Pineapple
Geo Coppens d'Eeckenbrugge & Freddy Leal (January 2001)
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Pineapple is among the most ancient American crops. It was domesticated and dispersed several thousands years ago by Amerindian peoples in the basins of the Orinoco and Amazon. This is indicated by the thorough knowledge of the plant that they accumulated, as well as its wide distribution and cultivation in pre-Columbian times. The native names nana and anana are the most common in South America. They were maintained in most languages of the countries where pineapple was introduced. The spanish "piña" and the english "pineapple" came from the comparison with the exotic pine cone. The Brazilian name "abacaxi" is derived from the guarani word for the maize ear.

In addition to the fresh fruit, pineapple had such varied uses as wine making, fiber production, emmenagogue, abortifacient, antiamoebic, vermifuge, correction of stomach disorders, and poisoning of arrow points. Modern studies have confirmed the emmenagogue and abortifacient effects of green pineapple.

Economic importance

Pineapple is the most important American fruit, and the third most important tropical fruit, after banana and mango (citrus fruits being produced mainly in subtropical areas). It is cultivated in all tropical and subtropical countries. Annual world production has tripled during the last 30 years and now exceeds 12 million tonnes. Most of this production (70%) is locally consumed as a fresh fruit. World trade mainly consists of processed products; of which 80% canned slices (1,065,000 t) and juice (215,000 t) is supplied by Thailand and the Philippines. The fresh fruit market (680,000 t) is dominated by the Philippines, Costa Rica, and Côte d'Ivoire which supplies 60 % of the European market, the leading importer with more than 226,000 t (FAO, 1994; Loeillet, 1996b). Thailand is the leading pineapple producer (16% of world production), followed by Brazil (13%) and the Philippines (13%), India (8.9%), and China (7.3%). Important American producers are Colombia (329,300t), Mexico (301,407), Costa Rica (260,000), U.S.A. (Hawaii and Puerto Rico; 301,000 t). About 70% of the world production and 96% of the pineapple used by the processing industries comes from one cultivar, ‘Smooth Cayenne’. The cultivar Queen has occupied a small specific niches of high quality and expensive fresh fruit. Recently, the pineapple industry has responded to the increasing demand for fresh pineapple in the temperate markets with the introduction of more attractive cultivars as ‘MD2’, also named ‘Golden Ripe’.

Portions of the pineapple plant and processing wastes in the form of shell and core materials, and centrifuge solids from juice production have been used as animal feeds.

Pineapple fiber is considered to be more delicate in texture than any other vegetal fiber. About 60 cm long, white and creamy and lustrous as silk, it easily takes and retain dyes. Numerous tests in Brazil, Florida, India and the Philippines have shown its exceptional resistance to salt, vapor, and traction. A small cottage industry still exists in the Philippines for high quality clothes from pineapple fiber. Pineapple fiber has also been processed into a paper of remarkable thinness, smoothness, and pliability.

Bromelain was originally only extracted from Hawaiian pineapple stems but now is manufactured in Taiwan, Brazil and Puerto Rico. The variability in the commercially produced product and its multiple active ingredients has limited successful development. Pineapple bromelain has been used commercially as meat tenderizing enzyme and a nutraceutical. Bromelain has shown activity with the interference of growth of malignant cells; the inhibition of platelet aggregation; fibrinolytic action; anti-inflammatory processes; and skin debridement.

Morphology

Ananas comosus is a herbaceous perennial whose terminal inflorescence gives origin to a multiple fruit (sorose). After maturation of the first fruit, the plant develops new shoots from axillary buds, so producing new growth axes capable of a new fruit production. The same plant may thus give a sequence of various production cycles. This vegetative reproduction is also dominant in wild pineapples where, in addition to lateral shoots, the crown and slips contribute to propagation as they resume rapid growth at fruit maturity. The long peduncle then bends because of their weight, the crowns and slips reach the ground and may root.

