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Foliar Trichomes and Their Systematic Relevance in Solanum (Solanaceae) Species from Southern Western Ghats, Kerala

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Foliar Trichomes and Their Systematic Relevance in Solanum (Solanaceae) Species from Southern Western Ghats, Kerala Rheedea Vol. 27(2) 119–131 2017 ISSN: 0971-2313 https://dx.doi.org/10.22244/rheedea.2017.27.2.24 Foliar trichomes and their systematic relevance in Solanum (Solanaceae) species from southern Western Ghats, Kerala V.S. Anil Kumar*, A.V. Sunila and K. Murugan Plant Biochemistry and Molecular Biology Laboratory, Department of Botany, University College, Thiruvananthapuram – 695034, Kerala, India. *E-mail: [email protected] Abstract Morphological features of foliar trichomes have been successfully employed in classification of diverse groups of plants. The present study evaluates the taxonomic value of trichomes in 17 taxa of Solanum L. using Scanning Electron Microscopy. Both glandular and non-glandular trichomes were observed on the leaves which showed variations in their density of distribution among the species as well as on the two surfaces of the leaves of the same species. The non-glandular trichomes with 1 to 14 arms often form a dense mat over the leaf surfaces, providing protection. The differential length of the radiating arms also helpful in delimiting the species. The non-glandular trichomes are of two kinds: long and short capitate or ovate forms. Species with sparse non- glandular trichomes showed comparatively more number of glandular trichomes. The micromorphology and distribution patterns of trichomes can be used in the taxonomic delineation of various Solanum species. Keywords: Adaptations, Glandular Hairs, Scanning Electron Microscopy, Stellate Hairs Introduction Solanaceae comprise 98 genera with c. 2700 species (Olmstead & Bohs, 2007). The family includes many Plants are adapted to survive under different economically important vegetable species which environmental conditions by morphological are also employed as biological model systems. modifications. Aerial parts of the plants develop Adedeji et al. (2007) analyzed the organographic modifications when they are exposed to distribution of the trichomes of Solanaceae environmental stress. Trichomes and epidermal species and reported two types of trichomes, features are the main foliar adaptations in which showed variations among the species even response to specific environments (Li et al., 2011). within the same genus. Phylogeny of species in Trichomes are the major morphological marker to the family was traced by Sarkinen et al. (2013) in analyze the interplay of various developmental terms of phytochemistry, environmental niche, events correlated with cellular functions (Edwin- geographical distribution, ecological parameters Wosu et al., 2012). Celep et al. (2011) revealed and the genomes. Comparing other groups, the the significance of trichome micromorphology studies on trichome micromorphology of Solanum of Lamium species. Trichome specificitiesis scanty. Thus, the present study is an attempt to effectively delineate taxa within genus or even analyze the ultrastructural features of trichomes in at family level. Secretory and non-secretory Solanum using scanning electron microscopy and trichomes provide insight on the nature of the their ecological significance. secreted phytochemical and their defense against herbivores. Lamiaceae, Brassicaceae, Verbenaceae and Cucurbitaceae display glandular trichomes Materials and Methods with potent bioactive molecules with immense Plant Materials economic importance. Hayat et al. (2009) and Dipa & Daniel (2011) provided the significance Leaves of 17 taxa of Solanum, viz., S. americanum of foliar trichomes in taxonomic discrimination Mill., S. torvum Sw., S. violaceum (Ortega) subsp. in Acanthaceae and Artemisia in Asteraceae, violaceum, S. violaceum (Ortega) subsp. multiflorum respectively. (C.B. Clarke) K.M. Matthew, S. wendlandii Hook.f., 120 Foliar trichomes and their systematic relevance in Solanum S. macrocarpon L., S. melongena L. var. insanum (L.) The number of radiating arms in most species is Prain, S. erianthum D. Don, S. mauritianum Scop., S. 9. Maximum number of radiating arms (up to 18) giganteum Jacq., S. exarmatum Anil et al., S. capsicoides was observed in the pluricellular stellate trichomes All., S. seaforthianum Andrews, S. mammosum L., S. of S. mauritianum (Fig. 1c). Solanum aculeatissimum aculeatissimum Jacq., S. pseudocapsicum L. and S. is found to be distinct from the rest of the species trilobatum L. were collected from Western Ghats in having stellate non-glandular trichomes along of Kerala were subjected to Scanning Electron with simple trichomes on the lower surface Microscopic examination, and the voucher (Fig. 1h). specimens were later deposited at TBGT. The radiating arms of S. giganteum were broader and shorter under SEM (Fig. 