Morphological characterization of manzano hot pepper (Capsicum pubescens Ruiz & Pav.) landraces in the central region of Veracruz state, México

O. R. Leyva-Ovalle1; P. Andrés-Meza1*; D. Del Valle-Hernández2; I. Meneses-Márquez3; J. Murguía-González1; M. E. Galindo-Tovar1; H. López-Sánchez4; R. Serna-Lagunes1; L. Del Rosario-Arellano5; H. E. Lee- Espinoza1; M. Sierra-Macías3; A. Espinosa-Calderón6

1. Facultad de Ciencias Biológicas y Agropecuarias. Universidad Veracruzana. Región Orizaba-Córdoba. Camino antiguo a Amatlán-Peñuela. Amatlán de los Reyes, Córdoba, Veracruz, México. CP. 94950. Tel: 271 7166410., Universidad Veracruzana, Facultad de Ciencias Biológicas y Agropecuarias, Universidad Veracruzana,

<city>Córdoba</city>
<state>Veracruz</state>
<postal-code>94950</postal-code>
, Mexico , 2. Maestría en Horticultura Tropical. Facultad de Ciencias Biológicas y Agropecuarias. Universidad Veracruzana., Universidad Veracruzana, Maestría en Horticultura Tropical, Facultad de Ciencias Biológicas y Agropecuarias, Universidad Veracruzana, Mexico , 3. Campo Experimental Cotaxtla. Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP). Km 34.5, carretera Veracruz-Córdoba. C.P. 91700. Medellín de Bravo, Veracruz, México., Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo Experimental Cotaxtla, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias,
<postal-code>91700</postal-code>
<state>Veracruz</state>
, Mexico ,
4. Colegio de Postgraduados. Campus Puebla. Km. 125.5, carretera federal México-Puebla (actualmente Boulevard Forjadores de Puebla), C.P. 72760, Puebla, Puebla, México. Tel: 222 285 07 38., Colegio de Postgraduados, Campus Puebla,
<postal-code>72760</postal-code>
<city>Puebla</city>
<state>Puebla</state>
, México ,
5. Doctorado en Ciencias Agropecuarias. Facultad de Ciencias Biológicas y Agropecuarias. Universidad Veracruzana. Región Orizaba-Córdoba. Camino antiguo Amatlán-Peñuela. Amatlán de los Reyes, Córdoba, Veracruz, México. CP. 94950. Tel: 271 7166410., Universidad Veracruzana, Doctorado en Ciencias Agropecuarias, Facultad de Ciencias Biológicas y Agropecuarias, Universidad Veracruzana,
<city>Córdoba</city>
<state>Veracruz</state>
<postal-code>94950</postal-code>
, Mexico ,
6. Campo Experimental Valle de México. Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias (INIFAP). 56250. Coatlinchán, Texcoco, Estado de México., Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Campo Experimental Valle de México, Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias,
<postal-code>56250</postal-code>
<city>Texcoco</city>
<state>Estado de México</state>
, Mexico

Correspondence: *. Corresponding Author: Andrés-Meza, P.: Facultad de Ciencias Biológicas y Agropecuarias. Universidad Veracruzana. Región Orizaba-Córdoba. Camino antiguo a Amatlán-Peñuela. Amatlán de los Reyes, Córdoba, Veracruz, México.CP. 94950.. Cel: 271 173 5239. E- mail: E-mail:


Abstract

The heterogeneity, diversity of climates, and selection of cultivars by farmers for the local adaptation have favored the maintenance of numerous species of chili peppers (Capsicum spp.) in Mexico. This high genetic diversity offers an excellent alternative for the search of native and semi-domesticated genotypes as a genetic reservoir for genetic improvement and a primary source of phytochemical compounds as a multifunctional food. The objective of the present work was the in situ characterization in 10 fruits from 22 landraces of C. pubescens Ruiz & Pav., in the central region of the state of Veracruz, Mexico, through eight morphological descriptors. The morphological characterization showed that the variables: weight of fruit, length of pedicel, length of placenta, and number of seeds per fruit presented high variation. The Principal Component Analysis of the eight morphological traits showed that the first three components were selected as the principal factors. These components accounted for 77.9 % of the total variation. The cluster analysis gave off six groups or population patterns that differed mainly by the climatic conditions where they thrive. The C. pubescens landraces analyzed in the present study, native to central Veracruz, showed high morphological variability, both inter and intra population. This shows the potential of the native C. pubescens germplasm, making it also a starting point to establish strategies for a genetic improvement program.

