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Phenotypical characterization of Portulaca umbraticola: A non-conventional edible ornamental crop

ABSTRACT.

Purslanes are frequently used in Brazil as potted plants and for landscaping because of their beautiful colored flowers. Despite their commercial importance, the genetic diversity within ornamental purslane remains largely unknown. Thus, a complete characterization of ornamental purslane resources is essential for its utilization since genetic variability is indispensable for the efficient development of new cultivars. This study aimed to evaluate genetic variability and recommend accessions to start a genetic breeding program for purslane (Portulaca umbraticola). Twenty purslane accessions were collected in public places in Areia City and Santa Rita City, Paraiba State, Brazil. The experiment was conducted in a greenhouse. An entirely randomized experimental design with 20 accessions and 5 replicates was utilized. Plants were evaluated for quantitative and qualitative traits. Data were submitted to analysis of variance and grouped by Scott-Knott’s criteria (p ≤ 0.05). The accessions were grouped using Tocher’s method based on the generalized Mahalanobis distance. Principal component analysis was also used to analyze the genetic divergence. There were significant differences (p ≤ 0.05) for all evaluated traits except for internode distance, plant height, and leaf length. The Scott-Knott criteria clustered the accessions into two groups for all traits except days to flowering (five groups). Tocher’s grouping based on Mahalanobis distance allowed the accessions to be separated into eight clusters. The genetic diversity observed in this study was supported by the consonance between the Tocher cluster method and the PC analyses. The traits that most contributed to genetic divergence among accessions were days to flowering (70.37%), flower length (6.10%), leaf width (3.89%), branch number (3.57%), and stem length (3.36%). Considering the criteria for use in outdoor spaces, wider plants with several large colored flowers, thicker stems, greener leaves, and earliness were interesting. Accessions 5, 7, 13, 14, 16, 18, and 20 should be selected as parents for further breeding programs. However, if the objective is to produce potted plants, dwarf plants with bigger leaves are desired, and accession 1 should be selected. This will create greater variability in segregating populations, which will increase the overall quality of ornamental use of this species. For those accessions with no reproductive verticils but with other excellent attributes, for example, high chlorophyll content breeding strategies, such as induced mutations or protoplast fusion, can be used.

Keywords:
ornamental plant; neglected plants; underutilized food plants; moss rose purslane; genetic variability; morphological traits

Introduction

Portulaca umbraticola Kunth belongs to the Portulacaceae family, which includes 10 genera and 258 species (The Plant List, 2020The Plant List. (2020). Portulacaceae. Retrieved on Feb. 12, 2021 from 12, 2021 from http://www.theplantlist.org/1.1/browse /A/Portulacaceae
http://www.theplantlist.org/1.1/browse /...
). They are mainly distributed in the west of North America, South America and Africa, with a smaller number of representatives in Europe and Asia (Coelho, Giulietti, Harley, & Yesilyurt, 2010Coelho, A. A. O. P., Giulietti, A. M., Harley, R. M., & Yesilyurt, J. C. (2010). Synonymies and typifications in portulaca (Portulacaceae) of Brazil. Kew Bulletin, 65(1), 37-43. DOI: https://doi.org/10.1007/s12225-010-9187-2
https://doi.org/https://doi.org/10.1007/...
).

Most Portulaca species have no major commercial importance, except Portulaca oleraceae (common purslane), Portulaca grandiflora (nine o’clock) and P. umbraticola (moss rose purslane or large-flowered purslane) (Ocampo & Columbus, 2012Ocampo, G., & Columbus, J. T. (2012). Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae). Molecular Phylogenetics and Evolution, 63(1), 97-112. DOI: https://doi.org/10.1016/j.ympev.2011.12.017
https://doi.org/https://doi.org/10.1016/...
). Most literature involving ornamental Portulaca is about P. oleraceae or P. grandiflora; however, some authors have affirmed that P. umbraticola is taking the floriculture industry by storm. Portulaca umbraticola is planted as a bedding plant or as a container plant for landscaping with morphological characteristics that resemble those of common purslane (P. oleracea L.) (Elias, 2018; Jia et al., 2017Jia, S., Yan, Z., Wang, Y., Wei, Y., Xie, Z., & Zhang, F. (2017). Genetic diversity and relatedness among ornamental purslane (Portulaca L.) accessions unraveled by SRAP markers. 3 Biotech, 7(4), 1-8. DOI: https://doi.org/10.1007/s13205-017-0881-8
https://doi.org/https://doi.org/10.1007/...
).

