Agronomic performance of quinoa cultivars in two locations of ColombiaABSTRACT
Quinoa is a pseudocereal of great importance because of its nutritional content, great adaptability and broad genetic variability. In Colombia, genetic studies on quinoa are scarce, and there are no certified varieties. Rather, varietal mixtures that affect grain quality and, therefore, postharvest and marketing processes grow in the field. The objective of this study was to carry out a morphoagronomic characterization of five quinoa cultivars under field and greenhouse conditions in Tunja-Boyacá using 21 qualitative and quantitative descriptors, estimating their variation with descriptive, multivariate, and cluster analyses. The results show a high phenotypic segregation among cultivars, where yield, diameter, and panicle length were the quantitative variables with greater variation both in the field and in the greenhouse. The multivariate analysis showed that 72.54% of the variance observed in the greenhouse and 75.20% in the field were explained by the first two components, where plant height, stem diameter, panicle density and color, and stem color were discriminating variables. In the field and in the greenhouse, three groups of cultivars were formed according to the yield and desirable morphoagronomic characteristics. Finally, under the present environmental conditions, the more promising quinoa materials for a breeding program were Blanca Subachoque and Tunkahuan.
Keywords:
Chenopodium quinoa; andean culture; morphoagronomic descriptors; phenotypic variation; pseudocereal; yield
RESUMO
A quinoa é um pseudocereal muito importante devido ao seu conteúdo nutricional, grande adaptabilidade e ampla variabilidade genética. Na Colômbia, os estudos genéticos sobre a quinoa são escassos e não existem variedades certificadas. Em vez disso, crescem no campo misturas varietais que afetam a qualidade dos grãos e, portanto, os processos pós-colheita e de comercialização. O objetivo deste estudo foi caracterizar morfoagronômicamente cinco cultivares de quinoa em condições de campo e casa de vegetação em Tunja-Boyacá, utilizando 21 descritores qualitativos e quantitativos, estimando sua variação com análises descritivas, multivariadas e de agrupamento. Os resultados mostram alta segregação fenotípica entre as cultivares, sendo a produtividade, o diâmetro e o comprimento da panícula as variáveis quantitativas com maior variação tanto no campo quanto na casa de vegetação. A análise multivariada mostrou que 72,54% da variância observada em casa de vegetação e 75,20% no campo foram explicadas pelos dois primeiros componentes, onde altura da planta, diâmetro do caule, densidade e cor da panícula e cor do caule foram variáveis discriminantes. No campo e em casa de vegetação, foram formados três grupos de cultivares de acordo com a produtividade e características morfoagronômicas desejáveis. Finalmente, nas atuais condições ambientais, os materiais de quinoa mais promissores para um programa de melhoramento foram Blanca Subachoque e Tunkahuan.
Palavras-chave:
Chenopodium quinoa; cultura andina; descritores morfoagronômicos; variação fenotípica; pseudocereal; colheita
Quinoa (Chenopodium quinoa Willd.) is a crop of great agronomic importance and food security, which is distributed in the Andean region. The main producing and exporting countries are Peru, Bolivia and Ecuador. This species is native to South America, its center of origin is the Andes of Bolivia and Peru, and, currently, it covers different thermal floors from sea level to 4000 meters above sea level, from southern Chile and Argentina to Colombia (Manjarres et al., 2021a). It is rich in essential amino acids, vitamins, minerals and protein (Lan et al., 2023).
In Colombia, quinoa is cultivated on a small scale in the departments of Boyacá, Cundinamarca, Cauca and Nariño; however, its cultivation is expanding because of its nutritional characteristics and production potential (Infante et al., 2018). The planting material contains varietal mixtures without certified varieties. Information on morphoagronomic characteristics is scarce, and there are differences in maturation times, plant height, and yield, among other characteristics, generating problems in grain quality, post-harvest process and commercialization (Chura et al., 2019; EL-Harty et al., 2021; Morillo et al., 2020).
