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RESEARCH ARTICLE
Collection and morphological characterization of Lagenaria
siceraria germplasm from the Mediterranean region
of Turkey
H. Yetis¸ir ÆM. S¸ akar ÆS. Serc¸e
Received: 27 December 2007 / Accepted: 26 March 2008 / Published online: 15 April 2008
ÓSpringer Science+Business Media B.V. 2008
Abstract The landraces of Lagenaria siceraria in
Turkey show great diversity for morphologic traits,
particularly in fruit size and shape even though
Turkey is not centre of genetic diversity for
L. siceraria. In Turkey, L. siceraria is used as
food, musical instruments and containers, according
to the type and shape of their fruits. Its diversity
has been gradually declining over the last 25 years.
With the aim of assessing variations in plant, fruit
and seed morphology among the L. siceraria
landraces, 15 field trips for collection of germplasm
to southern parts of Turkey (Mediterranean region)
were carried out in the period of 2003–2004. The
study also aimed at developing a representative
core collection of the material to guide future
studies and uses concerning its existing genetic
diversity in Turkey. A total of 182 accessions
(fruits and/or seeds) were collected. The seeds from
all the sample material were sown in green house
conditions at the experimental station of the Faculty
of Agriculture of the Mustafa Kemal University in
Hatay (Antakya), (36°1802200 N, 36°1303300 E,
82 m). In the spring of 2004, 162 out of the 182
sown seeds germinated and developed into viable
seedlings. These were further transplanted in the
field in 10 replicates. The morpho-agronomic
characterization was carried out following the
international standards for crop descriptors set by
Bioversity International. The data were subject to
both descriptive statistics and multivariate analysis
by Principle Component Analysis. The descriptive
statistics revealed that the whole collection exhibits
a great deal of morphological diversity and that the
subset core collection represents most of the
variability. The first three principle components,
calculated by using six quantitative traits, explained
26%, 21% and 17% (a total of 64%) of the total
variation. Among the studied accessions, no appar-
ently distinct patterns such as geographical origin
were detected. This may suggest that the accessions
have been introduced to Turkey from multiple
locations and/or their diversity had been distributed
almost evenly across the Mediterranean region of
this country. Based on our results from the
morphological characterization, 30 genotypes were
selected to develop a subgroup (core) collection in
order to represent most of the genetic diversity of
all accessions. The work presented here is the first
report about morphological characterization of
Turkish L. siceraria germplasm.
Keywords Bottle gourd Core collection
Cucurbitaceae Genetic diversity
Germplasm collection Lagenaria siceraria
Morphological characterization Rootstock
White flower
H. Yetis¸ir (&)M. S¸ akar S. Serc¸e
Department of Horticulture, Faculty of Agriculture,
University of Mustafa Kemal, Antakya, Hatay 31034,
Turkey
e-mail: yetisir1@yahoo.com
123
Genet Resour Crop Evol (2008) 55:1257–1266
DOI 10.1007/s10722-008-9325-y
Introduction
The species Lagenaria siceraria (Molina) Standl., a
member of the Family Cucurbitaceae, is commonly
known as the white-flowered bottle gourd. It is
cultivated as an annual monoecious, vigorous climber
species. The genus Lagenaria contains also five wild
species: Lagenaria brevifilora (Benth.) Roberty,
Lagenaria abyssinica (Hook. f.) C. Jeffrey, Lagena-
ria rufa (Gilg) C. Jeffrey, Lagenaria sphaerica E.
Mey. and Lagenaria guineensis (G. Don) C. Jeffrey
(GBIF 2007). Bottle gourd has a wide bihemispheric
distribution. Remains of L. siceraria were identified
from Egyptian tombs (dated back to about 3,000–
3,500 BC), caves of Thailand (10,000–6,000 BC), in
Mexico (7,000–5,000 BC), in Peru (4,000–3,000 BC)
and in China (500 AD) (Bose and Som 1986).