Purslane has flowers with diverse colors, such as yellow, white, red, orange, pink, purple and violet (Coelho et al., 2010Coelho, A. A. O. P., Giulietti, A. M., Harley, R. M., & Yesilyurt, J. C. (2010). Synonymies and typifications in portulaca (Portulacaceae) of Brazil. Kew Bulletin, 65(1), 37-43. DOI: https://doi.org/10.1007/s12225-010-9187-2
https://doi.org/https://doi.org/10.1007/...
; UPOV, 2019International Union for the Protection of New Varieties of Plants [UPOV]. (2019). Portulaca. Guidelines for the tests for distinctness, uniformity and stability. Wellington, NZ: UPOV.). Flower color variation and small plant size potentialize ornamental use for this species (Datta, 2021Datta, S. K. (2021). Breeding of ornamentals: Success and technological status. The Nucleus, 65, 107-128.; Franca & Maia, 2008Franca, C. A. M. D., & Maia, M. B. R. (2008). Panorama do agronegócio de flores e plantas ornamentais no Brasil. In 46th Congress. Rio Branco, AC: Sociedade Brasileira de Economia, Administração e Sociologia Rural (SOBER). DOI: https://doi.org/10.22004/ag.econ.113994
https://doi.org/https://doi.org/10.22004...
). Purslane can also be used as a food source because it has essential nutrients, phenolic compounds, flavonoids, carotenoids and antioxidants and has been considered medicinal (Alam et al., 2014Alam, A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Aslani, F., Hasan, M. M, … Uddin, K. (2014a). Evaluation of antioxidant compounds, antioxidant activities, and mineral composition of 13 collected purslane (Portulaca oleracea L.) accessions. Biomed Research Internacional, 2014(2), 1-10. DOI: https://doi.org/10.1155/2014/296063
https://doi.org/https://doi.org/10.1155/...
a; 2014bAlam, A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Uddin, K., Alam, M. Z., & Latif, M. A. (2014b). Genetic improvement of purslane (Portulaca oleracea L.) and its future prospects. Molecular Biology Reports, 41(11), 7395-7411. DOI: 10.1007/s11033-014-3628-1
https://doi.org/10.1007/s11033-014-3628-...
; 2014cAlam, A., Juraimi, A. S., Yusop, M. R., Hamid, A. A., & Hakim, A. (2014c). Morpho-physiological and mineral nutrient characterization of 45 collected Purslane (Portulaca oleracea L.) accessions. Bragantia, 73(4), 426-437. DOI: https://doi.org/10.1590/1678-4499.253
https://doi.org/https://doi.org/10.1590/...
).

Brazil has eight thousand ornamental plant producers. Most of them are small- and medium-sized producers that produce in an area of more than 15.6 thousand hectares (Ibraflor, 2022Ibraflor. (2022). O mercado de flores no brasil. Retrieved on Aug. 10, 2021 from 10, 2021 from https://www.ibraflor.com.br/numeros-setor
https://www.ibraflor.com.br/numeros-seto...
). In Brazil, small farmers produce purslane, making it difficult to quantify the commercialized amount. Other species occupy the place in the big chains, such as the roses, orchids, cacti, succulents, ornamental pepper, violets and lisianthus (Ibraflor, 2021Ibraflor. (2021). Após baque em 2020, setor de flores projeta aumento nas vendas com o Dia das Mães. Ibraflor. Retrieved on Aug. 10, 2021 from 10, 2021 from https://www.ibraflor.com.br/post/após-baque-em-2020-setor-de-flores-projeta-aumento-nas-vendas-com-o-dia-das-mães
https://www.ibraflor.com.br/post/após-ba...
).

