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Opuntia and Other Cacti: Applications and Biotechnological Insights

Article  in  Tropical Plant Biology · September 2010

DOI: 10.1007/s12042-010-9055-0

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Tropical Plant Biol. (2010) 3:136–150
DOI 10.1007/s12042-010-9055-0

Opuntia and Other Cacti: Applications

and Biotechnological Insights
Utkarsha U. Shedbalkar & Vinayak S. Adki &
Jyoti P. Jadhav & Vishwas A. Bapat

Received: 12 May 2010 / Accepted: 4 August 2010 / Published online: 24 August 2010
# Springer Science+Business Media, LLC 2010

Abstract The cactus family is unusual among tropical Abbreviations

plants. Cacti, known for their minimum water requirement, ACC 1-aminocyclopropane-1-carboxylate
have been grown extensively in arid lands, for food, feeds BA benzyladenine
and medicinal and therapeutic uses.Several food products DLG daily live-weight gain
have cacti as a main ingredient. Cacti biochemical analysis 2,4-D 2,4-dichlorophenoxyacetic acid
substantiate the high nutritive value of this plant family. DMI dry-matter intake
Tissue cultures, including micropropagation, callus, and FAP 6-furfurylaminopurine
cell suspension cultures have been established for numer- Kin kinetin
ous cacti species. Genetic engineering has opened oppor- LDL low density lipoprotein
tunities for gene isolation and integration of genes from LWG total live-weight gain
other sources for cacti improvement. Cacti might be a store MS Murashige and Skoog
house of stress tolerant genes for other crops. Since cacti NAA 1-naphtalenacetic acid
can be cultivated easily with minimum agriculture inputs, SF stem flour
they hold great potential for cultivation and farming on WF wheat flour
degraded lands and for at least partial remediation of degraded GA3 Gibberelic acid
lands. The present review outlines some of the older and more
recent research on the properties and applications for Opuntia
and other cacti especially as they might apply towards
agricultural sustainability. Introduction

Keywords Biochemical analysis . Cactus . Food products . Cacti are the most conspicuous and characteristic plants of
Genetic engineering . Medicines . Phytoremediation . warm arid regions. Cacti are remarkable for their diversity of
Tissue culture growth forms and their ability to not only grow, but to thrive
under environments recognized as stressful for most plant
species. Cacti can be used to prevent soil erosion and act as
an effective living fence for land reclamation while func-
Communicated by: Paul Moore tioning as a commercial crop with unique attributes. Cacti do
Utkarsha U. Shedbalkar and Vinayak S. Adki have equal contributions. not need much water and accordingly they exhibit unusual
U. U. Shedbalkar : J. P. Jadhav
physiological and morphological features (Scheinvar 1995;
Department of Biochemistry, Shivaji University, Le Houerou 2000). Cacti are known to contain several
Kolhapur 416 004, India useful chemical compounds having nutritional and medic-
inal desirable properties (Agurell 1969; Knishinsky 1971;
V. S. Adki : J. P. Jadhav : V. A. Bapat (*)
Department of Biotechnology, Shivaji University,
Rosemberg and Paul 1973; Bruhn and Agurell 1974; Bruhn
Vidyanagar, Kolhapur 416 004, India and Lindgren 1976; Ferrigni et al. 1984; Wang 1988; Frati
e-mail: [email protected] et al. 1990; Fernandez et al. 1992; Kazemini 1994; Sahelian
Tropical Plant Biol. (2010) 3:136–150 137

2001; Fernández-López et al. 2002; Alarcon-Aguilar et al. countries. Within the genus Opuntia, O. ficus-indica is the
2003; Galati et al. 2003; Oliveira and Da Machado 2003; most important for its production of edible fruits and
Gentile et al. 2004; Sirivardhana and Jeon 2004; Tesoriere cladodes, which are used as a vegetable and as animal
et al. 2004; Zou et al. 2005; Saleem et al. 2006). Cacti have forage. This particular cactus is a valuable resource
been exploited as a cheap, alternate source of food suitable especially during periods of drought when there is a shortage
for humans and feed for animals and are cultivated as of other herbaceous plants for forage (Scheinvar 1995; Le
ornamental crops (Estrada-Luna et al. 2008). Based on this Houerou 2000; Juárez and Passera 2002).
backdrop, the main objective of the present review is to Cacti have evolved to grow in to water scarce environ-
outline the significance of cacti highlighting opportunities ments. Cacti stems are swollen with water-storage tissues
for utilizing this group of plants in numerous applications. and are sometimes termed stem succulents. Cacti stems
Cacti exhibit many characteristics of a suitable crop model have a thick water proof epidermis covered with a waxy
to achieve productivity and sustainability with minimal cuticle and many are grooved with ribs so that the stem can
ecological or environmental impact to meet the growing expand and contract without damage to the surface tissues
demand for food and as a new source for meeting in the process of storing and utilizing water. These traits
nutritional health requirements and thus deserves further evolved to allow the stems to function as the main
investigations (Fig. 1). photosynthetic system. In fact, most cacti do not produce
The Cactaceae family includes approximately 130 leaves at all. The main exceptions are the species of
genera and 2,000 species. Different growth forms or habits Opuntia (prickly pears and chollas) that have rudimentary
of cacti are depicted in Table 1. Opuntia and Nopalea are leaves in their juvenile stages. However, even these juvenile
the most important genera due to their many applications leaves are cylindrical and fleshy, adapted to withstand water
(Hollis 1978; Hollis and Mejorada 1992; Flores-Valdez and loss and drought conditions. Cacti generally have extensive,
Osorio 1996). The prickly pear (Opuntia ficus indica) is shallow root systems, spreading just below the soil surface,
native of the semi-arid regions of South and Central so that they can absorb water, even the relatively small
America (Hegwood 1990; Hau 2002). This branched cactus amounts that moisten the soil surface during light rain
is a long-domesticated crop critically important in agricul- showers. Plants in the genus Opuntia prefer dry, hot areas
tural economies throughout the arid and semiarid parts of populated with perennial shrubs, trees, and creeping plants.
the world (Hau 2002; Griffith 2004). It is widely distributed Cacti can grow 3–19 m in height and can spread over an
in Europe, Southwestern United States, Northern Mexico, area of up to 40 m in diameter. Among cacti, Pachycereus
much of Latin America, South Africa and the Mediterranean pringlei is recorded as reaching a height of 19.2 m and

