genus hylocereus: beneficial phytochemicals, nutritional...

R E V I EW

Genus Hylocereus: Beneficial phytochemicals, nutritionalimportance, and biological relevance—A review

Sabrin Ragab Mohamed Ibrahim1,2 | Gamal Abdallah Mohamed3,4 |

Amgad Ibrahim Mansour Khedr5 | Mohamed Fathalla Zayed1,6 |

Amal Abd-Elmoneim Soliman El-Kholy7,8

1Department of Pharmacognosy and

Pharmaceutical Chemistry, College of

Pharmacy, Taibah University, Al Madinah AlMunawarah 30078, Saudi Arabia2Department of Pharmacognosy, Faculty of

Pharmacy, Assiut University, Assiut 71526,

Egypt3Department of Natural Products and

Alternative Medicine, Faculty of Pharmacy,

King Abdulaziz University, Jeddah, 21589,

Saudi Arabia4Department of Pharmacognosy, Faculty of

Pharmacy, Al-Azhar University, Assiut

Branch, Assiut 71524, Egypt5Department of Pharmacognosy, Faculty of

Pharmacy, Port Said University, Port Said

42526, Egypt6Department of Pharmaceutical Chemistry,

Faculty of Pharmacy, Al-Azhar University,

Cairo, Egypt7Department of Clinical and Hospital

Pharmacy, College of Pharmacy, Taibah

University, Al Madinah Al Munawwarah

30078, Saudi Arabia8Department of Clinical Pharmacy, Faculty

of Pharmacy, Ain-Shams University, Cairo

11566, Egypt

CorrespondenceSabrin Ragab Mohamed Ibrahim,Department of Pharmacognosy andPharmaceutical Chemistry, College ofPharmacy, Taibah University, Al Madinah AlMunawarah 30078, Saudi Arabia.Emails: [email protected];[email protected]

AbstractThe genus Hylocereus (family Cactaceae) includes about 16 species. Now its reputation is spreading

everywhere in the world due to its fruit (pitaya or pitahaya or dragon fruit), which is one of the most

popular and widely used functional foods in the world. The fruit is a wealthy provenance of vitamins,

minerals, antioxidants, and fiber. The ethno-pharmacological history of this genus indicated that it

possesses antioxidant, anticancer, antimicrobial, hepato-protective, antihyperlipidemic, antidiabetic,

and wound healing activities. Furthermore, it has been used for the treatment of cough, asthma,

hyperactivity, tuberculosis, bronchitis, mumps, diabetes, and cervical lymph node tuberculosis. Differ-

ent chemical constituents have been reported from this genus as betalains, flavonoids, phenolic

acids, phenylpropanoids, triterpenes, sterols, fatty acids, etc. The current review focuses on the uses,

botanical characterization, chemical constituents, nutritional importance, biological activities, and

safety of Hylocereus species. Also, biosynthetic pathways of betalains have been discussed.

Practical applicationsPitaya fruit is one of the most known fruit that is commercially grown in different countries of the

world for its nutritional advantages. It has acquired a wide acceptance for its pharmacological actions

against a variety of ailments. The present review revealed that pitaya contains various bioactive phyto-

constituents which might participate directly or indirectly in their highlighted biological effects.

Therefore, these compounds can be taken into account as favorable candidates for the development

of effective and novel pharmaceutical leads. Deep phytochemical studies of pitaya fruit and its phar-

macological effects, especially the mechanism of action of its constituents to clarify the relation

between traditional uses and pharmacological activities will obviously be the focus of further research.

K E YWORD S

Betalains, biological activities, Cactaceae, chemical constituents, Hylocereus, uses

Abbreviations: AA, ascorbic acid; ABTS, 2,20-azinobis-(3-ethylbenzothiazoline-6-sulfonic acid; ADM, adriamycin; AEDA, aroma extract dilution analysis; ALP, alkalinephosphatase; ALT, alanine transaminase; AST, aspartate transaminase; B16F10, mus musculus skin melanoma; Bcap-37, human breast cancer cell line; CCl4, carbon

tetrachloride; CO2, carbon dioxide; Cyt P 450, cytochrome P450; DAA, dehydroascorbic acid; DNA, deoxyribonucleic acid; DOPA, dopamine; DPPH, 2,2-diphenyl-1-

picrylhydrazyl; ESR, electron spin resonance spectroscopy; FACS, fluorescence activated cell sorting; FRAP, ferric reducing antioxidant power; FTC, ferric thiocyanate;

GSH, glutathione; H2O2, hydrogen peroxide; HDL-C, high density lipoprotein cholesterol; HepG2, liver cancer cells; HFD, high fructose diet; HO-1, heme oxygenase 1;

HT-29, human colonic adenocarcinoma; Huh7, human liver hepatoma; IC50, half maximal inhibitory concentration; IL-1b, interleukin-1b; LDL-C, low density lipoprotein;

MAS, marker-assisted selection; MGC-803, human gastric cancer cell line; MIC, minimum inhibitory concentration; mol-TEA/mol-compound, mol-trolox equivalent

activity/mol compound; NCI, National Cancer Institute; NF-jB, nuclear factor-jappa beta; NOAEL, no-observed-adverse-effect level; Nrf2, nuclear factor-erythroid-

derived 2-like 2; PC3, human prostate; PON1, paraoxonase 1; QTL, quantitative trait loci; SPME, solid phase micro-extraction; SRB, sulphorhodamine-B; TAC, total

antioxidant capacity; TBA, thiobarbituric acid; TEAC, trolox equivalent antioxidant capacity; TG, triglycerides; UDP-G, uridine diphosphate glucose; VEGF, vascular

endothelial growth factor; WRL68, embryonic normal liver cells.

J Food Biochem. 2018;e12491.https://doi.org/10.1111/jfbc.12491

wileyonlinelibrary.com/journal/jfbc VC 2018Wiley Periodicals, Inc. | 1 of 29

Received: 13 November 2017 | Revised: 29 November 2017 | Accepted: 6 December 2017DOI: 10.1111/jfbc.12491

http://orcid.org/0000-0002-6858-7560

1 | INTRODUCTION

The genus Hylocereus (A. Berger) Britton & Rose belongs to family Cac-

taceae. The species of this genus are vine cacti (climbing with aerial

roots) with three angled stems and glabrous large-scaled berry

(Montoya-Arroyo et al., 2014). It is grown ornamentally in gardens and

indoors for its big, fragrant, and night-blooming flowers. Now, its repu-

tation is spreading everywhere in the world due to its fruit. The plant is

grown in the tropical region but it must be conserved from subfreezing

temperatures and intensive solar radiation when planted under sub-

tropical states (Siddiq & Nasir, 2012). The members of family Cactaceae

became popular in Europe after American’s discovery. H. megalanthus

is native of Venezuela, Colombia, Bolivia, Peru, and Ecuador. H. undatus

is native of South America, Mexico, Martinica, and Colombia (Siddiq &

Nasir, 2012). However, H. trigonus is considered to be native of Brazil,

Uruguay, and Colombia. Hylocereus is cultivated in Nicaragua, Guate-

mala, Mexico, Costa Rica, Colombia, Peru, and Venezuela. Also, it is

come into China, Bahamas, Bermuda, Australia, United States (Florida

and California), India, Thailand, Taiwan, Malaysia, Philippines, Vietnam,

Cambodia, Indonesia, and Israel (Choo & Yong, 2011; Lim, Tan, Karim,

Ariffin, & Bakar, 2010). The common name of these fruits is pitaya or

pitahaya. Pitaya is often called “dragon fruit” in Asia. It is a medium-

large berry-bearing scales or bracts on the fruit skin, which resembles a

dragon (Wybraniec et al., 2001). The pulp is juicy and delicate with

abundant soft small seeds (Nerd & Mizrahi, 1998). Hylocereus genus

includes about 16 species (Choo & Yong, 2011; Royal Botanic Garden

Kew, 2016). They may be distinguished from each other by either the

color of soft fleshy center (mesocarp or endocarp) which has the seeds

and/or the pulpy skin’s color (exocarp). Morphologically, the observed

amount of seeds to fruit is low. The most vastly commercially grown

species are H. megalanthus (yellow Pitaya, white pulp with yellow skin),

H. polyrhizus (Red Pitaya, red pulp with pink skin), and H. undatus

(White Pitaya, white pulp with pink skin) as well as their varieties and

hybrids (Choo & Yong, 2011) (Figure 1; Table 1). In Central America,

special names have been allocated to Hylocereus genotypes based on

scales number, morphology, and shape. The most popular Hylocereus

genotypes are Rosa, San Ignacio, and Orejona in Nicaragua. Cisneros

and Tel-Zur (2012) stated that the molecular techniques used to iden-

tify the different genotypes of Hylocereus sp. are the molecular

markers, fluorescence activated cell sorting (FACS), marker-assisted

selection (MAS), and quantitative trait loci (QTL) (Cisneros & Tel-Zur,

2012). Currently, pitaya is a quite economical product for the conven-

tional producer because its cultivation requires little or no investment.

