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THE STRUCTURE of FLORAL ELEMENTS of Anchusa Officinalis L

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THE STRUCTURE of FLORAL ELEMENTS of Anchusa Officinalis L ACTA AGROBOTANICA Vol. 62 (1): 37–47 2009 THE STRUCTURE OF FLORAL ELEMENTS OF Anchusa officinalis L. CREATING ATTRACTANTS FOR INSECTS Mirosława Chwil, Elżbieta Weryszko-Chmielewska Department of Botany, University of Life Sciences in Lublin, ul. Akademicka 15, 20-950 Lublin, Poland e-mail: [email protected] Received: 10.02.2009 Abstract INTRODUCTION The present study involved the measurement of size and Anchusa officinalis L. (Boraginaceae) is the micromorphology of the floral elements of Anchusa offici- found in natural stands in western, central and east- nalis L. which are attractants for insects. The structure of the ern Europe. It occurs in xerothermic communities epidermis on the surface of the calyx, petals, throat scales, pistil (Szweykowscy, 2003). The name of the genus and nectary were analysed using light and scanning electron mi- croscopy (SEM). For light microscopy observations, semi-per- is derived from a Greek work ánchein, which means manent slides were prepared, which were treated with Lugol’s to choke, to strangle, and it is probably associated with iodine solution, Sudan III and fluoroglucine. the characteristic structure of the corolla in which the The dark violet lobes of the corolla of Anchusa officina- inlet to the tube is covered by strongly haired throat lis, with a velvety surface, and the throat scales, contrasting with scales (Rejewski, 1996). Common bugloss (An- them, belong to the most important optical attractants which lure chusa officinalis L.) is a biennial or perennial plant. In insects from large distances. The dark pink colouring of the sepals the first year, it only forms a rosette of lanceolate basal additionally increases the attractiveness of the flowers. The epi- leaves. In the next years, stiff foliated shoots grow, 60 dermis covering the calyx formed different-sized non-glandular – 90 cm in height and covered with bristly hairs. The trichomes as well as glandular trichomes. The glandular trichomes lower leaves develop petioles, whereas the upper ones were composed of a uni – or bicellular leg and a unicellular head. are sessile, entire. Cyme inflorescences develop on the The colour of the corolla petals was determined by anthocyanins accumulated in the epidermal cells and in the more deeply situ- branched stems (P odbielkowski and Sudnik- ated parenchyma. The velvety surface was formed by the coni- Wójcikowska, 2003). The plants flower from cal papillae, densely growing from the adaxial epidermis. The May to September (Rutkowski, 2006). pink-violet throat scales with white hairs, covering the inlet to the Disk, actinomorphic flowers are five-petalled. tube of the corolla, were found at the inlet to the corolla throat. The corolla is purple-violet or blue, less frequently The longest trichomes on the surface of the scales were located white or red (Szweykowscy, 2003). The upper in their lower and middle parts, whereas the shortest ones at their part of the corolla is flatly open and it is 15 mm in tips. The epidermis of the central part of the throat scales formed diameter, whereas the lower part forms a straight tube small papillae. The trichomes had thin cell walls, large vacuoles, 5 – 7 mm in length (Weryszko-Chmielewska numerous plastids and lipid droplets. The two-parted stigma of and Chwil, 2007). In terms of morphological adap- the pistil was covered by characteristic expanded outgrowths with tations to entomogamy, bugloss flowers are included wavy edges which performed the functions of structures facilitat- ing the capture of pollen grains. in the class of trumpet-shaped flowers. Such flowers As a result of the present study it was found that the are most frequently directed obliquely or vertically structures affecting the attractiveness of the flowers, through upwards; they seldom hang downwards. The stamens various light effects within the corolla of Anchusa officinalis, and pistils are hidden in the floral tube, at the bottom include the papillae on the corolla surface, trichomes of the of which nectar accumulates (S zafer and Wojtu- throat scales and the epidermal cells of the style. The trichomes siakowa, 1969; K ugler, 1970). Bugloss flowers of the scales can also be responsible for protecting pollen and are visited by honey bees, bumble bees as well as other nectar against rainfall. Hymenoptera and Diptera. Anchusa officinalis belongs to plants providing large amounts of nectar and pol- Key words: micromorphology, sepals, petals, scales, stamens, len to insects (Andersson 1988; Dreisig, 1995; stigma, pollen grains Koł towski, 2006). In our previous study we dem- 38 Mirosława Chwil, Elżbieta Weryszko-Chmielewska onstrated that the nectar of this species contains approx. (Figs 1A, B, D). The sepals of the calyx also have dark 60% of sugars (W eryszko-Chmielewska and pink colour, which