In Ibraflor’s assessment, there is a tendency for home office work and some consumption habits acquired during the pandemic to persist, which makes the sector invest in increased production for the coming years, especially the production of potted plants. This market grew during the pandemic to the detriment of ornamental plants for cutting, such as flowers and foliage (Globo Notícia, 2020Globo Notícia. (2020). Mercado de flores 'renasce' durante pandemia e projeta faturamento 5% maior em 2020. Retrieved on Aug. 10, 2021 from 10, 2021 from https://g1.globo.com/sp/campinas-regiao/noticia/2020/11/08/mercado-de-flores-renasce-durante-pandemia-e-projeta-faturamento-5percent-maior-em-2020.ghtml .
https://g1.globo.com/sp/campinas-regiao/...
; Pereira, Aguiar, & Pires, 2022Pereira, B. R., Aguiar, R. A., & Pires, L. L. (2022). Consumo de plantas ornamentais e os impactos iniciais da pandemia de Covid-19. Biodiversidade Brasileira-BioBrasil, 12(2), 1-11. DOI: https://doi.org/10.37002/biodiversidadebrasileira.v12i2.1875
https://doi.org/https://doi.org/10.37002...
).

The Ornamental plant’s global market is expanding (Ibraflor, 2021Ibraflor. (2021). Após baque em 2020, setor de flores projeta aumento nas vendas com o Dia das Mães. Ibraflor. Retrieved on Aug. 10, 2021 from 10, 2021 from https://www.ibraflor.com.br/post/após-baque-em-2020-setor-de-flores-projeta-aumento-nas-vendas-com-o-dia-das-mães
https://www.ibraflor.com.br/post/após-ba...
), and it is extremely dynamic and demands the constant release of new cultivars. Safdari and Kazemitabar (2009Safdari, Y., & Kazemitabar, S. K. (2009). Plant tissue culture study on two different races of purslane (Portulaca oleracea L.). African Journal of Biotechnology, 8(21), 5906-5911. DOI: https://doi.org/10.5897/AJB09.816
https://doi.org/https://doi.org/10.5897/...
) reported that millions of micropropagated plants of Portulaca are being afforded to the commercial ornamental market and the agricultural, clonally propagated crop market large-scale through many mircopropagation laboratories. Thus, to meet this market, ornamental plant breeding programs need to accelerate the production of new cultivars synchronized with the market’s demands (Filliettaz, 2007Filliettaz, A. (2007). Melhoramento genético de plantas ornamentais. Biológico, 69(2), 95.).

Despite its commercial importance, the genetic diversity within ornamental purslane remains largely unknown (Egea-Gilabert, Ruiz-Hernández, Parra, & Fernández, 2014Egea-Gilabert, C., Ruiz-Hernández, M. V., Parra, M. Á., & Fernández, J. A. (2014). Characterization of purslane (Portulaca oleracea L.) accessions: Suitability as ready-to-eat product. Scientia Horticulturae, 172, 73-81. DOI: https://doi.org/10.1016/j.scienta.2014.03.051
https://doi.org/https://doi.org/10.1016/...
; Jia et al., 2017Jia, S., Yan, Z., Wang, Y., Wei, Y., Xie, Z., & Zhang, F. (2017). Genetic diversity and relatedness among ornamental purslane (Portulaca L.) accessions unraveled by SRAP markers. 3 Biotech, 7(4), 1-8. DOI: https://doi.org/10.1007/s13205-017-0881-8
https://doi.org/https://doi.org/10.1007/...
; Setiawan, Aisyah, & Krisantini, 2016Setiawan, F. I. D., Aisyah, S. I., & Krisantini, K. (2016). Characterization of 13 accessions of purslane (Portulaca sp.) from Bogor, West Java, Indonesia. Journal of Tropical Crop Science, 3(3), 67-73. DOI: https://doi.org/10.29244/jtcs.3.3.67-74
https://doi.org/https://doi.org/10.29244...
). Thus, a complete characterization of ornamental purslane resources is essential for their utilization (Jia et al., 2017Jia, S., Yan, Z., Wang, Y., Wei, Y., Xie, Z., & Zhang, F. (2017). Genetic diversity and relatedness among ornamental purslane (Portulaca L.) accessions unraveled by SRAP markers. 3 Biotech, 7(4), 1-8. DOI: https://doi.org/10.1007/s13205-017-0881-8
https://doi.org/https://doi.org/10.1007/...
), since genetic variability is indispensable for efficient new cultivar development and for optimizing the choice of parents in a breeding program (Alam et al., 2015Alam, M. A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Arolu, I. W., & Latif, M. A. (2015). Genetic diversity analysis among collected purslane (Portulaca oleracea L.) accessions using ISSR markers. Comptes Rendus Biologies, 338(1), 1-11. DOI: https://doi.org/10.1016/j.crvi.2014.10.007
https://doi.org/https://doi.org/10.1016/...
).