Subsequently, it can be considered as an alternative crop with high

commercial potential (Guti�errez, Solís, Baez, & Flores, 2007). The fruits

have played a remarkable role as medicine, food, and ornamentally. The

fruit is a rich source of vitamins (B1, B2, B3, C, niacin, pyridoxine, and

cobalamin), minerals (calcium, potassium, phosphorus, sodium, iron, and

zinc), protein, fat, carbohydrate, and fiber (Halimoon & Abdul Hasan,

2010; Jaafar, Abdul Rahman, Mahmod, & Vasudevan, 2009). It is also

rich in phytoalbumins, flavonoids, phenolics, and betacyanins, which are

extremely valued for their antioxidant potential (Elfi Susanti, Utomo,

Syukri, & Redjeki, 2012; Jaafar et al., 2009). The flowers of H. undatus

have been utilized for treating cough, hyperactivity, tuberculosis, bron-

chitis, mumps, diabetes, and cervical lymph node tuberculosis for a long

time in the southern China folk medicine (Guti�errez et al., 2007; Wu

et al., 2011). Dragon fruit improves the digestion process due to its

fiber, which prevents cancer of the colon and diabetes, neutralizes toxic

materials as heavy metals, and reduces high blood pressure and levels

of cholesterol (Jaafar et al., 2009). The regular consumption of dragon

fruit can help against cough and asthma. H. undatus flowers and leaves

were traditionally utilized by the Mayas as cicatrizing agent, diuretic,

and hypoglycemic (Wybraniec et al., 2001). H. polyrhizus pulp has been

utilized for the manufacturing of red-violet colored ice cream, juices,

and lipstick (Choo & Yong, 2011). In Taiwan, the fruit has been used as

a food substitute for rice and as a dietary fiber source for diabetic per-

sons (Elfi Susanti et al., 2012). Pharmacological study displayed that

Hylocereus had various bioactivities as antioxidant, anticancer,

FIGURE 1 Photos of the most common Hylocereus species fruits

2 of 29 | IBRAHIM ET AL.

antimicrobial, hepato-protective, antihyperlipidemic, antidiabetic, and

wound healing. Extensive studies of the chemical components of Hylo-

cereus have led to the identification of different compounds as betalains,

flavonoids, phenolic acids, triterpenes, sterols, and fatty acids. In this

review, botanical characterization (Table 1), chemical constituents iso-

lated over the past few decades, nutritional importance, biological activ-

ities, and safety of the genus Hylocereus are reviewed (Figures 2–15;

Tables 2–4). Also, biosynthetic pathways of betalains have been

discussed.

2 | CHEMICAL CONSTITUENTS

Genus Hylocereus is a rich source of various classes of natural constitu-

ents with diverse structural types as betalains, flavonoids, phenolic

acids, terpenes, sterols, and fatty acids. The GC-MS analysis of H. poly-

rhizus stem MeOH extract revealed the existence of four major compo-

nents: terpinolene (3.69%), eucalyptol (6.54%), b-selinene (7.25%), and

5-cedranone (73.05%), representing 91.15% of the total oil composi-

tion (Ismail, Abdel-Aziz, Ghareeb, & Hassan, 2017). C�elis, Gil, and Pino

(2012) identified 121 volatiles from H. megalanthus, consisting of alco-

hols, terpenes, paraffin’s, acids, esters, ketones and other miscellaneous

compounds utilizing solvent extraction and subsequent concentration

(C�elis et al., 2012). Then, they carried out aroma extract dilution analy-

sis (AEDA) to identify nine odor-active compounds that could poten-

tially influence flavor (C�elis et al., 2012). Obenland et al. (2016) using

solid phase micro-extraction (SPME), identified nineteen aroma vola-

tiles from six varieties of Hylocereus (Cebra, Rosa, Lisa, San Ignacio,

Mexicana, and Physical Graffiti) grown in California, including alde-

hydes, alcohols, ketones, and hydrocarbons. It is noteworthy that alde-

hydes constituted more than 90% of the total volatile amount

(Obenland et al., 2016). The observed differences between the two

previous studies could be due to the differences in the analyzed pita-

haya tissues and the used extraction methods (Obenland et al., 2016).

In the present work, we have summarized the chemical constituents

that have been characterized in the literature from Hylocereus sp. over

TABLE 1 Botanical characteristics of the common Hylocereus species

Species name Botanical characteristics Reference

H. polyrhizus (F.A.C. Web.) Britton &Rose (syn. H. monacanthus) (redpitaya or red pitaya with red flesh)

Flowers are 25–30 cm long. Perianth is red, especially at the tips. Stigma isshort, lobed, and yellow in color. Fruit is 10–12 cm long and 130–350 gin weight. It is oblong and covered with scales with different size. It hasa red flesh with many small black seeds, pleasant flesh texture andsweeter taste. H. polyrhizus has two varieties; pink- and yellow-skinned.

Ariffin et al. (2009); Le Bellecet al. (2006); Lim (2012);Siddiq and Nasir (2012);Wybraniec et al. (2001)

H. venezuelensis Britton & Rose It is closely related to H. polyrhizus, but it has bifid stigma lobes. Le Bellec et al. (2006); Lim(2012)

H. undatus (Haw.) Britton & Rose(white pitaya or red pitaya withwhite flesh)

Stems are long and green. Flowers are very long (up to 29 cm). Perianthhas outer green (or yellow-green) and inner white segments. Fruit isrosy-red with 15–22 cm length and 300–800 g weight. It is oblong andcovered with large and long scales, which are red and green at the tips.It has a white flesh with many small black seeds, pleasant flesh texture,and a good taste. Its fruit is slightly to significantly less sweeter than thered-fleshed pitaya fruit.

Ariffin et al. (2009); Le Bellecet al. (2006); Lim (2012);Siddiq and Nasir (2012)

H. megalanthus (K. Schumann exVaupel) Ralf Bauer (syn Selenicer-eus megalanthus) (yellow pitaya)

Stems are green, robust, three-ribbed, 1.5 cm thick, with slightly undu-lating margins, white areoles bearing 1–3 yellowish spines, 2–3 mmlong. Flowers are nocturnal, large, white, and funnel-shaped, 32–38 cmlong. Perianth has outer green and inner white segments.Stigma is lobed and green in color. Fruit is ovoid, tuberculate, spiny, yel-low with numerous black seeds embedded in a sweet, juicy white pulp,and much smaller than the redpitaya. It is the sweetest varieties, withrelatively smaller sized fruits.

Ariffin et al. (2009); Lim(2012); Siddiq and Nasir(2012)

H. purpusii (Weing.) Britton & Rose Flower is 25 cm long with margins.Perianth has more or less reddish outer, golden middle, and white innersegments.Fruit is oblong covered with large scales. It is 10–15 cm in length and150–400 g in weight.It has red flesh with many small black seeds, and pleasant flesh texture,but not very pronounced.

Le Bellec et al. (2006)

H. ocamponis (S.D.) Britton & Rose It is closely related to H. purpusii. They can be distinguished only by theacicular and slender spines of H. ocamponis.


H. costaricensis (Web.) Britton &Rose

Stems are waxy white. Flower is nearly the same as H. polyrhizus. Fruit is10–15 cm in diameter and 250–600 g in weight. It is ovoid and coveredwith scales with different size. It has a red purple flesh with many smallblack seeds, pleasant flesh texture and good taste.


H. trigonus (Haw.) Saff Stem is slender, green with margins, not horny. Fruit is red, ovoid oroblong, becoming nearly smooth. It is 7–9 cm in diameter and 120–250 g in weight. It has white flesh with many small black seeds andpleasant flesh texture, but not a very pronounced flavor.


IBRAHIM ET AL. | 3 of 29

the past few decades and provided a summary of their biological prop-

erties, structures, molecular weights, molecular formulae, source, and

associated references (Figures 2–15; Tables 2–4).

2.1 | Betalains

Betalains are a class of hydrophilic nitrogen-containing pigments, which

have been reported from genus Hylocereus, especially from red pitaya.

They are divided into betaxanthins (yellow-orange pigments) and beta-

cyanins (red-violet pigments). They are capable of absorbing radiation

in the visible range between 476 and 600 nm. In contrast to anthocya-

nins, betalains have carboxyl functional groups instead of hydroxyl

functional groups (Al-Alwani, Mohamad, Kadhum, & Ludin, 2015). Beta-

nins possessed a wide range of biological activities as antioxidant, anti-

inflammatory, hypoglycemic, antiproliferative, cardioactive, radiopro-

tective, neuroprotective, diuretic, hypolipidemic, and osteoarthritis pain

reliever (Esatbeyoglu et al., 2014; Khan, 2016; Lugo-Radillo, Delgado-

Enciso, & Pena-Beltr�an, 2012). The details of their biological activities

had been discussed in previous reviews (Gandía-Herrero, Escribano, &

Garcìa-Carmona, 2016; Khan, 2016).

2.2 | Biosynthesis of betalains

They are derived from the L-tyrosine amino acid that is assumed to be

originated from arogenic acid in plants (Chung et al., 2015). There are

three main enzymes involved in the biosynthesis of betalains: 4,5-DOPA-

extradiol-dioxygenase, tyrosinase, and betanidin-glucosyltransferase.

