ISSN e:2007-4034 / ISSN print: 2007-4034

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Vol. 24, issue 2 May - August 2018

ISSN: ppub: 1027-152X epub: 2007-4034

Scientific article

Genetic diversity within wild species of Solanum

http://dx.doi.org/10.5154/r.rchsh.2017.08.030

Flores-Hernández, Luis Antonio 1 ; Lobato-Ortiz, Ricardo 1 * ; Sangerman-Jarquín, Dora María 2 ; García-Zavala, J. Jesús 1 ; Molina-Galán, José D. 1 ; Velasco-Alvarado, Mario de Jesús 1 ; Marín-Montes, Iván Maryn 1

  • 1Colegio de Postgraduados, Campus Montecillo. Carretera México-Texcoco km 36.5, Montecillo, Texcoco, México, C. P. 56230, MÉXICO.
  • 2Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo Experimental Valle de México. Carretera Los Reyes-Texcoco km 13.5, Coatlinchán, Texcoco, México, C. P. 56250, MÉXICO.

Corresponding author: rlobato@colpos.mx, tel. (595) 20 200 ext. 1534.

Received: August 18, 2017; Accepted: October 25, 2017

This is an open-access article distributed under the terms of the Creative Commons Attribution License view the permissions of this license

Abstract

Cultivated tomato (Solanum lycopersicum L.) has undergone a reduction in its genetic base as a result of the processes of modern domestication and breeding, which has been extensively documented by molecular markers in different genotypes, both nationally and internationally. Faced with this situation, some plant breeders in Mexico have proposed making use of native Mexican germplasm, as well as of the genetic variation present in wild species related to the cultivated tomato. The aim of this study was to characterize agronomically, under greenhouse conditions, accessions of five wild relatives of the cultivated tomato for their incorporation into breeding programs of this vegetable. In addition, it is expected to reduce its vulnerability to climate change and adverse biotic and abiotic factors. The species described were Solanum pennellii L., Solanum pimpinellifollium L., Solanum peruvianum L., Solanum chilense R. and Solanum habrochaites S. The accessions were evaluated under greenhouse conditions under a completely randomized experimental design with four replications. Twelve traits of agronomic interest were evaluated to describe the variation between the accessions of each one of the evaluated species, which were studied by analysis of variance and comparison of means. The results showed high significance among the accessions of each one of the evaluated species for all the traits. The above shows that there is a high potential in each of the accessions of the species studied to exploit them genetically in the improvement of the cultivated tomato.

Keywords:Solanum pennellii L.; Solanum pimpinellifollium L.; Solanum peruvianum L.; Solanum chilense R.; Solanum habrochaites S

Introduction

The wild relatives of the cultivated tomato (Solanum lycopersicum L.) are distributed in Ecuador, Peru, northern Chile and the Galapagos Islands (Peralta & Spooner, 2001), but Mexico is considered as its center of domestication (Jenkins, 1948; Peralta & Spooner, 2007; Rick & Fobes, 1975; Rodríguez et al., 2011). Wild tomatoes grow in diverse habitats, from those found at sea level to almost 3,300 masl (Rick, 1973; Taylor, 1986).

The species evaluated in this work have a wide distribution. Solanum pennellii L. grows in western Peru; its habitats are not evenly dispersed, but they are grouped along streams in the west of the country, usually between 500 and 1,500 masl (Rick, 1973). Solanum pimpinellifollium L. is distributed along the coasts of Peru and Ecuador and has been used frequently in tomato breeding. The natural distribution area of Solanum peruvianum L. is mainly Peru, where it ranges from the west coast in the Andes to northern Chile (Chetelat, Pertuze, Faundez, Graham, & Jone, 2009). Solanum chilense R. is found mainly in southern Peru to the north of Chile, from 0 to 3,000 masl. The last species studied, Solanum habrochaites S., is located from southwestern Ecuador to the southern part of Peru, between 500 and 3,300 masl (Spooner, Peralta, & Knapp, 2005).

The evolution of the wild relatives of the tomato to the cultivated one resulted in an increase in productivity, but at the same time to a reduced genetic base of the present varieties (Ladizinsky, 1998); therefore, cultivated varieties have been negatively affected by biotic and abiotic factors. To counteract this situation, the use of native germplasm or wild relatives is required for the introgression of new allelic combinations of tomatoes to increase their productivity, quality, resistance or tolerance to biotic and abiotic factors (Cervantes-Moreno, Rodríguez-Pérez, Carrillo-Fonseca, Sahagún-Castellanos & Rodríguez-Guzmán, 2014; Fernie, Tadmor, & Zamir, 2006; Foolad, 2007; Gur & Zamir, 2004; Hernández-Bautista, Lobato-Ortiz, Cruz-Izquierdo, García-Zavala, & Chávez-Servia, 2014; Hernández-Bautista et al., 2015; Marín-Montes, Rodríguez-Pérez, Sahagún-Castellanos, Hernández-Ibañez, & Velasco-García, 2016).

Some authors indicate that the genetic diversity obtained from the tomato’s wild relatives is 95 %, while in the cultivated tomato only 5 % is obtained (Miller & Tanksley, 1990). Currently, one of the strategies in tomato breeding is to use the diversity that was lost during the domestication processes of the current varieties (Zamir, 2001); this diversity must be found in its wild relatives. Therefore, the aim of this work was to characterize agronomically, under greenhouse conditions, accessions of five wild relatives of the tomato for their incorporation into breeding programs of this vegetable.

