Vol 20, No. 9;Sep 2013
Epidermal characteristics of 34 species of Brassicaceae growing in desert of Saudi Arabia.
Abdullah Doaigey*, Ihsan El-Habashy, Ahlam Al- Watban, Abd Elsalam Milagy, Abdulaziz Al Sahli, Manzer Siddiqui, Mohamed Al-Whaibi and Mohamed El-Zaidy Botany and Microbiology Dept., College of Science, King Saud Univ., P. O. Box 2455, Riyadh 11451, Saudi Arabia.
*Abdullah DOAIGEY (Corresponding author) Tel: 00966505154324
E-mail:
[email protected]
Abstract The aim of the study was to investigate the detail epidermal characteristics of the leaf and stem of 34 species of the family Brassicaceae which grow in different location of Saudi Arabia under desert conditions. The study was carried out using Light and Scanning Electron Microscope. The epidermal cell parameters, such as straight or sinuous, thin or thick anticlinal cell walls, smooth or warty cuticle, stomata and trichomes types were studied. Therefore, on the basis of these epidermal characteristics we divided these species into four groups on the basis of presence or absence of nonglandular trichomes. Group-1 represents 6 species and the leaves and stems of these species were glabrous, Group-2 represents 17 species and hairy leaves and stems were found in these species, Group-3 represents 5 species and leaves of this group were hairy and Group-4 also represents 6 species and stems were hairy. Also, species within each group were distinguished using other epidermal characters, such as cell shapes, anticlinal cell walls and stomata types. Key based on the epidermal characters is provided. Key words: Epidermal characters; Leaf epidermis; Stem epidermis; Trichomes; Stomata; Species; Brassicaceae.
1. Introduction Brassicaceae is an economically important family which comprises many economical plants. It is well represented in Saudi Arabia. (Chaudhary, 1999) reported74 species in 53 genera growing in different areas of the Saudi Arabia. The epidermal characters of the leaves and stems of the species of Brassicaceae family were not fully investigated. (Metcalfe and Chalk, 1950) mentioned 202
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thatplants belong to the Brassicaceae contain nonglandular, unicellular, unbranched or branched,T-shape or stellate trichomes and their cell walls with calcium carbonate, glandular trichomes rare, stomata commonly anisocytic type and secretory cells represented by myrosin cells. (Inamdar and Rao, 1983)reported nonglandular trichomes in the epidermis of 35 species of the family Brassicaceae, and classified them into unicellular, bicellular and multicellular on the basis of their number of cells. (Ahmad et al., 2010) separated some members of Brassicaceae by their epidermal characters, such as epidermal cells with slight sinuous anticlinal cell walls and polygonal to nearly linear in Brassica campestris while they are polygonal to nearly rounded and linear in Lepidium apita and polygonal to less undulating to clearly irregular at different areasof the organ epidermis in Raphanus sativus and in Sisymbrium irio. Also, (Tuoand Zhou, 2010) reported that the shapes of leaf epidermal cells of 3 species of Isatis and 5 single genera of Eulidieae were usually polygonal or irregular and stomata mainly anisocytic, rarely anomocytic.There were no significant differences in each other. (Sun and Li, 2007) reported the presence of stomata in both epidermises and their types mainly anisocytic, rarely anomocytic and occasionally paracytic.They concluded that the leaf epidermal features can serve as a criterion of distinguishing species in the genus Lepidium. In another study, (Sun and Li, 2008) reported different features of the upper and lower epidermis of leaf of some species of the genus sisymbrium for example upper epidermal cellsusually polygonal with straight or arched anticlinal cell walls, few slightly sinuous while the lower epidermal cells irregular in shape with sinuous or sinuate anticlinal cell walls. Stomata are mainly anomocytic, rarely anisocytic and occasionally paracytic in both epidermises of leaf. (Gostin, 2009) studied Erysimum wittmanii ssp. Wittmanii and reported that stomata anisocytic type is present in both side of epidermis of leaf; trichomes with three- armed, rarely twoarmed with thick, warty cell walls are also present. (Kadry, 2005) observed nonglandular and glandular trichomes in leaves or stems epidermises of 82 species of Brassicaceae, describing nonglandular as unicellular with 9 forms and glandular trichomes with two forms. The significance of leaf epidermal features in the systematic of many genera and species of different families of flowering plants has been reported (Stace, 1984; Adedeji & Dloh, 2004; Celka et al., 2006; Zou et al., 2008; Yasmin et al., 2009: Khan et al., 2011). Our work aims to study in details the epidermal characters of leaves and stems of 34 species of the family Brassicaceae growing under desert conditions in Saudi Arabia which might be used in the identification of the species.