Consequently, morphoagronomic characterization studies have been carried out in Colombia but they are insufficient. In the Department of Boyacá, research was carried out by Infante et al. (2018), who morphologically and botanically characterized cultivated varieties of Chenopodium quinoa. Manjarres et al. (2021a), evaluated the yield and phenological aspects of 30 quinoa genetic materials in Boyacá and established a selection index according to production, finding broad variability resulting from existing varietal mixtures and highlighting the need to produce certified varieties with the identification of elite materials that improve breeding programs for this species.
In the Department of Cundinamarca, selection processes have been initiated for the generation of certified seeds based on characteristics such as good yield and resistance to biotic and abiotic conditions, among others. In order to select cultivars with better yields and the potential for profitable production for quinoa growers in the Boyacá Department, the objective of this research was to morphoagronomically characterize five quinoa cultivars from the Department of Cundinamarca under controlled field conditions in Tunja-Boyacá as a first approach towards obtaining certified seed.
MATERIAL AND METHODS
Plant material
Five cultivars of quinoa (C. quinoa) from the department of Cundinamarca (Colombia); Aurora (AUR), Blanca de Jericó (BJ), Blanca Subachoque (BS), Púrpura (PUR) y Tunkahuan (TK), have undergone a selection process and were also registered with ICA (Instituto Colombiano Agropecuario). These cultivars were selected for this study and are the most cultivated in Boyacá and Colombia. The plant material used in the study is cultivated and complied with all existing national standards for the study of cultivated germplasm.
Morphoagronomic characterization of quinoa cultivars under controlled conditions and open field
The morphoagronomic characterization was carried out in two locations: the first was carried out in the open field at the La María experimental farm of the Pedagogical and Technological University of Colombia, located at 2691 meters above sea level, average temperature of 16.2°C and relative humidity of 71.8% and the second under controlled conditions, in the Puente Restrepo greenhouse of the Pedagogical and Technological University of Colombia, located in Tunja, with an average temperature of 12°C and a relative humidity of approximately 72%.
The cultivars were established in a completely randomized three-block design, with five replicates per cultivar. The plants were grown with a distance of 0.4 m between plants and 0.8 m between rows, and for each of the conditions evaluated in the agronomic management was conventional. For the morphoagronomic characterization, descriptors defined by Rojas & Padulosi (2013) were evaluated, which have already been evaluated and validated in other studies of quinoa genetic diversity (Table 1). For the evaluation of the characters associated with color, the RGB color scale was used.
Statistical analysis
The statistical analyses were carried out in three steps: (1) descriptive, (2) multivariate, and (3) clustering. For the analysis of the data obtained in the morphoagronomic characterization, the descriptive analysis was carried out for the qualitative and quantitative variables with the statistical software InfoStat 2020 (Di Rienzo et al., 2020). Then the assumptions for the parametric analyzes were verified and the analysis of variance was performed. (ANOVA). To determine significant differences between cultivars, a Tukey comparison test was made with p<0.05. These analyzes were performed using R Core Team (2020) software. For the qualitative variables, the frequency analysis was carried out in the statistical software InfoStat 2020 (Di Rienzo et al., 2020).
For the quantitative variables, the Pearson correlation and its significance were estimated, using the t test (The null hypothesis is H0: r = 0, with 5% significance). For the multivariate analysis, a hierarchical grouping in principal components (HCPC) was performed, with the algorithms included in the factoextra package of the R program, which were plotted in a two-dimensional plane using the FactomineR package. Factorial analysis of mixed data considering quantitative and qualitative variables was done with the factoextra package in the R program. In addition, a dendrogram was generated using Euclidean distance and Ward's minimum variance hierarchical clustering method with the FactoMineR package (Lê et al., 2008).
Morphoagronomic descriptors used for the characterization of C. quinoa cultivars from Cundinamarca in Tunja-Boyacá. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
RESULTS AND DISCUSSION
The climatic conditions of the study region were typical of tropical zones that are characterized by few fluctuations in the photoperiod and average temperatures. The minimum temperature during the development of the research ranged between 7.43 and 10.00°C, the maximum between 17.80 and 20.00°C with an average temperature between 13.43 and 14.75°C. The monthly precipitation was less than 2 mm and average relative humidity of 78%. The daily illumination period that the cultivars received during the experiment was approximately 12 hours. The previous environmental conditions are presented mainly because the expression of morphological characters is influenced by the environment, and it is necessary to correlate these factors with the diversity found in the variables evaluated in quinoa cultivars
Morphoagronmic characterization of cultivars of C. quinoa under controlled conditions
The quantitative and qualitative descriptors evaluated in the quinoa cultivars under controlled conditions presented a wide range of variation, also showing that the descriptors with the highest coefficient of variation were yield (36.30%), panicle diameter (29.24%), length of the panicle (24.19%) and number of teeth (23.48%). While the variables seed diameter (5.06%) and seed weight (9.94%), were those that presented a lower coefficient of variation (Table 2).
