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BROMELIACEAE(Pineapple family)
Members of this family, natives of tropical America and the West Indies, number some 1400 species in 44 genera. Typically, they are spiny rosette-forming plants, many of which grow terrestrially, but most of which are epiphytes in their natural environment. Because of the peculiar arrangement of their leaves, many of the rosette-forming species in the tropics are often found associated with a varied fauna including ants, spiders, and scorpions (Wheeler 1942). Several species produce edible fruits, the best known of which is the pineapple, from Ananas comosus (L.) Merr., which is an important article of commerce. Tillandsia usneoides L. is an unusual bromeliad. It hangs from branches in long grey festoons that resemble certain lichens. The plant is known by a variety of common names including long, Florida, Louisiana, or Spanish moss, old man's beard, and vegetable horsehair. It is used for stuffing pillows and similar such items (Willis 1973, Usher 1974). Members of this family are occasionally grown as house or greenhouse plants. Most commonly grown are Aechmea Ruiz & Pav., Billbergia Thunb., and Vriesea Lindl. species. The most likely hazard associated with members of this family is mechanical injury caused by the strong, sharp, curved spines that adorn the leaf edges of many species. In addition, persons coming into contact with the plant sap of some species may develop an irritant dermatitis from the proteolytic enzymes that are present. Tillandsia L. species in particular, when growing in the wild, may cause a pseudophytodermatitis when handled. This is because they provide nesting sites for ants which, depending on species, may bite or sting aggressively if the plant is disturbed. An herbarium specimen of this species was described as being a "small, wasp-inhabited plant" (von Reis & Lipp 1982).
In its natural habitat in the tropical forests of South America, Streptocalyx longifolius is found growing on arboreal ant nests ("ant gardens") together with other myrmecophytes including Philodendron myrmecophilum Engl. (fam. Araceae), Anthurium scolopendrinum var. poiteauanum (Kunth.) Engl. [= Anthurium gracile (Rudge) Lindl.] (fam. Araceae), Aechmea spicata Mart. ex Schult. & Schult.f. [= Aechmea mertensii (G.Mey.) Schult. & Schult.f.] (fam. Bromeliaceae), Peperomia nematostachya Link [= Peperomia macrostachyos (Vahl) A.Dietr.] (fam. Piperaceae), Codonanthe uleana Fritsch [= Codonanthopsis uleana (Fritsch) Chautems & Mat.Perret] (fam. Gesneriaceae), and Phyllocactus phyllanthus (L.) Link. [= Epiphyllum phyllanthus (L.) Haw.] (fam. Cactaceae) (Ule 1905). See also Aechmea mertensii (G.Mey.) Schult. & Schult.f. below.
The strong sharp recurved hooks on the leaves of Ananas magdalenae can inflict mechanical injury (Allen 1943).
Referring to earlier studies by Ule (1901, 1905, 1906a), Wheeler (1921) described how, in certain localities of Central and South American forests, balls of earth bristling with epiphytes in various stages of germination and growth inhabited by colonies of ants (hence "ant gardens") could be found attached to the branches of trees. The densely felted rootlets of the plants kept the particles of soil together and formed the walls of the galleries and chambers occupied by the insects. The bromeliad Aechmea spicata Mart. was included in a list of about 14 species from several families (see also Araceae, Cactaceae, Gesneriaceae, Moraceae, and Piperaceae) that had been found as constituents of these ant gardens, which were tenanted by a number of larger and smaller species of ant. Amongst the species of ant described were Camponotus femoratus (Fabricius, 1804), fam. Formicidae and Crematogaster cf. limata parabiotica Forel [= Crematogaster levior Longino, 2003], fam. Formicidae occurring in about 90% of the ant gardens and living together in parabiosis. Wheeler (1921) further noted that:
Schmidt (1985) further explored the dermatological hazards associated with these "super nettles" (myrmecophytes) and their potential to elicit a pseudophytodermatitis in plant collectors.
"Pineapple estate pyosis", occurring in workers who gather the fruits, probably represents an acarus infection with secondary bacterial infection (Simons 1953). The raw fruit, when eaten in large quantity, produces a burning sensation of the lips and mouth (Watt & Breyer-Brandwijk 1962) and can also produce angular stomatitis (Fasal 1945). Almost complete obliteration of the fingerprints by removal of the stratum corneum in workers employed in cutting up pineapples has been ascribed to the effects of pressure together with the keratolytic effect of bromelain, a protease found in the plant sap. A variety named Mauritius pineapple seemed to be most injurious. The nails were unaffected, in contrast to the nail damage that occurs in workers exposed to proteases of animal origin. Calcium oxalate and citric acid were excluded as causes of the skin changes. Moniliasis of the finger webs was occasionally observed in pineapple cutters (Polunin 1951). The proteases in this species are glycoproteins and are related in their properties to papain. The different parts of the plant yield slightly different proteases, and each may be further fractionated into its constituent proteolytic enzymes. Thus, stem bromelain, fruit bromelain, and leaf bromelain have been studied (El-Gharbawi & Whitaker 1963, Murachi 1970, Daley & Vines 1978). Pineapple fruits contain ethyl acrylate. Ethyl acrylate (4% in petrolatum) produced sensitisation in 10 of 24 subjects in a maximisation test (Opdyke 1975). A positive patch tests reaction to pineapple in a fruit handler (Morris 1954) cannot be interpreted in the absence of controls. The spines of the edible fruits can produce mechanical injury (Burkill 1935). Garduño et al. (1974) isolated the protease sylvestrisin from Bromelia sylvestris. It was found to consist of several component proteins. See also Ananas comosus (L.) Merr. above and Bromelia pinguin L. below. Bongiorno de Pfirter & Buttazzoni de Cozzarin (1976) isolated a protease named balansain from the fruit juice of this plant. Balansain was found to contain at least 6 constituent proteins, and to resemble papain in its properties. If eaten without peeling, the fruit irritates the lips (Morton 1981). Commonly referred to, incorrectly, as Bromelia hemisphaerica Lam., the fruit juice from this taxon has been reported to contain a proteolytic enzyme named by Agundia et al. (1977) as hemisphaericin-D, a dialysable polypeptide that evidently can polymerise to form the hemisphaericin elsewhere described by Garduño et al. (1974) and by León-Alcalá et al. (1989). See also Ananas comosus (L.) Merr. above and Bromelia pinguin L. below.