2.Materials and methods In the present experiment for the study of epidermal characteristics of plants species belonging to Brassicaceae family,almost of the samples taken from the Herbarium of KSU, Department of Botany and Microbiology, College of Science, King Saud University, where voucher specimens are deposited (Table 203
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1).In addition, some fresh samples were collected from the different location of the Saudi Arabia. For the study of epidermal characteristics, dried sample swere taken from herbarium. The leaves and stems were soaked in water for 2-5 days and then prepared in the same way as fresh material done. Samples of fully expanded leaves and stems of plant species were takenand surface preparation were made from leaves and stems of the materials either by stripping the epidermis and mounting in 50% v/v glycerol solution or by clearing leaf segments in chloral hydrate solution and then mounting in 50% v/v glycerol solution. Dry materials were soaked in water for 2-5 days and then prepared in the same way as fresh materials except scraping the epidermis when is needed. 5 samples were taken and 3 leaves (upper, middle and lower) of each sample were chosen and at least 5 epidermal strips of each leaf area (midrib region, intercostal region and leaf margin) were examined. Three epidermal strips of 5 stem samples (from the lower, middle and upper areas of the stem) were examined with the help ofPhoto Microscope III (LM). For the study of scanning electron microscope, the plant materials were fixed in 2.5% glutaraldehyde for 3 h then washed thoroughly with 0.1 M phosphate buffer. A series of ethanol mixtures (30, 50, 75, 90 and 100%) were applied to dehydrate the fixed materials and at the end 100% acetone were used for final dehydration of the samples. The epidermis characteristics of the sample were examined using scanning electron microscope (JSM- 6380- LA, JEOL) (Falk, 1980). Cell wall thickness of the epidermal cells and trichomes was measured according to (Doaigey et al., 1985). Whereas, “thick” > 6 um, “slightly thick” 2.5 - 6 um and “thin” less than 2.5 um.Number of stomata per area 0.23 mm2 and the measurements for length and width of epidermal cells andstomatawere takenwith the help of scale fitted in light microscope.Statistical analysis was madeusing JMP software. 3. Result and Discussion: The epidermal characters of leaves and stems of 34 species are summarized inTables2 and 3 respectively. 3.1. Epidermal cells The epidermal cells of upper and lower epidermis of leaves are polygonal, isodiametric or elongated in all species under investigation.Cells with straight anticlinal cell walls in 13 species, cells with sinuous anticlinal walls in 12 species and cells with straight and sinuous anticlinal cell walls in 9 species were recorded (Table,2; Figs. 1- 4). These observations are in general agreements with the results of other studies (Ahmed et al., 2010; Zhang et al., 2009; Tuoand Lin, 2010; Sun and Li, 2008). However, stem epidermal cells were polygonal, elongated with straight anticlinal cell walls in all 34 species except one species (Leptaleum filifolium) that had epidermal cells with rarely sinuous anticlinal walls. Moreover, epidermal cells with thin anticlinal cell walls occurred in 18 species (4 species out of them showed occasional epidermal cells with thick anticlinal cell walls), while epidermal cells with thick anticlinal cell walls were 204
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observed in16 species (Table 3; Figs.5, 6). Beaded anticlinal cell walls were observed in leaf epidermises in 17 species out of 34 species weather they had thin or thick (Table 2; Fig.4) while in stem, epidermal cells with beaded anticlinal cell walls were present in 8 species (3 out of them, cell walls were rarely beaded (Table3, Fig.5). This cell wall character was not studied in previous studies. Out of 34 plants species, 31 species exhibited smooth cuticlein both lower and upper epidermis of leaves as well as stems (Tables, 2 and 3; Figs. 1-6) while striated cuticle was present commonly in lower epidermis and rarely present in stem epidermis of one species (Coronopus niloticus) while it is rarely present in two species (Cardaria draba and Matthiola fruticulosa) (Tables, 2 and 3). The smooth cuticle may be considered as a special and common feature in these species that are growing naturally in Saudi Arabia.