Archaeological evidence suggests that humans have
collected and used Lagenaria for at least
12,000 years in both new and old worlds. A theory
of trans-oceanic drift of bottle gourd has been
suggested to explain its pre-Colombian distribution
in tropical America. It has been suggested that
Lagenaria had a pan-tropic distribution and was
independently domesticated in both old and new
worlds (Harris 1967). On the other hand, this theory
does not agree with Whitaker (1971), who affirmed
that bottle gourd was indigenous to tropical Africa
(south of Equator) and was dispersed by trans-
Oceanic drift or human transport to other parts of
the world. Such a theory speculates that all six
species are present in Africa, where a centre of
genetic diversity for L. siceraria is believed to exist,
although its wild progenitors have not been identified
(Whitaker 1971). This species might have dispersed
to the shores of the New World by ocean currents or
by human migration in pre-historic times (Morimoto
et al. 2005).
The bottle gourd fruits are generally eaten as a
vegetable in Africa and Asia. Immature fruits are
eaten after being cooked (boiling, frying or stuff-
ing), similarly to what is done with Cucurbita pepo.
The mature fruit is often scooped out and its dried
hard rind is used in making containers, bowls,
musical instruments, decorations or in some cases,
fishing floats. Shoots, tendrils and leaves are also
cooked and the seeds are removed for oil extrac-
tion. Seeds, tendrils and young leaves are also used
for some medical purposes (Herklots 1972;
Moerman 1998; Manandhar 2002). Another utiliza-
tion of L. siceraria is as rootstocks for watermelon
against soil-borne diseases and low soil tempera-
ture. Grafting of watermelon onto bottle gourd was
first performed in Korea and Japan in the late
1920s (Ashita 1927) and showed a high compati-
bility rate with watermelon (Lee 1994; Oda 1995;
Yetisir and Sarı2003).
Turkey is not the centre of genetic diversity for L.
siceraria, but the landraces of L. siceraria show great
diversity, particularly in fruit size and shape. In most
of small cities and villages of Turkey, a number of
different landraces have been commonly cultivated
for several uses such as food, music instrument,
decorative and containers, depending on the charac-
teristics of the fruit. It was possible to see landraces
of L. siceraria in many parts of Turkey about
25 years ago, but landraces have gradually disap-
peared because tools made of plastic have been
quickly spreading, thus replacing those made out of
bottle gourds. Currently, L. siceraria is grown only in
some small regions of Turkey for human consump-
tion and decorative purposes; however, there still
seems to be considerable diversity among L. siceraria
genotypes in southern and western Turkey. The
objectives of the present study were to survey and
analyze variations in plant, fruit and seed morphology
in L. siceraria in southern Turkey and establish a core
collection. This represents the first study on collec-
tion and characterization of Turkish L. siceraria
germplasm.
Materials and methods
In 2003 and 2004, 15 germplasm-collecting missions
were carried out in seven provinces (Adana, Antalya,
Gaziantep, Hatay, Mersin, Kahramanmaras¸, and
Osmaniye) of southern Turkey. A total of 182
accessions of fruit and/or seed were collected. The
sampling locations are shown in Fig. 1. All collected
accessions belong to L. siceraria. The samples were
provided by farmers or purchased from local seed
markets. The accessions were labeled with the traffic
plate number of the province and numbered accord-
ing to the order of collection (e.g., 01-01 means the
first accession collected from Adana province). The
traffic plate numbers of provinces are 01 for Adana,
1258 Genet Resour Crop Evol (2008) 55:1257–1266
123
07 for Antalya, 31 for Hatay, 27 for Gaziantep, 33 for
Mersin, 46 for Kahramanmaras¸, and 80 for
Osmaniye.
Seeds of the collected 182 accessions were sown on
March 17th 2004 in a mixture of peat and perlite (1:1)
in a greenhouse at the Department of Horticulture,
Faculty of Agriculture, Mustafa Kemal University.
The experimental site was located on 36°1802200 N,
36°1303300 E, 82 m. The yearly average rain fall for the
region varies between 570 and 1,160 mm and the soil is
described as sandy-loam having low amount of lime
(Kılıc1999). Seedlings were subsequently trans-
planted at the emergence of the first true leaves into
the field at 3 m between rows and 0.5 m within rows.