Betalamic acid is the chromophore of all betalains and the basic structure

for betalains biosynthesis. The biosynthetic pathway of betalains started

with the conversion L-tyrosine to L-DOPA (dopamine) by hydroxylation

through the tyrosinase enzyme (or polyphenoloxidase) (Figures 16 and

17). The extradiol cleavage of L-DOPA to produce 4,5-seco-DOPA

(an intermediate) was catalyzed by 4,5-DOPA extradiol-dioxygenase

(Sunnadeniya et al., 2016). Betalamic acid could be produced from

FIGURE 2 Biosynthetic pathways of betalains


4,5-seco-DOPA by the spontaneous intra-molecular condensation

between the enzymatically produced aldehyde group and the amino

group in L-DOPA. A variety of betalains have been formed by incorporat-

ing a betalamic acid unit in their structures. Betaxanthins are obtained by

spontaneous condensation reaction between the betalamic acid’s alde-

hyde group and the amino group of an amine to produce the correspond-

ing imine (Schliemann, Kobayashi, & Strack, 1999). Due to the variation

of the available amines in plants, it is difficult to determine the actual

number of plausible betaxanthins in nature. L-DOPA is transformed to

O-DOPA-quinone by tyrosinase in the lack of a reducing agent using

molecular oxygen (Chung et al., 2015). In presence of AA or reducing

agent, O-DOPA-quinone is converted back into L-DOPA (Gandía-

Herrero, Escribano, & García-Carmona, 2007). Then, the amino group of

the O-quinone undergoes an intra-molecular nucleophilic attack on the

ring. This leads to the formation of leuko-DOPA-chrome (cyclo-DOPA)

by spontaneous cyclization (Harris et al., 2012). Also, the conversion of

L-DOPA into cyclo-DOPA can be carried out by a cytochrome P450

(Hatlestad et al., 2012). Due to leuko-DOPA-chrome’s instability, it

undergoes spontaneous oxidation to DOPA-chrome accompanying with

the reduction of DOPA-quinone molecule back to L-DOPA. Furthermore,

DOPA-chrome will emerge to produce the brown polymers, character-

izing the enzymatic browning (Toivonen & Brummell, 2008). When

DOPA-chrome reacted with a reducing agent, it transformed back to

leuko-DOPA-chrome. Also, betanidin can be obtained from dopaxan-

thin and tyrosine-betaxanthin. Tyrosine-betaxanthin is produced by

the condensation of betalamic acid with L-tyrosine. Tyrosine-

betaxanthin and dopaxanthin are turned into dopaxanthin and

dopaxanthin-quinone, respectively, by the action of tyrosinase

(Gandía-Herrero et al., 2007). If dopaxanthin is to be maintained in

the existence of tyrosinase, a reducing agent is required to change

the O-quinone to the initial pigment. An intra-molecular nucleophilic

attack causes cyclization of dopaxanthin-quinone to betanidin in the

absence of a reducing agent. Cyclo-DOPA has been proposed to react

with betalamic acid to produce betanidin, which is the key intermedi-

ate in the production of betacyanins (Schliemann et al., 1999) (Figures

16 and 17). The condensation of L-DOPA with betalamic acid pro-

duces dopaxanthin. Betanidin-5-O-glucosyltransferase converts betani-

din into betanin by incorporating a glucose moiety to the 5-hydroxyl

FIGURE 3 Biosynthetic pathways of betalains continued


group (Sakuta, 2014). Conversion of betanin back to betanidin is pos-

sible due to the b-glucosidase activity (Zakharova & Petrova, 2000).

Also, it has been proposed that betanin is produced by the action of

a 5-O-cyclo-DOPA glucosyltransferase that stimulates the transmission

of sugar to cyclo-DOPA and subsequent condensation of the resulted

glucoside with betalamic acid. The formation of betanidin-quinone is

achieved from betanidin by tyrosinase enzyme. Ascorbic acid converts

betanidin-quinone to betanidin (Gandía-Herrero et al., 2007). The

remaining hydroxyl group of betanin can be oxidized by peroxidase

enzyme into betanin phenoxy radical (Gandía-Herrero & Gandía-

Carmona, 2013).

3 | NUTRITIONAL IMPORTANCE

Hylocereus polyrhizus is known as a wealthy source of minerals (e.g.,

potassium, sodium, phosphorus, iron, and calcium), vitamins (e.g., B1,

B2, B3, and C), betacyanins, protein, carbohydrate, fat, fiber, flavonoids,

polyphenols, phytoalbumin, and carotenes (Le Bellec, Vaillant, & Imbert,

2006). Jaafar et al. (2009) have stated that the nutritional composition

of H. polyrhizus is protein (0.159–0.229 g), moisture (82.5–83 g), fat

(0.21–0.61 g), vitamin C (8–9 mg/L), and crude fiber (0.7–0.9 g) (Jaafar

et al., 2009). Whereas, each 100 g of H. undatus contains protein

(1.1 g), fat (0.57 g), sorbitol (32.7 mg), vitamin C (3 mg), fiber (11.34 g),

FIGURE 4 Chemical structures of betalains (1–8) isolated form Hylocereus species


Ca (10.2 mg), P (27.5 mg), Mg (38.9 mg), K (3.37 mg), Fe (0.7 mg), Na

(8.9 mg), Zn (0.35 mg), fructose (3.2 mg), niacin (2.8 mg), b-carotene

(1.4 mg), lycopene (3.4 mg), and vitamin E (0.26 mg) (Ar�evalo-Galarza &

Ortiz-Hern�andez, 2004; Charoensiri, Kongkachuichai, Suknicom, &

Sungpuag, 2009; FAMA, 2006). Moreover, H. megalanthus fruit per

100 g edible portion contains water (85%), fat (0.1 g), energy (50 cal),

protein (0.4 g), carbohydrate (13.2 g), fiber (0.5 g), P (16 mg), Ca

(10 mg), Fe (0.3 mg), niacin (0.2 mg), and vitamin C (4 mg) (ICBF, 1992).

It is noteworthy that the high fiber content of different Pitahaya fruits

increases stool volume and protects from cancer. Hylocereus seeds oils

have earned attention due to their health significance which is related

to their comparatively high composition of endogenous antioxidants as

phenolics, tocopherols, and essential fatty acids (EFA) (Lim et al., 2010).

Ariffin et al. (2009) mentioned that linoleic and linolenic acids com-

prised a considerable proportion of the unsaturated fatty acids of H.

undatus and H. polyrhizus seed oil extracts (Ariffin et al., 2009). They

contained about 50% EFA, in which linoleic acid is in a greater ratio

than linolenic (48% C18:2 and 1.5% C18:3). In other study, Chemah,

Aminah, Noriham, and WanAida (2010) stated that the seeds are weal-

thy source of antioxidant and EFA with marked level of linoleic acid:

H. megalantus (660 g/kg), H. undatus (540 g/kg), and H. polyrhizus

(480 g/kg) (Chemah et al., 2010). Ariffin et al. (2009) stated that the

concentration of linoleic acid in Hylocereus seeds is greater than that in

canola, linseed, sesame or grapevine (Ariffin et al., 2009). Lim et al.

(2010) reported that H. undatus and H. polyrhizus seeds have a high

quantity of oil (18.33–28.37%) (Lim et al., 2010). Also, their total con-

tents of tocopherol were 36.7 and 43.5 mg/100 g, respectively (Lim

et al., 2010). These studies showed that pitaya’s seed oil has a high per-

cent of functional lipids and could be a new source of essential oil.

Wichienchot, Jatupornpipat, and Rastall (2010) reported that H. unda-

tus and H. polyrhizus pulps have glucose, fructose, and oligosaccharides

of different molecular weights, representing 86.2 and 89.6 g/kg,

respectively, total concentrations (Wichienchot et al., 2010). In yogurt,

H. undatus or H. polyrhizus pulp addition augmented lactic acid content,

milk fermentation rates, total phenolic content, and antioxidant activity

(Zainoldin & Baba, 2012).



4 | BIOLOGICAL ACTIVITIES

4.1 | Antioxidant activities

Phytonutrients are the secondary metabolites of plant origin, which

have health-boosting properties. The prominence of the antioxidant

constituents in maintaining health and protecting from cancer and cor-

onary heart disease is raising a significant interest among consumers,

food manufacturers, and scientists. Accordingly, the future’s trend is

directed to the functional food with particular health effects. In vitro

researches referred that phytonutrients as phenolic compounds may

possess a significant role, in addition to vitamins in the biological sys-

tems protection from the serious effects of oxidative stress (Kalt,

2005). Polyphenols or phenolics have gained a great attention due to

their physiological effects: antimutagenic, antioxidant, and antitumor.

They have been cited to be a powerful opponent to resist free radicals,

which are harmful to our foods systems and body (Nagai, Reiji, Hachiro,

& Nobutaka, 2003). Although phenolics do not have any nutritional

value, they may be fundamental to human health due to their

antioxidative potential. Phenolics are abundant components of the

plant that are primarily originated from phenylalanine through the phe-

nyl propanoid pathway (Hollman, Hertog, & Katan, 1996).

