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DOI: 10.2298/GENSR1303679Z
Original scientific paper
1
Institute for Vegetable Crops, Smederevska Palanka, Serbia
2
Faculty of Agriculture, University of Belgrade, Belgrade, Serbia
INTRODUCTION
Tomato (Lycopersicon esculentum Mill) is one of the most widely grown vegetables in
the world and, therefore, any factor influencing tomato yield has been attracted considerable
interest. Among environmental factors, drought is one of the major limiting factors of tomato
fruit growth and productivity (BENTON JONES, 1999). Furthermore, current climate change
scenarios in Europe predict that drought events during summer and heavy rainfalls during the
other seasons will become more frequent during the next decades (GAO and GEORGI, 2008).
Thus, a detailed understanding of the determinants of plant stress tolerance and subsequent
breeding for increased stress tolerance will play a major role in future agriculture (LOBELL et al.,
2008). Adaptation measures to mitigate the reduction of yield induced by drought include the
production and use of drought-resistant genotypes. The prerequisite to produce resistant
genotypes of tomato, as other vegetables, is a better understanding of the plant response and
adaptation to drought stress, the improvement of phenotyping, the selection of key-genes
involved in the resistance to drought and the evaluation of the impact of resistance on crop yield
and quality. These are very difficult tasks because drought tolerance is quantitative traits, with
complex phenotype and genetic control. The reaction of plants, including tomato, to drought is
the complex phenomenon where the plant response depends on genotypes, the type, duration or
intensity of drought and on phenological stage in which drought stress is experienced (CHAVES et
al., 2003).
The complexity of drought effects on plants and drought tolerance mechanisms explains
the slow progress in breeding tomato for drought conditions. The most commercial tomato
cultivars are drought sensitive at all stages of development, including seed germination, seedling
emergence, vegetative growth and reproduction. Furthermore, tomato stress tolerance is a
developmentally regulated state-specific phenomenon, such that tolerance at one stage of plant
development is independent of tolerance at other stages. Conventional breeding based on
phenotypic selection is further complicated by the fact that several types of abiotic stresses (as
high temperature or high irradiance, water and nutrient deficiency) in the field conditions can
influence plants reactions simultaneously and may have adverse impact on the accuracy and
repeatability of investigated phenotypic traits.
The different strategies have been suggested for selecting resistant and relatively
resistant genotypes to drought stress. According to ZAMIR (2001) development of exotic genetic
libraries consisting of marker-defined genomic regions taken from wild species and introgressed
to the background of elite crop varieties will provide a resource for the discovery and
characterization of genes that underlie traits of agricultural value. The potential sources of
drought tolerance in tomato are found in wild species, including L. chilense, L. peruvianum, L.
pennellii, L. pimpinellifolium, L. hirsutum, L. cheesmanii, L. chmielewskii and L. parviflorum
(FOOLAD, 2005). Using this approach GUR and ZAMIR (2004) could demonstrate that introgressed
tomato lines carrying three independent yield-promoting genomic regions produced significantly
higher yield than then control lines grown under drought conditions.
Breeding for resistance to drought of tomato is also complicated by the lack of fast and
reproducible screening techniques. Although a large number of different traits during vegetative
and reproductive growth phases have been employed to characterize the physiological and
genetic basis of drought tolerance in tomato, it is still difficult to identify drought tolerant
genotypes (FOOLAD, 2005). To differentiate the degree of drought resistance between different
genotypes, several selection indices have been suggested. One of them is the stress susceptibility
J. ZDRAVKOVIC et al: TOMATO CULTIVARS UNDER DROUGHT STRESS 681
index (SSI) which is a ratio of genotypic performance under stress and non-stress conditions
(FISCHER and MAURER, 1978). Efforts to overcome the problem of drought and to save water for
irrigation of tomatoes were, in the previous period, solved by applying the agro-technical
interventions (measures). Partial root drying (PRD) technique has been the subject of research of
the members of the research team in the previous period (STIKI et al. 2003, SAVI et al. 2004,
2006). However, there was a need to include this problem in selection programs of tomato, in
order to solve the global problem multidisciplinary: by affecting the physiological processes of
growth and creating tolerance in selected genotypes for decreased irrigation conditions
(ZDRAVKOVI et al 2010).
The aim of presented study was to use the stress tolerance index to screen the drought
tolerance of 40 tomato genotypes from domestic tomato population collected in Serbia.
Analysis was done on the base of two drought-associated traits (plant height and shoot/root ratio)
during vegetative phase of plant development. The identified genotypes with increased drought
tolerance will be use as starting material in the tomato drought tolerance breeding program.