Out of the 182 accessions, 162 emerged and success-
fully produced seedlings. Ten seedlings from each
accession were transplanted to the field. Soil was
amended with 100 kg N/ha, 100 kg P
2
O
5
/ha and
100 kg K
2
O/ha during soil preparations. Plants were
watered by drip irrigation. All observations were
carried out on 10 plants.
Each accession was classified for 18 qualitative traits
following the standards of the descriptor list published
by the Bioversity International for Cucurbitaceae
(2007). The class scores were also presented in Table 1.
Six qualitative characters were measured for each
treatment in replicates. Seed color was measured as
reflected in the CIELAB (L*a*b*) color space using a
Minolta model CR-300 Colorimeter (Minolta, Osaka).
Three readings were taken from each accession.
Lightness L* was determined with L* range from
0=black to 100 =white (McGuire 1992). Seed
weight was determined by weighting 100 mature seeds
from each accession. Hypocotyl length and diameter
were measured at the first true leaf stage of 50 seedling
from each accession. Ten internodes on the main stem
were measured and the mean was presented as inter-
node length. Days of flowering were determined based
on days after seed sowing.
The statistical analyses were carried out using SAS
(SAS Institute 2005). For the descriptive statistics,
TABULATE was employed using the unconverted
data. PRINCOMP was used to perform a principle
component (PC) analysis using six quantitative
variables. The distributions of qualitative variables
were calculated for each province.
Results
A great deal of the genetic diversity was observed
among the bottle gourd genotypes collected from the
Mediterranean region of Turkey. The morphological
variation was most apparent in fruit shape and fruit
size. The fruit shape varied from globular to spindle-
elongated. Some of the accession had necks with
Fig. 1 Map of the collection sites for bottle gourd (L. siceraria) genotypes in the Mediterranean region in Turkey
Genet Resour Crop Evol (2008) 55:1257–1266 1259
123
Table 1 Distribution of qualitative variables of bottle gourd (L. siceraria) genotypes collected from the Mediterranean region in
Turkey
Variable Class Score Adana Osmaniye Gaziantep K. maras¸ Antalya Hatay Mersin Total
Seed size Small 3 4 – – 3 15 8 3 33
Medium 5 4 2 3 3 19 26 23 80
Large 7 4 2 – 8 7 13 15 49
Seed margin Absent 0 7 1 3 5 14 35 9 74
Thin and uniform 1 1 1 – 6 13 10 22 53
Thin and irregular 2 4 2 – 3 14 2 10 35
Thick uniform 3 – – – – – – – 0
Thick irregular 4 – – – – – – – 0
Seed margin color Absent 0 7 1 3 5 14 35 9 74
White 1 – – – – – – – 0
Tan 2 – – – – – – – 0
Yellow 3 – – – – – – – 0
Orange 4 – – – – – – – 0
Brown 5 3 2 – 6 20 8 25 64
Grey 6 – – – – – – – 0
Black 7 – – – – – – – 0
Light brown 8 2 1 – 3 6 3 6 21
Dark brown 9 – – – – 1 1 1 3
Cotyledon size Small 3 1 – – – 6 1 6 14
Intermediate 5 11 4 3 13 34 46 35 146
Large 7 – – – 1 1 – – 2
Cotyledon color Light green 3 4 2 – 3 8 12 4 33
Intermediate 5 5 2 2 4 25 31 28 97
Dark green 7 3 – 1 7 8 4 9 32
Leaf shape Ovate 1 0
Orbicular 2 5 – 1 3 15 27 27 78
Reniform 3 – – – – 4 1 – 5
Retuse 4 – – – – – – – 0
Heart 5 7 4 2 11 22 19 14 79
Leaf size Small 3 2 2 – 2 3 2 2 13
Intermediate 5 4 2 1 10 26 30 27 100
Large 7 6 2 2 12 15 12 49
Leaf edge Smooth 1 11 1 3 11 38 39 37 140
Toothed 2 1 3 – 3 3 8 4 22
Leaf pubescence Small 3 6 – – 4 5 2 17
Intermediate 5 6 4 3 10 34 27 17 101
Large 7 – – – – 2 20 22 44
Pubescence of upper
surface of leaf
Small 3 10 3 3 13 19 – 2 50
Intermediate 5 1 1 – 1 21 25 17 66
Large 7 1 – – – 1 22 22 46
Flower size Small 3 1 – – 2 8 5 10 26
Intermediate 5 5 4 3 10 28 31 22 103
Large 7 6 – – 2 5 11 9 33
1260 Genet Resour Crop Evol (2008) 55:1257–1266
123
various shapes. A general view showing some of the
observed variations in fruit shape and size is
presented in Fig. 2. Similarly, Fig. 3shows variations
in seed shape, size and color.