Fig. 1 Work done

(unpublished) on Nopalea
cochenillifera (L.) Salm. Dyck.
at Department of Biotechnology,
Shivaji University, Kolhapur.
a: N. cochenillifera (L.)
Salm. Dyck. shrub on field.
b: N. cochenillifera (L.) Salm.
Dyck. flower. c: In vitro shoot
culture of N. cochenillifera (L.)
Salm. Dyck. d: In vitro callus
culture of N. cochenillifera (L.)
Salm. Dyck
138 Tropical Plant Biol. (2010) 3:136–150

Table 1 Different growth forms of Cacti 1976). Spines, 1–6 per areole, may be absent or very highly
Type of Cactus Photograph Example reduced, or on the margins of nearly all areoles. Spines are
Columnar cactus Carnegiea (Saguaro), Lophocereus erect to spreading, whitish, tan or brown, setaceous only, or
(Senita), Packycereus (Cardon),
setaceous and subulate, straight to slightly curved, basally
angular-flattened, or appear as small bristle-like deflexed
Stenocereus (Organpipe Cactus)
spines to 5 mm. Spines are constituted of 96% poly-
saccharides consisting of 49.7% cellulose and 50.3%
arabinan. The cellulose microfibrils, 0.4 mm length and
Barrel cactus Ferocactus (Fishhook Barrel)
6–10 mm in diameter and are loosely imbedded in an
Echinocactus (Many-headed Barrel) arabinan matrix. The latter is partly present as a solid gel,
partly woven tightly with the cellulose fibers (Malainine et
al. 2003). Functions of the spines include mechanical
protection from herbivores, reflection of light, and shading
Hedgehog cactus Echinocereus (Hedgehog Cactus)
of the cladode to reduce water loss and for condensing fog
(Anderson 2001).
Cacti flowers, petals and sepals are numerous in quantity
and color. The inner petals are generally yellow to orange
throughout, filaments and anthers are yellow; the style is
bright red, and the stigma lobes yellow. Fruits are oval,
Pincushion cactus Mammillaria (Fishhook Pincushion
pear-shaped, yellow to orange to purple, fleshy (pulpy),
Cactus). juicy, sweet, glabrous, usually spineless, but they may be
covered with spines or bristles. Areoles on fruit are 45–60
in number and evenly distributed. The seeds within the pulp
are disk-shaped, sub circular and have numerous colors but
Cholla cactus Opuntia (Chainfruit, Christmas,
are generally pale tan. (Chevallier 1996; Hocking 1997; Le
Staghorn, and Teddybear Cholla) Houerou 2000; Van Sittert 2002; Defelice 2004; Synman
2006; Saleem et al. 2006). O. ficus-indica seeds, as in many
other Opuntia species, have low germination capacity
mainly due to their hard lignified integuments, the most
Prickly pear cactus Opuntia (Prickly Pear), Nopalea inward of which is the funiculus that envelops the embryo
(cochineal nopal)
and obstructs radicle protrusion. Certain physical and
chemical treatments can overcome low seed germination.
Altare et al. (2006) found that chemical scarification of O.
ficus-indica seeds with concentrated H2SO4 or with
Schweizer reagent, followed by incubation of the treated
seeds in solutions of H2O2, under photoperiodic conditions
Source: http://wc.pima.edu/~bfiero/tucsonecology/plants/cactus_types.htm
of 14 h light per day gave the highest germination
percentage in the shortest time. Ochoa-Alfaro et al. (2008)
Carnegiea gigantea to a height of 15 m with a girth of studied the effect of salt stress, proline and polyamines on
3.1 m. Stems (cladodes) of cacti are branched, green, seed germination of O. streptacantha. Proline (1 mM)
broadly oblong to ovate to narrowly elliptic. About 7–11 stimulated higher germination compared to untreated seeds.
areoles are present on stems per diagonal row across the On the other hand, salinity reduced seed germination at the
midstem segment, rhombic to subcircular in shape and may higher concentrations of NaCl (50 and 75 mM).
be covered with brownish multicellular hair or trichomes Present review highlights and describes numerous
having wooly appearance. Areoles may be the origin of desirable characters of Opuntia and other cacti, which
flowers and depending on the species they give rise to constitute a major group of tropical plants. This group of
spines and/or glochids (small, detachable barb-tipped plant is a source for the isolation of several bioactive and
bristles) (Anderson 2001). Leaves are cylindrical in shape, nutritionally valuable compounds. Cacti have been used
and the plant is covered with glochids that are unique to regularly to produce health drinks and the extract of cacti is
Opuntia. Glochids are small and are present along the an effective remedy for treating various diseases. Cacti are
adaxial margin of the areoles and under an inconspicuous appropriate for phytoremediation and producing crops on
tuft of yellowish hair. This hair on aging turns brown and is degraded and arid lands. Significant results have been
composed of 100% crystalline cellulose (Pritchard and Hall achieved in cactus tissue culture and molecular markers
Tropical Plant Biol. (2010) 3:136–150 139