The cultivars did not present significant differences in yield, which was in a range of 79.68 g of seed per plant for Blanca Subachoque and 84.47 g of seed per plant for Tunkahuan. The descriptors evaluated in the panicle, such as number and diameter, did not present significant differences, registering an average of 15 panicles/plant and 64.5 cm in diameter, respectively. However, the length of the panicle showed significant differences, where the cultivar Púrpura exhibited the longest length (57.20 cm), while Tunkahuan the shortest (43.07 cm) (Table 2).
Seed weight was similar in all cultivars, with a range between 2.38 and 2.73 g, while seed diameter was between 2.24 and 2.41 mm. Regarding the evaluation of stem diameter, an average of 15.50 mm was obtained, with Púrpura being the cultivar with the largest diameter (17.05 mm) and Tunkahuan the one with the smallest (13.63 mm). The height of the plant was between values of 155.33 cm (Tunkahuan) and 204 cm (Blanca Subachoque) (Table 2).
Pearson's correlation analysis (p≤0.05) in the system under controlled conditions showed that characteristics such as yield obtained a high and positive correlation with seed diameter (r = 0.96), number of panicles (r = 0.73), stem diameter (r = 0.71) and weight of 1000 seeds (r = 0.57), however, it had a negative correlation with plant height (r = -0.44) and panicle diameter (r = - 0.77) (Figure 1).
Pearson correlation analysis (p≤0.05) between the quantitative variables of five quinoa cultivars under controlled conditions. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
The frequency analysis of the qualitative variables of the cultivars under controlled conditions showed that 100% of the cultivars have a cylindrical grain shape and green axils. The color of the beige episperm predominated in the five cultivars with frequencies between 60 and 92%. The grain edge with undulations was the most frequent, being 100% of the individuals of the Tunkahuan cultivar those that presented the edge with undulations. Blanca de Jericó and Aurora presented a light beige color of the pericarp, while in Púrpura and Tunkahuan, a golden color was observed. Regarding the color of calcium oxalates, the plants exhibited various colors, the most frequent being pink oxalates followed by white ones (Table 3).
The analysis of mixed factors, in which qualitative and quantitative variables were taken into account, showed that 72.54% of the total phenotypic variation observed was explained by the first two components CP1 (44.15%) and CP2 (28.39%) (Figure 2a), where the first component grouped the cultivars according to the qualitative characteristics such as the branched growth habit up to the second third, pink color of the axils, white episperm color, intermediate panicle density and quantitative characteristics such as yield, weight and seed diameter, panicle length and height; while the second component was characterized by qualitative variables such as the color of the pink flowering panicle, the color of the white and pink oxalates, the color of the green and yellow stem, and quantitative variables such as the diameter and number of panicles.
On the other hand, in Figure 2b we see that the variables most correlated with yield are grain weight and diameter, which agrees with the characteristics observed in Tunkahuan, a cultivar that obtained the highest yield under controlled conditions, with seeds of greater diameter and heavier weight; likewise, we observed that the morphological components (panicle length, height, number of teeth and stem diameter) were associated and were very similar to each other in Blanca Subachoque, Púrpura and Aurora. On the other hand, the number and diameter of panicles were correlated and were characteristics that allowed Blanca de Jericó to be placed in another cluster, since it was the cultivar with the largest diameter and number of panicles.