The distributions of qualitative traits, for each
province, were presented in Table 1. The mean and
standard deviations (SD) for the quantitative traits of
all accessions were presented in Table 2. All
quantitative traits varied considerably. For example,
seed color ranged from 46.9 to 62.7, while seed
weights were from 11.0 to 28.8. Similarly, the
qualitative traits were distributed into several dif-
ferent classes for each of the variables tested. For
example, for seed, cotyledon and leaf sizes, all three
classes (small, medium and large) were observed in
the collection. However, few variables missed
several classes. For example, seed margin color
had four classes, while there were 10 classes for
margin color.
Our results of the principle component analyses
for the qualitative traits indicated that the first three
PCs explained 26%, 21% and 17% (a total of 64%) of
the total variation (Table 3). Hypocotyl length and
diameter contributed most to the PC1, while seed
weight had the biggest contribution to the PC2. The
three dimensional presentation of all accessions,
grouped by their origin of provinces, is presented in
Fig. 4. In Figs. 3and 4groupings can be observed:
for example, the genotypes from Adana and Mersin
were generally grouped in the upper right side of
Fig. 4, while the genotypes from Kahramanmaras¸
were on the left side. However, the grouping based on
provinces was not apparent for all the accessions (as
in the case of the genotypes from Antalya and
Antakya distributed across all the sections of the
diagram).
After the evaluation of the 24 morphological traits
(both quantitative and qualitative), a subset collection
(core) of the bottle gourd genotypes was developed in
order to maximize the diversity in a smaller sample of
the accessions. Sampling size was taken into consid-
eration as well. In Table 2, the descriptive statistics
of the core collection are presented. The largest
differences between collections were in maximum
values of fruit shapes and cotyledon sizes and
minimum values of peduncle attachments and days
to flowering. None of the other traits varied greatly
between the whole and core collections. The
Table 1 continued
Variable Class Score Adana Osmaniye Gaziantep K. maras¸ Antalya Hatay Mersin Total
Place of female flower Main stem 1 – – – – – – – 0
Lateral stem 2 8 4 3 13 37 38 35 138
Both 3 4 – – 1 4 9 6 24
Peduncle transactional
shape
Round 3 3 3 – 2 5 1 3 17
Slightly angled 5 7 1 2 8 17 5 11 51
Angled 7 2 – 1 4 19 41 27 94
Peduncle attachment Easy 3 – – 1 1 – – – 2
Intermediate 5 12 4 2 13 41 47 41 160
Difficult 7 – – – – – – – 0
Stem end fruit shape Depressed 1 1 – 2 3 1 1 2 10
Flattened 3 3 1 – 4 2 11 5 26
Rounded 5 3 1 1 6 20 28 24 83
Pointed 7 5 2 – 1 18 7 10 43
Blossom end shape Depressed 1 2 – – 3 12 6 12 35
Flattened 3 7 2 1 5 10 24 11 60
Rounded 5 2 2 2 4 15 15 14 54
Pointed 7 1 – – 2 4 2 4 13
Variation in fruit shape Low 3 12 4 3 14 39 44 39 155
Intermediate 5 – – – – 2 – 1 3
High 7 – – – – – 3 1 4
Genet Resour Crop Evol (2008) 55:1257–1266 1261
123
comparisons of the core and whole collections are
presented in Fig. 5. As can be seen in Fig. 5, all of
the individuals of the core collections are well-
distributed among the whole collection, indicating
that the core collection is a satisfactory representative
of all the accessions.
