Pitahaya (Selenicereus Megalanthus Haw.): History
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Subjects: Agriculture, Dairy & Animal Science
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Ana Morillo-Coronado

The Dragon fruit, known as pitaya or pitahaya, belongs to the Cactaceae family, which originated from the southern and central regions of Mexico and America, and is separated in two genera: Hylocereus and Selenicereus. It is a nutritious and exotic fruit cultivated throughout the tropical and subtropical regions of the world. Pitahaya production has attracted interest in the United States, Australia, Southeast Asia, Israel and other regions. This fruit has gained considerable attention from consumers because it is a unique fruit that can tolerate drought stress and contains a considerable amount of nutrients. It is rich in polyphenols, vitamins, sugar, amino acids and betalain pigments.

  • germplasm
  • Selenicereus megalanthus
  • Morphological descriptors
  • Phenotypic variation
  • Genetic diversity

1. Introduction

The Dragon fruit, known as pitaya or pitahaya, belongs to the Cactaceae family, which originated from the southern and central regions of Mexico and America, and is separated in two genera: Hylocereus and Selenicereus. It is a nutritious and exotic fruit cultivated throughout the tropical and subtropical regions of the world. Pitahaya production has attracted interest in the United States, Australia, Southeast Asia, Israel and other regions [1]. This fruit has gained considerable attention from consumers because it is a unique fruit that can tolerate drought stress and contains a considerable amount of nutrients. It is rich in polyphenols, vitamins, sugar, amino acids and betalain pigments [2]. Furthermore, pitahaya fruit contains substantial amounts of unsaturated fatty acids (linoleic and linolenic) with broad applications in therapeutic and cosmetics preparations [3]. It has drawn worldwide attention because of its new flavor, color and attractive appearance, along with enormous health benefits [4]. It gained commercial potential in different countries as the result of consumer preference for new, exotic, and phytochemically rich fruits and its adaptability to new environments with abiotic stress tolerance, such as droughts and extreme temperatures [5].
Generally, differences in pitahaya germplasm can be easily shown with some distinguishing phenotypic characteristics, such as fruit size, fruit color, and number of spines at the areola that formed on branch/stem segments [6]. Nowadays, it is very difficult to separate species and varieties of the dragon fruit because of high intra and interspecific hybridization that has created some taxonomical confusion worldwide [7]. Morphological and genetic heterogeneity in many fruit characteristics, such as sweetness, size, shape, color, and bracts number, resulting from intra and inter-specific hybridization that makes it difficult to increase quality standards for the exportation market, posing serious problems when determining performance in handling and shelf-life [8].
Internationally, a large collection of pitahaya germplasm accessions is curated at the University of California South Coast Research and Extension Center (SCREC), Irvine, CA. The collection was first established in 2005 and includes seven varieties native to Nicaragua, with approximately 120 individual plants representing four different species (H. undatus, H. polyrhizus, H. costaricensis, and Hylocereus sp. Unnamed), two varieties native to Mexico with 34 individual plants representing two species (H. ocamponis and H. megalanthus), two varieties from San Diego with 34 individual plants representing two species (H. undatus and H. guatemalensis), and seven varieties from Florida, including 120 individual plants representing four species and hybrids (H. undatus, H. guatemalensis, H. megalanthus, and several putative hybrids identified as Hylocereus spp.). Additional accessions have been added intermittently to the collection, including 17 additional accessions from Nicaragua (within the H. polyrhizus/costaricensis group) [9]. These accessions were tentatively labeled H. costaricensis/polyrhizus because both species are commonly found in Nicaragua and throughout Central America. An additional variety was purchased from Mexico (17 individual plants representing H. ocamponis) and several other varieties were also sourced from Florida. Additional material has been obtained from local producers, and two plants per accession have been added to the collection. In total, the collection now includes 378 individual plants, potentially representing 54 varieties and seven different species [10].