Choo and Yong (2011) reported that the antiradical potential of H.

polyrhizus pulps and fruits peels (IC50s 9.93 and 11.34 mg/mL, respec-

tively) was higher than those of H. undatus peels and pulps (IC50s 14.61

and 9.91 mg/mL, respectively) in DPPH assay. These results were

attributed to their contents of polyphenols and ascorbic acid (Choo &

Yong, 2011). Five different Costa Rican genotypes of Hylocereus sp.

(Lisa, Orejona, Rosa, Nacional, and San Ignacio) and H. polyrhizus fruits

were evaluated for their antioxidant effects using TEAC assay. Lisa,

Nacional, and H. polyrhizus exhibited maximum TEAC values 36.1, 34.8,

and 30.5 mg/100 mL, respectively. While the remaining genotypes

showed lower TEAC values. The significant difference observed

between different genotypes was attributed to the difference in beta-

lains contents and their composition in the different types (Esquivel,

Stintzing, & Carle, 2007). Halimoon and Abdul Hasan (2010) reported

that the ethanolic extract of H. undatus exhibited the highest



scavenging activity (63.44%) of DPPH compared to the aqueous

(55.04%) and MeOH extracts (8.82%) (Halimoon & Abdul Hasan,

2010).

Khalili et al. (2009) mentioned that red pitaya extract showed

potent antioxidant activities with 76.10 and 72.9% in the FTC and TBA

assays, respectively (Khalili et al., 2009). Moreover, the supercritical

CO2 peel extracts of H. undatus and H. polyrhizus exhibited antioxidant

activities with IC50 values of 0.91 and 0.83 mg/mL, respectively (Luo,

Cai, Peng, Liu, & Yang, 2014). The ethanolic extract of H. undatus fruit

peel exhibited antioxidant activity with an IC50 value of 0.084 mg/mL

in DPPH and TEAC value of 0.685 mM/mg in ABTS assay (Okonogi,

Duangrat, Anuchpreeda, Tachakittirungrod, & Chowwanapoonpohn,

2007). Moreover, H. undatus juice at volumes 50–200 mL possessed

antioxidant activity range from 18.5 to 30% using DPPH assay com-

pared to ascorbic acid (Sudha, Baskaran, Ramasamy, & Siddharth,

2017). The pectin from dragon fruit peels had high antioxidant poten-

tial with IC50s 0.0063–0.0080 mg/mL compared to ascorbic acid (IC50

0.00502 mg/mL) (Zaidel, Rashid, Hamidon, Salleh, & Kassim, 2017).

Moreover, the MeOH extract of H. polyrhizus stem had antioxidant

effect with TAC (total antioxidant capacity) value 726.73 mg AAE/g

dry extract using phosphomolybdenum method (Ismail et al., 2017).

Tze et al. (2012) stated that the H. polyrhizus fruit powder exhibited

antioxidant activity with an IC50 value of 2.25 mg/L in the DPPH assay

(Tze et al., 2012). The fruit flesh and peels extracts of H. polyrhizus

fruits exhibited the highest radical scavenging and reducing potentials

in DPPH and FRAP assays, respectively, due to their high betacyanin

contents. The results referred that the flesh is a substantial source of

antioxidants with health benefits for human diet and peels as a valuable

manufacture by-product to be exploited for the formulation of nutra-

ceuticals and food applications (Tenore, Novellino, & Basile, 2012).

The peels and flesh extracts of H. polyrhizus fruits exhibited antiox-

idant activities with IC50 values of 118 and 22.4 mM vitamin C equiva-

lents/g for the DPPH assay and 28.3 and 175 mM TEAC/g for ABTS

assay, respectively (Wu et al., 2006). The MeOH extract of H. undatus

exhibited strong antioxidant activity with an IC50 193 lg/mL (Elfi

Susanti et al., 2012). This variation in the observed results of the



antioxidant activity may be attributed to geographical and seasonal var-

iations. Also, quantitative and qualitative variations in the phenolics,

betalains, and ascorbic acid contents between different species of Hylo-

cereus and within the genotypes of the same species have been

reported (Esquivel et al., 2007; Lim et al., 2010).

Compounds 2, 6, and 61 isolated from H. polyrhizus exhibited a

dose-dependent peroxyl radical scavenging capacity in concentration

range 25–100 nM. Also, they showed antioxidant capacities with TEAC

values of 3.31, 2.83, and 10.70 mol-TEA/mol, respectively. In addition,

they exhibited nitrogen radical scavenging activity with IC50s 17.51,

6.81, and 24.48 mM, respectively. These results indicated that these

betacyanins will be beneficial as natural pigments to give defense

against oxidative stress (Taira, Tsuchida, Katoh, Uehara, & Ogi, 2015).

Compound 2 exhibited a dose-dependent scavenging potential of galvi-

noxyl, DPPH, hydroxyl, and superoxide radicals in the spin trapping and

electron spin resonance spectroscopy (ESR) studies. Also, it prohibited



H2O2 produced DNA damage of HT-29 cell using Comet assay at dose

15 lM. Furthermore, the treatment of Huh7 cells with 2 (15 lM)

stimulated the transcription factor Nrf2 and led to the rise of PON1

transactivation, HO-1 protein levels, and cellular GSH. So, 2 acted as

an inducer of endogenous cellular enzymatic antioxidant defense

mechanisms and as a free radical scavenger (Esatbeyoglu et al., 2014;

Sakihama, Maeda, Hashimoto, Tahara, & Hashidoko, 2012). It was

reported that betacyanins as betanin, betanidin, betanidin, and phyllo-

cactin act as strong reducing agents (Khan, 2016).

4.2 | Anticancer activities

Polyphenolics, betalains, unsaturated fats, vitamins, minerals, and toco-

pherols commonly found in pitahaya fruits give chemo-protective

potentials to counter the oxidative stress and keep balance among anti-

oxidants and oxidants to make human health effects. An imbalance

caused by excess oxidants leads oxidative stress, resulting in damage of

protein and DNA and increasing the hazard of degenerative diseases as

cancer (Luo et al., 2014; Wu et al., 2006).

The supercritical CO2 peel extracts of H. undatus and H. polyrhizus

exhibited cytotoxic activities toward Bcap-37, PC3, and MGC-803 cancer

cell lines with inhibitory ratios of 62.4 and 63.5%, 60.7 and 67.3%, and

55.2 and 78.9%, respectively, at 0.7 mg/mL compared to ADM (% inhibi-

tions 97.2, 99.3, and 98.1, respectively, at 0.1 mg/mL). Moreover, they

showed concentration-dependent antiproliferative effects with IC50 val-

ues 0.61 and 0.64, 0.45 and 0.47, and 0.43 and 0.73 mg/mL, respectively,

toward the three tested cancer cell lines. It is noteworthy that the inhibi-

tory potential of H. polyrhizus was stronger than that of H. undatus partic-

ularly toward MGC-803 cells. These activities of the pitaya peel extracts

were extremely possibly due to the presence of pentacyclic triterpenoids

and steroids, which have been known to possess anticancer activities

(Luo et al., 2014). The H. polyrhizus stem MeOH extract exhibited in vitro

cytotoxic activity toward breast (MCF-7) and liver (HepG-2) carcinoma

with IC50s 2.8 and 4.2 mg, respectively, using sulphorhodamine-B (SRB)

assay (Ismail et al., 2017). Compounds 99, 100, and 106 isolated from H.

polyrhizus and H. undatus peels exhibited cytotoxicity toward Bcap-37,

PC3, and MGC-803 cells with IC50 values of 65.4, 74.4, and 73.2, 79.3,

58.2, and 78.4, and 56.9, 43.8, and 51.9 mM, respectively. While, com-

pound 105 was found to be less active with an IC50 >100 mM compared

to ADM (IC50s 1.09, 1.34, and 0.83 mM, respectively) (Luo et al., 2014).

The peel extract of H. polyrhizus exhibited stronger antiproliferative activ-

ity than its flesh extract toward B16F10 melanoma cells with an IC50

25.0 mg of peel matter (Wu et al., 2006).



4.3 | Antimicrobial activities

The antibacterial activities of the EtOH, CHCl3, and hexane extracts of

H. polyrhizus and H. undatus peels were evaluated toward Bacillus cer-

eus, Staphylococcus aureus, Listeria monocytogenes, Enterococcus faecalis,

Salmonella typhimurium, Escherichia coli, Yersinia enterocolitica, Klebsiella

pneumonia, and Campylobacter jejuni using disc diffusion and broth

micro-dilution methods. The results showed that the chloroform

extract exhibited good antibacterial activity toward all tested patho-

gens. In addition, all extracts prohibited the growth of all bacteria with

MICs in the range of 1.25–10.0 mg/mL (Nurmahani, Osman, Abdul

Hamid, Mohamad, & Pak, 2012).