Discussion
In this study, L. siceraria landraces were collected
from the Mediterranean coastal region of Turkey. The
162 accessions collected from 15 trips, the collection
adequately represented the bottle gourd population
present in the region. We measured/observed many
morphological traits on these accessions to reveal the
amount of genetic diversity. Our results demonstrated
that although Turkey is not the centre for origin, there
is still a great deal of genetic diversity. It is
interesting to note that Sensoy et al. (2007) also
found large amounts of genetic variations in melon
genotypes (Cucumis melo L.) in Turkey, which was
proposed as a secondary gene centre of genetic
diversity for the species.
We evaluated the entire collection for 24 mor-
phological traits in a homogenous experimental field
at the Amik Plain in Hatay, Turkey. The previous
observations reported fruit size and shape as the
Fig. 2 Picture of the
diversity in fruit size and
shape for bottle gourd
(L. siceraria) genotypes
which were collected from
the Mediterranean region in
Turkey and evaluated in a
common field
1262 Genet Resour Crop Evol (2008) 55:1257–1266
123
most apparently distinguished morphological traits
were (Sakar 2004). Our results corroborated these
findings as we found fruit size to be an important
discriminating character for the species.
Furthermore, other important morphological char-
acters that represented the diversity among the bottle
gourd accessions collected from Mediterranean
region of Turkey included fruit and blossom end
shapes, seed weight and size, seed margin and seed
margin color, and stem.
In our analyses, although there were some group-
ings based on the provinces, overall, distinct and
apparent geographical patterns were not detected. All
the 24 morphological traits were also analyzed using
correspondence analysis with similar conclusions
(data not presented). Although the provinces of
Turkey are artificial geographical areas, they may
not necessarily represent distinct ecological regions.
For example, the neighboring provinces of Adana and
Osmaniye are small and similar in ecological condi-
tions. Antalya and Mersin are also neighbors with
much larger land areas. The sampling of two
accessions from east of Antalya and west of Mersin
could be closer to one another than the two locations
within Antalya or Mersin. Given that Turkey is not
the centre of origin, it is possible that the bottle gourd
have been introduced to Turkey from multiple
locations. Finally, we have observed that bottle gourd
genotypes have been well-mixed in the Mediterra-
nean region. When we questioned farmers about the
origins of the accessions, we have been informed that
the accessions were brought from other areas. Since
bottle gourd has been utilized in so many different
ways, they are most likely to be dispersed to wide
ranging sites through the movement of people over a
long period of time. A sample for multiple introduc-
tions of Lagenaria accessions can be presented from
Italy. There are two major races of Lagenaria
Fig. 3 Picture of the diversity in seed size and shape for bottle
gourd (L. siceraria) genotypes which were collected from the
Mediterranean region in Turkey and evaluated in a common field
Table 2 Mean, standard deviation and range of the whole and
core groups of bottle gourd (L. siceraria) genotypes collected
from the Mediterranean region in Turkey
Mean ±SD Range
Variable Whole Core Whole Core
Seed color 53.7 ±3.2 54.5 ±3.3 46.9–62.7 49.1–62.7
Seed weight 19.8 ±3.3 19.5 ±3.0 11.0–28.8 13.5–25.3
Hypocotyl
length
3.9 ±0.7 3.9 ±0.7 2.2–5.6 2.6–5.5
Hypocotyl
diameter
3.1 ±0.3 3.1 ±0.3 2.5–4.0 2.6–3.9
Internode
length
15.1 ±2.3 14.9 ±2.3 9.1–20.0 9.1–20.0
Days to
flowering
76.3 ±1.2 76.0 ±3.2 65.0–82.0 65.0–82.0
Table 3 Mean, standard deviation and range of the whole and
core groups of bottle gourd (L. siceraria) genotypes collected
from the Mediterranean region in Turkey
Variable PC1 PC2 PC3
Seed color -0.43 -0.44 0.34
Seed weight 0.36 0.63 0.16
Hypocotyl length 0.60 –0.19 0.10
Hypocotyl diameter 0.51 –0.34 0.21
Internode length -0.20 0.39 0.75
Days to flowering -0.16 0.32 –0.48
Eigen value 1.57 1.3 1.03
Difference 0.26 0.27 0.11
Proportion 0.26 0.21 0.17
Genet Resour Crop Evol (2008) 55:1257–1266 1263
123
siceraria in Italy; L. siceraria ssp. siceraria for
making bottles and as vegetable and L. siceraria ssp.
asiatica (Kobiakova) Heiser with very long fruits
used as vegetable. Although ssp. siceraria exists
since classical times ssp. asiatica has been introduced
only in recent centuries (Hammer et al. 1999).