3.2. Stomata The stomata are commonly anisocytic type in the leaf epidermises of all species but intermixed with anomocytic type in 21 species out of 34 species (Table 2; Figs. 1- 4). These results are in agreement with the findings of (Tuoand Zhou, 2010) and (Sun and Li 2007, 2008) except paracytic stomata were not observed in all the studied 34 species. The occurrence of anisocytic stomata in the leaf upper and lower epidermis of 13 species out of 34 species support the results reported by (Gostin, 2009) in her study on Errysimum wittmanii ssp. wittmanii. On the other hand, in stem epidermis, anisocytic stomata were observed in all 34 species, intermixed with anomocytic stomata in 17 species, but the occurrence of anomocytic stomata in 7 out of these species is rare, The stem epidermis of these 17 species has anisocytic stomata only (Table,3; Figs. 5.6). However, the presence of anisocytic stomata in the stem and leaf upper and lower epidermises of 13– 17 species out of 34 species and the presence of intermixed with anomocytic type in the rest of studied species confirm what is commonly found in the members of the family Brassicaceae. 3.3. Trichomes Among the 34 species, only 6 species showed glabrous leaves and stems. However, 28 plant species showed nonglandular unicellular trichomes, and out of 28, 17 species had nonglandular unicellular trichomes in both leaves and stem epidermises and 6 species showed unicellular trichomes only in leaves and remaining 5 species had nonglandular unicellular trichomes in stems (Tables 2, 3). Glandular trichomes were not observed in any epidermal cell of leaves and stemsof all the species studied, but generally, nonglandular, unicellular trichomes occur in many forms. 3.3.1. Nonglandular, unicellular, unbranched However, in the present study, nonglandular, unicellular, unbranched were 205
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observed in the leaf epidermis of 10 species with four forms: 1) thin warty cell walls in one species (Coronopus niloticus); 2) thin smooth cell walls in one species (Horwoodia dichsoniae). 3) Thick warty cell walls in 4 species), and 4) thick smooth cell walls in 4 species (Table 2, Fig. 8). Also, we observed these trichomes in the stem epidermis of 11 species having three forms: 1) thin smooth cell walls in one species (Horwoodia dichsoniae), 2) thick warty cell walls in 9 species (Table 3, Figs.7, 15), and 3) thick smooth cell walls in one species (Schimpera arabica) (Table 3). 3.3.2. Nonglandular, unicellular, branched Nonglandular, unicellular, branched were observed in the leaf epidermis of 13 species with four forms:1 ) Two branches, T– shaped with thick warty cell walls occur in 5 species (Table 2, Fig. 16 ), 2 ) slightly Y-shaped with thick warty cell walls occur in one species ( Malcolmia africana) (Table 2; Fig. 10 ), 3 ) many branches with thick warty cell walls in 5 species (Table 2, Figs 4, 13, 14), and 4) many branches with smooth cell walls in 2 species ( Matthiola longipetala and M. livida), (Table 2 ). Nonglandular, unicellular, branched were observed in the stem epidermis of 11 species in three forms: 1) T-shaped with warty cell walls in 3 species, 2) slightly Y-shaped with thick warty cell walls in one species (Malcolmia africana), and 3) many branches with warty cell walls in 7 species (Table 3). Similar results were reported by (Inamdar and Rao, 1983) and (Metcalfe and Chalk, 1950). They reported that nonglandular unicellular unbranched trichomes (simple trichomes) are present in the stems or leaves of the species, e.g. Eruca sativa Mill., Erucaria hispanica (L.) Druce and Leptalium filifolium (Willd.) Dc, but in the present study stems and leaves of these three species were glabrous. Moreover, our investigation shows that the stems and leaves of the species, Moricandia sinaica and Zilla spinosa (Turra) were glabrous. This result is similar to the finding of (Kadry, 2005).