studies, which may contribute to germplasm improvement. defense mechanisms leads to oxidative modification in
Cloning and isolation of unique genes from cacti may prove cellular membranes or molecules causing degenerative
useful for engineering into other crops for imparting stress diseases including aging, cancer, arteriosclerosis, and heart
tolerance. diseases (Kehrer 1993; Fridovich 1998; Kourounakis et al.
1998). Antioxidants as in cactus pears can interfere with
oxidation process by scavenging free radicals, chelating
Significance of the Plant free catalytic metals and by acting as oxygen scavengers,
contributing to health maintainance by acting against
Health Benefits of Cacti oxidative damages of ROS and RNS. Sirivardhana and
Jeon (2004) demonstrated antioxidative effects of cacti
Cacti (Opuntia) have been used for centuries as common fruit extract on lipid peroxidation inhibition in oils and
vegetables and medicines by Native Americans to treat a emulsion model systems. Characterization of the antiox-
variety of ailments and disorders (Knishinsky 1971; Kay idative properties proved that cacti antioxidants are heat
1996; Cornett 2000; Tesoriere et al. 2004). The nutritional stable offering a natural source for the stabilization of food
properties of the fresh cladodes have long been known oils (Sirivardhana and Jeon 2004).
(Frati et al. 1990; Fernandez et al. 1992). The bark is a tonic The effects of dried cladode powder of O. ficus-indica
and diuretic, whereas, the fruits have cooling and tonic var. Saboten were investigated on gastric lesion and ulcer
properties. The young buds and the milky juice are models in rats by Lee et al. (2002). Significant inhibition in
astringent. Currently, prickly pear cacti extracts are avail- indomethacin induced gastric lesion was noted without any
able in powder or pill forms and used as a dietary anti ulcer activity.
supplement. Prickly pear are available as food items in Prickly pear has been used in traditional medicines for
North American supermarkets catering to Hispanic custom- its hypoglycemic and hypolipidemic actions. Ancient
ers. Prickly pear cacti are a rich source of flavanoids, Mexican Indian tribes used prickly pear cacti to sooth
including kaempferol and its methyl ether, quercetin and its inflamed insect bites. Prickly pear cacti have been applied
methyl ether, narcissin, dihydrokaempferol (aromadendrin, topically to heal superficial wounds such as cuts and
6), dihydroquercetin, and eriodictyol (Table 2). Flavanoids, scrapes. Reports indicate that in addition to being able to
as those in cactus pear fruit, are known for their affirmative heal minor cuts and wounds, prickly pear cacti may protect
health-benefits (Knishinsky 1971; Iwashina et al. 1984; the immune system and prevent oxidative stress by acting
Wang 1988; Goycoolea and Cárdenas 2003; Tesoriere et al. as scavengers of free radicals (Tesoriere et al. 2004).
2004). Cacti are reported to produce alkaloids, which is a Prickly pear antioxidant actions could protect mammalian
characteristic feature of Cactaceae family (Agurell 1969; cells and organs and slow the aging process, illness and
Rosemberg and Paul 1973; Bruhn and Agurell 1974; Bruhn disease (Kuti 2004). Additionally, prickly pear cacti could
and Lindgren 1976; Gabermann 1978; Ferrigni et al. 1984). lower low density lipoprotein (LDL) and cholesterol levels
Cactus pear also contains antioxidants, primarily pectin, helping to check high blood pressure (Fernandez et al.
carotenes, betalains, ascorbic acid, quercetina and quercetin 1992). Prickly pear cacti are also well known for their anti-
derivatives (Table 2). Mitochondrial β-oxidation of fatty inflammatory effects (Sahelian 2001). The young buds, like
acids is an important source of energy, especially during the milky juice, are astringent and are useful for treating
fasting period and during metabolic stress. Steps of β- dysentery and diarrhea. Concentrated juice in combination
oxidation of long-chain fatty acids are catalyzed by long- with the fruit is reported to be an aphrodisiac and is also
chain 3-hydroxyacyl-CoA dehydrogenase (LCHAD) (a part believed to be of value in spermatorrhoea and gonorrhea.
of multienzyme complex associated with the inner mito- The milky juice is useful as an external application in pains
chondrial membrane) and the trifunctional protein (MTP). and bruises and for the treatment on rheumatism and
Metabolites accumulating in LCHAD and MTP deficien- lumbago. An infusion of the bark (1 in 10) is said to have
cies induce lipid and protein oxidative damage and decrease specific properties in the treatment for diabetes. A decoc-
the antioxidant defenses in rat brain in vitro (Tonin et al. tion of the bark, which contains about 10% of tannin, is
2010). Reactive oxygen species (ROS) and reactive used as an astringent lotion in leucorrhoea. The leaves are
nitrogen species (RNS) are continuously formed as part of heated and applied as a poultice to abscesses to promote
normal cellular functions associated with aerobic respira- suppuration and discharge of pus (Galati et al. 2003). The
tion, pathological process and toxic exposure during cell leaves after they have turned yellow are given in decoction
life. ROS and RNS can damage a wide range of essential with rice as a diaphoretic. The root fibers in the form of
biomolecules including nucleic acids (DNA/RNA), pro- decoction with or without the addition of honey are
teins, lipids and carbohydrates (Halliwell and Gutteridge supposed to resemble sarsaparilla in action and be useful
1990). The imbalance between ROS and antioxidant in treating gonorrhoea. An infusion of the small branches is
140 Tropical Plant Biol. (2010) 3:136–150

Table 2 Active components of Cactus species Chemical Structure Applications Reference

cacti
compound

O. ficus-indica A. Flavanoids Anticancer Kuti, 2004;

O. lindheimeri Kaempferol Lee et al.,

O. streptacantha (aromadendrin, 6), 2003

O. ficus-indica dihydroquercetin,

var. Saboten and eriodictyol

O. ficus-indica . Anticancer Kuti and

Quercetin Antitumour Galloway,

1994

Opuntia dillenii Narcissin Antiprotozoal Qiu et al.,

2007

Opuntia ficus- B. Antioxidants Against Goycoolea

indica var Ia O. Pectin constipation and Cardenas,

ficus-indica var and 2003

IIa O. spp. diarrhoea,

(Blanca I) demulcent,

O. spp (Blanca stabilizer, in

II)O. amylaceaa wound

O. megacanthaa healing

O. steptracanthaa preparations

O. robustaa

Opuntia ficus- Carotenes (α and β) Nutritional Ramadan

indica L supplement and Morsel,

2003

Opuntia ficus- Betalains Anticancer Gentile et al.,

inadica L, agent, radical 2004;