The in vitro antimicrobial potential of the extracts and fractions

from the flesh and peels of H. polyrhizus was evaluated toward two

yeasts, four molds, and 13 bacteria species, which are known to be

foodborne pathogens causing gastrointestinal, respiratory, urinary, and

skin disorders. It is noteworthy that the polyphenolic fractions showed

a broad antimicrobial spectrum toward all human pathogenic and/or

food spoilage bacteria (B. cereus, E. faecalis, S. aureus, L. monocytogenes,

E. coli, Salmonella typhi Ty2, Proteus mirabilis, Proteus vulgaris, Pseudomo-

nas aeruginosa, Y. enterocolitica, Enterobacter cloacae, K. pneumonia, and

Enterobacter aerogenes), moulds (Fusarium oxysporum, Botrytis cinerea,

Cladosporium herbarum, and Aspergillus flavus (ATCC 15517), and yeasts

(Candida albicans and Rhizoctonia solani). However, the nonfractionated

extracts revealed a very low or no activity (Tenore et al., 2012). The

acetone extract (conc. 70%) of Hylocereus peel had a high antibacterial

effect toward Salmonella typhi using agar diffusion assay (Escobar,

G�omez, Bautista, & P�erez, 2010). These studies mentioned that beta-

cyanins, flavonoids, phenolic acids, tannins, and terpenoids might be

responsible compounds for the antimicrobial activity (Nurmahani et al.,

2012; Tenore et al., 2012). The stem MeOH extract of H. polyrhizus

possessed strong antimicrobial activity against S. aureus, P. aeruginosa,



C. albicans, Aspergillus niger, and F. oxysporum with inhibition zones 29,

29, 29.5, 17.5, and 29.5 mm and 9.5, 11, 10, 8, and 16.5 mm, respec-

tively, using cup agar and disk diffusion methods, respectively (Ismail

et al., 2017).

4.4 | Antihyperlipidemic and antidiabetic activities

Consumption of vegetables and fruits lessen the incidence of cancer

and cardiovascular diseases (Stintzing, Schieber, & Carle, 2002).

Wybraniec et al. (2001) suggested that the high ingestion of vegetables

and fruits (5–7 serving/day) decreases the incidence of coronary heart

disease, attenuates the insulin resistance and dyslipidemia, and pre-

vents atherosclerosis (Omidizadeh, 2009; Wybraniec et al., 2001). It is

believed that these effects could be produced through the useful com-

bination of antioxidants, micronutrients, fiber, and phytochemical con-

tents in food (Wybraniec et al., 2001).

Daily oral administration of 1.17, 0.87, and 0.5% red pitaya to rat

feed with cholesterol-rich diet showed a significant reduction in the

total plasma cholesterol levels (59.06, 56.72, and 49.14%, respectively)

after 5 weeks of supplementation. Moreover, it had potential in



increasing HDL-C and decreasing LDL-C and TG levels. Thus, the food

supplementation of red pitaya may be helpful in the prohibition of dys-

lipidemia and cardiovascular disease (Khalili et al., 2009). Stintzing et al.

(2002) stated that the mucilage from the pulp of H. polyrhizus exerted a

positive influence on cholesterol metabolism (Stintzing et al., 2002). It

was reported that oligosaccharides obtained from white-flesh dragon

fruit decreased insulinemia and caloric intake in comparison to digested

carbohydrates. Therefore, they may be appropriate for inclusion as

food supplements in the products designed for the overweight and

diabetic individuals (Wichienchot et al., 2010). Consumption of red pit-

aya attenuated dyslipidemia and insulin resistance caused by HFD in

rats (Omidizadeh, 2009). It was reported that the fresh H. polyrhizus

fruit juice significantly reduced the hypertriglyceridemia, insulin resist-

ance, and atherosclerotic changes caused by fructose supplement in

rats. Its anti-insulin resistant effect could be referred to its polyphenols,

soluble dietary fiber, and antioxidant contents. Moreover, the antioxi-

dant content is fundamental to improve dyslipidemia and atherogenesis

in insulin-resistant rats. In addition, the soluble dietary fiber sole could



not reverse independently the hyper-insulinemia side effects (Omidizadeh

et al., 2014). Sudha et al. (2017) reported that white dragon fruit juice

had a-amylase inhibitory activity ranging from 1.033 to 32.436% at conc.

25–100 mL using starch-agar gel diffusion assay (Sudha et al., 2017).

Also, the lipase inhibitory capacity of H. undatus juice was assessed

using a Rhodamine agar plate assay. The results revealed that the

juice (conc. 25–100 mL) exhibited antilipase activity 6.125–46.939%

(Sudha et al., 2017).

4.5 | Wound healing activities

Application of the aqueous extracts of the leaves and flowers of H.

undatus topically in wounded-diabetic rats produced a significant

wound healing activity. While the fruit pulp aqueous extract had

less activity. H. undatus caused increase in the tensile strength,

hydroxyproline, DNA collagen content, total proteins, and better

epithelization thereby facilitating healing. This plant property vali-

dated its uses for the treatment of injuries in traditional medicine

(Perez, Vargas, & Ortiz, 2005).

4.6 | Anti-anemia and anti-inflammatory activities

Widyaningsih, Setiyani, Umaroh, Sofro, and Amri (2017) stated that the

red dragon fruit juice had significant effect on pregnant women’s

hemoglobin and erythrocyte levels in the seventh day of intervention

and had no effect on hemoglobin and erythrocyte levels in the 14th

day of intervention (Widyaningsih et al., 2017). Thus, its juice can be an

alternative treatment for pregnant women’s anemia. The red dragon

fruit rind extract at doses 0.25–1 mg/g bodyweight decreased

interleukin-1b (IL-1b) level, vascular endothelial growth factor (VEGF)

expression, and endometriosis in mice via decreasing nuclear factor-

jappa beta (NF-jB) activity (Eka, Hendarto, & Widjiati, 2017).

4.7 | Micro-vascular protective activities

Compounds 103 and 104 isolated from H. undatus leaves possessed

protective effects toward the skin vascular permeability increase in rab-

bits. They showed 53.5 and 70.1% reduction in the leakage of Evans

blue, respectively, at 50 mg/kg compared to troxerutin (64.5%) at the

FIGURE 13 Chemical structures of phenolic compounds (70–79) isolated form Hylocereus species


same doses. The results indicated that they increased capillary resist-

ance and reduced permeability (Guti�errez et al., 2007).

4.8 | Hepato-protective activities

The methanolic extract of H. polyrhizus fruits at 300 mg/kg body

weight exhibited significant protection of the liver against CCl4

induced hepatotoxicity in rats compared to silymarin. The results

indicated that the oral intake of H. polyrhizus fruits extract promoted

the defense status toward liver injury. The effect was due to the

phenolics and tocopherols contents which have a strong effect in

reducing the oxidative stress that enhances the cardiac and nephro-

logical damages including hepatic injury (Islam et al., 2013). Ramli,

Brown, Ismail, and Rahmat (2014) reported that red pitaya juice sup-

plementation for 8 weeks decreased ALT and ALP but gave rise to a

significant increase in AST in rats fed with a high-carbohydrate and

HFD (Ramli et al., 2014). This provides scientific evidence that the

juice of red pitaya may provide a protection toward the damage of

the liver, which could be attributed to the presence of multiple bio-

active compounds which may act synergistically. The consumption

of red pitaya supplemented diet prevents or treats the paracetamol

induced hepatotoxicity in rats and other associated deleterious

effects. This hepato-protective potential could be related to poly-

phenols, flavonoids, alkaloids, amino acids, steroids, and vitamins

(Ramli et al., 2014).

4.9 | Prebiotic effects

White-flesh dragon fruit’s oligosaccharides showed prebiotic effects.

They were used as a carbon source for the cultivation of two probiotic

strains: Lactobacillus delbrueckii BCC13296 and Bifidobacterium bifidum

NCIMB 702715. They stimulated their growth by increasing their num-

bers from 9.02 3 107 to 6.17 3 109 cell/mL within 48 hr for L. del-

brueckii and from 1.70 3 108 to 2.51 3 109 cell/mL within 72 hr for B.

bifidum (Thammarutwasik et al., 2009).

5 | ROLES OF HYLOCEREUS IN FOODINDUSTRY

People usually consume dragon fruits directly or processed into

juice. Therefore, the peel is the main byproduct of dragon fruits. The

pectic-like substance of H. polyrhizus peel and mesocarp could be

used in the food manufacture as a thickening agent (Stintzing et al.,

2002). The aqueous extract of mesocarp and the pulp juice of H. pol-

yrhizus could act as a coloring agent for low acid food commodities

(Stintzing & Carle, 2004; Stintzing et al., 2002). Tze et al. (2012)

reported that the pitaya fruit powder produced from whole pitaya

fruit has potential to use as a natural coloring agent and a health

supplement (Tze et al., 2012). Also, white-flesh dragon fruit oligosac-

charides have been included as food supplements in various food

products, for example, prebiotic and dairy products (Wichienchot

et al., 2010). Furthermore, the addition of red and white dragon

fruits into yogurt enhanced the lactic acid content, milk fermentation

rate, antioxidant activity, and total phenolics content in yogurt

(Zainoldin & Baba, 2009). The betalains, fruit pigments from red

dragon fruit are utilized as natural food colorants in different areas

of the food manufacturing (Choo & Yong, 2011). In pharmaceutical

industries, the amylase enzyme encapsulated in Arabic gum-chitosan

matrix hold complete bio-catalytic effect and possessed a consider-

able rise in the pH and temperature stabilities in comparison to the

free enzyme (Amid, Manap, & Zohdi, 2014). Additionally, the peels

could be a substantial source of novel pectinases for using in a vari-

ety of industrial and biotechnological implementations due to their

broad specificity to substrate with high stability under overdone

conditions. Also, pitaya peel could be utilized as a rich and cost-

efficient source for producing valuable types of enzymes, which

have a wide range of uses in beverage, fruit, and textile industries,

paper and pulp making, and tea and coffee fermentation (Zohdi &

Amid, 2013).