Received: 2017 October 27; Accepted: 2018 June 29

revbio. 2020 May 31; 5: e388
doi: 10.15741/rebbio.05.e388

Keywords: Keywords: Capsicum pubescens Ruiz & Pav., Manzano chili pepper landraces, morphological characterization.

Introduction

The genus Capsicum is originally from Mesoamerica and Mexico is the main center of genetic diversity. Recently, studies have been carried out with the objective of identifying the morphotypes of C. pubescens in the south of the State of Mexico (Ayala et al., 2017); however, in Veracruz, and especially in the central zone of the state where different morphotypes of C. pubescens are grown, they have not been studied. These morphotypes represent a valuable reservoir of germplasm that could be useful to improve this species. Moreover, documenting its morphological variability could be a viable alternative for economic development, which would allow to promote it as a crop with adequate genetic management in order to obtain quality fruits (Sánchez et al., 2010).

In recent years, the market foe Capsicum has grown exponentially; in 2014, the worldwide production was over 4.3 million ha-1 (FAOSTAT, 2017). Every year, over 140 000 ha of fresh chili peeper are grown in Mexico. The area used in Veracruz is approximately 4008 ha-1, with a mean yield of 6.4 t ha-1 (SIAP, 2016). C. pubescens production is reported to be done in backyard gardens and areas no greater than five ha, in association with fruit trees. It is mainly a rainfed crop and its yields are relatively low, 5 to 7 t/ha/year (Barrios-Puente et al., 2014).

The heterogeneity, diversity of climates, and selection of cultivars by farmers for the local adaptation have favored a high variability of morphotypes and subtypes of chili peppers (Pickersgill, 1997; Hernández et al., 1999: Lefebvre, 2004; Portis et al., 2006). In Mexico, “Manzano” or “Wax” chili pepper is the fifth most important species of the Capsicum genus grown due to the economic influx that it generates as well as its use in traditional dishes, which makes it highly demanded (Pérez & Castro, 2008).

Cultivation and production of Manzano chili pepper are limited to the high and cold areas of the country, at altitudes ranging from 1300 to 2400 m. The main producing areas in Mexico are Tacámbaro and Zitácuaro in Michoacán, Altotonga, Zongolica, and Huatusco in Veracruz, Teziutlán in Puebla, Coatepec Harinas and Tenancingo in the State of Mexico, and less so in Chiapas and Oaxaca (Espinoza & Villa, 2008). Given the importance of and demand for this species in Mexico and the United States of America, a collection of Manzano chili pepper fruits was done in two contrasting zones in the central region of the state of Veracruz. The morphological characterization in fruits will help to identify superior landraces and will give the breeders information to establish strategies for genetic improvement aimed at producing high-yield varieties and hybrids, commercial quality fruits, tolerance to pests and diseases, and showing high adaptability. Currently, there are few studies on the genetic diversity in Manzano chili pepper landraces in Mexico, especially in the central region of the state of Veracruz. Therefore, the present research work was focused on the morphological characterization of fruits from 22 landraces collected in two contrasting regions in the central region of the state of Veracruz.

Materials and Methods

Exploration communities. A total of 22 collections of Manzano chili pepper were done (Figure 2), most of them from plants over one year of age. The collection routes were through pathways, backyards, family gardens, and coffee and banana plantations. The collections were done in two totally contrasting regions, these being: region A, made up of the municipalities of Zongolica, Ixhuacán, Mixtla de Altamirano, and Tehuipango; and region B with the municipalities of Calcahualco, Alpatlahuac, Chocamán, Coscomatepec, and Huatusco, Veracruz. Region A has a Cf climate with constant rainfall, while region B has a Cw climate with a dry winter (Díaz et al., 2006). In both regions, the inter and intra population phenotypic variability was checked (Figure 2), as reported by Pérez et al. (2004). The Manzano chili pepper collection and ex situ morphological characterization were done from November 2016 to March 2017. The exploration communities were selected based on previous studies done by Pérez et al. (2004), as well as on information given by merchants and citizens of the explored municipalities. In each collection site, the coordinates were registered with a Garmin Etrex® GPS, and noted in a map to identify the sampling locations (Figure 1).


[Figure ID: f1] Figure 1.

Geographical location of the collection sites of 22 Manzano chili pepper landraces in the central region of Veracruz.



[Figure ID: f2] Figure 2.

Manzano chili pepper landraces showing great morphological variability in some fruits. Photograph: David Del Valle Hernández and José Luis Del Rosario Arellano.