Opuntia stricta, scavenger, Fernández-

Opuntia undulata antioxidant Lopez et al.,

2002

O. ficus-indica Ascorbic acid Antioxidant Kuti, 2004

O. lindheimeri

O. streptacantha
Tropical Plant Biol. (2010) 3:136–150 141

useful in haemoptysis. The tender ends of the hanging roots Cactus pear fruit juice has been used as a health drink.
are given for obstinate vomiting. Saenz and Sepúlveda (2001) reported technological charac-
Prickly pear cacti have shown promise for diabetes teristics and main difficulties in producing high-quality
treatment. Clinical trials indicate that cacti help to stabilize cactus pear juice. The authors stated that acidity, pigments,
blood sugar levels, and are effective for the treatment of aroma, and other components play important roles in cactus
type-II (adult onset) diabetes (Alarcon-Aguilar et al. 2003). pear juice processing. The fruit juice of red cactus pear
Research has shown that cacti can help to reduce the effects contains betalain and green cactus pear contains chlorophyll.
of excessive alcohol consumption if used prior to drinking Heat treatment of the juices affects cactus pear juice color,
alcoholic beverages. (Wiese et al. 2004). but the purple juice is more stable. Lengthy thermal
In Chinese medicine, cactus fruits are considered weak treatments impart an unpleasant taste and undesirable aroma
poisons and are used for treatment of inflammation, pain in the juice. Blends with other fruit juices (e.g. pineapple)
and as detoxification agents for snake bites (Wang 1988). could enhance the quality of the juices. Pineapple juice and
Aqueous extracts of prickly pear cacti were found to inhibit the addition of citric acid decreased the pH of the blends and
in vitro cell growth effectively in several different immor- increased their microbiological stability. Cactus pear juice
talized and cancer cell cultures including ovarian, cervix, has also been used as a good feedstock for fermentation
and bladder cancer cells and suppressed tumor growth in a using Saccharomyces cerevisiae as the fermentation micro-
nude mouse of ovarian cancer models. The mechanism of organism (Turker et al. 2001). Turker and co-workers (2001)
anticancer effect of cactus pear extracts is not understood reported that the fermentation process did not affect the
(Zou et al. 2005). thermo stability of the betalain pigments obtained from the
Prickly pear cacti are well recognized for their wound cactus pear. Fruit pulp of the red cactus pear (O. ficus-indica)
healing properties and anticancer effects (Zou et al. 2005). has been used for ice cream production (El-Samahy et al.
Currently, extensive medical research using prickly pear 2009). The pulp was concentrated up to 30° brix and then
cacti is focused on the management and treatment of type II added at four levels (0, 5, 10 and 15%) to basic ice cream
diabetes, hyperlipidemias, obesity, investigation of antiviral mix. The rheological properties of all ice cream mixes before
properties and use in treatment of benign prostate hyper- and after aging reported the flow of mixes is non-Newtonian
trophy. Galati et al. (2003) studied the effect of O. ficus- with pseudo plastic behavior. The specific gravity and weight
indica (L.) Mill cladodes in the wound-healing process. per gallon of produced ice cream samples increased pulp
Cladodes have been used in Sicilian folk medicine as a additions. Sensory evaluation of the produced ice cream
cicatrizant. In a study by Galati and co-workers (2003), samples showed the sample with 5% cactus was very
a base cream containing 15% lyophilized cladodes and a desirable and close to the control sample (El-Samahy et
commercial ointment was used on wounds produced on the al. 2009).
backs of rats to evaluate healing activity of prickly pear. The relationship between dietary fiber and control of
After 3 days of treatment, both fibers and fibroblasts in the cholesterol for the prevention of illnesses such as diabetes
derma were properly arranged. The dermal vessels were and obesity is well known (Hollingsworth 1996;
reduced in lumen and the keratinocytes showed prolifera- Grijspaardt-Vink 1996; Sloan 1994). Dietary fiber could
tion areas. After 5 days of treatment, the wound healing also contribute to the prevention or treatment of illnesses
process was almost complete and the piliferous bulbs were like gastrointestinal disorders that are associated with
recovered leading to complete epithelization. Evidently, the deficient fiber intake. Saenz (1997) reported the potential
O. ficus-indica treatment accelerated wound healing, use of cactus cladodes as a new source of fiber in human
probably by involving the proliferation and migration of diet. Based on cactus pear (O. boldinghii) cladodes having
the keratinocytes in the healing process (Galati et al. 2003). high fiber content and potential health benefits bakery
products have been produced and evaluated from composite
Food Industry Applications of Cactus Pear flours of wheat (WF) and cactus pear stems (SF) (Moreno-
Álvarez et al. 2009). Cladodes were dried, ground and
The use of natural sweeteners other than sucrose is a sifted to make breads using pure wheat flour (WF, control)
priority area for the food industry. Saenz et al. (1998) and four formulations of WF and SF: I (SF 0%; control), II
demonstrated that cactus fruit can be used as a raw (SF 5%), III (SF 10%), IV (SF 15%) and V (SF 20%). The
material in the creation of natural sweeteners. Cactus proximate composition of breads presented significant
sugar syrup was found to be more acidic than traditional differences (p<0.05) and all formulations were micro-
sugars. Sawaya et al. (1983) attempted to use prickly pear biologically stable. In the farinaceous test, flours composed
fruits in the manufacturing of jam but preferred the of formulations II and III showed the best baking behavior.
combinations of citric acid and tartaric acid (1:1) as an Sensory evaluation showed that formulation III (SF 10%)
acidifying agent. was the most acceptable for its color, odor and flavor, while
142 Tropical Plant Biol. (2010) 3:136–150