4. Conclusion: The growing 34 species (belong to the Brassicaceae) in different locations of the Saudi Arabia were divided into four groups on the basis of epidermal characteristics with reference to the presence and absence of non-glandular trichomes: Group-1:Leaves and stems glabrous include 6 species representing 6 genera. Group-2: Stems and leaves with nonglandular trichomes includes 17 species belong to 12 genera. Group-3: Leaves with Nonglandular trichomes includes 5 species belonging to 5 genera. . 206
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Group-4: Stems with Nonglandular trichomes includes 6species representing 4 genera. Species within each group could also be further separated on the basis of other epidermal characters, such as cell shapes, anticlinal cell walls and stomata types. Key based on epidermal characters of leaves and stems of 34 species of the family Brassicaceae growing wildly in Saudi Arabia.
1´Leaves and stems glabrous. ------------------------------------------------------------------------------- 2 1´´ Leaves and stems with nonglandular trichomes. --------------------------------------------------- 6 1´´´ Leaves or stems with nonglandular trichomes. ---------------------------------------------------- 17 2´Leaf epidermal cell polygonal elongated with thin anticlinal cell walls.
---------------------- 3
2´´Leaf epidermal cells polygonal isodaimetric with thin anticlinal cell walls. ------------------- 4 3´, Straight, thin anticlinal cell walls, stomata anisocytic type. ------------------- (1) Zilla spinosa 3´´ Sinuous, thin, anticlinal cell walls, stomata anisocytic intermixed with anomocytic type. ------------------------------------------------------------------------------------------ (2) Leptaleum filifolium 4´ straight, thin beaded anticlinal cell walls. ------------------------------------------------------------- 5 5´ Stomata anisocytic type. ------------------------------------------------------------ (3) Eruca sativum 5´´, Stomata anisocytic intermixed with anomocytic type. ------------------- (4) Erucaria hispanica 4´´ Sinuous thin beaded anticlinal cell walls, stomata anisocytic type. --------- (5) Isatis lusitanica 4´´´Straight thick anticlinal cell walls stomata anisocytic type. ------------- (6) Moricandia sinaica 6´Nonglandulartrichomesunicellulerunbranched. ----------------------------------------------------- 7 6´´Nonglandular trichomes unicellular branched. ----------------------------------------------------- 8 6´´Nonglandular trichomes unicellular unbranched and branched intermixed .---------------------------------------------------------------------------------------------------------------(7) Malcolmia africana 7´Trichomes with thin cell walls. ------------------------------------------- (8) Horwoodia dichsoniae 7´´Trichomes with thick cell walls. ------------------------------------------------------------------------ 9 9´Smooth cell walls. -------------------------------------------------------------- (9) Corrichtera annua 9´´ Warty cell walls. --------------------------------------------------------------------------------------10 10´ Stomata anisocytic type. ------------------------------------------------------------------------------11
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10´´ Stomata anisocytic intermixed with Anomocytic type. ----------------------------------------12 11´Leaf epidermal cells with straight, thin anticlinal cell walls. --------- (10) Brassica tournefortii 11´´ Leaf epidermal cells with sinuous, thick anticlinal cell walls. ----------- (11) Cardaria draba 12´, Leafepidermal cells with straight, thin anticlinal cell walls. -------------- (12) Diplotaxis acris 12´´ Leaf epidermal cells with straight, thick anticlinal cell walls. ----------- (13) Diplotaxis harra 8´Trichomes, T – shaped branches, with thick warty cell walls. ------------------------------------13 13´ Leaf epidermal cells with straight, thin anticlinal cell walls. ---------- (14) Farsetia aegyptiaca 13´´Leaf epidermal cells with straight or sinuous, thin anticlinal cell walls. --------------------------------------------------------------------------------------------------------------- (15) Labularia maritima 13´´´ Leaf epidermal cells with sinuous, thin anticlinal cell walls. ----- (16) Notocera sparviflora 8´ Trichomes, many