Vegetative propagules are classified according to their position on the plant. Suckers appear on the earthed part of the stem. Stem shoots, which appear on the aerial part, are more frequent. Slips appear on the peduncle. They are often grouped near the base of the fruit. Sometimes, they are produced from the basal eyes of the fruit (collar-of-slips). Crown can be also used for planting. Some plants may lack a crown or, on the contrary, present multiple crowns. Also, crownlets may grow at the base of the main crown or from some of the upper fruitlets.

The adult plant is 1 to 2 m high, and 1 to 2 m wide. The main morphological structures to be distinguished are the stem, the leaves, the peduncle, the multiple fruit or syncarp, the crown, the shoots and the roots. The sword-like concave leaves allow the plant to collect water in the rosette, where it can be absorbed by the aerial roots present along the stem or through the epidermis of their sheath. Leaf color ranges from light or dark green to dark red or purple according to the cultivar and conditions. Both sides are covered by peltate trichomes, particularly the abaxial one, which is densely furfuraceous and silvery, thus having a high reflectance. In addition, the trichomes function as one-way valves, playing an important role in the plant capacity to improve and maintain its water status, so their main role is to protect the plant from excessive transpiration and intense sunlight. More generally, the thick cuticle, the water-storage tissue, the disposition of the stomata, the trichomes, as well as the CAM metabolism, all contribute to the remarkable water economy of pineapple. The leaf margins are usually thorny, however certain cultivars are partially or totally inermis. In some smooth cultivars, the lower epidermis is folded over the leaf edge and extended over the upper surface, so producing a narrow silvery stripe, a trait called 'piping'. Pineapple leaves are highly fibrous.

Primary roots are only found in very young seedlings. They die rapidly and are replaced by the adventitious roots. These form a short and compact system at the stem base, with numerous strong roots and limited branching.

The peduncle and inflorescence develop from the apical meristem. The stage of inflorescence emergence is called "red heart" because of the five to seven reddish peduncle bracts at its base. These bracts are shorter and narrower than the ordinary leaves. The peduncle elongates after flower formation. Its length varies widely with the botanical varieties or even cultivars. In addition to its bracts, it bears, in many cultivars, a variable number of slips (up to a dozen or more) which can be positioned more or less regularly between the stem and the fruit, at the axis of the peduncle bracts, or grouped just beneath the fruit. These slips may constitute an appreciated source of planting material in extensive cultivation systems.

The inflorescence consists of less than 50 (in some wild clones) to more than 200 (in some cultivars) individual flowers; it is capped by a crown, composed of numerous short leaves (up to 150) on a short stem. Flowers are hermaphroditic and trimerous. The abundant nectar production and the tubular corolla are particularly adapted to hummingbird pollination.

The entire blossom develops parthenocarpically into a berry-like fruitlet. In the cultivated pineapple, growth from blossoming inflorescence to mature fruit results in a twenty-fold increase in weight. The edible part of the fruit consists chiefly of the ovaries and of the bases of sepals and bracts and of the cortex of the axis. The fruit shell is mainly composed of sepal and bract tissues and the apices of the ovaries.

Reproductive biology and cultivation cycle

Pineapple reproduction is predominantly asexual. However sexual reproduction is still functional and can be exploited in breeding. Allogamy is mainly ensured by a self-incompatibility system. Thus fertilization is a rare event under monoclonal cultivation and the fruit develops parthenocarpically.

Pineapple is considered a short-day plant. The susceptibility of the plant to relatively long and cool nights depends on the cultivar, the plant size and age, and the planting material. Flowering may be provoked by other kinds of stress or, artificially, by hormonal induction with ethylene–producing chemicals. Depending on the local climatic conditions, the time interval between planting and first harvest varies from 12 months in equatorial areas to 36 months in subtropical areas. The generative phase, between floral induction and harvest lasts from 140-150 days in Ivory Coast to 280-300 days in Queensland or South Africa.

In most commercial plantings, the plants are not allowed to produce more than two to three crops, due to a reduction in fruit size and uniformity. Then a new plantation must be regularly established. This may be done with the same lateral shoots of the preceding crop, or with other vegetative propagules such as the fruit crown or, in many cultivars, slips produced along the peduncle.