the texture of formulation II (SF 5%) was the most Cactus pear cladodes were used as a supplemental feed
acceptable making prickly pear cladode flour a viable for animals including small goats (Fuentes-Rodrίguez
addition for making better bakery products. 1997) and for finishing lambs. Aranda-Osorio et al.
Orange-yellow cactus pear fruit pulp has been used to (2008), proved positive for dry-matter intake (DMI), total
produce a dehydrated cactus pulp sheet and pasteurized live-weight gain (LWG) daily live-weight gain (DLG), feed
cactus pear fruit juices (El-Samahy et al. 2007). A conversion and profitability. Lambs fed with a high ratio of
preliminary study was carried out to evaluate the effects cactus pear were more efficient in converting feed to LWG.
of drying temperature (60 and 70°C) and pulp layer The inclusion of cactus pear reduced feed cost about 65%
thickness (5, 10, and 15 mm) on the dehydration rate of for T30% reducing LWG cost about 64.3% which may
cactus pear pulp. Cactus pear pulp was prepared for offer an alternative to traditional feeds for growing-
dehydration by adding different proportions of sucrose (0, finishing lambs without affecting animal performance while
1, 2, 3, 4, 5, and 10%). The prepared pulps were dehydrated reducing production costs (Fuentes-Rodrίguez 1997).
in an air oven, and then evaluated organoleptically. The
most accepted sheets were those prepared with 2% and 3% Use of Cacti for Degraded Lands
sucrose. For cactus pear juices, pulp was mixed with a
sugar solution at 15° brix and pH 5 in a ratio of (1:1) before Land erosion and degradation coupled with levels of water
thermal treatment. The prepared juice was divided into insufficient for most agriculture systems are serious prob-
three parts, the first was pasteurized directly at 95°C, the lems posing threat to agricultural sustainability.
second was treated with 100 ppm sodium benzoate and then The arid and semi-arid areas of North Africa are
pasteurized at 95°C, and the third was sterilized directly at becoming deserts (Alary et al. 2007). Most research and
121°C. Chemical, microbiological and sensory character- development projects in these areas aim at developing
istics of the three produced juices were evaluated during alternative technologies to reduce land degradation and
storage at room (28± 5°C) and refrigeration (8± 2°C) favor sustainable economic activities. The ‘spineless
temperatures for 6 months. All produced juices were cactus-alley cropping system’ is an interesting approach
microbiologically stable for the storage period and the for the low rainfall areas of North Africa since it limits land
pasteurized juices were best organoleptically. degradation by the use of a perennial crop for the
Nopalitos, the prepared pads of cladodes, are very production of cheap and drought resistant sources of feed
perishable with a storage life of only 1 day at room and biomass in the inter rowspaces between food crops. A
temperature and 6 days when packaged in polyethylene bio-economic model has been developed to identify the
bags and stored at 5°C (Guevara et al. 2001). The quality of conditions for applying the ‘spineless cactus-alley cropping
nopalitos (O. ficus-indica) during cold storage suffers system’ in an agro-pastoral community of Central Tunisia.
weight loss and chilling injury. These disorders have been Findings suggested that extension services will play a
reduced in several fruits and vegetables, including cladodes crucial role in creating awareness among farmers of the
or nopalinas, by covering the product with edible coatings. impact of this technology on yields and income diversifi-
Rodrίguez-Félix et al. (2007) reported the effect of edible cation (Alary et al. 2007).
coatings on nopalitos of Opuntia sp. during cold storage. The
effect of application of two edible coatings (Semperfresh® Use of Cacti for Phytoremediation
and another formulation based on carboxymethylcellulose,
CMC, named Wax 1) on the quality of nopalitos of two Phytoremediation is an effective technology for cleaning up
cultivars stored at 5°C and 10°C for 30 days was evaluated. of the environmental hazardous pollutants. Since cacti are
Guevara et al. (2001) demonstrated that O2 concentration so sturdy they could be ideal plants for decreasing the load
decreased up to 8.6 kPa and CO2 concentration increased up of toxic pollutants in the environment. Even if cacti absorb
to 6.9 kPa could significantly decrease quality deterioration and store toxic heavy metals and other organic pollutants in
and thus increase storage life when nopalitos were stored at their cell systems; they would normally be safe to use since
5°C for up to 30 days. grazing animals will generally avoid eating non-processed
Cerezal and Duarte (2004) studied the effect of a cacti; so that as a non edible plant, cacti would pose little
combination of common food additives as sodium bisul- danger of entering into the food chain. Cacti would also be
phite (0, 50 and 100 ppm), phosphoric acid (50% v/v), efficient scavagers of pollutants since they have very
citric acid (50% v/v), ascorbic acid (500 ppm), calcium shallow and massive roots that can spread out widely near
chloride (120 ppm) and potassium sorbate (1,000 ppm), on to the soil surface. For example, a young Saguaro cactus
the sensory quality and storage of nopalitos canned in glass only 12 cm tall had a root system covering an area 2 m in
jars. Best preservation by one of the combinations allowed diameter and no roots more than 10 cm deep, thus capable
for wide distribution beyond the area of production. of effectively mining a large part of any contaminated soil.
Tropical Plant Biol. (2010) 3:136–150 143

In vitro cultures of cacti were used to test the effect of Cactus pear cladodes have been used in many building
degradation and/or biotransformation of dye precursors by construction applications in Mexico, since ancient times.
plants (Golan-Goldhirsh et al. 2004). Cacti were shown to New applications of cladodes in hydraulic cementation
produce protective compounds, such as alkaloids, from a materials was found based on knowledge of that former
family of aromatic hydrocarbons (Oliveira and Da Machado use. Hernández-Zaragoza et al. (2008) found that mortar
2003). Thus, it was hypothesized that the metabolic made with cladodes as an additive had a greater compres-
pathways and enzymes related to biotransformation of sive strength than mortars made without the additive and
xenobiotic aromatics would more likely to be found in suggested that cactus pear acts as a retardant of slowing of
these plants (Golan-Goldhirsh et al. 2004). the setting time in mortar manufacturing process.
Synthesis of metal-binding peptides, known as phyto- An electrochemical cell was fabricated by Deshpande
chelatins, is one defense mechanism used by higher plants and Joshi (1994) from cactus cladodes acting as an
in exposure to non-essential metal/metalloids or toxic electrolyte. The discharge characteristics revealed that,
concentrations of essential elements. Phytochelatins syn- at a current drain of 100 φA, the cell gave an optimum
thesis has been evaluated in cladodes of O. ficus in relation energy density of 175 mWh/kg. The power generated by
to plant and soil levels of Cd, Pb, Cu and Ag (Figueroa et these cells was sufficient to run a piezoelectric buzzer
al. 2007). The feasibility of genetic engineering to assure and a LCD calculator for a few hours. This work opens
higher rates of phytochelatin induction has been explored up a new interdisciplinary area for physicists, botanists
for phytoremediation purposes (Wojas et al. 2008; Guo et and electro-chemists.
al. 2008; Couseloa et al. 2010).