6 | ECONOMY OF HYLOCEREUSSP. PRODUCTION

Hylocereus is among the most important commercial tropical fruits

in the World (Lim et al., 2010). A great attention has been given to

it due to the promising high net profit depending on growing

Asian-United State population. Its known health significances linked

to its potential antioxidant capacities. However, its publicity at

high-end restaurants is because of its unequaled taste, prettiness,

FIGURE 14 Chemical structures of phenolic compounds (80–95)isolated form Hylocereus species


TABLE 2 List of betalains isolated from Hylocereus species

No.Compoundname Source

Molecularformula

Molecularweight Reference

1 Betanidin 5-O-b-so-phoroside

Fruit of H. polyrhizus, H. ocam-ponis, H. undatus

C30H37N2 O18 713 Wybraniec, Nowak-Wydra, Mitka,Kowalski, and Mizrahi (2007)

Fruit of H. polyrhizus Tenore et al. (2012); Wybraniec et al.(2009)

Fruits of Hylocereus sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio

Esquivel et al. (2007)

Mesocarp of H. polyrhizus Stintzing et al. (2002)

2 Betanin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus

C24H27N2 O13 551 Wybraniec et al. (2007)

Fruits of H. polyrhizus, H. un-datus, H. costaricensis, H.purpusi

Fruit of H. polyrhizus Stintzing, Conrad, Klaiber, Beifuss,and Carle (2004); Taira et al.(2015); Tenore et al. (2012); Wy-braniec and Mizrahi (2004); Wy-braniec et al. (2009); Wybraniec,Nowak-Wydra, and Mizrahi (2006)



(Continues)

FIGURE 15 Chemical structures of sterols and terpenes (96–110) isolated form Hylocereus species


TABLE 2 (Continued)


Molecularformula


Mesocarp of H. polyrhizus Stintzing et al. (2002)Fruit pulp of H. polyrhizus Wybraniec et al. (2001)Juice of H. polyrhizus Herbach, Stintzing, and Carle (2004,

2005)

3 Isobetanin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



Fruit of H. polyrhizus Stintzing et al. (2004); Taira et al.(2015); Tenore et al. (2012);Wybraniec and Mizrahi (2004);Wybraniec et al. (2006, 2009)



Mesocarp of H. polyrhizus Stintzing et al. (2002)Fruit pulp of H. polyrhizus Wybraniec et al. (2001)Juice of H. polyrhizus Herbach et al. (2004, 2005)

4 20-O-Apiosyl-betanin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus


Fruit of H. polyrhizus Wybraniec et al. (2009)Fruits of Hylocereus. sp. geno-types: Lisa, Nacional, Orejona,Rosa, and San Ignacio


5 20-O-Apiosyl-isobeta-nin



Fruit of H. polyrhizus Wybraniec et al. (2009)Fruits of Hylocereus sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio


6 Phyllocactin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



Fruit of H. polyrhizus Stintzing et al. (2004); Taira et al.(2015); Tenore et al. (2012);Wybraniec and Mizrahi (2004);Wybraniec et al. (2009)




7 Isophyllocactin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



Fruit of H. polyrhizus Stintzing et al. (2004); Tenore et al.(2012); Wybraniec et al. (2009)




8 40-Malonyl-betanin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus


(Continues)


TABLE 2 (Continued)


Molecularformula


Fruit of H. polyrhizus Wybraniec et al. (2009)Fruit of H. polyrhizus Tenore et al. (2012)

9 40-Malonyl-isobetanin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus


Fruit of H. polyrhizus Wybraniec et al. (2009)

10 Hylocerenin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



Fruit of H. polyrhizus Stintzing et al. (2004); Tenore et al.(2012); Wybraniec and Mizrahi(2004); Wybraniec et al. (2009)

Fruits of Hylocereus. sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio



11 Isohylocerenin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



Fruit of H. polyrhizus Stintzing et al. (2004); Tenore et al.(2012); Wybraniec et al. (2009)

Fruits of Hylocereus. sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio



12 20-O-Apiosyl-phyllo-cactin




13 20-O-Apiosyl-isophyl-locactin




14 500-O-E-Feruloyl-20-apiosyl-betanin



15 500-O-E-Feruloyl-20-apiosyl-isobetanin



16 500-O-E-Sinapoyl-20-apiosyl-betanin



17 500-O-E-Sinapoyl-20-apiosyl-isobetanin



18 500-O-E-Feruloyl-20-apiosyl-phyllocac-tin



19 500-O-E-Feruloyl-20-apiosyl-isophyllo-cactin



20 Isobetanidin 5-O-b-sophoroside

Fruit of H. polyrhizus C30H37N2 O18 713 Wybraniec et al. (2009)Fruits of Hylocereus sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio


(Continues)


TABLE 2 (Continued)


Molecularformula


21 Betanidin-60-O-malo-nyl-5-O-b-sophoroside

Fruit of H. polyrhizus C33H39N2 O21 799 Wybraniec et al. (2009)

22 Isobetanidin-60-O-malonyl-5-O-b-so-phoroside


23 40-O-(300-Hydroxy-300-methyl-glutaryl)betanin


24 40-O-(300-Hydroxy-300-methyl-glutaryl)isobetanin

Fruit of H. polyrhizus C30H35N2 O17 695 Wybraniec et al. (2009)Mesocarp of H. polyrhizus Stintzing et al. (2002)

25 Betanidin-5-O-(60-O-3-hydroxy-butyryl)-b-glucoside

Fruit of H. polyrhizus C28H33N2 O15 637 Esquivel et al. (2007); Tenore et al.(2012)

Mesocarp of H. polyrhizus Stintzing et al. (2002)Juice of H. polyrhizus Herbach et al. (2004)

26 2-Decarboxy-betanin Fruit of H. polyrhizus C23H27N2 O11 507 Wybraniec et al. (2006)

Juice of H. polyrhizus Herbach et al. (2005)

27 17-Decarboxy-beta-nin

Fruit of H. polyrhizus C23H27N2 O11 507 Wybraniec and Mizrahi (2004);Wybraniec et al. (2006)

Juice of H. polyrhizus Herbach et al. (2004, 2005)

28 2,17-Bidecarboxy-be-tanin


29 2-Decarboxy-phyllo-cactin

Fruit of H. polyrhizus C26H29N2 O14 593 Wybraniec et al. (2006)Juice of H. polyrhizus Herbach et al. (2004, 2005)

30 2,17-Bidecarboxy-phyllocactin


31 2-Decarboxy-hylo-cerenin


32 2,17-Bidecarboxy-hy-locerenin


33 17-Decarboxy-isobe-tanin

Juice of H. polyrhizus C23H27N2 O11 507 Herbach et al. (2004)

34 15-Decarboxy-beta-nin

Juice of H. polyrhizus C23H27N2 O11 507 Herbach et al. (2004, 2005)

35 Neobetanin Fruits of Hylocereus sp.genotypes: Lisa, Nacional,Orejona, Rosa, and SanIgnacio

C24H25N2 O13 549 Esquivel et al. (2007)


36 2-Decarboxy-neobe-tanin


37 2-Decarboxy-neobe-tanidin 5-O-(60-O-malonyl)-b-gluco-side


38 17-Decarboxy-neo-betanin


39 2,17-Bidecarboxy-neobetanin


(Continues)


TABLE 2 (Continued)


Molecularformula


40 2-Decarboxy-isophyl-locactin




42 2,17-Bidecarboxy-neobetanidin 5-O-(60-O-malonyl)-b-glucoside


43 2,17-Bidecarboxy-neobetanidin 5-O-(60-O-3-hydoxy-3-methyl-glutryl)-b-glucoside


44 17-Decarboxy-phyl-locactin

Fruit of H. polyrhizus C26H29N2 O14 593 Wybraniec and Mizrahi (2004); Wy-braniec et al. (2006)



Fruit of H. polyrhizus C29H35N2 O15 651 Wybraniec and Mizrahi (2004)Juice of H. polyrhizus Herbach et al. (2005)

46 Gomphrenin I (Beta-nidin-6-O-b-gluco-side)



47 Isogomphrenin I (Iso-betanidin-6-O-b-glucoside)



48 Isobetanidin-5-O-(60-O-3-hydroxy-butyryl)-b-glucoside(Isobutyrylbetanin)



49 15-Hydroxybetani-din-5-O-b-gluco-side


50 15-Hydroxyisobetani-din-5-O-b-gluco-side


51 Neobetanidin 5-O-b-glucoside, bi-decarboxylated,dehydrogenated

Juice of H. polyrhizus C22H23N 2O9 459 Herbach et al. (2005)

52 2-Decarboxy-neobe-tanidin 5-O-(60-O-3-hydoxy-3-methyl-glutryl)-b-glucoside


53 2,15,17-Tridecar-boxy-neobetanidin5-O-(60-O-3-hy-doxy-3-methyl-glu-tryl)-b-glucoside


54 15-Decarboxy-phyl-locactin


55 Betanidin-5-O-(60-acetyl)-b-glucoside


56 Indicaxanthin Fruit of H. polyrhizus, H. ocam-ponis, H. undatus



(Continues)


and variation (Lobo & Bender, 2008). It is commercially grown from

northern Costa Rica to Nicaragua, where �3,000 tons are producedannually on 420 ha. In 2006 in Florida, less than 50 acres were

planted (Steele & Crane, 2006) and in 2010 the production has

increased sixfold to be around 320 acres (Evans & Huntley, 2011).