Evaluated descriptors. Ten fruits were collected each from five plants in each location; all the sampled fruits were placed in paper bags and taken to the Genetic Resources Management and Conservation Unit of the Biological and Agricultural Science School of the Universidad Veracruzana, where they were characterized and the seeds were taken out. Eight morphological variables were evaluated in the sampled fruits, in accordance to the descriptors for Capsicum by the IPGRI (1995). These descriptors were: length of fruit (cm), width of fruit (cm), weight of fruit (g), length of pedicel (mm), thickness of fruit (mm), number of locules, length of placenta (mm), and number of seeds per fruit. These descriptors were chosen based on the recommendations of the International Plant Genetic Resources Institute (IPGRI, 1995) for mature Capsicum fruits.

Statistical methods. With the measurements of each variable per collection, a principal component analysis (PCA) was done, which included the 22 collections and the 8 variables, standardized to μ= 0 and σ2= 1. The estimation of the principal components was done with the correlation matrix so that the variables involved in the analysis were all of the same importance. From the correlation matrix, a cluster analysis was done through the Ward method; the measurement of unlikelihood was the Euclidean distance between groups. In both cases, the SAS/STAT® ver. 9.0 (SAS Institute Inc, 1990) software was used.

Results and Discussion

Even though in this study color was not evaluated as a morphological descriptor, it is worth mentioning that in both regions A and B, there were fruits ranging in hue from opaque red to intense red MV4, MV8, and MV13) (Figure 2) and from lemon-yellow to orange (MV5 and MV17) (Figure 2). The first landraces (red fruits) are at risk of genetic erosion since their spice levels are low (50 Scoville); they generally have one or two pear-shaped locules, which makes them undesirable for local cuisine and therefore are not grown that often (Ayala et al., 2017). Contrastingly, the yellow landraces are more commercialized locally and in the region. These landraces generally have three or four pear-shaped locules and are more appreciated by the consumers since they are spicier (hotter) (Pérez et al., 2004).

The analysis of variance detected highly significant differences (p ≤ 0.05) between collections in all variables (Table 1). The highest variation coefficients (25.8 to 28.5 %) were obtained in number of seeds per fruit, weight of fruit, and length of placenta. This indicates high variability in the behavior of the landraces found in situ.

Table 1.

Average, variation coefficient (VC) and mean square (MS) of each traits in 22 accesions of Manzano hot pepper in the central region of Veracruz. 2016.


Traits Average CV (%) CM
Fruit length [cm] 5.2 14.3 0.7**
Fruit width [cm] 3.6 13.7 0.5**
Fruit weight [g] 24.8 28.3 7.0**
Fruit pedicel length [mm] 30.4 13.7 4.2**
Fruit wall thickness [mm] 4.3 17.4 0.8**
Number of locules 2.4 21.1 0.5**
Placenta length [mm] 2.0 28.6 0.6**
Number of seeds per fruit 54.1 25.9 14.0**

TFN1* and ** significantly different at a level (p≤0.05) and (p≤0.01), respectively.


The principal component analysis (PCA) showed that the first three components make up 77.9 % of the total morphological variability (Table 2). PC1 made up 42.18 of the total variation, where the variables with the greatest contributions were weight of fruit, length of pedicel, thickness of fruit, and number of seeds per fruit. This component was made up of features that determine the size of the fruit and which allowed us to distinguish the landraces with small fruits (Table 2).

Table 2.

Vectors and eigenvalues of the analysis of principal components and descriptive value of the total variance in Manzano hot pepper (Capsicum pubescens Ruiz & Pav.) in the central region of Veracruz.


Traits PC1 PC2 PC3
Fruit length [cm] -0.569 -0.646 -0.022
Fruit width[cm] -0.330 0.312 -0.862
Fruit weight[g] -0.890 -0.215 0.109
Fruit pedicel length [mm] -0.711 0.272 0.091
Fruit wall thickness [mm] -0.627 -0.531 -0.055
Number of locules -0.475 0.732 -0.109
Placenta length [mm] 0.084 0.690 0.199
Number of seeds per fruit -0.659 0.461 0.360
Eigenvalues 3.374 2.099 0.763
Variance explained (%) 42.175 26.239 9.539
Cumulative variance (%) 42.175 68.414 77.953

TFN2†= Variables that contribute to the component; CP= principal component.


PC2 made up 26.24 % of the total variation. The variables regarding number of locules, length of placenta, and number of seeds per fruit contributed positively to this component, while length of fruit did so negatively. With this second component, we were able to distinguish the Manzano chili pepper landraces with greater internal dimension (Table 2). Moreover, the fruits with a higher number of locules proved to have a higher number of seeds. Nevertheless, this is not true for all landraces; some landraces with a single locule had twice as many seeds as those fruits with three or four locules. Pérez et al. (2004) mention that that yellow, apple-shaped fruits with three or four locules are generally preferred over red or orange, pear-shaped fruits which generally have one or two locules.