Additional Potential Benefits Chemical and Biochemical Composition of Cactus

Researchers tested different types of milk for clotting time Cactus pear fruit contains about 85% water, 11–14% ferment-
caused by extracts from cacti fruits (Texeira et al. 2000). able sugar, 1.8% crude fiber, 0.1% lipids, 0.21–1.5% protein,
Screening was done in fruit extracts of cacti for various and 0.4–1.6% (w/w) ash (Lakshminarayana et al. 1979;
enzymes such as arylamidases, lipases, proteinases and Hernandez 1980; Joubert 1993). It also contains calcium
glucosidases. Amongst these enzymes, proteinases are the and vitamin C up to 60 and 30 mg/100 g and vitamin A
most important plant enzymes used in food, pharmaceuti- 50 IU. These studies were confirmed by Salim et al. (2009).
cal, detergent, leather and wood industries (Mantell and Mc Based on these data, Turker et al. (2001) used cactus pear
Kee 1985). Several bioassays by Han et al. (2001) indicated fruit juice as a feedstock for yeast fermentation.
inhibitory activity of monoamine oxidase from methanolic Cladodes of prickly pear contain minerals (mainly
extracts of cacti fruits. Four compounds isolated were tested potassium, magnesium, calcium and iron), vitamin A in
for in vitro monoamine oxidase inhibitory activities and the the form of β-carotene at levels comparable to those in
authors concluded that industrial applications for these spinach and high vitamin C. Cladodes are reported to
molecules may be found through further investigations contain all dietary essential and nonessential amino acids in
(Han et al. 2001). significant proportion (Kazemini 1994) (Table 3). Nitrogen

Table 3 Free L amino acids and

vitamins in Opuntia ficus-indica Amino acida mg per 100g fresh weight Amino acida mg per 100g fresh weight
cladodes
Alanine 0.6 Phenylalanine 1.7
Arginine 2.4 Serine 3.2
Asparagine 1.5 Threonine 2.0
Asparaginic acid 2.1 Tryosine 0.7
Glutamic acid 2.6 Tryptophane 0.5
Glutamine 17.3 Valine 3.7
Glycine 0.5 Vitaminb per 100 g fresh weight
a Histidine 2.0 β-Carotene 11.3–53.5 μg
Source: Bruckner and
Westhauser 2003 Isoleucine 1.9 Niacine 0.46 mg
b
Source: Rodrίguez-Félix and Leucine 1.3 Riboflavine 0.60 mg
Cantwell 1988; Pimienta-Barrios Lysine 2.5 Thiamine 0.14 mg
1993; Teles et al. 1994; Guevara Methionine 1.4 Total vitamin C 7–22 mg
et al. 2001
144 Tropical Plant Biol. (2010) 3:136–150

tends to be more concentrated in young cladodes than in Table 4 Chemical composition of fruit pulp and seeds of Opuntia
ficus-indica
mature ones (Nobel 1983). Cultivated cactus pear plants
tend to accumulate more calcium in their cladodes than Component Fruit pulp (fresh Seeds (dry
most cultivated cacti species. Calcium accumulation weight basis) weight basis)
increases with age of the tissue. Two cactaceous powders
Watera 85.60 5.3
were shown by X-ray diffractometry, spectroscopic, thermic,
Proteina 0.21 16.6
nitrogen and water adsorption methods to contain calcium
Lipida 0.12 17.2
oxalate and consist of globular or cubosome particles (Corro-
Fibera 0.02 49.6
Hernández and Elizalde-González 2003). It was suggested
that the complex nature of cacti powders might be used to Pectina 0.19 –
adsorb a range of chemical substances varying in size and Asha 0.44 3.0
polarity. Tovar-Puente et al. (2007) demonstrated differences Vitamin Cb 22 –
in the type of calcium oxalate crystals among Cactaceae β carotenec tr –
subfamilies and suggested the possibility of using the Cab 28 16
characteristic type of calcium oxalate crystals to classify Mgb 28 75
species of this family. In addition, differences in chemical Kb 161 163
composition of cladodes among species of cactus pear have Nab 0.8 68
been reported (Magallanes-Quintanar et al. 2004). Pb 15.4 152
Physical and spectral property methods have been used Feb 1.5 9
to identify eight flavonoid compounds synthesized in cacti a
%
cladodes and fruits (Oliveira and Da Machado 2003; Moffat b
mg per 100 g
et al. 1986; Lee et al. 2003). Fernández-López et al. (2002) c
IU
investigated the presence of betalain pigments in three
tr traces
species of cactus fruits. Identity of this pigment was
Source: Sawaya et al. 1983; Salim et al. 2009
confirmed by mass spectral methods, but the authors failed
to detect the presence of neobetanin. Some of the
components of cacti fruit pulp and seeds are summarized
in Table 4. netic responses of O. ficus-indica on Murashige and Skoog
(MS) medium (Murashige and Skoog 1962). This work
demonstrated that BA at the highest concentration (5.0 mg/l)
Plant Biotechnology Research on Cacti gave the highest number of shoot per explant (26.5±1.74)
after 3 months of culture. The combinations of NAA and the
Methods for propagation of succulent plants are by seeds, cytokinins BA and Kin did not increase the number of shoots
cuttings, or grafting (Krulik 1980). Cacti can be propagated per explant. The effect of two different media on in vitro
from seeds, but seed propagation has problems of genetic growth of O. ficus-indica explants was studied by Aliyu and
segregation, slow growth, and a long juvenile stage Mustapha (2007). Medium A contained MS basal salt and
compared to asexually propagated material (Mohamed- vitamins supplemented with 5% sucrose and 1% BA;
Yasseen et al. 1995). Tissue culture can be an excellent tool medium B contained vitamins and MS salt supplemented
for cacti micropropagation and has been used on species in with 3% sucrose, 1.25 mg/l BA, and 0.25 mg/l IAA. There
the genera Cereus, Equinocereus, Ferocactus, Mammil- were no significant differences in shoot height or days for
laria, and Opuntia (Escobar et al. 1986; Machado and shoot emergence between the two media over a period of
Prioli 1996; Pérez-Molphe et al. 1998; Juárez and Passera 30 days.
2002). Malda et al. (1999) demonstrated in vitro culture as Estrada-Luna et al. (2008) established conditions for
a potential method for the conservation of endangered micropropagation of the ornamental prickly pear cactus O.
plants, including cacti, that possess crassulacean acid lanigera Salm-Dyck through axillary shoot development
metabolism. Massive in vitro production of new propagules from isolated areoles and studied the effects of sprayed
resulted in regenerated plants for two endangered cacti, gibberelic acid (GA3) after transplantation to ex vitro
Obregonia denegrii Fric and Coryphantha minima Baird. conditions. Escobar et al. (1986) studied the effect of BA
Khalafalla et al. (2007) demonstrated the micropropaga- on axillary bud development in O. amyclaea and reported
tion of O. ficus-indica to combat desertification in arid and the best response at 10 μM. Llamoca-Zárate et al. (1999a)
semi arid regions. These workers studied the effects of proposed 2.2 μM BA while Mohamed-Yasseen et al. (1995)
benzyladenine (BA) and kinetin (Kin) alone or in combi- reported 8.8 μM BA as optimal for O. ficus-indica
nation with 1-naphtalenacetic acid (NAA) on morphoge- micropropagation.
Tropical Plant Biol. (2010) 3:136–150 145