Conventionally, morphological traits had been used to differentiate plant germplasm/species and to elucidate their genetic relationship [11]. The great morphological diversity between Hylocereus and Selenicereus species has been reported at the intraspecies and intravarietal levels, as a result of their coevolutionary process with the environment, which makes the production of new varieties extremely difficult. This led the International Union for the Protection of New Varieties of Plants [12] to develop a guide to document how new varieties of pitahaya are determined. However, as mentioned above, intraspecies/intravarietal morphological differences between vegetative clones and hybridization within this group lead to genetic mosaics between new lines and make identification between varieties extremely difficult.
Mainly because pitahaya species have easily hybridized since the late 1980s and early 1990s, breeders in the United States, Israel, and Southeast Asia have developed several hybrids [13], including crosses between either H. guatamalensis or H. megalanthus with H. undatus as the other parent, which have resulted in plants with great adaptability and high fruit quality [9].
In Colombia, the germplasm bank of yellow pitahaya and its wild relatives is located in the facilities of the National University of Colombia, Palmira. It has 300 introductions, both cultivated and wild, of yellow pitahaya (S. megalanthus 238 accessions) and red pitahaya (H. undatus, H. costaricensis, Hylocereus spp. 36 accessions). All of these are properly coded; however, not all of them are characterized [14].
In this country, knowledge of this crop mainly comes from the empirical processes of farmers who are motivated by the prices that fruits can fetch in some of the months of the year; however, increased penetration into international markets requires research on the processes of propagation, obtaining elite material, and resistance to biotic and abiotic factors, among other topics [15]. However, one of the bigger limitations is the broad morphological variation seen in the vegetative structures, which leads to confusion in identifying each species, with a lack of consensus [16], where classification is mainly based on the number of areola ribs, the contour of the stem, the relative firmness of the stem and the size and color of the fruits; in addition, various studies on domesticated cactus species have demonstrated variations in fruit characteristics related to the domestication process, resulting in a lack of a taxonomic database [1].
Studies on morphological characterization of germplasm worldwide have shown great variation in the evaluated accessions that can be conserved and used in the genetic improvement of the species; for example, a study evaluated morphological, biochemical and molecular characterizations of four dragon fruit (Hylocereus spp.) genotypes grown in Andaman and on Nicobar Island and revealed the presence of a considerable amount of genetic variations that could be used as key traits for distinguishing three different species [8]. Four dragon fruit genotypes: H. polyrhizus, H. megalanthus and two H. hybrids resulted in different plant growth and development. H. polyrhizus had the best plant growth; whereas, H. megalanthus had the lowest plant growth. Therefore, the red variety was more suitable for cultivation in Pangandaran [17]. The differences in the anatomical structure and morphology of the plants could cause differences in its optimal growth location. Morphological and agronomic characteristics can be used to determinate genetic variation in a single population.
Despite the productive potential, a limiting factor in the development of this crop in Colombia is the incidence of pest and diseases, low fruit quality, the technological level, the associativity and the lack of cultivated material, generating significant losses in yield [18][19]. Different research institutions and universities have tried to find a solution to these problems [14][18]. These studies have shown that there is no certified planting material and that only a few are grown by farmers, generating vulnerability to different phytosanitary problems, for which it is necessary to carry out genetic studies that lead to the identification of elite materials that meet the needs of the productive chain of yellow dragon fruit in Colombia since the genetic base of germplasm resources is limited.