Table 1. The effects of drought (D) and full irrigation (FI) on shoot/root ratio and plant height in
investigated tomato genotypes
a) Determinant genotypes
Genotypes Shoot/root ratio Plant height (cm) Plant height reduction
FI D FI D (%)
b)Indeterminate genotypes
Genotypes Shoot/root ratio Plant height (cm) Plant height reduction
(%)
FI D FI D
G120 5,46 12,43 97,2 66,8 31,28
G121 7,08 10,59 89,6 56,6 36,83
G122 6,90 12,06 77,4 50,76 34,42
G123 4,36 9,82 78,8 58,8 25,38
G124 6,76 5,89 93,8 65 30,70
G125 6,12 6,50 80,6 59,2 26,55
G126 4,55 4,21 118,6 94 20,74
G127 6,89 12,18 101,4 83 18,15
G128 6,50 6,45 106,6 75 29,64
G129 4,18 9,02 111 87,8 20,90
G130 2,13 7,09 141,8 105,8 25,39
G131 1,91 5,18 124,4 87,8 29,42
G132 2,22 4,41 137,8 100,6 27,00
G133 1,85 5,39 141,2 105 25,64
G134 2,07 6,93 116 81 30,17
G135 2,62 4,51 121 95,6 20,99
G136 3,30 7,78 130,8 80 38,84
G137 3,56 7,27 111,8 80 28,44
G138 2,76 6,21 121 89,4 26,12
G139 2,29 3,38 106,2 89,6 15,63
G140 6,09 15,52 126,4 78,25 38,09
G141 5,78 6,57 128,32 106,6 16,93
Stress
intensity -0,708 0,265
(SI)
The evaluation of reaction to drought of investigated genotypes was also done by using
of SSI index. There are a number of ways to determine stress intensity, but this one has been
established as the most useful in the comparative analysis among the four indicators of stress
effect (ILKER et al., 2011). Through this index, more or less tolerant genotypes to drought can be
specified. When the amount of SSI is less, the tolerance of genotype to drought stress will be
higher. Graph 1 show the SSI values for plant height which was in the range from 0 to 1.6. The
SSI values between 0.4 and 0.6 of several genotypes (G102, G104, G109, G110, and G139)
indicated that these genotypes can be considered highly tolerant to drought caused stress. Higher
SSI values (0.6 to 0.8 ranges) which were still within the limits can be considered as values for
684 GENETIKA, Vol. 45, No.3,679-689, 2013
moderate drought tolerance. These values were calculated for following genotypes: G105, G126,
G127, G129, G135 and G141.
Fig 1. The stress susceptibility index calculated on the base of plant height values.
The SSI values for the shoot/root ratio where in the range from 1 to 4 (Graph 2). The
values between the -0.5 and 0.5 were considered as values indicating the genotypes with the
tolerance to drought (G105, G107, G110, G119, G124, G125, G126, G128, and G141). The SSI
values for four genotypes (G110, G124 and G126) were negative, which indicated that under
drought conditions, the development of the root system in these genotypes was more intensive
than shoot growth.
Fig. 2: The stress susceptibility index calculated on the base of shoot/root ratio values.
J. ZDRAVKOVIC et al: TOMATO CULTIVARS UNDER DROUGHT STRESS 685
Plant growth rate is generally inhibited by soil drying, but many results confirmed that
root growth is less influenced by the soil water deficit than that of the shoot. The maintenance of,
or even promotion of, root growth during soil drying can provide several advantages, such as a
better exploitation of soil nutrients and water when environmental conditions are less favorable
(SHARP et al., 2004). The increase in root dry mass in our experiments was most likely
attributable to stimulated secondary root initiation and root growth. The similar trend of shoot
and root growth and dry matter accumulation has been recorded by other authors in tomato
plants grown under drought or deficit irrigation conditions (MINGO et al., 2004: PROKIC and
STIKIC, 2012). There is also some consistency in results concerning the roots characteristics.
Drought resistance of tomato related wild species L. chilense was attributed to its deep vigorous
root system. In contrast the results for the drought-tolerant L. pennellii accession LA716
showed a limited and shallow root system. In the basis of drought tolerance of this genotype is
the specific succulent leaf structure with allows reduction of transpiration during the drought
period (FOOLAD, 2005).
The final estimation of genotypes to be included in the selection program for drought
tolerance involved the use of cluster method. The aim of the analyses was to classify different
genotypic groups in similar classes. The dendrograms of plant height mean values for genotypes
grown under the optimal water regime showed the distribution of genotypes into two groups (Fig
3). The formation of three (I, IIa and IIb) groups indicates the existence of divergence of
population selected for the tomato breeding program for drought tolerance. The largest number
of genotypes under the irrigation regime belonged to cluster I (17 genotypes), then to cluster IIa
(14), and to cluster IIb (9).
G101
G103
G128
G126
G134
G129
G137
G139
G127
G105
G114
G121
G120
G124
G130
G133
G136
G132
Genotypes
G131
G140
G135
G141
G138
G102
G115
G118
G123
G111
G104
G106
G116
G107
G122
G125
G109
G117
G110
G119 Linkage Distance
G112
G113
Fig 3. Dendogram of genetic distance of genotypes (Euclid distance) based on plant height of genotypes
under full irrigation conditions.