Materials and methods

This research was carried out at the Colegio de Postgraduados, Montecillo Campus, Texcoco, State of Mexico (19° 27’ North latitude and 98º 54’ West longitude, 2,246 masl), in the greenhouses of the graduate school’s Program for the Conservation and Improvement of the Genetic Resources of the Tomato in Mexico. In total, 39 accessions of five wild species related to the cultivated tomato (Figures 1, 2, 3, 4 and 5, Table 1), provided by the Tomato Genetics Resource Center (TGRC) of the University of California, Davis, USA, were evaluated.

Figure 1. S. pimpinellifollium L. accession LA0373

Figure 2. S. peruvianum L. accession LA1365

Figure 3. S. habrochaites S. accession LA2409

Figure 4. S. pennellii L. accession LA1272

Figure 5. S. chilense R. accession LA2759

Table 1. Species, accessions and origin of collections evaluated in this research.

Species Accession Origin of the collection Accession Origin of the collection
S. pennellii L. LA2580 Valle de Casma, Ancash, Perú LA1272 Pisaquera, Lima, Perú
LA0716 Atico, Arequipa, Perú LA1277 Trapiche, Lima, Perú
LA1367 Santa Eulalia, Lima, Perú
S. pimpinellifollium L. LA1584 Jayanca de La Vina, Lambayeque, Perú LA0373 Culebras núm. 1, Ancash, Perú
LA1689 Castilla núm. 1, Piura, Perú LA0442 Sechin, Ancash, Perú
LA1237 Atacames, Esmeraldas, Ecuador LA1576 Manchay, Alta Lima, Perú
LA1593 Puente Chao, La Libertad, Perú
S. peruvianum L. LA2172 Cuyca, Cajamarca, Perú LA0446 Atiquipa, Arequipa, Perú
LA1982 Huallanca, Ancash, Perú LA1346 Casmiche, La Libertad, Perú
LA1677 Fundo Huadquina, Topara, Ica, Perú LA1336 Atico, Arequipa, Perú
LA1973 Yura, Arequipa, Perú LA1274 Pacaibamba, Lima, Perú
LA1360 Pariacoto, Ancash, Perú LA1365 Caranquilloc, Ancash, Peru
LA0103 Cajamarquilla, Lima, Perú LA2152 San Juan #1, Cajamarca, Perú
S. habrochaites S. LA2409 Miraflores, Lima, Perú LA1223 Alausi, Chimborazo, Ecuador
LA1731 Río San Juan, Huancavelica, Perú LA1777 Rio Casma, Ancash, Perú
LA2650 Ayabaca, Piura, Perú GH0810
LA2158 Río Chotano, Cajamarca, Perú LA2167 Cimentario, Cajamarca, Perú
S. chilense R. LA2930 Guatacondo, Tarapaca, Chile LA1958 Pampa de la Clemesi, Moquegua, Perú
LA2750 La Despreciada, Antofagasta, Chile LA2778 Chapiquina, Tarapaca, Chile
LA1960 Río Osmore, Moquegua, Perú LA2748 Soledad, Tarapaca, Chile
LA2759 Tarapaca, Chile

A completely randomized experimental design was used with four replicates of ten plants each. Sowing was carried out on May 28, 2014, and the transplant (to 12-L polyethylene bags), 36 days after sowing. As substrate, volcanic sand (red tezontle) was used. The plants were irrigated with the nutrient solution proposed by Steiner (1984) at 25 % during the vegetative stage, at 50 % in flowering and at 100 % during fruit ripening. Additionally, Confidor® (imidacloprid) and Ampligo® (50 lambda cyhalothrin + 100 chlorantraniliprole) were used for the control of whitefly (Bemisia tabaci Gennadius), Captan® 50 plus (carboxamide) and Ridomil Gold® (metalaxyl-m + chlorothalonil) for late blight (Phytophthora infestans), and Amistar® (azoxystrobin) for early blight (Altenaria solani).

According to the tomato descriptors manual of the International Plant Genetic Resources Institute (IPGRI, 1996), 12 traits were evaluated: days to flowering (DF), days to maturity (DM), leaf length (LL, cm), leaf width (LW, cm), stem diameter (SD, cm), number of flowers per cluster (FC), cluster length (CL, cm), fruit weight (FW, g), fruit length (FL, cm), fruit width (FWi, cm), total soluble solids (TSS, °Brix) and number of seeds per fruit (SF). For the measurement of SD, FL and FWi, a Truper® digital standard and millimeter Vernier caliper was used. The LL, LW, CL traits were measured with a Truper® model FH-3M flexometer. The FW was obtained with an Ohaus® model SP2001 digital scale. An ATAGO® model PAL-1 digital refractometer with a range of 0.0 to 53.0 °Brix was used to evaluate TSS.

Average, range, coefficients of variation and standard deviation were calculated for each variable. Likewise, analysis of variance and Tukey’s range test (P ≤ 0.05) were performed with the Statistical Analysis System package (SAS Institute Inc., 2002). These tests were carried out in order to determine if there are significant statistical differences within the species evaluated and identify those accessions that had the highest and lowest parameters.

Results and discussion

The traits with the greatest variation among species were DF, FW and SF, with 45.57, 62.30 and 62.57 %, respectively. By contrast, those with the smallest variation were DM, FL, FWi and TSS with 14.70, 16.78, 18.07 and 17.82 %, respectively (Table 2).

Table 2. Descriptive statistics among and within species for the traits evaluated.

DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
General m 26.96 82.64 24.63 14.54 0.95 18.97 23.09 2.53 1.34 1.57 7.14 94.52
sd 12.29 12.15 7.48 4.19 0.19 5.62 8.22 1.57 0.22 0.28 1.27 59.14
r 42.00 50.00 32.00 17.70 0.98 22.00 34.40 7.10 1.23 1.40 4.80 217.30
cv 45.57 14.70 30.36 28.81 19.88 29.60 35.60 62.30 16.78 18.07 17.82 62.57
S. pennellii L. m 33.30 88.80 20.54 12.42 1.19 13.78 25.32 2.58 1.34 1.76 8.10 197.00
sd 1.37 8.53 2.44 0.81 0.07 2.18 1.17 0.72 0.05 0.05 1.49 31.95
r 3.60 20.00 5.60 1.90 0.17 5.80 3.00 2.00 0.10 0.10 3.20 84.00
cv 4.13 9.60 11.87 6.50 5.48 15.84 4.63 27.87 4.09 3.11 18.42 16.22
S. pimpinellifollium L. m 13.86 62.57 19.27 11.79 0.98 15.86 14.21 1.40 1.21 1.34 7.11 26.44
sd 1.86 13.29 2.07 1.98 0.15 3.72 5.34 0.27 0.07 0.16 1.06 8.91
r 5.00 32.00 5.40 5.50 0.36 10.00 14.00 0.70 0.20 0.40 2.80 22.00
cv 13.45 21.24 10.74 16.81 15.28 23.43 37.57 19.34 5.68 12.05 14.84 33.71
S. habrochaites S. m 43.00 92.88 36.49 20.76 0.97 21.75 16.88 2.94 1.43 1.71 7.91 84.25
sd 7.25 4.79 6.43 4.25 0.07 3.24 2.80 1.76 0.21 0.28 1.41 29.97
r 25.00 15.00 18.40 12.00 0.23 8.00 7.50 5.40 0.70 0.90 3.80 80.00
cv 16.86 5.16 17.61 20.48 7.16 14.90 16.59 59.94 14.89 16.35 17.84 35.57
S. peruvianum L. m 19.08 83.17 21.77 12.50 0.80 19.25 28.57 3.43 1.43 1.63 6.62 123.33
sd 6.93 3.33 3.35 1.66 0.11 6.80 8.74 1.99 0.30 0.27 0.90 32.40
r 26.00 11.00 12.30 5.80 0.37 22.00 30.80 7.00 1.10 0.80 3.20 89.00
cv 36.34 4.00 15.37 13.26 13.56 35.31 30.58 57.82 21.11 16.37 13.67 26.27
S. chilense R. m 30.71 85.71 24.27 15.21 1.00 22.14 28.06 1.59 1.20 1.37 6.47 51.71
sd 9.20 3.15 5.09 2.76 0.27 5.46 3.58 0.20 0.15 0.29 1.09 17.90
r 25.00 9.00 15.10 7.60 0.82 13.00 10.40 0.60 0.43 0.70 3.00 61.00
cv 29.94 3.67 20.96 18.13 27.04 24.66 12.76 12.31 12.44 21.35 16.85 34.60
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; m = arithmetic mean; sd = standard deviation; r = range; cv = coefficient of variation.

In Solanum pennellii L. the traits with the greatest variation were FW, TSS, SF and FC with 27.87, 18.42, 16.22 and 15.48 %, respectively (Table 2). Likewise, significant statistical differences were observed (P ≤ 0.05) among accessions in DM, LL, SD, FW, SF and TSS (Table 3). These differences are the product of the ecological niche and adaptability of the accessions to each of the environments. In spite of the above, the accessions of S. pennellii were the ones with the least variation with respect to the other species, which agrees with the findings reported by Rick and Tanksley (1981), who found that S. pennellii L. has stable and less variable characteristics between individuals and accessions.

Table 3. Comparison of means among accessions of Solanum pennellii L. for 12 traits.

Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2580 33.3 az 86 b 22.6ab 13.1 a 1.22 ab 16.9 a 26 a 1.7 c 1.4 a 1.7 a 6.6 b 192 ab
LA0716 34.6 a 97 a 18 b 11.2 a 1.25 a 14.8 a 25.4 a 3.7 a 1.4 a 1.8 a 6.5 b 149 b
LA1272 31 a 98 a 18.8 ab 13 a 1.21 ab 13.2 a 24.6 a 2.6 b 1.3 a 1.8 a 9.3 a 233 a
LA1367 34 a 78 c 19.7 ab 12 a 1.19 ab 12.9 a 23.8 a 2.5 bc 1.3 a 1.8 a 9.7 a 218 a
LA1277 33.6 a 85 b 23.6 a 12.8 a 1.08 b 11.1 a 26.8 a 2.4 bc 1.3 a 1.7 a 8.4 a 193 ab
DMSH 6.3 6.4 4.9 4.3 0.15 6.2 8 0.7 0.35 0.23 1.4 44
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).

Accessions LA2580 and LA0716 showed self-compatibility by presenting less variability with respect to the rest of the evaluated accessions, which presented self-incompatibility, in DF (2.7 %), SD (1.7 %), CL (1.6 %) and TSS (1 %) (Table 3). This agrees with what was reported by Mercer and Perales (2010), who indicate that the genetic variation of individuals is influenced by the type of reproduction, since individuals who have autogamy (self-compatibility) systems have less variation within the population and more between populations.

As for the quality of the fruit, Fernie et al. (2006) indicate that the increase in TSS in S. pennellii L. is the result of an increase in sucrose and glucose. The comparison of means showed that LA1272 and LA1367 have the highest amount of TSS (9.3 and 9.7, respectively) (Table 3). Therefore, these accessions can be used to improve the fruit quality of the elite tomato varieties.