Its main season is summer (June to September). Twelve to eighteen

months is the time from planting until the beginning of harvesting.

Its yields range from � 20 to 60 lb/plant (Gunasena, Pushpakumara,& Kariyawasam, 2006). Additionally, Hylocereus a perennial crop has

a lifespan of 20–30 years, assuring that with appropriate concern,

the crop can supply a stable income (Gunasena et al., 2006). The

crop also showed certain desirable agronomic features as the rela-

tive ease of propagation. Thus, reduction of the expense usually

connected with the buying of extra planting material, the simple

agronomic practices needed once the crop has been settled, and

the short turn around period of growing compared with other tropi-

cal fruits. Furthermore, its cultivation would be lucrative over a 20-

year delineation horizon (Evans & Huntley, 2011). Moreover, it is a

drought-tolerant, so it is being grown in particular areas to replace

certain crops as avocados and citrus (Gunasena, Pushpakumara,

Kariyawasam, & Hardesty, 2015). A study performed by Evans and

Huntley (2011) on an orchard of pitaya in South Florida revealed

that the cost of the establishment would be $15,136/acre, or

$75,680/5-acre of an orchard. Total investing costs are evaluated

at $109,830 (without the land cost). The total values of operating

mature plant are evaluated to be $10,127/acre, with an average

cost of $1.35/pound and a market yield of 19,000 pounds/acre.

Total profit is determined to be $25,650/acre, leading to a net

profit � $15,523/acre. That illustrates a very convenient profit incomparison with other tropical fruits, as avocados and mangoes

with a medium profit of � $1,500/acre (Evans & Huntley, 2011).

7 | SAFETY AND TOXICITY STUDIES OFHYLOCEREUS SP

The oral administered extract of H. polyrhizus fruit is relatively safe.

Acute and subchronic toxicity studies of H. polyrhizus fruit showed that

the administration of the MeOH extract of H. polyrhizus orally at doses

of 1,250, 2,500, and 5,000 mg/kg/day to female and male rats for 28

days did not show any mortality and adverse effects. Thus, its lethal

oral dose is more than 5,000 mg/kg and the NOAEL of the extract for

both female and male rats is 5,000 mg/kg/day for 28 days (Hor et al.,

2012). H. polyrhizus pulp and peel extracts are considered nontoxic

with NOAEL of more than 5 g/kg for pulp extracts and 3 g/kg for peel

extracts, administered intra-peritoneal in mice. The NOAEL via oral

administration for both pulp and peel extracts in mice were more than

TABLE 2 (Continued)


Molecularformula


57 g-Aminobutyric acid-betaxanthin




58 Isoindicaxanthin Fruit of H. polyrhizus C14H17N2 O6 309 Wybraniec et al. (2009)

59 Portulacaxanthin II(tyrosine-bx)


60 Isoportulacaxanthin II(tyrosine-isobx)


61 Betanidin Fruits of Hylocereus sp. geno-types: Lisa, Nacional, Orejo-na, Rosa, and San Ignacio


62 Neobetanidin, bi-decarboxylated,dehydrogenated

Juice of H. polyrhizus C16H13N 2O4 297 Herbach et al. (2005)

63 Miraxanthin V (dopa-mine-bx)


64 Isoleucine-Bx Fruit of H. polyrhizus C15H21N2 O6 325 Wybraniec et al. (2009)

65 Isoleucine-isoBx Fruit of H. polyrhizus C15H21N2 O6 325 Wybraniec et al. (2009)

66 Leucine-Bx (vulgax-anthin IV)


67 Leucine-isoBx (iso-vulgaxanthin IV)


68 Phenylalanine-Bx Fruit of H. polyrhizus C18H19N2 O6 359 Wybraniec et al. (2009)

69 Phenylalanine-isoBx Fruit of H. polyrhizus C18H19N2 O6 359 Wybraniec et al. (2009)


TABLE 3 List of phenolic compounds isolated from Hylocereus species

No. Compound name SourceMolecularformula


Flavonoids

70 Dihydroquercetin Flowers of H. undatus C15H12O7 304 Wu et al. (2011)71 Dihydrokaempferol Flowers of H. undatus C15H12O6 288 Wu et al. (2011)72 Kaempferol-3-O-b-D-glucopyranoside Flowers of H. undatus C21H20O11 448 Yi et al. (2012)73 Kaempferol-3-neohespedridosoide Flowers of H. undatus C27H30O15 594 Wu et al. (2011)74 Kaempferol-3-O-b-D-robinobioside Flowers of H. undatus C27H30O15 594 Yi et al. (2012)75 Kaempferol-3-O-b-D-rutinoside Flowers of H. undatus C27H30O15 594 Yi et al. (2012)

Fruit of H. polyrhizus Tenore et al. (2012)76 Quercetin-3-O-b-D-rutinoside Flowers of H. undatus C27H30O16 610 Wu et al. (2011)

Fruit of H. polyrhizus Tenore et al. (2012)77 Isorhamnetin-3-O-b-D-robinobioside Flowers of H. undatus C28H32O16 624 Yi et al. (2012)78 Kaempferol-3-O-b-D-glucopyranoside Flowers of H. undatus C22H22O12 478 Yi et al. (2012)

Fruit of H. polyrhizus Tenore et al. (2012)79 Isorhamnetin-3-O-b-D-rutinoside Flowers of H. undatus C28H32O16 624 Yi et al. (2012)

Fruit of H. polyrhizus Tenore et al. (2012)

Phenolic acids and phenylpropanoids

80 P-Hydroxybenzoic acid Fruit of H. polyrhizus C7H6O3 138 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

81 Protocatechuic acid Fruit of H. polyrhizus C7H6O4 154 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

82 Vanillic acid Fruit of H. polyrhizus C8H8O4 168 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

83 Caffeic acid Fruit of H. polyrhizus C9H8O4 180 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

84 Gallic acid Fruit of H. polyrhizus C7H6O5 170 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

85 Syringic acid Fruit of H. polyrhizus C9H10O5 198 Tenore et al. (2012)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

86 Trans-3,4-dimethoxycinnamic acid Flowers of H. undatus C11H12O4 208 Wu et al. (2011)87 Trans-Ferulic acid Flowers of H. undatus C10H10O4 194 Wu et al. (2011)88 P-Coumaric acid (88) Fruit of H. polyrhizus C9H8O3 164 Tenore et al. (2012)

Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)89 Phthalic acid, 6-ethyloct-3-yl 2-ethylhexyl ester Peel fruits of H. polyrhizus, H. undatus C26H42O4 418 Luo et al. (2014)90 1,2-Benzenedicarboxylic acid, mono

(2-ethylhexyl) esterPeel fruits of H. polyrhizus, H. undatus C16H22O4 278 Luo et al. (2014)

91 Undatuside A Flowers of H. undatus C19H26O10 414 Wu et al. (2011)92 Undatuside B Flowers of H. undatus C20H28O10 428 Wu et al. (2011)93 Undatuside C Flowers of H. undatus C20H28O10 428 Wu et al. (2011)94 Benzyl-b-D-glucopyranoside Flowers of H. undatus C13H18O6 270 Wu et al. (2011)95 Phenylethyl-b-D-glucopyranoside Flowers of H. undatus C14H20O6 284 Wu et al. (2011)

TABLE 4 List of sterols, triterpenes, fatty acids, aliphatic, and miscellaneous compounds isolated from Hylocereus species



Sterols and triterpenes

96 Campesterol Peel fruits of H. polyrhizus, H. undatus C28H48O 400 Luo et al. (2014)Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)

97 Stigmasterol Peel fruits of H. polyrhizus, H. undatus C29H48O 412 Luo et al. (2014)98 g-Sitosterol Peel fruits of H. polyrhizus, H. undatus C29H50O 414 Luo et al. (2014)

Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)99 b-Sitosterol Peel fruits of H. polyrhizus, H. undatus C29H50O 414 Luo et al. (2014)

Seed oil of H. undatus, H. polyrhizus Lim et al. (2010)100 Stigmast-4-en-3-one Peel fruits of H. polyrhizus, H. undatus C29H46O 410 Luo et al. (2014)101 Ergosta-4,6,8(14),

22-tetraen-3-onePeel fruits of H. polyrhizus, H. undatus C28H40O 392 Luo et al. (2014)