Many cacti produce an excess of auxin under in vitro growth was observed on MS medium supplemented with
culture conditions, and this stimulates callus production 0.9 μM 6-furfurylaminopurine (FAP), 2.3 μM 2,4-D,
(Clayton et al. 1990). Garcίa-Saucedo et al. (2005) 1.0 μM 4-amino 3,5,6-trichloropicolinic acid, 400 mg/l casein
developed a micropropagation system for three Opuntia hydrolysate and 3% sucrose. The same medium without agar
lines used as vegetables in Mexico. Balch et al. (1998) was used for establishing cell suspensions.
developed a micropropagation system for 21 species of Callus tissue of Cereus peruvianus was established and
Mexican cacti (Astrophytum, Cephalocereus, Coryphantha, used for alkaloid production (Oliveira and Da Machado
Echinocactus, Echinocereus, Echinofossulocactus, Ferocactus, 2003). Morphologically undifferentiated callus cells of
Mammillaria, Nyctocereus, Stenocactus) using explants from Cereus peruvianus cultured in the original medium and in
seedlings germinated in vitro or shoot segments of juvenile medium supplemented with tyrosine were used as the
2–3 year old greenhouse plants. Several workers have alkaloid source. Culture medium supplemented with tyro-
achieved success in micropropagation of species including sine showed higher levels of alkaloids than the original
Astrophytum myriostigma, Mammillaria carmenae, M. medium. Alkaloid levels were almost twice as high in
prolifera, and Trichocereus spachianus (Vyskot and Jara callus tissues as in shoots of C. peruvianus plants. The ratio
1984), Echinocereus engelmanii, E. pectinatus, Ferocactus of alkaloid concentration between mature plant shoots and
covillei and F. wislizenii (Ault and Blackmon 1985), F. morphologically undifferentiated cells of callus tissue was
acanthodes (Ault and Blackmon 1987), Mammillaria san- 1:1.7. Since increased alkaloid production can be induced
angelensis (Martínez-Vázquez and Rubluo 1989), Medi- by factors such as tyrosine, the levels of tyrosine or other
ocactus coccineus (Infante 1992) and Cereus peruvianus conditions of the culture medium may be responsible for
(Machado and Prioli 1996), Turbinicarpus laui (Rosas et al. the higher levels of alkaloid production by the callus
2001). The effect of different growth regulators on the tissues.
morphogenesis of O. polycantha, a non-edible Opuntia, has
been determined by Mauseth and Halperin (1975) and
Mauseth (1976). Genetic Engineering and Molecular Biology Research
De Medeiros et al. (2006) worked to establish a protocol on Cactus
for in vitro culture and plant regeneration of Notocactus
magnificus, an ornamental species called the blue cactus Isolation of relatively pure DNA from cacti has proven to
that is native to Brazil. These workers observed callus be notoriously difficult because cacti contain high amounts
formation in Notocactus magnificus when the explants were of polysaccharides and secondary metabolites that form
cultured on MS medium supplemented with sucrose at 2% insoluble complexes with nucleic acids during extraction
(w/v), 2,4-D 0.5 μM, BA 4.4 μM, thiamine HCl 0.4 mg/l (Guillemaut and Marechal-Drouard 1992). The cactus
and i-inositol 100 mg/l. The highest number of shoots polysaccharides, visually apparent by their viscous, glue-
was obtained when MS medium was supplemented with like texture, make the extracted DNA difficult to pipette
BA 22.2 μM, sucrose 3% (w/v) and agar 0.6% (w/v). and hard to amplify by the polymerase chain reaction
In vitro spontaneous rooting of shoots was observed (PCR) (Porebski et al. 1997). De la Cruz et al. (1997)
after 8 months on MS medium. In vitro rooted shoots modified their earlier protocol (De la Cruz et al. 1995) to
developed into normal plants under glasshouse culture reduce the problems of polysaccharide contamination of
conditions. cactus DNA by minimizing the amount of tissue extracted.
Advantages of the propagation systems outlined above A modified CTAB method for DNA isolation from
include (1) the time required for production of cacti shoots by epiphytic/climbing cacti of the genera Hylocereus and
microprogagation is quite short compared to the time required Selenicereus was reported by Tel-Zur et al. (1999). In this
for germination of seedlings (Ault and Blackmon 1985) and procedure, roots were the tissue source for DNA due to
(2) micropropagated cacti from axillary buds are genetically roots having lower levels of polysaccharides relative to
stable compared to other methods of in vivo propagation other tissues and thus resulting in the DNA extracts having
(Vyskot and Jara 1984; Machado and Prioli 1996). lower viscosity. Later cactus DNA extraction techniques
Relatively little work has been reported on the included the use of DNAzol for the extraction of total DNA
establishment of callus and cell suspension cultures of and use of and TRIazol for RNA from cactus pear cultivars
cacti. Llamoca-Zárate et al. (1999b) developed a method (Esparza et al. 2004). DNA prepared this way was
for establishment of callus and cell suspension culture of consistently amplified in RAPD reactions to give reproduc-
O. ficus-indica. Friable callus cultures were initiated when ible profiles. Molecular analysis of cactus pear cultivars
cotyledons and hypocotyls of O. ficus-indica were used as RAPD markers showed no differences among cactus fruit
explants. Explants from cotyledons produced significantly cultivar, however, several cactus pear proteins reacted to
more callus than those from hypocotyls. Optimum callus anti-bean and anti-HeLa polyvalent antibodies showing
146 Tropical Plant Biol. (2010) 3:136–150

