Journal of Experimental Botany, Vol. 60, No. 9, pp. 2461–2464, 2009 doi:10.1093/jxb/erp166 PLANT CULTURE Symbolism of plants: examples from European-Mediterranean culture presented with biology and history of art Riklef Kandeler1 and Wolfram R. Ullrich2* 1 2 Institute of Botany, University of Natural Resources and Applied Life Sciences, Gregor Mendelstr. 33, 1180 Wien, Austria Institute of Botany, Darmstadt University of Technology, Kirchbergweg 6, D-64287 Darmstadt, Germany JULY: Lotus On hot summer days we like to see our ponds and small eutrophic lakes covered with the white, pink or yellow flowers of water lilies. Everybody knows how Claude Monet became so fascinated by his ‘nymphe´as’ that he painted them in all situations, but the Nymphaeaceae have played a role in symbolism for over 4000 years. Even more important in Asian culture, particularly in Buddhism, has been the Indian lotus, Nelumbo nucifera (Nelumbonaceae). In literature and mythology, the term lotus (in Greek lotos) has been used to cover both families, relating the high symbolic and cultural importance of the lotus in great parts of the world. Biology, water lilies (Nymphaeaceae) All the nymphaeas live in still or slow water and have a condensed shoot apical meristem underwater close to the substrate. The stems, petioles, and leaves are characterized by wide and efficient tracts of aerenchyma and the leaves, and usually the flowers, float on the water surface due to their high intercellular air content. Their ‘homobaric’, i.e. widely interconnected, intercellular system enables them to send oxygen-enriched air by ‘pressurized ventilation’ from the younger leaves to the subaquatic parts which live in an almost anaerobic stagnant environment (Dacey, 1981; Armstrong et al., 1996; Grosse, 1996; Colmer, 2003). In the Nile delta, two species predominate, Nymphaea caerulea and N. lotus. N. caerulea has entire or obtuse-dentate leaves and the blue petals are pointed (Fig. 1, left), N. lotus has sharply dentate leaves and the white petals are broadly rounded at the end (Fig. 1, right). Both of them release a pleasant fragrance. Nymphaeas have long had a particular significance as intoxicants for shamans because of their alkaloid and glycoside content. They contain compounds similar to atropin and papaverin (nupharin, nymphalin, ellagic acid) (Roth et al., 1994). Symbolism: nymphaea flowers in pharaonic Egypt The flowers of the blue water lily emerge out of the water in the morning and recede back into the water at noon. Thus they became a symbol of the sun rising out of the night, as well as the continual renewal of life. The Egyptian symbol of eternal life, the ‘ankh’, possibly a magic knot in older depictions (Lurker, 1987), was composed of three plant stems or petioles knotted together, very likely from lotus. The sacredness of lotus plants in pharaonic Egypt was possibly also due to their use as drugs by priests and shamans who used the plant as an intoxicant for trances and to make contact with the other world. As a state symbol, intertwined lotus and papyrus plants were used to symbolize the union of upper and lower Egypt. However, more familiar was the use of both Nymphea species for funeral rites. Close to Saqqara, south of Cairo, in limestone relief on a tomb (2500 BC), the flowers of both lotus species were reproduced rather precisely (Fig. 2). Within the tomb, lotus flowers and various food items were placed close to the feet of the deceased, apparently to feed both spirit and body in the after-life. In an older limestone relief (c. 2600 BC), probably from the area of Memphis (also in lower Egypt), the deceased is shown on a boat-trip with the water surface being almost completely covered with the blue lotus. More than 1000 years later, in the grave of Tut-ankhAmun (Valley of the Kings, Egypt 1347–1338 BC) Howard Carter found wreaths of dry flowers some of which were N. caerulea (Newberry, 1973) on the second and the innermost coffin. The innermost coffin contained a large flower collar with rows of lotus petals, a symbol of revival and reanimation (Lurker, 1987). Hence, in ancient Egypt, the lotus, most commonly N. caerulea, was used widely as religious and