Thus, this study aimed to evaluate genetic diversity and recommend accessions to start a breeding program for moss rose purslane (Portulaca umbraticola).

Material and methods

The experiment was conducted at Laboratório de Biotecnologia e Melhoramento Vegetal of the Centro de Ciências Agrárias, Universidade Federal da Paraíba, Areia, Paraíba State, Brazil. Twenty accessions of P. umbraticola Kunth were used, numbered 1-20 (Figure 1), and collected from public places at Areia and Santa Rita, Paraíba State, Brazil. Accessions were asexually propagated using three cuttings of approximately 15 cm in plastic pots with commercial substrate (Plantmax®). They were submitted to the best agronomic practices, including irrigation, fertilization and pesticide application for pest and plant disease control. Fertilizers were supplied monthly in the dosages recommended for vegetables. The plants received a daily water supply until they reached field capacity in the pots through micro-sprinklers installed on the greenhouse benches. They were also in full sunlight for the location 6°58'18.7" S and 35°43'15.0" W.

The evaluated characteristics were flower number, flower length, stem diameter, internode distance, branch number, canopy width, plant height, stem length, chlorophyll b, chlorophyll a, leaf length, leaf width and days to flowering. Data measurements were made with a digital caliper (Leetools®) and a graduated rule. Chlorophyll contents were measured with a digital chlorophyll meter (ClorofiLOG -FALKER®). The evaluated traits were recorded 83 days after planting.

The qualitatively evaluated traits were corolla type (pentamerous and multi), presence or absence of reproductive verticils, leaf shape, leaf tip shape and leaf base shape. The accessions were also visually evaluated using the color chart from the Royal Horticultural Society for flower color, corolla base color, anther color, style color, stem color, and leaf color.

The experimental design was completely randomized, with 20 treatments (accessions) and five replicates. The data obtained was submitted to analysis of variance and the means were grouped by Scott-Knott’s criteria (p ≤ 0.05). Genetic parameters and their estimators were analyzed for each quantitative trait using the following expressions (Cruz, Carneiro, & Regazzi, 2014Cruz, C. D., Carneiro, P. C. S., & Regazzi, A. J. (2014) Modelos biométricos aplicados ao melhoramento genético (v. 2) (3. ed). Viçosa, MG: UFV.):

a) Heritability: h2=σ²Gσ²F

b) Phenotypic variance: (σ²F) =MStk

c) Environmental variance: (σ² E ) =MSrk

d) Genotypic variance: (σ² G ) = (MSt ˗MSr) k

e) Coefficient of genetic variation: (CVg) = σG2m×100

f) Coefficient of genetic variation to coefficient of environmental variation ratio: CVg/ CVe CVgCVe=σG2σ2

where MSt = treatments mean square; k = replicate number; and MSr = residual mean square.

Mahalanobis distance was employed to determine the degree of diversity among accessions, and groups were formed following Tocher’s method (Rao, 1952Rao, C. R. (1952). Advanced statistical methods in biometric research. New York, NY: Wiley.). Divergence analysis was also performed using principal component analysis (PCA). The relative importance of the variables was determined using the method described by Singh (1981Singh, D. (1981). The relative importance of characters affecting genetic divergence. Indian Journal of Genetics and Plant Breeding, 41(1), 237-245.). Qualitative data were determined by frequency distribution. All quantitative analyses were performed using GENES software (Cruz, 2018Cruz, C. D. (2018). Programa genes (Windows): aplicativo computacional em genética e estatística. Viçosa, MG: UFV.).