102 Cholesterol Seed oil of H. undatus, H. polyrhizus C27H46O 386 Lim et al. (2010)103 Taraxast-20-ene-3a-ol Leaves of H. undatus C30H50 O 426 Guti�errez et al. (2007)104 Taraxast-12,20(30)-

dien-3a-olLeaves of H. undatus C30H48 O 424 Guti�errez et al. (2007)

105 a-Amyrin Peel fruits of H. polyrhizus, H. undatus C29H48O 412 Luo et al. (2014)106 b-Amyrin Peel fruits of H. polyrhizus, H. undatus C29H48O 412 Luo et al. (2014)107 Terpinolene Stem of H. polyrhizus C10H16 136 Ismail et al. (2017)

(Continues)


TABLE 4 (Continued)



108 Eucalyptol Stem of H. polyrhizus C10H18O 154 Ismail et al. (2017)109 b-Selinene Stem of H. polyrhizus C15H24 204 Ismail et al. (2017)110 5-Cedranone Stem of H. polyrhizus C15H24O 220 Ismail et al. (2017)

Fatty acids and aliphaticcompounds

111 Myristic acid Seed oil of H. undatus, H. polyrhizus C14H28O2 228 Ariffin et al. (2009); Liaotrakoon,Clercq, Hoed, and Dewettinc(2013); Lim et al. (2010)

112 Palmitic acid Peel fruits of H. polyrhizus,H. undatus C16H32 O2 256 Luo et al. (2014)Seed oil of H. undatus, H. polyrhizus Ariffin et al. (2009); Liaotrakoon

et al. (2013); Lim et al. (2010)Seed oil of H. polyrhizus Villalobos-Guti�errez, Schweiggert,

Carle, and Esquivel (2012)113 Margaric acid Seed oil of H. undatus, H. polyrhizus C17H34O2 270 Liaotrakoon et al. (2013)114 Stearic acid Seed oil of H. undatus, H. polyrhizus C18H36O2 284 Ariffin et al. (2009); Liaotrakoon

et al. (2013); Lim et al. (2010)Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)

115 Arachidic acid Seed oil of H. undatus, H. polyrhizus C20H40O2 312 Liaotrakoon et al. (2013); Lim et al.(2010)

Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)116 Behenic acid Seed oil of H. undatus, H. polyrhizus C22H44O2 340 Liaotrakoon et al. (2013)117 Lignoceric acid Seed oil of H. undatus, H. polyrhizus C24H48O2 368 Liaotrakoon et al. (2013)118 Oleic acid Peel fruits of H. polyrhizus, H. undatus C18H34O2 282 Luo et al. (2014)

Seed oil of H. undatus, H. polyrhizus Ariffin et al. (2009); Liaotrakoonet al. (2013); Lim et al. (2010)

Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)119 Palmitoleic acid Seed oil of H. undatus, H. polyrhizus C16H30O2 254 Ariffin et al. (2009); Liaotrakoon

et al. (2013); Lim et al. (2010)Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)

120 Cis-Vaccenic acid Seed oil of H. undatus, H. polyrhizus C18H34O2 282 Ariffin et al. (2009)Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)

121 Erucic acid Seed oil of H. undatus, H. polyrhizus C22H42O2 338 Liaotrakoon et al. (2013); Lim et al.(2010)

122 Gadoleic acid Seed oil of H. undatus, H. polyrhizus C20H38O2 310 Liaotrakoon et al. (2013)123 Hexadecadienoic acid Seed oil of H. undatus, H. polyrhizus C16H28O2 252 Liaotrakoon et al. (2013)124 Linoleic acid Peel fruits of H. polyrhizus, H. undatus C18H32 O2 280 Luo et al. (2014)

Seed oil of H. undatus, H. polyrhizus Ariffin et al. (2009); Liaotrakoonet al. (2013); Lim et al. (2010)

Seed oil of H. polyrhizus Villalobos-Guti�errez et al. (2012)125 2-Chloroethyl linoleate Peel fruits of H. polyrhizus, H. undatus C20H35ClO2 342 Luo et al. (2014)126 Linolenic acid Seed oil of H. undatus, H. polyrhizus C18H30O2 278 Ariffin et al. (2009); Liaotrakoon

et al. (2013); Lim et al. (2010)127 Eicosatrienoic acid Seed oil of H. undatus, H. polyrhizus C20H34O2 306 Liaotrakoon et al. (2013)128 Arachidonic acid Seed oil of H. undatus, H. polyrhizus C20H32O2 304 Liaotrakoon et al. (2013)129 1-Nonadecene Peel fruits of H. polyrhizus, H. undatus C19H38 266 Luo et al. (2014)130 17-Pentatriacontene Peel fruits of H. polyrhizus, H. undatus C35H70 490 Luo et al. (2014)131 Octacosane Peel fruits of H. polyrhizus, H. undatus C28H58 394 Luo et al. (2014)132 Eicosane Peel fruits of H. polyrhizus, H. undatus C20H42 282 Luo et al. (2014)133 Tetratriacontane Peel fruits of H. polyrhizus, H. undatus C34H70 478 Luo et al. (2014)134 1-Tetracosanol Peel fruits of H. polyrhizus, H. undatus C24H50O 354 Luo et al. (2014)135 Heptacosane Peel fruits of H. polyrhizus, H. undatus C27H56 380 Luo et al. (2014)136 11-Hexacosyne Peel fruits of H. polyrhizus, H. undatus C26H50 362 Luo et al. (2014)137 Octadecanal Peel fruits of H. polyrhizus, H. undatus C18H36O 268 Luo et al. (2014)138 Nonacosane Peel fruits of H. polyrhizus, H. undatus C29H60 408 Luo et al. (2014)139 Octadecane Peel fruits of H. polyrhizus, H. undatus C18H38 354 Luo et al. (2014)140 Docosane Peel fruits of H. polyrhizus, H. undatus C22H46 310 Luo et al. (2014)

Miscellaneous compounds

141 (R)-(2) Citramalicacid

Flowers of H. undatus C6H10O4 146 Wu et al. (2011)

142 (R)-(2) Citramalicacid-1-methyl ester


143 (R)-(2) Citramalicacid-4-methyl ester


144 a-Tocopherol Seed oil of H. undatus, H. polyrhizus C29H50O2 430 Liaotrakoon et al. (2013); Lim et al.(2010)

145 b-Tocopherol Seed oil of H. undatus, H. polyrhizus C28H48O2 416 Lim et al. (2010)

(Continues)


5 g/kg. Moreover, H. polyrhizus pulp and peel extracts were nontoxic in

WRL68 and HepG2 in vitro. The peel extract caused cell death in

HepG2 cells with a high IC50 (4.2 mg/mL), which is considered nontoxic

according to the NCI. Intake of exaggerated amounts of H. polyrhizus

fruit resulted in pseudo-hematuria which is a harmless reddish discolor-

ation of the feces and urine (Shakir, 2009).

8 | CONCLUSION

Currently, the awareness of consumer for healthy food products is

growing and food researchers have been looking for beneficial sources

of healthy components. Antioxidants from a natural source are more

idealistic as food due to their free radical scavenging effects.

TABLE 4 (Continued)



146 g-Tocopherol Seed oil of H. undatus, H. polyrhizus C28H48O2 416 Liaotrakoon et al. (2013); Lim et al.(2010)

147 d-Tocopherol Seed oil of H. undatus, H. polyrhizus C27H46O2 402 Liaotrakoon et al. (2013); Lim et al.(2010)

148 Squalene Peel fruits of H. polyrhizus, H. undatus C30H50 410 Luo et al. (2014)149 Trichloroacetic acid,

hexadecyl esterPeel fruits of H. polyrhizus, H. undatus C18H33Cl3O2 386 Luo et al. (2014)

150 Hexadecyl oxirane Peel fruits of H. polyrhizus, H. undatus C18H36O 268 Luo et al. (2014)151 6-Tetradecanesulfonic

acid, butyl esterPeel fruits of H. polyrhizus, H. undatus C18H38 O3S 344 Luo et al. (2014)

FIGURE 16 Chemical structures of fatty acids and aliphatic compounds (111–140) isolated form Hylocereus species


Additionally, they are safer and healthier than synthetic ones. Pitaya

fruit is one of the most known fruits that is commercially grown in dif-

ferent countries of the world for its nutritional advantages. It has

acquired a wide acceptance for its pharmacological actions against a

variety of ailments. Recently, many studies have shown it to exhibit dif-

ferent bioactivities, some of which justified its uses in various cultures.

The present review focused on the pharmacological activities and nutri-

tional benefits of pitaya fruit. It contains bioactive phytoconstituents

which might participate directly or indirectly to the highlighted biologi-

cal effects in this review. These compounds can be taken into account

as favorable candidates for the evolution of effective and novel phar-

maceutical leads. Deep phytochemical studies of pitaya fruit and its

pharmacological effects, especially the way of action of its constituents

to clarify the relation between traditional uses and pharmacological

activities will obviously be the focus of further research.

CONFLICT OF INTEREST

We wish to confirm that there are no known conflicts of interest

associated with this publication and there has been no significant

financial support for this work that could have influenced its

outcome.

ORCID

Sabrin Ragab Mohamed Ibrahim http://orcid.org/0000-0002-6858-

7560

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