branches with thick warty cell walls. ------------------------------------------14 14´Leaf epidermal cells polygonal isodaimetric with straight, anticlinal cell walls. ------------15 14´´ Leaf epidermal cell polygonal elongated with thin anticlinal cell walls. -------------------16 15´ Thick anticlinal cell walls, Stomata anisocytic intermixed with Anomocytic type. ----------------------------------------------------------------------------------------------- (17) Anastatica hierochuntica 15´´Thin beaded anticlinal cell walls Stomata Anomocytic intermixed with rarely anisocytic type. ---------------------------------------------------------------------------------------- (18) Matthiola livida ------------------------------------------------------------------------------------ (19) Matthiola longipetala 16´, Sinuous, thin beaded anticlinal cell walls, stomata anisocytic intermixed with rarely anomocytic type. ------------------------------------------------------------- (20) Matthiola fruticulosa 16´´Sinuous, thin occasionally beaded anticlinal cell walls, stomata anomocytic intermixed with rarely anisocytic type. --------------------------------------------------------- (21) Morettia conescens 16´´´ Sinuous, thin unbeaded anticlinal cell walls, stomata anomocytic intermixed with anisocytic type. ----------------------------------------------------------------------------- (22) Morettia parviflora 8´´ T-shaped and many branched intermixed with thick smooth cell walls.- (23) Farsetia burtonae 17´Leaves hairy. ------------------------------------------------------------------------------------------18 17´´, Stem hairy. ------------------------------------------------------------------------------------------19 18´Nonglandulartrichomesunicellulerunbranched. ----------------------------------------------------20
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18´´Nonglandulartrichomesunicelluler branched. ------------------------------------------------------21 20´With thin smooth cell walls. ---------------------------------------------- (24) Coronopus niloticus 20´´With thick smooth cell walls. ------------------------------------------------------------------------22 21´T – shaped branched with thick warty cell walls. ------------------------- (25) Lepidium sativam 21´´Many branched with thick warty cell walls. ---------------------------- (26) Eremobium lineare 22´ Leaf epidermal cell walls thin beaded, stomata anisocytic type. ------ (27) Savignya parviflora 22´´ Leaf epidermal cell walls straight, thick beaded, stomata anisocytic intermixed with anomocytic type. -------------------------------------------------------------- (28) Schouwia thebaica 19´Nonglandulartrichomes, unicellular unbranched. -------------------------------------------------23 19´´Nonglandulartrichomes, unicellular branched, T - shaped branched, with thick warty cell walls. --------------------------------------------------------------------------- (29) Farsetia longisiliqua 23´With thick smooth cell walls. --------------------------------------------- (30) Schimpera arabica 23´´With thick warty cell walls. ------------------------------------------------------------------------24 24´Leaf epidermal cells polygonal elongated. --------------------------------------------------------25 24´´ Leaf epidermal cells polygonal isodaimetric with thin sinuous or straight, rarely beaded anticlinal cell walls. --------------------------------------------------------------- (31) Sinapis arvensis 25´With straight, thick beaded anticlinal cell walls, stomata anisocytic type. ------------------------------------------------------------------------------------------------------------ (32) Sisymbrium orientale 25´´With sinuous thick anticlinal cell walls. ----------------------------------------------------------26 26´Stomata anisocytic intermixed with rarely anomocytic type. ----- (33) Sisymbrium erysimoides 26´´Stomata anisocytic type. ------------------------------------------------------ (34) Sisymbrium irio.
References Adedeji, O., Dloh, H.C. (2004). Comparative foliar anatomy of ten species in the genus Hibiscus Linn.in Nigeria. New Bot. 31, 147 – 180. Ahmad, K., Khan, M.A., Ahmad, S., Nazir, N. A., (2010).Taxonomic diversity in epidermal cells of some sub-tropical plant species. Int. Agric. Biol. 12, 115 – 118.