Figure 1
Portulaca umbraticola UFPB accessions: 1 (A), 2 (B), 3 (C), 4 (D), 5 (E), 6 (F), 7 (G), 8 (H), 9 (I), 10 (J), 11 (K), 12 (L), 13 (M), 14 (N), 15 (O), 16 (P), 17 (Q), 18 (R), 19 (S), and 20 (T).

Results and discussion

There were significant differences (p ≤ 0.05) for all evaluated traits except internode distance, plant height, and leaf length (Table 1).

The heritability values were higher than 60% for flower length (72.54%), stem diameter (64.14%), branch number (72.86%), stem length (75.83%), leaf width (66.45), and days to flowering (98.56%). The remaining traits presented values lower than 60% (Table 1). Lower values of heritability were presented for stem diameter, number of branches, plant height, and chlorophyll in a study with P. oleraceae (Talei, Labbaf, & Naji, 2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
). High values of heritability indicate that differences among accessions were due to genetic variation and not to environmental factors (Cruz et al., 2014Cruz, C. D., Carneiro, P. C. S., & Regazzi, A. J. (2014) Modelos biométricos aplicados ao melhoramento genético (v. 2) (3. ed). Viçosa, MG: UFV.).

The coefficient of genetic variation to the coefficient of the environmental variation ratio (CVg/CVe) was lower than 1 for all traits except days to flowering (Table 1). When the values of the CVg/CVe are higher than 1, this indicates a better chance of genetic gain for this trait. According to Cruz et al. (2014Cruz, C. D., Carneiro, P. C. S., & Regazzi, A. J. (2014) Modelos biométricos aplicados ao melhoramento genético (v. 2) (3. ed). Viçosa, MG: UFV.), this parameter should be considered together with the heritability values as an indicator of selection success in breeding programs. Then, it is possible to select for earliness in the evaluated population.

Table 1
Analysis of variance summary: phenotypical variance (σ² F ), environmental variance (σ² E ), genotypic variance (σ² G ), heritability (h2), coefficient of genetic variation to environmental variation ratio (CVg/CVe), and coefficient of environmental variation (CV%) for 13 morphoagronomic traits with different accessions of Portulaca umbraticola.

The Scott-Knott criteria clustered the accessions into two groups for all traits except days to flowering (five groups) (Table 2). The accessions with a major number of flowers were 13, 15, 10, 18, 12, 9, 20, 14, and 6. Genotypes 14, 12, 20, 19, 1, 6, 11, 4, 18, 15, 5, and 13 showed larger flowers (Table 2). In an ornamental plant breeding program, it is important to select accessions with larger flowers and an increased number of flowers per plant (Datta, 2021Datta, S. K. (2021). Breeding of ornamentals: Success and technological status. The Nucleus, 65, 107-128.). Thus, accessions 6, 12, 13, 14, 15, 18, and 20 can be selected to improve these characteristics (Table 2).

Regarding stem diameter, accessions 8 and 17 presented minor values of 0.41 and 0.40 cm, respectively (Table 2). Despite all the other accessions presenting thicker stems than these, they had thinner stems than the P. oleraceae genotypes studied by Talei et al. (2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
). A higher stem diameter provides better support for the plant and facilitates asexual propagation by cutting.

Accessions 10, 17, 20, 14, 16, 18, 13, 2, and 5 presented higher branch numbers, with mean values varying from seven to eleven (Table 2). The accessions with major canopy widths were 7, 4, 3, 5, 15, 2, 19, and 6, with values varying from 48.2 to 52.8 cm (Table 2). For the variable stem length, accessions 16, 1, 17, 10, and 8 presented the lowest mean values. All other accessions formed a cluster with major stem length means varying from 53.8 to 65 cm (Table 2). The accessions with major values for these three traits should be used as bedding plants since they presented numerous long and wider branches, allowing empty spots to be filled in the garden or outdoor spaces. Talei et al. (2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
) reported no differences among genotypes of P. oleraceae for branch number. Despite the lack of significance of plant height, it is important to highlight that the values varying from 15.4 to 36.0 cm (data not shown), showing that the accessions of P. umbraticola were smaller than those of P. oleraceae studied by Talei et al. (2020).