1a). The stellate Ultrastructural analysis using trichomes in S. violaceum subsp. violaceum were scanning electronic also comparatively shorter than all others (Fig. 1f). microscopy (SEM) Solanum violaceum subsp. multiflorum is distinct from others in having single elongated slender pointed central arm in their stellate trichomes (Fig. The ultramorphological analysis was carried out 1e). In all other species with stellate trichomes, the using Scanning Electron Microscopy with healthy slender radiating arms were more or less similar expanded leaf blades, fixed in 3% glutaraldehyde. in size. The number of arms in stellate trichomes The samples were kept for 24 hrs and subsequently exhibit variations on both surfaces of the same washed in a 0.05 M potassium phosphate buffer leaves in the species investigated. In S. erianthum, for 30 minutes. After that the leaf blades were the stellate trichomes on dorsal leaf surface have dehydrated in ethyl alcohol series followed by 1,3,4,5 and 8 arms (Fig. 2a), a feature not noticed critical point drying using CO2 (Bozzolla & Russel, in other species whilst those on lower surface have 1992) and mounted on metallic stubs. Further, 8–10 arms (Fig. 1b). Similar stellate trichomes have the specimens are coated with gold in the Sputter been reported in S. leprosum (Toledo Picoli et al., coater (ION 25 SEM). Analysis was done using the 2013) and suggested that these differences are scanning electron microscope (SU 6600, Hitachi due to ontogenic processes starting at different and Zeiss EVO 18) and images were captured in times. In the present study, similar aged leaves different magnifications. A minimum of ten leaves of S. erianthum have been used, but the species per taxon have been analyzed and in all cases, showed the difference with respect to the number fully expanded leaves are chosen. Very young and of arms in trichomes on the two surfaces. In S. older leaves have been excluded from the analysis. mauritianum, the dorsal surface had 6–15 radiating arms for the stellate trichomes (Fig. 1c) and Results and Discussion the lower leaf surface had mostly 6 arms (Fig. 2b). In S. melongena var. insanum (Fig. 1d) and Leaves of Solanum species show wide range S. giganteum (Fig. 1a), both the upper and lower of variation in micromorphology in terms of surfaces of the leaves had stellate trichomes with 9 distribution, nature and structure. Glandular arms. It has been noted that in S. violaceum subsp. and non-glandular trichomes were noticed in the multiflorum, the stellate trichomes were having Solanum species. The non-glandular trichomes may one much elongated slender pointed central arm be stellate or simple while the glandular trichomes with 3 basal protruding arms on the upper surface may be short or long-stalked. The number of arms and with 8 basal protruding arms on the lower in the stellate trichomes showed variations and surface (Figs. 1e & 2c). Interestingly, the elongated similarly the number of cells in the stalk and head of arms were found to have micropappillae or the glandular trichomes also displayed variations echinate ornamentations (Fig. 2d). According to among the species and sometimes on the two sides Barthlott (1981) the cuticular micropappillae are of the same leaf. Among the studied taxa, stellate in continuation of the cuticular folding present trichomes are found in S. giganteum, S. erianthum, S. on the surface of the surrounding epidermal cells. mauritianum, S. melongena var. insanum, S. violaceum Werker (2000) reported that the outer surface of subsp. multiflorum, S. violaceum subsp. violaceum the foliar trichomes in plants including members and S. torvum (Fig. 1a–g). All these species also of Lamiaceae may be smooth or exhibit micro possess stellate non-glandular trichomes on both ornamentations such as micropappillae, warty, upper and lower surfaces of the leaves. However, reticulate and seriate. In S. aculeatissimum, the density, number and nature of radiating arms lower leaf surface has stellate trichomes with 4 show considerable variations among these taxa. basal arms particularly along the veins and also has V.S. Anil Kumar et al. 121 Fig. 1. Non-glandular trichomes of Solanum taxa: a. Upper leaf surface of S. giganteum Jacq.; b. Lower leaf surface of S. erianthum D. Don.; c. Upper leaf surface of S. mauritianum Scop.; d. Upper leaf surface of S. melongena (L.) var. insanum (L.) Prain; e. Upper leaf surface of S. violaceum Ortega subsp. multiflorum (C.B. Clarke) K.M. Matthew; f. Upper leaf surface of S. violaceum Ortega subsp. violaceum; g. Upper leaf surface of S. torvum Sw.; h. Lower leaf surface of S. aculeatissimum Jacq. 122 Foliar trichomes and their systematic relevance in Solanum Fig. 2. Non-glandular trichomes of Solanum taxa: a. Upper leaf surface of S. erianthum D. Don.; b. Lower leaf surface of S. mauritianum Scop.; c. Lower leaf surface of S. violaceum Ortega subsp. multiflorum (C.B.
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