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Celka, Z., Szkudlarz, P., Bierezonoj, U. (2006). Morphological variation of hairs in MalvaalceaL. (Malvceae). Biodiver. Res. Conserv. 3, 258 – 261. Chaudhary, S.A. (1999). Flora of the Kingdom of Saudi Arabia,Vol.1, National Agriculture & water Research Center, Ministry of Agriculture &Water, Kingdom of Saudi Arabia. Doaigey, A.R., Gawad, H.A., Meligy, A.M. (1985). Morphology and types of trichomes in some species of Lamiaceae, Proce. Saudi Biol. Soc. 8, PP. 189-203. Gostin, I.N. (2009). Anatomical and micro morphological particularities of vegetative organs in endemic Errysimum wittmanii Zaw. ssp. Wittmanii. Tom. XVI, 74 – 79. Inamdar, J.A., Rao, N.V. (1983). Light and scanning electron microscopic on trichomes of some Brassicaceae. Feddes Rep. 94, 183- 190. Kadry, A. Kh. (2005). Morphological studies on trichomes of Brassicaceae in Egypt and taxonomic significance. Acta Bot. Croat. 64(1), 57 – 73. Khan, Y.K., Khan, M.A., Ahmad, M., Shah, G.M., Zafer, M., Niamat, R., Munir, M., Abbasi, A.M., Fazal, H., Mazri, P., Seema, N. (2011). Foliar epidermal anatomy of some ethnobotanically important species of genus Ficus Linn. J. Mid.Pl. Res. 5(9), 1627-1638. Metcalfe, C.R., Chalk, L. (1950). Anatomy of Dicotyledons. Vol. 1 Clarendon Press, Oxford. Stace, C.A. (1984). The taxonomic importance of leaf surface. In: Heywood, V.H. and D.M. Moore (eds.), Current Concepts in plant Taxonomy, pp: 67 – 94. London, Academic Press, London. SUN, Z. Y., LI, F. Z. (2008). Studies on the leaf epidermal micromorphology of Sisymbrium (Brassicaceae) from China. Bull. Bot. Res, 28(1), 20-24. Sun, Z.Y., Li, F.Z. (2007). Studies on the leaf epidermal features of Lepidium (Brassicaceae) from China. Zhongyaocai= Journal of Chinese medicinal materials, 30(7), 780-785. Tuo, Zhong-yun, Zhou, Gui-Ling. (2010). Micromorphologic characteristic of leaf epidermis of eight species in Brassicaceae (Cruciferae). Journal of Xinjiang Agricultural University. 1. Yasmin, G., Khan M.A., Shaheen N., Hayat, M.Q. (2009). Micromorphological investigation of foliar anatomy of genera Aconogonon and Bistorta of family Polygonaceae, Int. J. Agric. Biol. 11, 285 – 289. 210
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Zhang, Y., He, P., Zhang, C., He, j. (2009). Leaf epidermal features of 21 medicinal plants of Euonymus in China. Zhong Guo Zhong Yao ZaZhi. 12, 14937. Zou, P., Liao, J., Zhang D. (2008). Leaf epidermal micromorphology of Cercis (Fabaceae: Caesalpinioideae). Bot. J. Linn. Soc. 158, 539 - 547.
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Table1: Sources of plant materials used in this study. Specimen
KSU
No.
Species
1
Anastatica hierochuntica L.
2
Brassica tournefortii Gouan.
√
19919
3
Cardariadraba (L.) Desv.
√
20773
Dry
Fresh √
herbarium
Collector
Place
Ihsan El-Habashy
Riyadh
no. 285
Al-Turi
&
J.
Al-Baha
Thomas J. Thomas Migahid&
Al-Kharj El-
Magmaa
4
Carrichtera annua (L.) DC.
√
2987
5
Coronopus niloticus (Del.) Spreng.
√
14370
J. Thomas
Dillam
6
Diplotaxis acris (Forssk) Boiss
√
13106
Abu Haila, A.
Wadi El-Jaffi
7
Diplotaxis harra (Forssk) Boiss
√
13117
Ihsan El-Habashy
Riyadh
√
13209
Doaigey
8
Eremobium
aegyptiacum
(Spreng)
Boiss
Sheikh
Riyadh
9
Eruca sativa Mill
√
13243
Ihsan El-Habashy
Riyadh
10
Erucaria hispanica (L.) Druce
√
1338
Doaigey
Riyadh
11
Farsetiaa egyptiaca Turra
√
13355
Ihsan El-Habashy
Riyadh
12
Farsetia burtonae Oliv.
√
13392
Idris, S.
Riyadh
13
Farsetia longisiliqua Decne
√
13417
El-Sheikh
Bashawat
14
Horwoodia dichsoniae Turrill
13462
Ihsan El-Habashy
Riyadh
15
Isatis lusitanica L.
√
19457
J. Thomas
Al-Kharj
16
Lepidium sativum L.