may contribute to increasing the at- Chwil, 2007). tractiveness of the flowers (Fig. 1B). Attractants found in insect-pollinated flowers Calyx can be divided into signal and food attractants (K u - The calyx in the flowers of A. officinalis was ca. gler, 1970; Banaszak, 1986). The signal, i.e. 6 mm long and it was composed of 5 sepals fused in the so-called secondary, attractants include the following: basal part and sharp-pointed at the tip. Non-glandular and flower shape, size, odour, colour, glossy elements, sig- glandular trichomes were found both on the abaxial sur- nals and colour indicators. The food attractants, which face and on the edges of the unfused parts of the sepals include nectar, pollen and food hairs, are a reward for (Figs 1 D, 2 A-C). The spiky non-glandular trichomes insects visiting flowers, that is, these are primary at- were live, uni- or bicellular, and they had different sizes tractants. (Figs 2 A, B). The smaller trichomes had a swollen basal The aim of this study was to determine the struc- part, whereas the larger-sized bristly hairs were surround- ture of floral elements of Anchusa officinalis which are ed at the base by 6 up to 8 cells forming a massive base. important in pollination ecology and which contribute The length of the non-glandular trichomes was within to the reproduction success of this species. The micro- a range of 172 – 778 μm. The glandular trichomes with morphology of sepals, petals, throat scales, elements of a length of approx. 70 μm were composed of a uni- or the pistil and nectary was investigated. Special atten- bicellular leg and a unicellular head (Fig. 2C). Numerous tion was paid to the structure of the trichomes covering anisocytic stomata were found on the abaxial surface of the surface of the throat scales. the sepals. Most of the epidermal cells and the guard cells showed irregular striation of the cuticle (Figs 2 C-E). MATERIALS AND METHODS Corolla The study was carried out in the years 2007 - The corolla length reached ca. 10 mm, whereas its 2008. The plants of Anchusa officinalis L. from which diameter 8-9 mm. The length of the corolla tube was 5-7 inflorescences were sampled came from the Botanical mm, and the tube diameter approx. 2.5 mm. The colour of Garden of the Maria Curie-Skłodowska University in the corolla petals was determined by anthocyanins con- Lublin, Poland. tained both in the epidermal cells and in the more deeply The micromorphology of the following floral situated parenchyma. The velvety surface of the petals was formed by the conical papillae, densely growing elements: sepals, petals, stamens, pistil and nectary, was from the adaxial epidermis, which had an average height observed using light and scanning electron microscopy of 25 μm and a basal width of 24 μm. The surface of the (SEM). The structure of the epidermis on the surface of papillae was covered with a thin, slightly striated cuticle the sepals, petals, throat scales as well as on the surface of (Figs 3B-D). The lower, abaxial epidermis of the petals the pistil and nectary was analysed. For light microscopy was smooth (Fig. 3A). The vacuoles of the papillae were observations, semi-permanent slides were prepared in stained by pink anthocyanins. There were two layers of glycerol-gelatine, which were stained with Lugol’s iodine parenchyma between the epidermis layers. The thickness solution, Sudan III or fluoroglucine. On the cross sections of the petal was ca. 75 μm. Hairy throat scales occurred of the perianth, observations of cells were made for the at the inlet to the corolla tube, forming 5 epipetalous pro- presence of starch and fatty substances as well as for the tuberances. The throat scales were pink-violet coloured, cell wall structure. Measurements were made of the floral whereas the long trichomes of the scales were directed elements and epidermis formations on their surface: non- to the central part of the corolla tube, and those closing its glandular and glandular trichomes as well as papillae. inlet were white coloured (Figs 1C, D; 4A, B). The plant samples used for scanning electron In the flowers of Anchusa officinalis, the throat microscopy analysis were fixed in glutaraldehyde, next scales, strongly contrasted with the colour of the corolla, dehydrated in acetone and, after critical point drying in perform, inter alia, the role of well visible colour sig- liquid CO , coated with gold using a Sputter Coater. 2 nals, indicating the location of pollen and nectar in the The floral elements were viewed under a TESLA BS- flower. The throat scales with a length of approx. 1.7 300 microscope. mm were ligulate in shape (Figs 4C; 5A) and they were structures composed of several layers of cells. The long- RESULTS est trichomes were located in the lower and middle parts The colour of the flower belongs to the most im- of the scales, whereas the shortest ones at their tips (Figs portant optical attractants, luring pollinators from large 1C, 4C, D; 5A, B).
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