ and
‘Verdolino’ have a yellow and light green coat color, respectively, while Lamon common beans are
borlotto types. Moreover, Dinelli et al. [113] evidenced a great variation of flavonoid content not only
among the tested landraces (0.19–0.84 g/kg of seed fresh weight) but also for each landrace sampled in
different years. The coefficient of variation observed in the period 2001–2003 ranged from ±18% to
±50%. Further experimental works is needed for a deep understanding of physiological mechanisms
influencing the expression of these compounds.
Diversity 2010, 2
853
A recent study has shown that different parts of dry beans exhibit antioxidant activity [114]. The
anti-radicalic activity of dry beans belonging to ‘Zolfino’ (Tuscany), ‘Poverella’, ‘Verdolino’ and
‘A’Marozzo’ (Basilicata) landraces, grown in three different years and in different geographic areas,
was quantify by Heimler et al. [115]. Authors recorded a wide range of antioxidant activity (from 39 to
2,810 EC50 for ‘Verdolino’ and ‘Poverella’, respectively) and attributed these results mainly to
landraces and environmental conditions more than to total phenolics and total flavonoids detected in
dry seeds.
With regards to the investigated physical seed properties (i.e., cooking time, hydration index,
swelling capacity and splitting of seeds during cooking) the results are strictly related to the
commercial value of landraces. Very recently, a significant variation of hydration rate among Italian
common bean germplasm has been described. The screening of 62 landraces belonging to different
market classes revealed that they behaved differently in terms of hydration rate. Based on these data
the tested landraces were classified into three hydration groups characterized by slow, intermediate
and fast water uptake [116].
Finally, it should be mentioned that any scientific study has been reported in the literature regarding
the tests on consumer approval of Italian common beans though the landraces are generically
emphasized as preferred with respect to modern cultivars because of their sweet taste, soft texture, and
cooked bean flavor.
5. Conclusions
In accordance with the literature presented in this review, Italian common bean germplasm is
characterized by a high degree of genetic diversity. This is the consequence of five centuries of
uninterrupted cultivation coupled to a capillary diffusion of this crop in all the regions. Although the
overall number of Italian common bean accessions stored ex situ is remarkable, the lack of systematic
investigations of these collections appears evident. Unfortunately, the chance of a systematic
monitoring of the Italian common landraces is very low as a consequence of the lack of appropriate
financial resources devoted to this aim. Taking into account the above mentioned studies, only a
multidisciplinary approach can be fully effective to characterize each landrace and to help the efforts
in planning adequate safeguard actions.
Moreover, available data demonstrated that wide differences between landraces and cultivars are
detectable at several levels. As an example, if landraces are predominantly climbing, bush cultivars are
widely used in the advanced agricultural systems by reason of their more appropriate features for
mechanical harvesting. Similarly, the seed coat variation (color and pattern type) detected among
landraces contrasts with the large predominance of white and borlotto types among the cultivars. This
condition is a direct consequence of market demand, because white seed type is preferred for dry seed
consumption while borlotto type is required as snap seed. From a practical standpoint, the safeguard of
landraces assumes a strategic value to be able to answer multiple questions, though the implementation
of adequate actions, and depends on the availability of economic resources and facilities as well as the
sharing of goals among scientific, governmental, and local institutions.
Acknowledgements
Diversity 2010, 2
854
Thank to A. Dell’Aquila for discussion and critical revision of the manuscript. Study partially
supported by Ministry of Agriculture Food and Forestry Policies with funds released by C.I.P.E
(Resolution 17/2003).
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