√
3297
Mustafa, H.
Taif
17
Leptaleum filifolium (willd.) DC.
Al Farhan & J.
Raudhat
Thomas
khuraim
18
Lobularia maritima (L.) Desv.
14809
J. Thomas
Riyadh
19
Malcolmia africana (L.) R. Br.
√
14838
Ihsan El-Habashy
Riyadh
20
Matthiola fruticulosa (L.) Maire
√
-
Doaigey
Riyadh
21
Matthiola livida (Del.) DC.
√
13515
Doaigey
Riyadh
22
Matthiola longipetala (Vent.) DC.
√
19521
Doaigey
Riyadh
23
Morettia canescens Boiss.
√
13527
Doaigey
√
10199 √
√
Riyadh El-
Hail
24
Morettia parviflora Boiss.
25
Moricandia sinaica (Boiss) Boiss.
√
13567
Ihsan El-Habashy
26
Notoceras bicorne (Aiton) Amo
√
19529
Doaigey
27
Savignya parviflora (Del.) Webb
√
13594
Ihsan El-Habashy
Riyadh
28
Schimpera arabica Hochst. & Steud
√
13633
Doaigey
Riyadh
29
Schouwia thebaica Webb
√
13662
Basalah, M.
Khurais Rd.
30
Sinapis arvensis L.
√
31
Sisymbrium erysimoides Desf.
32
Sisymbrium irio L.
33
Sisymbrium orientale L.
34
Zilla spinosa (L.)
212
13532
Migahid& Sheikh
Riyadh Wadi
Al-
Haier
1642
Chaudary, Sh.
Al-Harra
√
554
Ihsan El-Habashy
Riyadh
√
290
Ihsan El-Habashy
Riyadh
13685
Migahid
Jabal Soodah
12241
Ihsan El-Habashy
Riyadh
√ √
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Vol 20, No. 9;Sep 2013
Table 2: Leaf Lamina epidermal characters. Epidermis cell shape
Character
Epidermal cell wall UE
s
Cuticle
Stomata
Nonglandular unicellular Trichome
LE
PI
PE
PI
PE
Str
Sin
T
Th
B
Sm
Stri
Ani
Ano
TWT
TWTh
UBr
Br
BrT
BrM
SW
WW
+
-
+
-
+
-
[+]
+
+
+
-
+
(+)
-
+
-
+
-
+
-
+
Brassica tourneforti
+
-
+
-
+
-
+
-
-
+
-
+
-
-
+
+
-
-
-
-
+
Cardaria draba
-
+
+
-
-
+
-
+
-
+
(+)
+
-
-
+
+
-
-
-
-
+
Coronopus niloticus
+
-
+
-
-
+
+
-
-
-
+
+
+
+
-
+
-
-
-
+
-
Corrichtera annua
-
+
-
+
+
(+)
+
-
-
+
-
+
+
-
+
+
-
-
-
+
-
Diplotaxis acris
+
+
+
-
+
-
+
-
(+)
+
-
+
(+)
-
+
+
-
-
-
-
+
Diplotaxis harra
+
-
+
-
+
-
-
+
+
+
-
+
(+)
-
+
+
-
-
-
-
+
Eremobium lineare
(+)
+
(+)
+
+
+
+
-
+
+
-
+
(+)
-
+
-
+
-
+
-
+
Eruca sativa
+
-
+
-
+
-
+
-
(+)
+
-
+
-
-
-
-
-
-
-
-
-
Erucaria hispanica
+
-
+
-
+
-
+
-
+
+
-
+
(+)
-
-
-
-
-
-
-
-
Farsetia aegyptiaca
-
+
-
+
+
-
+
-
-
+
-
+
(+)
-
+
-
+
+
-
-
+
Farsetia burtonae
+
-
+
-
-
+
+
-
-
+
-
+
-
-
+
-
+
+
(+)
+
-
Farsetia longisiliqua
+
-
+
-
-
+
+
-
-
+
-
+
-
-
-
-
-
-
-
-
-
Horwoodiadichsoniae
+
-
+
-
+
-
-
+
+
+
-
+
-
+
-
+
-
-
-
+
-
Isatis lusitanica
+
-
+
-
-