The proteolytic activity of the wild fruits of Bromelia karatas has been found to be greater than that of Ananas comosus var. comosus. This activity has been ascribed to a mixture of proteases named karatasin (Garduño et al. 1974, Villanueva-Alonzo et al. 2019). See also Ananas comosus (L.) Merr. above and Bromelia pinguin L. below. Bromelia karatas is often grown as a barrier hedge, impassable by animals and humans (Morton 1981). A protease named palmerin consisting of several component proteins has been reported from this species (Garduño et al. 1974, León-Alcalá et al. 1989). See also Ananas comosus (L.) Merr. above and Bromelia pinguin L. below.
This species is often planted for hedges, its spiny leaves serving well to repel animals (Standley 1937a). Recurved hooks on the leaf margins can produce mechanical injury (Allen 1943, Oakes & Butcher 1962). The fruit is eaten in Central America without ill effects other than mild irritation around the mouth on the upper part of the lips (Asenjo et al. 1944). The fruit juice is pleasantly acid, but produces a burning sensation when applied to the skin or lips, followed by peeling after a few hours (Asenjo & Capella de Fernandez 1942). Small painful ulcers readily developed within a period of 3–6 h on the fingertips of laboratory workers who handled the pulp of the fruit without rubber gloves. This was apparently the result of a combination of mechanical irritation caused by calcium oxalate raphides (of average size 108 µm in length and 5 µm in width) and subcutaneous injection of proteolytic enzyme (Asenjo et al. 1944). A protease named pinguinain may be isolated from the fruit. In a Puerto Rican variety, pinguinain has been found to be a single compound (Toro-Goyco & Matos 1964), whilst in a Cuban variety is has been found to consist of at least two components (Messing 1961). Pinguinain is a glycoprotein that resembles papain in its properties, but is unusual in being very resistant to heat denaturation and has an optimum activity at 65°C (Toro-Goyco et al. 1968).
In its natural habitat in the tropical forests of South America, Nidularium myrmecophilum is found growing on arboreal ant nests ("ant gardens") together with other myrmecophytes including Philodendron myrmecophilum Engl. (fam. Araceae), Ficus myrmecophila Warb. [= Ficus paraensis (Miq.) Miq.] (fam. Moraceae), Markea formicarum Dammer (fam. Solanaceae), Ectozoma ulei Dammer [= Hawkesiophyton ulei (Dammer) Hunz.] (fam. Solanaceae), and Codonanthe formicarum Fritsch (fam. Gesneriaceae) (Ule 1905, Wheeler 1921). See also Aechmea mertensii (G.Mey.) Schult. & Schult.f. above.
The bases of the leaves are furnished with sharp and dangerous spines (Smith LB & Standley PC in Standley 1937a). This is a genus of about 168 species found in highland regions of South America (Mabberley 1997). Many are cultivated as ornamentals (Hunt 1968/70). In most species, the leaves have spiny margins. The following list of spiny-leaved species is representative (Smith LB in Macbride 1936, Smith LB & Standley PC in Standley 1937a, Hunt 1968/70):
Ule (1906a) included Tillandsia exigua in a list of bromeliad ant-plants. See also Tillandsia L. below. In their natural habitat, the species listed below provide nesting sites for ants (Ule 1906a, Ule 1906b, Wheeler 1942, Benzing 1970) thus rendering them potential dermatological hazards for plant collectors (Schmidt 1985):
Wheeler (1942) noted that Tillandsia species constitute the great bulk of the epiphytic flora in the West Indies. Ants live only in those species of Tillandsia in which the leaf-bases are more or less spoon-shaped and compactly overlapping. However, they very probably occasionally also live among the roots of these plants in so-called "ant-gardens". See also Aechmea mertensii (G.Mey.) Schult. & Schult.f. above.
in the semiarid region of Guatemala, ants were found to be the predominant (92%) insect taxon inhabiting Tillandsia xerographica growing epiphytically, mainly on drought-resistant trees with rough bark such as Pereskia lychnidiflora DC. [= Leuenbergeria lychnidiflora (DC.) Lodé] (fam. Cactaceae). See also Tillandsia L. above. However, the blood-sucking reduviid bug Triatoma ryckmani Zeledón & Ponce, 1972, fam. Reduviidae has also been found to inhabit this epiphyte. In their study, Marroquin et al. (2004) found 53 specimens of Triatoma ryckmani in thirty Tillandsia xerographica. Accidental introduction of Triatoma ryckmani to houses and to other countries may be an unfortunate consequence of the illegal commercialisation of Tillandsia xerographica because this reduviid bug is a carrier of a parasite (Trypanosoma cruzi Chagas, 1909, fam. Trypanosomatidae) that causes Chagas disease otherwise known as American trypanosomiasis, a potentially fatal infectious disease. References
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