ceremonial icon. * To whom correspondence should be addressed: E-mail: [email protected] ª The Author [2009]. Published by Oxford University Press [on behalf of the Society for Experimental Biology]. All rights reserved. For Permissions, please e-mail: [email protected] 2462 | Plant Culture Fig. 1. (Left) Blue Egyptian lotus, Nymphaea caerulea. (Right) White Egyptian lotus, N. lotus. Botanical Garden, University of Vienna (Kandeler). Fig. 2. Limestone relief in a grave near Saqqara, Egypt, c. 2500 BC. On the left can be seen Nymphaea caerulea and N. lotus (Lange and Hirmer, 1967). Biology: Indian lotus The white or pink flowers of Nelumbo nucifera emerge out of the water like the peltate leaves (Figs 3, 4). The floral axis is in the form of a solid inverted cone on whose flat upper side the carpels recede into holes during the ripening process (Fig. 3). Indian lotus is widely distributed in stagnant waters on the Indian subcontinent and in South-East Asia. It was also introduced into Egypt at around the time of the Persian hegemony (5th century BC) (Germer, 1985), although it no longer grows there today. In recent years, Nelumbo is again in the news, as the inspiration for a modern engineering innovation. The waxes of its leaf cuticle have a very efficient water- and dirt-repellent structure and this ‘lotus effect’ Fig. 3. Indian lotus, Nelumbo nucifera. Botanical Garden, University of Bonn (Ullrich). (Fig. 4) is now imitated on superior glass and for other ‘stay-clean’ surfaces (Barthlott and Neinhuis, 1997). Symbolism: Indian lotus In Hinduism since antiquity the Indian lotus has been a particularly sacred plant (Beuchert, 2004). It was regarded as the first-born of creation and as the magic womb of the universe and the gods. In Buddhism, the Indian lotus also has a central significance (Beuchert, 2004). Buddha strode over seven lotus flowers when he was born and with the Bodhisattvas (future Buddhas) and various saints he was often depicted as sitting on lotus flowers (Fig. 5) or on the flat top of the gynaecium (Fig. 3) (Majupuria and Joshi, Plant Culture | 2463 Fig. 4. Indian lotus plant, peltate leaves with a structured hydrophobic cuticle, responsible for the famous lotus effect. Botanical Garden, University of Bonn (Ullrich). 1988). So important has the lotus been to Indian culture that it has been adopted as the flower of India, to represent not only divinity and enlightenment, but also fertility due to its fecundity and the design of its gynaecium, long life and knowledge due to its longevity, and wealth associated with its many culinary uses. Once introduced into Egypt, the Indian lotus replaced the native species in the cult of Isis in the 1st century BC. Isis became not only the protective goddess and divine mother but also the ruler of the underworld and queen of heaven (Helck, 1979), the Indian lotus flower being one of her most important attributes (Witt, 1997). As the Christian church grew it, too, adopted the lotus as a potent symbol. At the Christian Council of Ephesus in 431 AD, St Mary was designated as ‘Theotokos’ (the Bearer of God). The subsequent flush of church building dedicated to St Mary incorporated the image of the lotus. A good example is Panagia Theotokos (now Panagia Achiropiitos) in Thessaloniki, 450–460 AD. In the central arcade sit two dishes from which lotus leaves, flowers, and fruits are sprouting around the central golden cross (Fig. 6). Fig. 5. Buddha sitting on a pedestal formed from a stylized lotus flower, in his hands the typical beggar’s bowl. Bronze sculpture, T’ang period (Chinese), 7th to 9th century AD. (Hájek and Forman, 1954). Fig. 6. Mosaic in Panagia Theotokos, Thessaloniki, central arcade (450–460 AD). Two dishes with leaves and flowers of Indian lotus and two ears of grain each. (Kandeler). References Armstrong W, Armstrong J, Beckett PM. 1996. Pressurized ventilation in emergent macrophytes: the mechanism and 2464 | Plant Culture mathematical modelling of humidity-induced convection. Aquatic Botany 54, 121–135. Hájek L, Forman W. 1954. Chinesische Kunst in tschechoslowakischen Museen. Prague: Artia. Barthlott W, Neinhuis C. 1997. 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