686 GENETIKA, Vol. 45, No.3,679-689, 2013
G101
G131
G126
G139
G127
G129
G138
G128
G140
G134
G136
G137
G130
G133
G141
G132
G135
G102
Genotypes
G114
G109
G123
G121
G125
G110
G120
G103
G104
G105
G124
G106
G107
G118
G119
G111
G122
G117
G115
G116
G112 Linkage Distance
G113
Fig 4. Dendogram of genetic distance of genotypes (Euclid distance) based on plant height of genotypes
under drought conditions.
The dendrograms resulted from cluster analysis of genotypes under drought conditions
stress conditions was not diametrically different from the dendogram of plants under the optimal
water regime. All genotypes were also located into two larger groups (I and II), and each cluster
contains less two smaller groups (Ia,Ib and IIa, IIb, respectively), Fig 4. However some
genotypes were distributed into different clusters (G124, G120, G121,G114, G105 and G103)
which only indicate that they responded alike by plant height to drought impact. The largest
number of genotypes is found in clusters Ib and IIb (12 genotypes), Ia (11 genotypes), and
smallest group IIa (5 genotypes).
Of special importance in drought tolerance breeding program is phenotypic
characterization of useful germplasm. In the literature there is a limiting data concerning this
issue. More data are available for tomato resistance to another stress factors as salt or
temperature (FOOLAD, 1999). The majority of available selection criteria for screening or
breeding tomatoes for drought resistance come from the research done with tomato wild
relatives. In these experiments different tolerance indices were used: including dry weight (DW)
of shoot and root, root length, root morphology, leaf rolling, flower and fruit set, fruit weight,
fruit yield, WUE, recovery after re-watering, stomatal resistance, plant survival, leaf water
potential, leaf osmotic potential, osmoregulation, transpiration rate, photosynthetic rate,
enzymatic activities etc. (FOOLAD, 2005). However, these traits should be further checked in
tomato inbred lines or cultivars under drought conditions. There are also results done by
genotypes which not belong to the exotic germplasm. These results indicated that in these
genotypes, genes responsible for drought resistance may be found (MANOJ and UDAY, 2006,
2007). Recently, WAHB-ALLAH et al. (2011) have screened drought resistance of four tomato
cultivars based on several vegetative and fruit traits (plant height, stem diameter, leaf and fruit
J. ZDRAVKOVIC et al: TOMATO CULTIVARS UNDER DROUGHT STRESS 687
dry matter, fruit setting and yield) under drought conditions. Significant differences among
genotypes were found for almost all of these traits (except stem diameter), suggesting that they
could be taken into account when selecting for tomato drought tolerance.
Phenotypic analysis of genotypes for plant height has shown a high level of
heterogeneity and thus, may represent a desirable source of genetic variability. Dendograms
constructed for optimal irrigation regime and for drought conditions revealed two population
groups. These groups are already known as the groups of determinate and indeterminate types of
shoot growth habit. Their heterogeneity within a sub-cluster indicated diversity and possibility to
choose suitable genotypes.
CONCLUSIONS
The results of stress susceptibility index and grouping of genotypes on the base of plant
height and shoot/root ratio, lead to the conclusion that 11 genotypes (G102, G104, G107, G109,
G110, G119, G125, G128 and G141) exhibited better drought tolerance characteristics than
others from domestic population. Thus, they and could be suitable initial genetic material in
the selection program for tomato drought tolerance. However, broader conclusions can be
obtained only after molecular analyses of these genotypes.
ACKNOWLEDGEMENT
This study was supported by Serbian Ministry of Education and Science (project TR 31005 and
TR 31059).
Received July 22th, 2013
Accepted October 05th, 2013
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J. ZDRAVKOVIC et al: TOMATO CULTIVARS UNDER DROUGHT STRESS 689
Izvod
Ispitivanje je izvreno na 41genotipu paradajza poreklom iz populacije domaih i odomaenih
genotipova prikupljenih iz Srbije, a pripadaju kolekciji paradajza Instituta za povrtarstvo u
Smederevskoj Palanci. Skrining kolekcije imao je za cilj da se izvri izbor genotipova tolerantnih
na suu u vegetativnoj fazi intenzivnog porasta biljaka, ime bi se zapoeo program selekcije na
dobijanje rekombinovanih genotipova prema ovom abiotskom faktoru. Kriterijumi za skrining
bili su divergentnost genotipova za visinu biljke i odnos izdanak/ koren u uslovima: optimalnog
reima navodnjavanja i sue. Za ocenu divergentnosti koriena je klaster analiza sa
Euklidovom distancom kao merom udaljenosti, sa kompletnom vezanou gena za formiranje
grupa. Tolerancija na suu izraena je indeksom osetljivosti na stres (SSI-stress susceptibility
index). Razliiti rezultati su dobijeni na osnovu skrininga genotipova koji su gajeni u optimalnim
uslovima i u sui. Kao mera osetljivosti na stres prema indeksu osetljivosti definisani su
genotipovi sa razliitim nivoom tolerancije na suu. Na osnovu izvedenih analiza izdvojeno je 10
genotipova (G102, G104, G107, G109, G110, G119, G125, G126, G128 i G141) koji e
predstavljati bazu za dobijanje rekombinovanih genotipova i poetak selekcije na otpornost na
suu.