Meanwhile, PC3 contributed 9.54 % of the variation and is directly related with fruit width. This component helped to differentiate pear-shaped fruits with a lower number of locules. These results coincide with studies done by Chavez & Castillo (1999) who found that the first three components explained 91.86 % of the variation observed between Manzano chili pepper landraces; however, in our study, the higher variability is inter-population. Latournerie et al. (2002) found 79.5 % of the morphological phenotypical variability in collections of wild chili peppers and Habanero chili peppers in Yaxcaba, Yucatan, mainly originating in interspecific variation, resulting from a wide phenotypical diversity. Narez et al. (2014) found 73.27 % of the total variation in collections of wild chili peppers in the state of Tabasco, Mexico.

The cluster analysis grouped the 22 collections into six morphologically different groups (Figure 3). Group 1 corresponds to collections MVCO1, MVTE13, MVCA2, MVCO15, and MVHU16 and was characterized for having the highest length of pedicel, ranging from 28.7 to39.6 mm, as well as the highest number of locules, 2 to 3, respectively. The altitude range in the collection sites varied from1365 to 1762 m (Figure 1). Despite grouping landraces from geographically isolated zones, the fruits share similar intra-population morphological characteristics.


[Figure ID: f3] Figure 3.

Hierarchical classification of 22 Manzano hot pepper landraces collected in the central region of Veracruz, obtained using the Ward method.


Group II was made up of the landraces MVCA5, MVAL7, MVCA6, MVHU22, MVAL8, which were characterized for having the greatest number of seeds per fruit, with means ranging from 54.8 up to 86.1 seeds per fruit, respectively. In this sense, the MVAL8 landrace had the highest number of seeds per fruit at 86.1 while the MVTU20 landrace had the lowest number of seeds per fruit at 34.2, respectively.

Group III was integrated with landraces MVCA3 and MVHU18. Both of these landraces are characterized for having wider fruits which varied from 5.8 to 7.4 cm in MVCA3 and from 3.3 to 6.6 cm in MVHU18, respectively. Also, it is important to mention that these two landraces had the highest frequency in number of locules (>70 %), which is an important characteristic for consumers (Pérez et al., 2004).

Group IV was made up of only the MVCA4 landrace from the municipality of Calcahualco. It was characterized for having the highest means regarding length, width, weight, and thickness of the fruit wall with 6.33 cm, 4.29 cm, 48.54 g, and 5.93 mm, respectively. This landrace represents an important genetic reservoir susceptible of being used in a genetic improvement program to obtain varieties and hybrids with agricultural potential in the medium and long terms.

Group V was integrated by landraces MVAL9 and MVCH14, characterized for having the lowest values in weight and length of fruit, 16 g and 4.14 cm, respectively.

Lastly, group VI was integrated by collections MVZO10, MVHU17, MVMI12, MVTE11, MVTU21, MVHU19, and MVTU20, characterized for having low values regarding width and length of placenta in fruits, averaging 3.06 and 1.84 cm, respectively. The geographical range varied from 1373 to 2369 m, which changes the paradigm of many authors who state that this crop can only be found at altitudes >1500 m; in other papers, they state that altitude is a variable correlated with environmental conditions where specific phenotypes thrive, being low temperatures which decrease the characteristics of the fruit (Pérez et al., 2004; CONABIO, 2010; Meckelmann et al., 2015; Espinosa & Ramírez, 2016).

Conclusion

The C. pubescens landraces collected in the central part of the state of Veracruz show high interpopulation morphological variability in every location where they thrive and wide intra-population variation between contrasting locations. The MVCA4 landrace showed the best characteristics in fruit, and represents an important phenotypical reservoir susceptible to initiate a morphotypes improvement program with agricultural potential and high heritability. We suggest continuing with exploratory studies to identify more C. pubescens landraces with commercially important characteristics that can be included in hybridation and germplasm conservation programs.


fn1Cite this paper: Leyva-Ovalle, O. R., Andrés-Meza, P., Del Valle-Hernández, D., Meneses-Márquez, I., Murguía-González, J, Galindo-Tovar, M. E., López-Sánchez, H., Serna-Lagunes, R., Del Rosario-Arellano, L., Lee- Espinoza, H. E., Sierra-Macías, M., Espinosa-Calderón, A. (2018). Morphological characterization of manzano hot pepper (Capsicum pubescens Ruiz & Pav.) landraces in the central region of Veracruz state, México. Revista Bio Ciencias 5, e388. doi: https://doi.org/10.15741/revbio.05.e388

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