conserved epitopes. The length of the cDNA generated characterization of plants in the germplasm banks reduced
from specific Hsp70 RNA primers was ≈300 bp from all the time for selection of elite genotypes. Morphological
cactus pear cultivars which confirmed gene identity. description may not be sufficient to separate related
Analysis of the genetic variability is of paramount accessions or species, as in the case of some cactus pear
importance to group genotypes for germplasm conservation (Opuntia spp.) native to Mexico. The reader is reminded
and characterization for more adequate management and that the use of DNA markers has been limited by large
optimized use of collections. Souto Alves et al. (2009) quantities of polysaccharides extracted with the DNA
attempted to evaluate the level of genetic diversity in cactus hindering subsequent attempts at PCR analyses. Seed
clones by molecular markers to gather information for DNA is cleaner by lacking these compounds, so character-
future breeding programs. Eleven clones of the Breeding ization of seeds can contribute molecular data related to
program of Empresa Pernambucana de Pesquisa Agro- genetic variability.
pecuária—IPA, state of Pernambuco, Brazil, were evaluated Expression of ripening-related genes in fruit of prickly
by ISSR (Inter Simple Sequence Repeat) markers. The pear (Opuntia sp.) was studied by Collazo-Siques et al.
average linkage (UPGMA; Unweighted Pair Group Method (2003). cDNA fragments were cloned and partially charac-
with Arithmetic mean) clustering revealed low genetic terized to gain an understanding of the possible role of
diversity in the cactus pear clones, suggesting that genetic ethylene in nonclimacteric fruit ripening. These cloned
differences among them are minimal. The ISSR markers genes showed high homologies with 1-aminocyclopropane-
however did prove useful for developing polymorphic 1-carboxylate (ACC) synthase and ACC oxidase from other
cultivar-specific markers in cactus pear. plants. The corresponding genes were accordingly named
Wang et al. (1998) analyzed eight Opuntia cacti opaccs-1 and opacco-1, after Opuntia ACC synthase-1 and
consisting of five fruit varieties from Mexico and Chile, Opuntia ACC oxidase-1, respectively. Southern analysis
two ornamental Texas accessions, and one vegetable suggested the presence of at least one copy of each gene, as
accession from Mexico. Phenotypic and molecular analyses well as other related homologous sequences in the Opuntia
distinguished ornamental, vegetative, and fruit market genome. Northern analysis of the opaccs-1 gene showed
accessions of O. ellisiana, O. lindheimerii, O. cochinelli- enhanced expression in ripening fruit tissues, whereas,
fera, O. hyptiacantha, and O. ficus-indica from each other, opacco-1 expression was highly induced in ripened tissues
and suggested significant differences among accessions of compared to green fruits and mature cladodes. These results
different market classes. Differences among fruit cultivars are in agreement with an active metabolic role of ethylene
were smaller and the results demonstrated the potential for during non climacteric prickly pear fruit ripening.
use of RAPD (Random Amplified Polymorphic DNA) A system for genetic transformation of an elite prickly
markers to distinguish cacti from one another. pear cactus (O. ficus-indica L., cultivar Villa Nueva) by
RAPD markers detected DNA polymorphism in callus Agrobacterium tumefaciens was developed by Silos-Espino
tissue of cacti for classification of Opuntia cacti. DNA et al. (2006). Direct bacterial infection of areole meriste-
polymorphisms in callus tissues of Cereus peruvianus matic tissue was accomplished with a hypodermic syringe.
Millwere detected by using RAPD markers (Mangolin et Transgenic plants were obtained by selection on media
al. 2002). There was a higher level of genetic variability in containing 100 mg/l kanamycin. Transient and stable GUS
callus tissue maintained with the highest kinetin (8.0 mg/l) (β-glucuronidase) activities were observed on kanamycin-
versus 2, 4-D (4.0 mg/l) concentration. RAPD analysis was resistant shoots and regenerated plants. PCR and Southern
also used to estimate the genetic relationship among five blot analysis confirmed that the regenerated plants growing
Hylocereus and nine Selenicereus species (Tel-Zur et al. under selection were genetically transformed. The trans-
2004). The principal coordinate analysis (PCO) based on gene copy number in the genome ranged from two to six.
RAPD data clearly distinguished between three groups, The transformation frequency obtained was 3.2%. The
namely, Hylocereus species, S. megalanthus and the rest of method described by Silos-Espino et al. (2006) may be
the Selenicereus species studied. PCO thus strongly useful for routine transformation and introduction of
supported the notion that the tetraploid S. megalanthus is agronomically important genes in prickly pear cactus.
an exception among the Selenicereus group and that the
origin of this species was allopolyploid, rather than
autopolyploid. Prospects
Twenty-two cactus pear varieties were characterized with
RAPD and ISSR markers using DNA from seeds of Society must employ all available tools to meet the
Opuntia spp. (Luna-Paez et al. 2007). The RAPD and challenges of increasing food supplies for nutritional and
ISSR marker profiles revealed differences between the health security. One such tool can be the rich opportunities
varieties and species under study. Molecular marker that exist for food, beverages, drugs, and genes from
Tropical Plant Biol. (2010) 3:136–150 147

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