a) Mixed factorial analysis taking into account the quantitative and qualitative variables evaluated in five quinoa cultivars under controlled conditions. b) Correlation analysis between the quantitative variables evaluated, where it is observed that the variables diameter and weight of the seeds are more correlated with yield. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
The cluster analysis shows the grouping of cultivars into three clusters (Figure 3). The descriptors that generated the groupings were panicle density, plant height, weight of 1000 seeds, color and stem diameter. Tunkahuan is found in the first cluster, a cultivar characterized by its intermediate grain edge, intermediate and compact panicle density, seed diameter of 2.41 mm, weight of 1000 seeds of 2.73 g, lower plant height (155 cm) and smaller stem diameter (13.63 cm). In the second group, Blanca de Jericó is located, which was characterized by exhibiting glomerulate panicles, beige perigonium color, white oxalate color, green streaks and by presenting seed diameter below average (2.29 mm) and the lowest weight of 1000 seeds (2.38 g). In the third group are the materials Aurora, Púrpura and Blanca Subachoque, which exhibited pink striaes, green stems, panicles with intermediate density, yields between 79 and 84 g/plant, approximately 20 teeth on the leaves, weight of 1000 seeds between 2.49 and 2.55 g, stem diameters between 15.6 cm and 17 cm and for being the materials with the highest plant height.
Morphoagronomic characterization of cultivars of C. quinoa in the open field
The morphoagronomic characterization of the five quinoa cultivars in the open field showed a high phenotypic variation. The descriptors yield (36.25%), panicle length (26.62%) and panicle diameter (25.52%), were the ones with the highest coefficient of variation, while grain diameter (5.44%) and weight of 1000 seeds ( 8.04%), were the ones that showed less variation (Table 4).
Cluster analysis. Three groups of cultivars formed according to the qualitative and quantitative variables evaluated under controlled conditions are shown. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
The yield of the cultivars in the open field presented significant differences between the cultivars, with Tunkahuan (339.48 g/plant) having the lowest yield, compared to Blanca Subachoque, which obtained the highest yield with 587.04 g/plant. The descriptors associated with the panicle did not present significant differences; the average diameter of the panicle was 96.85 cm; the panicle length presented values between 28.57 and 41.13 cm, with Tunkahuan being the cultivar with the shortest panicle length and Blanca Subachoque with the longest, respectively. The seeds had an average diameter of 2.33 mm, while the weight of 1000 seeds was between 3.2 and 3.3 g (Table 4).
The height of the plant presented an average of 131.23 cm, with a difference of 46.41 cm in height between the tallest and the shortest cultivars, Tunkahuan and Blanca Subachoque respectively (Table 4). The diameter of the stem presented significant differences, with values that oscillated between 15.93 cm and 21.98 cm. Tunkahuan was the material with the smallest diameter and Blanca de Jericó presented the largest diameter, respectively. Pearson's correlation analysis of the cultivars evaluated in the open field showed a high and positive correlation between yield and variables such as seed weight and diameter with r = 0.98 and r = 0.96 respectively, seed weight and grain diameter (r = 0.92), the length of the panicle and the height of the plant (r = 0.98), likewise the length of the panicle and the diameter of the panicle (r = 0.98) (Figure 4).
Pearson correlation analysis (p≤0.05) between the quantitative variables of the five quinoa cultivars under field conditions. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
The analysis of the qualitative variables showed that 100% of the cultivars presented cylindrical grain shape and green branch axils. The white color of the episperm predominated in the Blanca Subachoque and Púrpura cultivars, while Aurora presented a beige color in 87% of the individuals. The most frequent grain edge was smooth, which was observed more frequently in Aurora (93%), Blanca de Jericó (80%) and Blanca Subachoque (53%), while in Tunkahuan, the grain edge was 100% intermediate. In the cultivars Aurora and Blanca Subachoque, the color of the pericarp was beige, while in the cultivars Púrpura and Tunkahuan yellow was more frequent with 71 and 64% respectively. The cultivars exhibited calcium oxalates with colors from pink to white, pink being the most common among the individuals, except for Blanca Subachoque, which presented white oxalates in 86% (Table 5).