The way we evaluated the accessions may have
also contributed to this conclusion. For example, to
assess the fruit shape, we used an evaluation scale
modified from the Bioversity International’s Cucur-
bitaceae (2007) descriptors’ list. Although the list
utilized in our study had more groups, both lists
considered the fruit shape to be categorical data. A
quantitative approach employing fruit dimensions
would have better represented this trait.
When an ideal core collection is compared to the
whole collection, their means and SDs are expected
to be similar (Van Hintum et al. 2000). For many of
Fig. 4 Principle
component analysis plot of
first three principle
components, depicting
relationship among
genotypes collected from
Mediterranean region in
Turkey and evaluated in a
common field. The analysis
was conducted using six
quantitative, morphological
traits
Fig. 5 Principle
component analysis plot of
first three principle
components, depicting
distribution of core
collection among all
genotypes collected from
Mediterranean region in
Turkey and evaluated in a
common field. The analysis
was conducted using six
quantitative, morphologic
traits
1264 Genet Resour Crop Evol (2008) 55:1257–1266
123
the traits tested, all descriptive statistics were com-
parable between the whole and core collections. Our
results indicate that the core collection composed in
our study is valid and adequately represents the whole
collections.
We characterized L. siceraria genotypes using
morphological traits and also studied L. siceraria
molecularly. Molecular markers have proven useful
in assessing land race diversity in L. siceraria.
Decker-Walters et al. (2001), for example, charac-
terized 31 landraces and 43 cultivars by RAPD
markers. The accessions were from Africa, Asia and
the New World. The results revealed that the
accessions from these geographical regions were
distinct and commercial cultivars had a diverse
genetic base. Decker-Walters et al. (2004) also
studied a wild population of L. siceraria from
Zimbabwe by both nuclear DNA, by RAPD, and
chloroplast DNA by sequencing to search for poly-
morphism. Their results made it possible for them to
draw the conclusion that the population was a part of
a genetically distinct and wild lineage of L. siceraria.
Assessment of our collection by molecular markers
would be valuable to further characterize the genetic
background of the Turkish bottle gourds. Moreover,
if the Turkish genotypes were studied with African
and Asian accessions simultaneously, it may be
possible to trace the origin of the L. siceraria
genotypes found in Turkey. Being located in the
intersections of Euro-Siberian, Irano-Turanian and
Mediterranean Phytogeographic regions, Turkey is an
important genetic diversity centre as well as an
immigration road as a bridge between the three
continents (Tan and Tan 1998). Therefore, it may be
possible that L. siceraria may have been introduced
to Turkey from Africa and Asia separately.
Other than the direct usage, the major utilization of
bottle gourd is as a rootstock for watermelons
because bottle gourd is resistant to many soil-borne
diseases and can tolerate some undesirable soil
conditions (Lee 1994). There are several bottle gourd
rootstocks bred in Japan and Korea for this purpose.
In future work, we will evaluate the rootstock
potential of the Turkish bottle gourd core collection
composed from the present study. First, the compat-
ibility of the bottle gourd genotypes, with various
watermelon cultivars, will be tested. Then, using the
compatible combinations, resistance/tolerance of the
core collection’s genotypes will be evaluated against
salinity, drought, and water logging. It is likely that
we may identify some genotypes more suitable to
Turkey conditions than the present commercial bottle
gourd–watermelon rootstock developed outside of
Turkey. The seeds of genotypes collected and char-
acterized in this study are preserved in the cold
storage of Aegean Agricultural Research Institute
(AARI) in _
Izmir at -20°C for long term storage.
They are also stored at 4°C in Horticultural Depart-
ment of Mustafa Kemal University. Sample seeds can
be obtained from either from the corresponding
author or AARI.
Acknowledgements We gratefully acknowledge the
financial supports from the Scientific and Technological
Council of Turkey (TOGTAG-3216) and Mustafa Kemal
University.
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