Main commercial cultivars

'Cayenne Lisse' or 'Smooth Cayenne', ('Maipuri', 'Kew', 'Sarawak', 'Esmeralda', 'Claire', 'Typhoon', 'Saint Michel'):

The ovoid medium-sized fruit (1.5 to 2.5 kg) of 'Smooth Cayenne' is held on a short and strong peduncle. It ripens progressively, turning yellow from the base to the top, which is reflected in a strong internal maturity gradient too. The flesh is pale yellow, soft and juicy, with considerable variation in sugar (from 13° to 19°Brix) and acidity, depending on environmental conditions, mainly rain and temperature, and low ascorbic acid content. 'Smooth Cayenne' juice is light yellow, turbid, with a high sugar content. The plant is a poor producer of shoots and slips. The production cycle is longer than for most other cultivars. 'Smooth Cayenne' is sensitive to many known pests (fruit borers, mites, symphillids, nematodes) and diseases (mealybug wilt, fusariosis, fruitlet core rot, butt rot), and to internal browning. However, it is considered tolerant to Phytophthora sp. and resistant to fruit collapse, caused by Erwinia chrysanthemi Burkbolder.

‘Cayenne Baronne de Rothschild’: spiny mutant of 'Smooth Cayenne'.

'Singapore Spanish' ('Singapore Canning', 'Ruby', 'Red Pine', 'Nanas Merah', 'Nangka', 'Gandol', 'Betek', and 'Masmerah'):

'Singapore Spanish' is second to ‘Smooth Cayenne’ in importance for canning. It is cultivated in South Asia, particularly in Malaysia. Fruits are small (around 1 kg, heavier in 'Masmerah'), cylindrical, dark purple turning copper orange when ripening, with a golden yellow flesh. Sugar and acidity are low (10 to 12°Brix) and the taste is poor. However, they give a juice of good color and quality. Leaf spininess is highly variable from clone to clone, from complete spininess to very few spines. The plant is vigorous and produces many slips (about two to six) and shoots. Multiple crowns are frequent. 'Singapore Spanish' is tolerant to Phytophthora. It shows severe chlorosis when exposed to high manganese concentration in the soil. It is also sensitive to fruit collapse and to nematodes.

‘Selangor Green’ ('Green Pine', 'Green Spanish', 'Nanas Hijau', 'Selassie'): derived from a mutation suppressing anthocyanins in all the organs of 'Singapore Spanish'. The leaves and inflorescence are uniformly green, the petals pale yellow, and the fruit yellow at maturity. Other traits have not been altered.

‘Queen’ ('Victoria', 'Mauritius', ‘Moris’, 'Malacca', 'Red Ceylon', 'Buitenzorg', 'Ripley Queen', 'Alexandra', 'McGregor'):

This cultivar is widely distributed, more particularly in the Southern Hemisphere, for the fresh fruit market. The small and very spiny plant gives a small fruit (0.5 to 1 kg), with a full yellow shell and small prominent eyes. The golden yellow pulp is crispy and sweet (14 to 18°Brix), with an excellent flavor and long shelf life. 'Queen' is a robust cultivar, showing more general tolerance to stress, pests and diseases than 'Smooth Cayenne'. On the other hand, it is susceptible to Phytophthora and fruit collapse, and highly susceptible to chilling and internal browning, particularly if harvested before maturity, and to fruitlet core rot and butt rot.

'Española Roja' ('Red Spanish', 'Black Spanish', 'Key Largo', Havannah' or 'Habana', 'Cubana', 'Cowboy', 'Bull Head', 'Cumanesa', 'Native Philippine Red'):

Widely cultivated in the Caribbean basin, 'Española Roja' gives a medium, barrel-shaped, orange fruit (1.2 to 2 kg). The flesh is firm, pale, aromatic, and sweet, with moderate sugar content (around 12°Brix) but low acidity. The medium-sized plant is spiny or half-spiny. Smooth clones have been selected. The plant regularly produces some slips and suckers. 'Española Roja' is vigorous and tolerant to high temperature, drought, internal browning, butt rot, wilt, and Phytophthora, but not to high manganese in the soil and nematodes. It is highly susceptible to the South American lepidopter Strymon basilides (Geyer