2. Discussions on Yellow pitahaya

Yellow pitahaya is a popular crop in Colombia because of its nutritional value and productive potential. Recently, farmers have begun growing different genotypes in the producer municipalities. The morphological characteristics of these genotypes have yet to be elucidated [19]. Colombia has developed vastly with the cultivation of several dragon fruit genotypes but the morphological characteristics and adaptation to different altitudes have not been clearly studied [18]. Morphological and agronomic characteristics can be used to measure genetic diversity in a particular individual population. The plant phenotype is a form of plant adaptation to environmental conditions [17].
This study analyzed the characteristics of 11 yellow pitahaya genotypes in five municipalities in the Department of Boyacá to find out which one is optimal since one of the main problems of the study area is that there is no such variety. The planting material was introduced to the region, perhaps through the exchange of seeds between producers in the country. It was stated by [19] that there were several morphological characteristic that can be described to distinguish different types of dragon fruit species. Eleven genotypes of yellow pitahaya have shown different plant characteristic. It was stated by [20] that the main differences of Hylocereus species were the size and color of the fruits and also the shape and number of spines. This statement corresponded with the current analysis of eleven Selenicereus genotypes, in which the differences in the stem shape and number of spines were observed. Similar research was conducted by [21][22], who stated that the number of spines was a reliable characteristic to describe Hylocereus. The four dragon fruits grown in Pangandaran have similarities in the number of spines per areola bur differ in the shape and color of the spines [17].
In the case of qualitative traits, most of the genotypes exhibited an elongated fruit shape (Gen1, Gen5, Gen6 and Gen7), except Gen2 and Gen9 with a round fruit shape (Table 1). None of the genotypes had wax but there was variation for each of the categories of the morphological descriptors evaluated. It was stated by [8] that cladode, floral and fruit characteristics of H. megalanthus, such as margin ribs of cladode, waxiness, sepal color, color of ring at base of reproductive organs in flower, fruit shape, position towards peel, pulp color, peel color and seed size are visible taxonomic traits to distinguish this species from two Hylocereus spp.: H.undatus and H. costariscensis.
Table 1. Qualitative descriptors used in the morphoagronomic characterization of the yellow pitahaya genotypes in the Department of Boyacá, Colombia.
Descriptors Category Gen 1 Gen 2 Gen 3 Gen 4 Gen 5 Gen 6 Gen 7 Gen 8 Gen 9 Gen 10 Gen 11
Fruit shape (FS) Elongated 100 --- 90 70 100 100 100 40 --- 60 80
Round --- 100 10 30 --- --- --- 60 100 40 20
Surface texture of the cladodes (STF) Smooth 100 100 50 100 50 20 100 90 100 100 90
Rough --- --- 50 --- 50 80 --- 10 --- --- 10
Presence of wax (PW) Absence 100 100 100 100 100 100 100 100 100 100 100
Shape of the margin between areolas (SMA) Concave 80 40 70 80 20 20 30 90 80 60 90
Convex --- --- --- --- --- --- --- --- --- --- 10
Right 20 60 30 20 80 80 70 10 20 40 ---
Areola coloring (AC) Light grey 100 --- 20 10 50 100 30 --- --- 70 60
Dark grey --- 100 80 90 50 --- 70 100 100 30 40
Thorns color (TC) Dull brown --- --- 40 20 10 --- --- 10 10 --- 10
Bone brown --- --- 10 --- --- --- 10 --- --- --- ---
Light brown 100 --- --- 10 60 100 60 50 80 70 30
Brown --- --- 40 --- --- --- --- 30 10 --- ---
Dark brown --- 100 10 70 30 --- 30 10 --- 30 60
Pigmentation at the tips and margins of vegetative shoots (PTM) Absence 20 40 10 30 --- 60 --- --- --- --- ---
Light 60 60 40 40 50 20 40 10 100 50 50
Intense 20 --- 50 30 50 20 60 90 --- 50 50
The values indicate the percentage of introductions that present the respective category.
Cladode characteristics such as cladode width (mm), distance between areoles (mm), number of spines, length of areoles (mm), margin ribs of cladode and waxiness could be used to identify Hylocereus spp. and Megalanthus spp. [8][17]. In this study, the genotypes showed the shape of margin between areolas between concave and right, contrary to that reported in other studies on different species of Hylocereus, where the predominant forms were convex, with the presence of wax [8]. The cladode width and distance between areoles corresponded well with earlier studies from México [23], India [8], Puerto Rico and Colombia [6][18][19][21].
It was stated by [24] that pitahaya is a dispersed crop species in the tropics and subtropics that presents high polymorphism. The species has undergone human selection through the action of collecting fruits, which promoted the diversity of fruits in shape, size, color and organoleptic quality. Today, more than one species of pitahaya is known. Morphological and genetic heterogeneity in many fruits characteristics such as sweetness, size, shape, color, and bracts number by this intra and inter-specific hybridization [25] makes it difficult to increase the quality standards for the exportation market because it poses serious problems in determining their performance in handling and shelf-life. Fruit morphology, such as size, color of peel and pulp and fruit color are the main taxonomic evidence to differentiate several Hylocereus spp. and exhibit the external quality of fruits [26].