On the other hand, among accessions of Solanum pimpinellifollium L., the traits with the greatest variation were DM (21.24 %), FC (23.43 %), CL (37.57 %) and SF (33.71 %) (Table 2). Likewise, there were significant differences (P ≤ 0.05) between the means of collections, with the exception of DF and FW (Table 4). Rick and Chetelat (1995) indicate that in S. pimpinellifollium L. the type of inflorescence, stem diameter, and days to flowering and maturity are very similar to those of the cultivated tomato, which makes this wild species the most used in tomato hybridization. In addition, both species are self-compatible and have red fruit, with the shape and size of the fruits being a relevant factor in genetic improvement (Rick & Forbes, 1975).

Table 4. Comparison of means among accessions of S. pimpinellifollium L. for 12 traits.

Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA1584 16 az 81 a 22 a 13.1 ab 1.1 a 15 a-c 21.4 a 1.3 a 1.2 ab 1.2 b 6.5 c 37.3 a
LA1689 15 a 82 a 21 ab 13.2 ab 1.1 a 15 a-c 21.6 a 1.2 a 1.2 ab 1.2 b 6.6 bc 37. 7 a
LA1237 15 a 54 cd 20 ab 14.5 a 1.2 a 12 c 7.6 c 1.8 a 1.2 ab 1.6 a 6.5 c 17.7 de
LA1593 13 a 57 bc 16.7 b 9 b 0.85 b 13 bc 10.7 bc 1.1 a 1.1 b 1.2 b 6.7 bc 15.7 e
LA1576 15 a 60 b 16.6 b 9.7 ab 0.84 b 20 ab 13.3 b 1.2 a 1.2 ab 1.3 b 9.3 a 21.7 cd
LA0373 12 a 50 d 18.6 ab 11.2 ab 0.94 b 22 a 13.4 b 1.7 a 1.3 a 1.5 ab 7.7 b 24.7 bc
LA0442 11 a 54 cd 20 ab 11.8 ab 0.84 b 14 a-c 11.5 bc 1.5 a 1.3 ab 1.4 ab 6.5 c 30.3 b
LSD 5.4 4.7 4.2 4.8 0.18 6.6 5.7 0.83 0.25 0.28 1.2 6
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).

Galiana-Balaguer, Roselló, and Nuez (2006) concluded that the TSS content in S. pimpinellifollium L. is high. In general, all the accessions of S. pimpinellifollium L. evaluated in the present work had higher TSS than those commonly presented by 'Saladette'-type hybrids, which oscillate between 3.9 and 5.2 °Brix (Bonilla-Barrientos et al., 2014; Hernández-Leal et al., 2013). Rodríguez, Pratta, Zorzoli, and Picardi (2006), when studying a population of recombinant lines derived from the cross between S. lycopersicum cv. Caimanta and the accession LA722 of S. pimpinellifollium L., found an increase of 1.6 °Brix and 19 days of shelf life with respect to the female parent. Therefore, accession LA1576, which presented 9.3 °Brix (Table 4), can be an alternative to improve the internal quality of tomato fruits.

With respect to Solanum peruvianum L., the traits with the greatest variability were DF, FC, CL, FW, SF and FL, with 36.34, 35.31, 30.58, 57.82, 21.11 and 26.27 %, respectively (Table 2). This variability is due to its reproduction system (allogamy). Given that cross-pollination is required in individuals, due to their self-incompatibility, they have greater variation compared to those with autogamy (Rick, 1988). The above can be observed in Table 2, where, with the exception of DM, the traits have coefficients of variation greater than 13 %. Accession LA1982 had later flowering (40 days) and ripening (89 days), greater leaf length (30.2 cm) and width (16 cm), and greater stem diameter (0.99 mm) and cluster length (41 cm) (Table 5); this suggests that LA1982 can be exploited in breeding programs of cultivated tomatoes.

Table 5. Comparison of means among accessions of S. peruvianum L. for 12 traits.

Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2172 16 cdz 82 b-d 19.4 ef 11.9 a 0.89 a-c 9 e 11.2 d 1.2 e 1.2 f 1.3 d 8.8 a 83 c
LA1982 40 a 89 a 30.2 a 16 a 0.99 a 20 bc 41 a 2.5 c-e 1.2 d-f 1.5 cd 7.1 bc 129 b
LA1677 18 b-d 80 cd 18.3 f 11.6 a 0.79 c-f 31 a 22.1 c 8.2 a 2.2 a 1.9 ab 6.7 b-d 126 b
LA1973 18 b-d 85 ab 21.3 c-e 13.4 a 0.82 b-e 18 b-e 27 c 1.8 de 1.2 f 1.4 cd 6 de 77 c
LA1360 21 b 80 cd 23.2 bc 13.4 a 0.92 ab 25 ab 39 ab 2.5 c-e 1.1 f 1.4 cd 7.5 b 137 ab
LA0103 21 bc 81 b-d 23 b-d 14.2 a 0.79 c-f 17 b-e 26.8 c 2.7 c-e 1.5 b-e 1.4 cd 6.3 c-e 79 c
LA0446 14 d 83 bc 20.7 e 12.1 a 0.71 e-g 15 c-e 24.5 c 3.9 c 1.5 b-d 1.7 bc 7.1 bc 90 c
LA1346 15 d 88 a 20 ef 10.6 a 0.72 e-g 20 bc 28 c 3.1 cd 1.5 bc 1.9 ab 6.5 c-e 152 ab
LA1336 15 d 78 d 21.2 c-e 11. 6 a 0.62 g 19 b-d 25.8 c 6.1 b 1.7 b 2.1 a 5.9 de 155 ab
LA1274 17 b-d 82 b-d 21 de 11.4 a 0.67 fg 17 b-e 31 bc 3.6 c 1.4 c-f 1.7 bc 6.1 de 166 a
LA1365 17 b-d 85 ab 25 b 13.6 a 0.86 b-d 30 a 42 a 3.9 c 1.5 b-e 1.9 ab 5.8 de 148 ab
LA2152 17 b-d 85 ab 17.9 f 10.2 a 0.76 d-f 10 de 24.5 c 1.7 de 1.2 ef 1.4 d 5.6 e 138 ab
LSD 6 4.7 2.2 6 0.13 9.2 9.2 1.6 0.26 0.29 0.9 30
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit. LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).