Considering chlorophyll b, accessions 17, 10, 13, 18, 20, 8, and 9 presented higher mean values (Table 2). All accessions showed higher values of chlorophyll a, except accession 20 (Table 2). However, Talei et al. (2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
) reported no significant differences in chlorophyll content among the purslane genotypes of P. oleraceae. The accessions with larger leaves were 16, 13, 11, 7, 5, 19, 4, 6, 17, 10, 12, 18, 15, and 20 (Table 2). Traits’ mean values are important to help in the decision-making process of plant selection and in choosing accessions with favorable traits in a breeding program. Datta (2021Datta, S. K. (2021). Breeding of ornamentals: Success and technological status. The Nucleus, 65, 107-128.) underlined the importance of pigments as different economic characteristics of ornamental plants. It is important to highlight traits such as canopy width and chlorophyll b leaf width for P. umbraticola species since it can be consumed as a traditional orphan crop (Kumar et al., 2021Kumar, A., Sreedharan, S., Singh, P., Achigan-Dako, E. G., & Ramchiary, N. (2021). Improvement of a traditional orphan food crop, Portulaca oleracea L. (purslane) using genomics for sustainable food security and climate-resilient agriculture. Frontiers in Sustainable Food Systems, 5, (711820), 1-16. DOI: https://doi.org/10.3389/fsufs.2021.711820
https://doi.org/https://doi.org/10.3389/...
). According to Alam et al. (2014Alam, A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Aslani, F., Hasan, M. M, … Uddin, K. (2014a). Evaluation of antioxidant compounds, antioxidant activities, and mineral composition of 13 collected purslane (Portulaca oleracea L.) accessions. Biomed Research Internacional, 2014(2), 1-10. DOI: https://doi.org/10.1155/2014/296063
https://doi.org/https://doi.org/10.1155/...
a) and Alam et al. (2014cAlam, A., Juraimi, A. S., Yusop, M. R., Hamid, A. A., & Hakim, A. (2014c). Morpho-physiological and mineral nutrient characterization of 45 collected Purslane (Portulaca oleracea L.) accessions. Bragantia, 73(4), 426-437. DOI: https://doi.org/10.1590/1678-4499.253
https://doi.org/https://doi.org/10.1590/...
), purslane can be consumed and can also be used as medicinal plants for bioactive compounds, such as phenolic compounds, flavonoids, carotenoids, essential minerals and antioxidants.

Accessions 16, 3, 7, 2, 17, and 14 presented earliness. They bloomed from 15 to 16 days after transplanting, while other accessions showed values varying from 17 to 25 days to flowering. According to Datta (2021Datta, S. K. (2021). Breeding of ornamentals: Success and technological status. The Nucleus, 65, 107-128.), earliness is a desired trait for ornamental plants. The faster the plants bloom, the less time will be needed until commercialization. Additionally, the breeding program, based on the hybridization method, can be accelerated.

Table 2
Quantitative trait means of 20 purslane accessions (P. umbraticola).

Tocher’s grouping based on Mahalanobis distance allowed the accessions to be separated into eight clusters (Table 3). Most of the individuals were included in group I (2, 3, 7, 4, 16, and 15), followed by group II (12, 19, 11, 20, and 13). Accessions 9, 18, 10, and 6 comprised group 3. Accessions 1, 5, 8, 14, and 17 formed clusters IV, V, VI, VII, and VIII, respectively (Table 3). Group I presented the major mean value for branch number and the second major value for canopy width. The second cluster showed the major mean value for internode distance, leaf length, leaf width and days to flowering. Group III exhibited a greater mean flower number. Cluster IV had a major mean leaf width and a minor mean plant height. Group V presented a major mean for canopy width and stem length. Cluster VI showed minor values for canopy width and stem length. Group VII presented bigger flowers, thicker stems and were the most precocious plants. The major means of plant height and chlorophyll a and b were presented by cluster VIII. Alam et al. (2015Alam, M. A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Arolu, I. W., & Latif, M. A. (2015). Genetic diversity analysis among collected purslane (Portulaca oleracea L.) accessions using ISSR markers. Comptes Rendus Biologies, 338(1), 1-11. DOI: https://doi.org/10.1016/j.crvi.2014.10.007
https://doi.org/https://doi.org/10.1016/...
) grouped 45 purslane accessions into seven clusters using ISSR markers. Thus, we demonstrated that it is possible to discriminate between different accessions based on morphological traits. Several authors opined that the evaluation of genetic diversity within a purslane population is indispensable to germplasm conservation, exploitation and establishment of breeding programs (Alam et al., 2014c; Alam et al., 2015; Talei et al., 2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
).