 

'Pérola' ('Pernambuco', 'Branco de Pernambuco', 'Abacaxi', 'Abakka', 'Eleuthera', 'Jupi'):

This is the main Brazilian cultivar. The fruit is small to medium (0.9 to 1.6 kg), ovoid (when small) to conical, and green, with a little yellow in the centre of the mature eyes. The flesh is soft, white and juicy with an exquisite aroma. Its sugar (13° to 16°Brix) and ascorbic acid contents are high. As with most conical fruits, it shows a strong gradient of maturation from base to top. The plant is medium size and vigorous, with dark green, erect and spiny leaves. In some strains, long and erect basal slips surround the fruit. 'Pérola' is a very robust cultivar, showing resistance to Phytophthora and tolerance to drought, mealybug wilt, and nematodes. However, it is threatened because of its high susceptibility to fusariosis that severely affects the Brazilian production.


'Perolera' ('Lebrija', 'Motilona', 'Capachera', 'Tachirense'):

This is an important cultivar of the northeastern Andes of Colombia and Venezuela. The fruit is large (1.5 to 3 kg), yellow to orange, with an irregular cylindrical shape, borne on a long peduncle, which makes it susceptible to fruit lodging and sunburn, particularly in hot lowlands. The flesh is pale yellow to yellow, firm and sweet (although Brix is only around 12°). Numerous crownlets protrude from the base of the crown and the upper eyes. Slips are also numerous (from 4 to 11). The leaves are completely smooth because the lower epidermis is folded over the leaf edge, a trait named "piping". 'Perolera' is resistant to fusariosis and susceptible to fruitlet core rot and to the fly Melanoloma canopilosum Hendel.

‘Manzana’ ('Bumanguesa') is cultivated in the same area, as well as in the Western Andes of Colombia. It is said to be a sport from 'Perolera', from which it differs in the more regular, globular to cylindrical, shape, and in the large flat eyes of its fruit, which is of an attractive dark to bright red colour when produced in tropical highlands. It is susceptible to the nematode Pratylenchus neglectus but tolerant to Meloidogyne incognita Kofoid & White. 

Other traditional cultivars of lesser regional importance are sometimes described. 'Cabezona' is a vigorous spiny triploid producing yellow to orange fruits of more than 3 kg with white pulp. 'Monte Lirio' ('Cambray', 'Milagreña') is a smooth ("piping") cultivar with a mid-size fruit and sweet white pulp, found from Mexico to Ecuador. 'Black Antigua' is an old spiny cultivar producing a small to medium fruit with a dark green shell turning yellow to orange at maturity. Its golden yellow pulp is firm and delicious. The Peruvian 'Samba' and 'Roja Trujillana' produce red fruits and show resistance to Strymon basilides (Bello et al., 1997b). Certain Amazonian cultivars as 'Gigante de Tarauacá' or Cabeça de Onça' yield enormous fruits up to 15 kg. Others are distinguished by an original red or purple color of their fruit, as the Amazonian ‘Cabeça de Arara’ and 'Roxo de Tefé', the Trinidadian 'Mundo Nuevo Red', and the Venezuelan 'Morada'. Further evaluation of the germplasm recently collected in the basins of the Amazon and the Orinoco will provide many new traits of interest for varietal diversification.


MD2' ('Golden Ripe', 'Extra Sweet', ‘Maya Gold’):

‘MD2’ is a hybrid developed by the Hawaiian Pineapple Research Institute. It gives a medium to large (1.3-2.5 kg) cylindrical, square-shouldered fruit, with large flat eyes, and an intense orange-yellow colour. The clear yellow pulp is sweet, compact, and fibrous. It is high in sugar (15-17°Brix) and ascorbic acid but lower in total acid than 'Smooth Cayenne'. 'MD2' is resistant to internal browning, but susceptible to fruitlet core rot, and more sensitive to Phytophthora than 'Smooth Cayenne'.