Chetelat et al. (2009) reported that the number of seeds of the evaluated accessions of S. peruvianum L. varies between 22.5 and 50 seeds per fruit. These values are much lower compared to those obtained in this research, which varied between 77 and 166.

Most of the traits evaluated in the Solanum habrochaites S. accessions had coefficients of variation greater than 14 % (Table 2). The traits with the greatest variation were FW (59.9 %), SF (35.37 %) and LW (20.48 %). Among accessions there were significant statistical differences (P ≤ 0.05) in the traits evaluated, except for FC (Table 6). Five accessions of this species were characterized as self-compatible, while the other three were self-incompatible, so their propagation is through cross-pollination; this generates greater diversity (Peralta & Spooner, 2001). Only LA1223 produced fruit without the need to manually pollinate.

Table 6. Comparison of means among accessions of S. habrochaites S. for 12 traits.

Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2409 44 bcz 85 e 30.4 cd 16.7 de 1.01 ab 26 a 20.9 a 1.9 d 1.4 b-d 1.5 de 8.6 b 74 cd
LA1731 46 a 93 cd 39 b 22.9 a-c 0.95 ab 25 a 14 cd 2.5 cd 1.5 b 1.8 b 8.5 b 56 de
LA2650 47 b 92 cd 47.1 a 25.9 ab 0.98 ab 18 a 13.4 d 2.8 bc 1.4 bc 1.8 b 6.5 c 57 de
LA2158 28 d 90 d 30.1 cd 18.3 c-e 0.87 b 20 a 14.8 b-d 7.1 a 1.9 a 2.3 a 9.1 b 129 a
LA2167 53 a 95 bc 36.1 bc 21.1 b-d 0.94 b 20 a 18.7 a-c 1.7 d 1.3 d 1.4 e 6.6 c 109 b
LA1223 43 bc 90 d 28.7 d 14.7 e 0.98 ab 25 a 15.6 b-d 2 d 1.3 cd 1.5 de 10.3 a 88 c
LA1777 44 bc 100 a 39 b 19.8 c-e 0.91 b 18 a 17.9 a-d 3.3 b 1.4 bc 1.7 bc 7 c 112 ab
GH0810 39 c 98 ab 41.5 ab 26.7 a 1.1 a 22 a 19.7 ab 2.2 cd 1.2 d 1.7 cd 6.7 c 49 e
LSD 4.7 3.8 6.5 5.6 0.15 10.7 5.1 0.8 0.17 0.14 1.1 18
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit width (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).

Carter, Gianiagna, and Sacalis (1989), in a study of tolerance to the Colorado beetle (Leptinotarsa decemlineata Say), concluded that the leaves of S. habrochaites S. contain zingiberene, a compound that promotes partial tolerance to this insect. In the accessions evaluated in the present work, LA2650 had the largest leaves, so this represents an alternative for tolerance to this pest, by associating leaf size with greater zingiberene production.

On the other hand, accession LA1777 has been widely used in tomato breeding, since it has alleles that increase fruit yield and TSS, detected on chromosomes 1 and 4, respectively (Bernacchi et al., 1998; Monforte & Tansksley, 2000). However, it was found that LA2158 had greater weight and size, statistically different from those of LA1777, so it could be a better alternative for the breeding program’s objectives.

Among the seven accessions of Solanum chilense R., it was found that the traits with the greatest variation were DF, LL, SD, FC, FWi and SF (Table 2) with significant differences (P ≤ 0.05) in most of them, except in FW (Table 7). LA1960 presented the highest value in number of seeds per fruit (82), while LA2748 and LA2759 were superior in the number of flowers per cluster (28). These traits are important for determining gene flow, and therefore the genetic and evolutionary structure of the species (Barrett, 2008).

Table 7. Comparison of means among accessions of S. chilense R. for 12 traits.

Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2930 40 az 86 a 23.3 bc 14.8 ab 0.88 bc 15 d 34. 4 a 1.6 a 1.1 cd 0.9 b 5.5 c 21 c
LA1960 40 a 88 a 23.6 bc 10.6 b 1 b 19 b-d 24 b 1.7 a 1.4 a 1.6 a 8.5 a 82 a
LA2759 29 b 87 a 15.1 a-c 12.7 ab 0.96 bc 28 a 30 ab 1.7 a 1.3 ab 1.5 a 7.2 ab 56 b
LA1958 37 a 87 a 27.7 a-c 18.2 a 0.9 bc 25 ab 29 ab 1.2 a 0.97 d 1 b 6.7 bc 50 b
LA2778 30 b 85 a 30.2 a 16.6 ab 0.78 c 24 a-c 24.7 b 1.6 a 1.2 bc 1.5 a 6.1 bc 53 b
LA2748 24 b 88 a 28.3 ab 17.8 ab 0.91 bc 28 a 28.3 ab 1.8 a 1.1 cd 1.5 a 5.7 c 46 b
LA2750 15 c 79 b 21.7 c 15.8 ab 1.6 a 16 cd 26 b 1.5 a 1.3 bc 1.6 a 5.6 c 54 b
LSD 5.6 3.4 6.5 7.2 0.2 8 7.1 0.8 0.17 0.15 1.3 14
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).