Table 3
Tocher’s grouping of 20 Eleven O’clock P. umbraticola accessions. Means, minimum and maximum values of 13 evaluated traits.

The two first canonical variables explained 81.733% of the total variance (Table 4). The genetic diversity results observed in this study were supported by the consonance between cluster and PC analyses, except for accessions 1 and 14, which were grouped together with the accessions of major group I (Table 3; Figure 2). The data support the variability presented among accessions in this study. The lowest values for the two first principal components were presented by Alam et al. (2015Alam, M. A., Juraimi, A. S., Rafii, M. Y., Hamid, A. A., Arolu, I. W., & Latif, M. A. (2015). Genetic diversity analysis among collected purslane (Portulaca oleracea L.) accessions using ISSR markers. Comptes Rendus Biologies, 338(1), 1-11. DOI: https://doi.org/10.1016/j.crvi.2014.10.007
https://doi.org/https://doi.org/10.1016/...
) (57.7%) and Talei et al. (2020Talei, D., Labbaf, N., & Naji, A. (2020). Genetic diversity of purslane populations of Iran based on some morphological and biochemical traits. Journal of Medicinal Plants and By-Products, 2, 227-234. DOI: https://doi.org/10.22092/JMPB.2020.123123
https://doi.org/https://doi.org/10.22092...
) (55.5%) in studies with P. oleraceae. Arunachalam (1981Arunachalam, V. (1981). Genetic distance in plant breeding. Indian Journal of Genetics & Plant Breeding, 41(2), 226-236. ) considered joining the distance and principal component analysis a good procedure if the first two components accumulated at least 70% of the total variation.

Table 4
Variance estimates (eigenvalues) associated with principal components of the 13 quantitative traits evaluated in Portulaca umbraticola accessions.

Figure 2
Dispersion of 20 accessions of Portulaca umbraticola based on the first three principal components (PCA).

Through Singh’s method (1981Singh, D. (1981). The relative importance of characters affecting genetic divergence. Indian Journal of Genetics and Plant Breeding, 41(1), 237-245.), it was determined that 5 out of 13 evaluated characteristics contributed to 87.29% of genetic divergence. However, nine traits contributed to only 12.71% of genetic divergence (Table 5). The traits that most contributed to genetic divergence among accessions were days to flowering (70.37%), flower length (6.10%), leaf width (3.89%), branch number (3.57%), and stem length (3.36%) (Table 5). The variables that least contributed to genetic divergence were chlorophyll a, canopy width, stem diameter, plant height, flower number, internode distance, leaf length and chlorophyll b. According to Arunachalam (1981Arunachalam, V. (1981). Genetic distance in plant breeding. Indian Journal of Genetics & Plant Breeding, 41(2), 226-236. ), the variables that contribute a low percentage of the total variability can be discarded in future studies. However, Rêgo et al. (2003Rego, E. R., Rêgo, M. M., Cruz, C. D., Cecon, P. R., Amaral, D. S. S. L., & Finger, F. L. (2003). Genetic diversity analysis of peppers: A comparison of discarding variable methods. Crop Breeding and Applied Biotechnology, 3(1), 19-26. DOI: https://doi.org/10.12702/1984-7033.v03n01a03
https://doi.org/https://doi.org/10.12702...
) highlighted the importance of choosing the proper method to avoid discarding important variables.

Table 5
Relative contribution of quantitative characteristics of major importance to the genetic divergence of Portulaca umbraticola by the method proposed by Singh (1981Singh, D. (1981). The relative importance of characters affecting genetic divergence. Indian Journal of Genetics and Plant Breeding, 41(1), 237-245.) based on the generalized distance of Mahalanobis.