'Josapine' is a hybrid between 'Johor' ('Singapore Spanish' x ‘Smooth Cayenne’) and 'Sarawak' ('Smooth Cayenne') released by the Malaysian Agriculture Research and Development Institute (MARDI) for the fresh fruit. It fruits very early, allowing an annual cycle in Malaysia. The vigorous plant produces two to three shoots. Leaves are spiny only at their tip. The fruits (1.1-1.3 kg) are cylindrical with a dark purple peel ripening to attractive orange-red. The flesh is deep yellow with strong aroma and sugar content between 17° and 22°Brix. 'Josapine' fruits have good storage-life and are resistant to black heart disorder or internal browning caused by low temperatures.

‘RL41’('Scarlett') is a hybrid between 'Smooth Cayenne' and 'Manzana' developed by CIRAD-FLHOR. Fruits are 1.4 to 2 kg, orange to red, with regular cylindrical shape. The eyes are medium to large and flat. The flesh is firm, golden yellow with high sugar content (15-18°Brix). Acidity is similar to 'Smooth Cayenne' but higher in ascorbic acid content. Core is thinner and flesh less fibrous. The crown is lighter and erect. The plant is compact with erect, smooth ("piping") leaves. The peduncle is long but resistant to lodging. 'RL41' has a short production cycle and responds well to floral induction. It is susceptible to fruitlet core rot.

References

Antoni, M. G. and Leal, F. 1981. Clave para la identificación de las variedades comerciales de piña. Rev. Fac. Agron. (Maracay), Alcance 29: 13-24.
Bartholomew, D. P. and Malézieux, E. 1994. Pineapple. In: Schaffer, B. and Andersen, P.C. (ed.). Handbook of environmental physiology of food crops. Subtropical and tropical crops. CRC Press, Boca Ratón, pp. 243-293.
Benzing, D. H. 1980. The biology of the bromeliads. Mad River Press Inc, Eureka. CA, 305pp.
Collins, J. L. 1960. The pineapple, botany, utilisation, cultivation. Leonard Hill Ltd, London, 294p.
Coppens d'Eeckenbrugge, G., Leal, F. and Duval, M.F. (1997) Germplasm resources of pineapple. Hort. Rev., 21: 133-175.
Duval, M.-F., Coppens d'Eeckenbrugge, G., Ferreira, F. R., Cabral, J. R. S. and Bianchetti, L. de B. 1996. First results from joint EMBRAPA-CIRAD Ananas germplasm collecting in Brazil and French Guyana. Acta Hort., 425: 137-144.
Leal, F. 1990. Complemento a la clave para la identificación de las variedades comerciales de piña Ananas comosus (L.) Merrill. Rev. Fac. Agron. (Maracay), 16 (1) : 1-11.
Leal, F. and Amaya, L. 1991. The curagua (Ananas lucidus, Bromeliaceae) crop in Venezuela. Econ. Bot. 45 (2) : 216-224.
Leal, F. and Antoni, M. G. 1981. Descripción y clave de las variedades de piña cultivadas en Venezuela. Rev. Fac. Agron. (Maracay), Alcance 29 : 51-79.
Leal, F. and Coppens d'Eeckenbrugge, G. 1996. Pineapple. In: Janick, J. and Moore, J.N. (ed.). Fruit Breeding, Wiley Publishing Co, New York, pp. 565-606.
Matos, A. P. d. 1995. Pathological aspects of the pineapple crop with emphasis on the fusariosis. Rev. Fac. Agron. (Maracay) 21 (3-4) : 179-197.
Montinola, L.R. 1991. Piña. Amon Foundation, Metro Manila, Philippines, 232pp.
Py, C., Lacoeuilhe, J.-J. and Teisson, C. 1984. L'ananas, sa culture, ses produits. G.P. Maisonneuve & Larose, Paris, 561pp.
Rohrbach, K.G. and Schmitt, D.P. (1994) Pineapple. In: Ploetz, R.C., Zentmyer, G.A., Nishiyima, W.T. and Rohrbach, K.G. (eds.) Compendium of tropical fruit diseases. A.P.S. Press, St Paul, MI, pp45-55.

Links

http://agrss.sherman.hawaii.edu/pineapple/pineappl.htm

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