The species S. chilense R. is self-incompatible (Breto, Asins & Carbonell, 1993), which promotes cross-pollination, giving rise to a wide genetic diversity among and within the accessions. This explains the fact that the evaluated accessions of this species have eight traits with a coefficient of variation greater than 20 %; on the contrary, a self-compatible species such as S. pimpinellifollium L. presented only four traits with coefficients of variation greater than 20 %. Rick (1988) indicates that S. chilense R. has a wide diversity, since cross-pollination is required; therefore, these accessions represent an ample source of genes, not only for the traits evaluated, but also in the resistance to viral diseases (Griffiths & Scott, 2001, Stamova & Chetelat, 2000).

On the other hand, Chetelat et al. (2009) found 20 to 50 seeds in collections of S. chilense, S. peruvianum and S. pennellii made in the Atacama Desert in northern Chile, a range in which the values of the species evaluated here are found.

Conclusions

The accessions within species showed wide variation, which makes them a promising germplasm to be used in the development of breeding programs. In this sense, the accessions with better characteristics were LA1272, LA1367, LA1576 and LA177, which can be a source of new allelic versions to improve the fruit, while LA1982 and LA2650 can help improve the archetype of the cultivated tomato.

The traits with the greatest variation among species were DF, FW and SF, while those with the smallest variation were DM, FL, FWi and TSS. The species with the greatest differentiation were S. peruvianum L., S. chilense R. and S. habrochaites S., because they had higher coefficients of variation compared to the rest of the species evaluated.

Solanum pennellii L. presented the lowest coefficients of variation among accessions in most of the variables evaluated.

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Figures:

Figure 1. S. pimpinellifollium L. accession LA0373
Figure 2. S. peruvianum L. accession LA1365
Figure 3. S. habrochaites S. accession LA2409
Figure 4. S. pennellii L. accession LA1272
Figure 5. S. chilense R. accession LA2759

Tables:

Table 1. Species, accessions and origin of collections evaluated in this research.
Species Accession Origin of the collection Accession Origin of the collection
S. pennellii L. LA2580 Valle de Casma, Ancash, Perú LA1272 Pisaquera, Lima, Perú
LA0716 Atico, Arequipa, Perú LA1277 Trapiche, Lima, Perú
LA1367 Santa Eulalia, Lima, Perú
S. pimpinellifollium L. LA1584 Jayanca de La Vina, Lambayeque, Perú LA0373 Culebras núm. 1, Ancash, Perú
LA1689 Castilla núm. 1, Piura, Perú LA0442 Sechin, Ancash, Perú
LA1237 Atacames, Esmeraldas, Ecuador LA1576 Manchay, Alta Lima, Perú
LA1593 Puente Chao, La Libertad, Perú
S. peruvianum L. LA2172 Cuyca, Cajamarca, Perú LA0446 Atiquipa, Arequipa, Perú
LA1982 Huallanca, Ancash, Perú LA1346 Casmiche, La Libertad, Perú
LA1677 Fundo Huadquina, Topara, Ica, Perú LA1336 Atico, Arequipa, Perú
LA1973 Yura, Arequipa, Perú LA1274 Pacaibamba, Lima, Perú
LA1360 Pariacoto, Ancash, Perú LA1365 Caranquilloc, Ancash, Peru
LA0103 Cajamarquilla, Lima, Perú LA2152 San Juan #1, Cajamarca, Perú
S. habrochaites S. LA2409 Miraflores, Lima, Perú LA1223 Alausi, Chimborazo, Ecuador
LA1731 Río San Juan, Huancavelica, Perú LA1777 Rio Casma, Ancash, Perú
LA2650 Ayabaca, Piura, Perú GH0810
LA2158 Río Chotano, Cajamarca, Perú LA2167 Cimentario, Cajamarca, Perú
S. chilense R. LA2930 Guatacondo, Tarapaca, Chile LA1958 Pampa de la Clemesi, Moquegua, Perú
LA2750 La Despreciada, Antofagasta, Chile LA2778 Chapiquina, Tarapaca, Chile
LA1960 Río Osmore, Moquegua, Perú LA2748 Soledad, Tarapaca, Chile
LA2759 Tarapaca, Chile
Table 2. Descriptive statistics among and within species for the traits evaluated.
DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
General m 26.96 82.64 24.63 14.54 0.95 18.97 23.09 2.53 1.34 1.57 7.14 94.52
sd 12.29 12.15 7.48 4.19 0.19 5.62 8.22 1.57 0.22 0.28 1.27 59.14
r 42.00 50.00 32.00 17.70 0.98 22.00 34.40 7.10 1.23 1.40 4.80 217.30
cv 45.57 14.70 30.36 28.81 19.88 29.60 35.60 62.30 16.78 18.07 17.82 62.57
S. pennellii L. m 33.30 88.80 20.54 12.42 1.19 13.78 25.32 2.58 1.34 1.76 8.10 197.00
sd 1.37 8.53 2.44 0.81 0.07 2.18 1.17 0.72 0.05 0.05 1.49 31.95
r 3.60 20.00 5.60 1.90 0.17 5.80 3.00 2.00 0.10 0.10 3.20 84.00
cv 4.13 9.60 11.87 6.50 5.48 15.84 4.63 27.87 4.09 3.11 18.42 16.22
S. pimpinellifollium L. m 13.86 62.57 19.27 11.79 0.98 15.86 14.21 1.40 1.21 1.34 7.11 26.44
sd 1.86 13.29 2.07 1.98 0.15 3.72 5.34 0.27 0.07 0.16 1.06 8.91
r 5.00 32.00 5.40 5.50 0.36 10.00 14.00 0.70 0.20 0.40 2.80 22.00
cv 13.45 21.24 10.74 16.81 15.28 23.43 37.57 19.34 5.68 12.05 14.84 33.71
S. habrochaites S. m 43.00 92.88 36.49 20.76 0.97 21.75 16.88 2.94 1.43 1.71 7.91 84.25
sd 7.25 4.79 6.43 4.25 0.07 3.24 2.80 1.76 0.21 0.28 1.41 29.97
r 25.00 15.00 18.40 12.00 0.23 8.00 7.50 5.40 0.70 0.90 3.80 80.00
cv 16.86 5.16 17.61 20.48 7.16 14.90 16.59 59.94 14.89 16.35 17.84 35.57
S. peruvianum L. m 19.08 83.17 21.77 12.50 0.80 19.25 28.57 3.43 1.43 1.63 6.62 123.33
sd 6.93 3.33 3.35 1.66 0.11 6.80 8.74 1.99 0.30 0.27 0.90 32.40
r 26.00 11.00 12.30 5.80 0.37 22.00 30.80 7.00 1.10 0.80 3.20 89.00
cv 36.34 4.00 15.37 13.26 13.56 35.31 30.58 57.82 21.11 16.37 13.67 26.27
S. chilense R. m 30.71 85.71 24.27 15.21 1.00 22.14 28.06 1.59 1.20 1.37 6.47 51.71
sd 9.20 3.15 5.09 2.76 0.27 5.46 3.58 0.20 0.15 0.29 1.09 17.90
r 25.00 9.00 15.10 7.60 0.82 13.00 10.40 0.60 0.43 0.70 3.00 61.00
cv 29.94 3.67 20.96 18.13 27.04 24.66 12.76 12.31 12.44 21.35 16.85 34.60
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; m = arithmetic mean; sd = standard deviation; r = range; cv = coefficient of variation.
Table 3. Comparison of means among accessions of Solanum pennellii L. for 12 traits.
Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2580 33.3 az 86 b 22.6ab 13.1 a 1.22 ab 16.9 a 26 a 1.7 c 1.4 a 1.7 a 6.6 b 192 ab
LA0716 34.6 a 97 a 18 b 11.2 a 1.25 a 14.8 a 25.4 a 3.7 a 1.4 a 1.8 a 6.5 b 149 b
LA1272 31 a 98 a 18.8 ab 13 a 1.21 ab 13.2 a 24.6 a 2.6 b 1.3 a 1.8 a 9.3 a 233 a
LA1367 34 a 78 c 19.7 ab 12 a 1.19 ab 12.9 a 23.8 a 2.5 bc 1.3 a 1.8 a 9.7 a 218 a
LA1277 33.6 a 85 b 23.6 a 12.8 a 1.08 b 11.1 a 26.8 a 2.4 bc 1.3 a 1.7 a 8.4 a 193 ab
DMSH 6.3 6.4 4.9 4.3 0.15 6.2 8 0.7 0.35 0.23 1.4 44
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).
Table 4. Comparison of means among accessions of S. pimpinellifollium L. for 12 traits.
Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA1584 16 az 81 a 22 a 13.1 ab 1.1 a 15 a-c 21.4 a 1.3 a 1.2 ab 1.2 b 6.5 c 37.3 a
LA1689 15 a 82 a 21 ab 13.2 ab 1.1 a 15 a-c 21.6 a 1.2 a 1.2 ab 1.2 b 6.6 bc 37. 7 a
LA1237 15 a 54 cd 20 ab 14.5 a 1.2 a 12 c 7.6 c 1.8 a 1.2 ab 1.6 a 6.5 c 17.7 de
LA1593 13 a 57 bc 16.7 b 9 b 0.85 b 13 bc 10.7 bc 1.1 a 1.1 b 1.2 b 6.7 bc 15.7 e
LA1576 15 a 60 b 16.6 b 9.7 ab 0.84 b 20 ab 13.3 b 1.2 a 1.2 ab 1.3 b 9.3 a 21.7 cd
LA0373 12 a 50 d 18.6 ab 11.2 ab 0.94 b 22 a 13.4 b 1.7 a 1.3 a 1.5 ab 7.7 b 24.7 bc
LA0442 11 a 54 cd 20 ab 11.8 ab 0.84 b 14 a-c 11.5 bc 1.5 a 1.3 ab 1.4 ab 6.5 c 30.3 b
LSD 5.4 4.7 4.2 4.8 0.18 6.6 5.7 0.83 0.25 0.28 1.2 6
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).
Table 5. Comparison of means among accessions of S. peruvianum L. for 12 traits.
Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2172 16 cdz 82 b-d 19.4 ef 11.9 a 0.89 a-c 9 e 11.2 d 1.2 e 1.2 f 1.3 d 8.8 a 83 c
LA1982 40 a 89 a 30.2 a 16 a 0.99 a 20 bc 41 a 2.5 c-e 1.2 d-f 1.5 cd 7.1 bc 129 b
LA1677 18 b-d 80 cd 18.3 f 11.6 a 0.79 c-f 31 a 22.1 c 8.2 a 2.2 a 1.9 ab 6.7 b-d 126 b
LA1973 18 b-d 85 ab 21.3 c-e 13.4 a 0.82 b-e 18 b-e 27 c 1.8 de 1.2 f 1.4 cd 6 de 77 c
LA1360 21 b 80 cd 23.2 bc 13.4 a 0.92 ab 25 ab 39 ab 2.5 c-e 1.1 f 1.4 cd 7.5 b 137 ab
LA0103 21 bc 81 b-d 23 b-d 14.2 a 0.79 c-f 17 b-e 26.8 c 2.7 c-e 1.5 b-e 1.4 cd 6.3 c-e 79 c
LA0446 14 d 83 bc 20.7 e 12.1 a 0.71 e-g 15 c-e 24.5 c 3.9 c 1.5 b-d 1.7 bc 7.1 bc 90 c
LA1346 15 d 88 a 20 ef 10.6 a 0.72 e-g 20 bc 28 c 3.1 cd 1.5 bc 1.9 ab 6.5 c-e 152 ab
LA1336 15 d 78 d 21.2 c-e 11. 6 a 0.62 g 19 b-d 25.8 c 6.1 b 1.7 b 2.1 a 5.9 de 155 ab
LA1274 17 b-d 82 b-d 21 de 11.4 a 0.67 fg 17 b-e 31 bc 3.6 c 1.4 c-f 1.7 bc 6.1 de 166 a
LA1365 17 b-d 85 ab 25 b 13.6 a 0.86 b-d 30 a 42 a 3.9 c 1.5 b-e 1.9 ab 5.8 de 148 ab
LA2152 17 b-d 85 ab 17.9 f 10.2 a 0.76 d-f 10 de 24.5 c 1.7 de 1.2 ef 1.4 d 5.6 e 138 ab
LSD 6 4.7 2.2 6 0.13 9.2 9.2 1.6 0.26 0.29 0.9 30
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit. LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).
Table 6. Comparison of means among accessions of S. habrochaites S. for 12 traits.
Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2409 44 bcz 85 e 30.4 cd 16.7 de 1.01 ab 26 a 20.9 a 1.9 d 1.4 b-d 1.5 de 8.6 b 74 cd
LA1731 46 a 93 cd 39 b 22.9 a-c 0.95 ab 25 a 14 cd 2.5 cd 1.5 b 1.8 b 8.5 b 56 de
LA2650 47 b 92 cd 47.1 a 25.9 ab 0.98 ab 18 a 13.4 d 2.8 bc 1.4 bc 1.8 b 6.5 c 57 de
LA2158 28 d 90 d 30.1 cd 18.3 c-e 0.87 b 20 a 14.8 b-d 7.1 a 1.9 a 2.3 a 9.1 b 129 a
LA2167 53 a 95 bc 36.1 bc 21.1 b-d 0.94 b 20 a 18.7 a-c 1.7 d 1.3 d 1.4 e 6.6 c 109 b
LA1223 43 bc 90 d 28.7 d 14.7 e 0.98 ab 25 a 15.6 b-d 2 d 1.3 cd 1.5 de 10.3 a 88 c
LA1777 44 bc 100 a 39 b 19.8 c-e 0.91 b 18 a 17.9 a-d 3.3 b 1.4 bc 1.7 bc 7 c 112 ab
GH0810 39 c 98 ab 41.5 ab 26.7 a 1.1 a 22 a 19.7 ab 2.2 cd 1.2 d 1.7 cd 6.7 c 49 e
LSD 4.7 3.8 6.5 5.6 0.15 10.7 5.1 0.8 0.17 0.14 1.1 18
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit width (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).
Table 7. Comparison of means among accessions of S. chilense R. for 12 traits.
Accession DF 1 DM LL LW SD FC CL FW FL FWi TSS SF
LA2930 40 az 86 a 23.3 bc 14.8 ab 0.88 bc 15 d 34. 4 a 1.6 a 1.1 cd 0.9 b 5.5 c 21 c
LA1960 40 a 88 a 23.6 bc 10.6 b 1 b 19 b-d 24 b 1.7 a 1.4 a 1.6 a 8.5 a 82 a
LA2759 29 b 87 a 15.1 a-c 12.7 ab 0.96 bc 28 a 30 ab 1.7 a 1.3 ab 1.5 a 7.2 ab 56 b
LA1958 37 a 87 a 27.7 a-c 18.2 a 0.9 bc 25 ab 29 ab 1.2 a 0.97 d 1 b 6.7 bc 50 b
LA2778 30 b 85 a 30.2 a 16.6 ab 0.78 c 24 a-c 24.7 b 1.6 a 1.2 bc 1.5 a 6.1 bc 53 b
LA2748 24 b 88 a 28.3 ab 17.8 ab 0.91 bc 28 a 28.3 ab 1.8 a 1.1 cd 1.5 a 5.7 c 46 b
LA2750 15 c 79 b 21.7 c 15.8 ab 1.6 a 16 cd 26 b 1.5 a 1.3 bc 1.6 a 5.6 c 54 b
LSD 5.6 3.4 6.5 7.2 0.2 8 7.1 0.8 0.17 0.15 1.3 14
1DF = days to flowering; DM = days to maturity; LL = leaf length (cm); LW = leaf width (cm); SD = stem diameter (cm); FC = number of flowers per cluster; CL = cluster length (cm); FW = fruit weight (g); FL = fruit length (cm); FWi = fruit width (cm); TSS = total soluble solids (°Brix); SF = number of seeds per fruit; LSD = least significant difference. zMeans with the same letters within each column do not differ statistically (P ≤ 0.05).