Two panicle forms were observed, the glomerulate one being the most common among the cultivars, except in Tunkahuan where 50% of the individuals had an intermediate form. Regarding panicle density, in Blanca Subachoque, Púrpura and Blanca de Jericó, the lax form followed by the intermediate form predominated, while for Aurora and Tunkahuan the intermediate form predominated; in the Tunkahuan cultivar a segregation of 71% of panicles was observed with intermediate density and 29% compact.
The factorial analysis of mixed data, in which all the quantitative and qualitative variables were taken into account, showed that the contribution of the variables to the first two components was 75.20%. The variables that contribute positively to CP1 (53.05%) were yield, weight of 1000 seeds, plant height and stem diameter (quantitative variables); as well as the density of the loose panicle, color of the panicle in pink flowering and color of the light beige and golden pericarp (qualitative variables). While for CP2 (22.15%), the variables that contributed the most were the diameter of the seed and the color of the beige episperm. According to the correlation analysis, it is observed in CP1 that variables such as stem diameter and plant height are positively correlated. Similarly, yield variables such as diameter, length and number of panicles were associated (Figures 5a and 5b).
The second group was made up of Blanca Subachoque, which was characterized by grains with smooth, intermediate, and wavy edges, beige and white oxalate color, white flowering panicle, higher than average yield (587.04 g/plant), height of plant of 147 cm, and greater length and diameter of panicle. In the third group, Blanca de Jericó, Aurora and Púrpura were located, characterized by pink calcium oxalates, glomerulated panicles and green stems, in the same way, for yields between 507 g/plant and 530 g/plant, diameter stem length between 20 and 21 mm, number of panicles between 14 and 16 panicles/plant, panicle length between 35 and 38 cm and panicle diameter between 95 and 100 cm.
The morphoagronomic characterization of the five quinoa cultivars from Cundinamarca evaluated under controlled conditions and in the open field in the Department of Boyacá-Colombia for the selection of cultivars that improve quinoa production in the Cundiboyacense region showed broad variability between the cultivars, which resulted from the fact that farmers in the Andes maintain mixtures of different varieties in a crop as a strategy against attacks by pests and pathogens (Morillo et al., 2021). The latitude and altitude where quinoa cultivars are developed generate changes in the growth and development of the plants, as well as in the phenotypic expression of their morphological characteristics. In quinoa, there are four environmental factors that affect its development: temperature, photoperiod, water balance and radiation. Changes are reflected in the differences in yield, grain diameter and weight, and plant height, among other characteristics (Reguera et al., 2018). The behavior of the quantitative descriptors evaluated in the open field and under controlled conditions in this study showed that, under greenhouse conditions, the five evaluated cultivars presented lower yields, weights of 1000 seeds, panicle diameters, and stem diameters. Only plant height was higher under controlled conditions.
a. Mixed factorial analysis taking into account the quantitative and qualitative variables evaluated in five quinoa cultivars in the open field b. Correlation analysis between the quantitative variables evaluated, where it is observed that the variables plant height and panicle length are correlated with yield. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
Cluster analysis. Three groups of materials formed according to the qualitative and quantitative variables evaluated in the open field are shown. Tunja, Boyacá, Universidad Pedagógica and Tecnológica de Colombia, 2021-2022.
Genotype and environment influence the determination of morphoagronomic traits of plants, especially in yield components (Manjarres et al., 2021a). The decrease in yield under controlled conditions can be attributed, among other variables, to temperature since it plays a role in the duration of development, panicle, flowering and grain formation, which are the more critical states (García et al., 2020a). Likewise, Chura et al. (2019) highlighted that temperature influences yield because of thermoperiodicity, but yield is also subject to factors such as nutrition, plant density, water supply, solar radiation and climatic factors that can act independently or together.
In quinoa, the characteristics that determine productivity are mainly panicle length, plant height, weight of 1000 seeds and yield. Gómez et al. (2022) showed that plant height is strongly dependent on variety; however, characteristics such as temperature, light hours, and radiation have a direct influence on development. Panicle length in quinoa plants depends on growth and differentiation processes that occur in the apical meristem (García et al., 2020b). These processes are determined by genotype, type of inflorescence and soil conditions, among others (Temel & Keskin, 2020). Panicle lengths between 12 and 40 cm were similar to reported by Manjarres et al. (2021b). In the morphological characterization of quinoa genotypes carried out by EL-Harty et al. (2021), under Middle Eastern conditions, the panicle length ranged between 17 and 32 cm because of saline conditions.