Concerning the variable corolla type, 55% of the accessions presented pentamerous flowers and 45% presented multiple petals (Figure 3A). Of the accessions, 55% presented reproductive verticils and 45 did not (Figure 3B). Although accessions 8 and 17 did not present reproductive verticils, they showed minor mean values for canopy width and major mean values for chlorophyll content and belonged to different clusters; they can be selected if the strategy used in the breeding program is induced mutations or protoplast fusion. These strategies can create new variability and increase the attributes of ornamental use of this species.

Figure 3
Corolla type (A), reproductive verticils (B), flower color (C), corolla base color (D), and style color (E) of Portulaca umbraticola.

Regarding flower colors, 30% were white, 25% yellow, 20% pink, 10% pink/yellow, 5% red, 5% hot pink and 5% orange (Figures 3C and 5). Each of these flowers had differences in their corolla base color, being 50% yellow, 20% white, 10% pink/yellow, 10% green, 5% pink and 5% orange (Figures 3D and 5). These accessions also have different style colors, presenting 40% yellow, 10% white, and 5% red; the others 45% did not present style (Figures 3E and 5). Datta (2021Datta, S. K. (2021). Breeding of ornamentals: Success and technological status. The Nucleus, 65, 107-128.) emphasized flower color as a desired commercial characteristic of ornamental plants.

For stem color, 65% of the accessions were pinkish (Figure 5S), 30% green and 5% pink (Figure 4A). The accessions presented 55% green leaves and 45% green-pink leaves (Figure 4B). However, leaf shape had 55% accessions with cuneiform leaves, 35% with spathulate leaves, and 10% linear leaves (Figure 4C). Leaf tips also presented divergences, being classified 65% as obtuse and 35% as rounded (Figure 4D). According to Souza, Pedrosa, Moreira, Rêgo, and Unêda-Trevisoli (2022Souza, J. D. S., Pedrosa, L. M., Moreira, B. R. D. A., Rêgo, E. R. D., & Unêda-Trevisoli, S. H. (2022). The more fractal the architecture the more intensive the color of flower: A Superpixel-Wise analysis towards high-throughput phenotyping. Agronomy, 12(6), 1-16. DOI: https://doi.org/10.3390/agronomy12061342
https://doi.org/https://doi.org/10.3390/...
), purplish glossy stems in yellow-flower individuals can act as confounders for the processing of images and modeling in analysis toward high-throughput phenotyping. However, this trait can be considered a differential descriptor in new purslane varieties.

Figure 4
Stem color (A), leaf color (B), leaf shape (C), and leaf tip shape (D) of Portulaca umbraticola.

Figure 5
Accessions A-1, B- 2, C-3, D-4, E-5, F-6, G-7, H-8, I-9, J-10, K-11, L-12, M-13, N-14, O-15, P-16, Q-17, R-18, S-19, and T-20 of Portulaca umbraticola. Scale: 2.82 cm.

Conclusion

There was a high level of genetic diversity among accessions of P. umbraticola in this study, suggesting that morphological traits were effective in the detection of differentiation in this species. Plants belonging to different clusters should be selected as parents and used in the hybridization-breeding program to produce recombinant genotypes of P. umbraticola. This species has big, colored flowers, which has made this plant a unique candidate for ornamental breeding programs for both outdoor environments and potted plants. Considering the criteria for use in outdoor spaces, wider plants with several large colored flowers, thicker stems, greener leaves and earliness are interesting. Accessions 5, 7, 13, 14, 16, 18, and 20 should be selected as parents for further breeding programs. However, if the objective is to produce potted plants, dwarf plants with bigger leaves are desired, and accession 1 should be selected. This will create greater variability in segregating populations, which will increase the overall quality of ornamental use of this species. For accessions with no reproductive verticils but with other excellent attributes, for example, high chlorophyll content breeding strategies, such as induced mutations or protoplast fusion, can be used.

Acknowledgments

We would like to acknowledge the Coordination for the Improvement of 458 Higher Education Personnel (CAPES) for financial support (code 0001) and the Plant Biotechnology-459 Laboratory of the Federal University of Paraíba for infrastructural support

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Publication Dates

  • Publication in this collection
    01 Mar 2024
  • Date of issue
    2024

History

  • Received
    31 Jan 2022
  • Accepted
    11 Nov 2022
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