The presence of a high number of natural pollinators such as honey bees in the field plays a major role in fruit set in pitahaya fruits [27]. Total soluble solids, being the most desirable characteristic for consumer preference, is measured as Brix, which can be affected by a set of factors, such as genetic, climatic, soil, and management, among others [6]. In the present study, the SS ranged between 12.43% and 16.97%, representing better fruit quality that evidenced the earlier report that SS values between 11 and 15% have good market preference [18].
For fruit weight, most of the genotypes had values above 100 g, a desirable characteristic for the market. Individuals 2 and 9 presented the best characteristics for the market, such as fruit weight, pulp weight, fruit diameter and a high content of soluble solids, which would be of interest for distribution.
The multivariate and cluster analyses showed that the cladode and fruit characteristics showed higher variability among the morphological traits (Figure 1a,b). Although many authors find positive results, morphological and agronomic characteristics used to measure genetic diversity in certain populations of individuals often do not allow identification of discrete taxonomic groups since most plant characteristics are influenced by environmental factors, exhibiting continuous variation and a high degree of phenotypic plasticity [28]. This study demonstrated that the natural populations of pitahaya from the five departments showed high variation in the morphological characteristics that were evaluated in situ. Although the five studied sites have similar climatic conditions, the same variables should be evaluated in accessions established in a single environment, thereby confirming whether the variation might have been affected by uncontrolled environmental factors in this observational study, such as different soil types [29].
Plants 10 02255 g005 550
Figure 1. (a) Variation in width of ribs of pitahaya filocladodes characterized by farms: 1, 3, 4, 5, and 6 in the municipality of Miraflores, Russian village 2 municipality of Berbeo village Batatal; 7 municipality of Paéz, Yamuntá village. (b) Variation in pitahaya fruits characterized by farms: (1), (3), (4), (5), and (6) in the municipality of Miraflores vereda Rusa (2) municipality of Berbeo, Batatal district; (7) municipality of Paéz, Yamuntá village.
Studies in other countries have found high variation in characteristics of agronomic importance, even within the same species of Hylocereus spp. and Megalanthus spp. [6][8][21], which is favorable for future breeding studies. The availability of this information would be of great assistance in developing an appropriate method for the cultivation of a particular species [29][30]. Several studies related to the diversity of dragon fruit have been reported. A study was reported by [31] on variations in dragon fruits based on morphology, isozyme, and vitamin C content in the area of Pasuruan (East Java), Sukoharjo, Klaten (Central Java), and Bantul sub-districts (Yogyakarta). A study was reported by [26] on pollination methods on fruit set and fruit characteristics in several Pitaya clones, which aimed to improve pollination efficiency, fruit quality, and yield by determining pollination agro-management requirements. The dragon fruit plants planted in Pasuruan, Sukaharjo and Bantulhave had significant differences in stem morphology between the different species/varieties [10]. The variations in stem morphology such as curvature of the stem, margin hardness (presence of sclerenchyma), distance between areoles, number of spines, rib height, rib thickness, length, and color of the stem are important for species differentiation [21].
Apart from differences in species or accessions, differences in fruit morphology can be related to changes in the physiological level of dragon fruit at various stages of fruit development [31]. The main differences among several Hylocereus species were the size and color of the fruits and the number and form of the spines [20]. Also, the variety and flowering time have a large influence on the physio-morphological traits of dragon fruits [4][32].
In Colombia, the morphoagronomic characterization studies of these two species have obtained results similar to those reported in this study, highlighting the existence of genetic variability that can be used in conservation and genetic improvement programs that lead to the identification of elite materials, where genotypes 2 and 9 could be a good alternative. However, it is necessary to complement these morphological characterization studies with biochemical and molecular data that better discriminate the germplasm given the limitations of this type of descriptor. Research on yellow pitahaya should be intensified and extended by emphasizing its value chain and production aspects for a long-term perspective.

This entry is adapted from the peer-reviewed paper 10.3390/plants10112255

References

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