The correlation analyses showed that the yield of the evaluated cultivars depended to a great extent on the diameter and weight of the seeds and the density of the panicle. These results agree with those reported by Gómez et al. (2022), who found a positive correlation between yield and the weight of 1000 grains and seed diameter when evaluating quinoa progenies from simple crosses in Puno-Perú. Additionally, characteristics such as the size and weight of quinoa seeds can be selected in the early stages of a breeding program, which could have important, short-term genetic advances (Bazile, 2022).
Stem diameter is a characteristic that allows uniform, erect growth of plants and is affected by the distance between plants in a row, fertilization, crop conditions and variety (Chura et al., 2019). Studies have shown that quinoa stem diameters are between 1 and 8 cm (Mazón et al., 2019). In this study, the stem diameters of cultivars evaluated in the field were greater (20 mm) than those in the greenhouse (13.63 and 17.05 mm). Research on highland quinoa has shown thick stems (10 and 14 cm), similar to those obtained in this study, since type of quinoa has stems that resist strong winds that are typical of these regions (Chura et al., 2019).
Qualitative characteristics, by nature, are more stable than quantitative ones since they are controlled by few genes and are expressed in all environments. In genetic improvement programs, they allow easy and fast discrimination between phenotypes since combinations of these characteristics differentiate genotypes (Manjarres et al., 2021a). However, in this study, although the materials come from a selection process, there was variation in the characteristics, such as episperm color, color of streaks and calcium oxalates. These segregations have also been documented by Morillo et al. (2020) in a characterization of 19 quinoa materials in Boyacá and by Manjarres et al. (2021b) in a phenotypic characterization of quinoa to select promising materials.
In Colombia, white quinoas are preferred but in countries, such as Bolivia and Peru, black and brown varieties are desirable for gourmet cuisine. Morillo et al. (2022) found variations in grain color, where 48% of the materials exhibited beige colorations and 18% had yellow grains, which are widely accepted in the Colombian market. Montes et al. (2018) also reported a broad range of colors in quinoa materials evaluated in the Department of Cauca, particularly variations in the panicle color resulting from the fact that it is covered by a vesicular-granular pubescence that is rich in calcium oxalate, with white, pink and purple tones that color the panicle of each variety. Although qualitative variables constitute a fundamental tool for determining adaptation strategies of plants and are used as varietal descriptors, these characteristics varied in this study within individuals of the same cultivar. That is, despite selection processes, quinoa materials still segregate many morphological characteristics, as reported in studies on morphoagronomic characterizations in the Department of Boyacá (Manjarres et al., 2021a,b; Morillo et al., 2022).
The results suggested that differences between observed phenotypic characteristics can be influenced by genotype, adaptability and phenotypic plasticity of each genetic material (Reguera et al., 2018). In addition, because they are controlled by genes with pleiotropic or epistatic effects that modify the expression of the phenotype, quinoa cultivars have great adaptability to different agroclimatological conditions (Sellami et al., 2021), which can be considered an adaptive response of some quinoa genetic materials that have been cultivated for several years. It should be noted that this variability in the evaluated descriptors is the basis of breeding programs because, if there is no variability, selection cannot occur since all individuals respond in the same way to the evaluated conditions. Therefore, the existence of phenotypic variability associated with qualitative or quantitative morphological characteristics allows for the selection of materials that respond to the needs of farmers, producers and consumers (Morillo et al., 2022).
In the morphoagronomic characterization of five C. quinoa materials in Tunja, Tunkahuan and Blanca Subachoque were the materials with the best agronomic characteristics. However, there is still high heterogeneity in the characteristics within their individuals, making it necessary to continue the seed purification processes through proper selection and pollination control. Nonetheless, these materials can be promising as parental lines for future breeding programs aimed at identifying genotypes adapted to market needs
ACKNOWLEDGEMENTS
The Universidad Pedagógica y Tecnológica de Colombia (UPTC) and Group: Competitividad, Innovación y Desarrollo Empresarial (CIDE).
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