+
+
-
+
+
-
+
-
-
-
-
-
-
-
-
-
Labularia maritima
-
+
-
+
+
(+)
+
-
-
+
-
+
+
-
+
-
+
+
-
-
+
Lepidium sativam
-
+
-
+
-
+
+
-
-
+
-
+
-
-
+
-
+
+
-
-
+
Leptaleum filifolium
-
+
-
+
(+)
+
+
-
-
+
-
+
+
-
-
-
-
-
-
-
-
Malcolmia Africana
+
-
+
-
-
+
+
+
-
+
-
[+]
+
-
+
+
+
+
-
+
-
Matthiola fruticulosa
-
+
-
+
-
+
+
-
(+)
+
(+)
+
[+]
-
+
-
+
-
+
-
+
Matthiola livida
+
-
+
-
+
+
+
-
-
+
-
[+]
+
-
+
-
+
-
(+)
+
-
Matthiola longipetala
+
-
+
-
+
+
+
-
(+)
+
-
[+]
+
-
+
-
+
-
+
+
-
Morettia conescens
-
+
-
+
-
+
+
-
(+)
+
-
[+]
+
-
+
-
+
-
+
-
+
Morettia parviflora
-
+
-
+
-
+
+
-
-
+
-
+
+
-
+
-
+
-
+
-
+
Moricandia sinaica
+
(+)
+
(+)
+
-
-
+
-
+
-
+
-
-
-
-
-
-
-
-
-
Notocera bicorne
-
+
-
+
-
+
+
-
(+)
+
-
+
+
-
+
-
+
+
-
-
+
Savignya parviflora
-
+
-
+
+
(+)
+
(+)
(+)
+
-
+
-
-
+
(+)
-
-
-
+
-
Schimpera arabica
-
+
-
+
+
(+)
+
-
+
+
-
+
(+)
-
-
-
-
-
-
-
-
-
+
-
+
-
+
-
+
-
+
-
+
(+)
-
-
-
-
-
-
-
-
Sisymbrium irio
-
+
-
+
-
+
-
+
(+)
+
-
+
-
-
-
-
-
-
-
-
-
Sisymbrium orientale
-
+
-
+
+
-
-
+
+
+
-
+
-
-
-
-
-
-
-
-
-
Sinapis arvensis
+
-
+
-
(+)
+
+
-
(+)
+
-
+
+
-
-
-
-
-
-
-
-
Schouwia thebaica
-
+
-
+
+
-
-
+
(+)
+
-
+
+
-
+
(+)
-
-
-
+
-
Zillaspinosa
-
+
-
+
+
-
+
-
-
+
-
+
-
-
-
-
-
-
-
-
-
Species Anastatica hierochantica
Sisymbrium erysimoides
UE = Upper epidermis ;LE = Lower epidermis; PI = Polygonal Isodiametric; PE = Polygonal elongated ; Str = Straight anticlinal walls; Sin = Sinuous anticlinal walls; T
= Thin walls; Th = Thick walls; B = Beaded ; Sm = Smooth cuticle ; Stri = Striated cuticle; Ani =
Anisocytic stomata ; Ano = Anomocytic stomata ;UBr = Unbranched trichomes ;TW= Thin Warty ; TWT= Thin Warty walls Br= Branched trichomes ; BrT = T- shaped ; BrM = Many branched ; SW = Smooth walls ; WW = Warty walls; +
= Present ; - = Absent ; (+) =
Rarely present; ]+[ = Numerous
213
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Table 3: Stem, epidermal characters. Epidermis Character Species
s
cell shape
Epidermal cell wall
Cuticle
Stomata
Nonglandular unicellular Trichome
PI
PE
Str
Sin
T
Th
B
Sm
Stri
Ani
Ano
TWT
TWTh
UBr
Br
BrT
BrM
SW
WW
Anastatica hierochantica
-
+
+
-
-
+
+
+
-
+
-
-
+
-
+
-
+
-
+
Brassica tourneforti
-
+
+
-
-
+
-
+
-
+
-
-
+
+
-
-
-
-
+
Cardaria draba
-
+
+
-
-
+
(+)
+
-
+
-
-
+
+
-
-
-
-
+
Coronopus niloticus
-
+
+
-
-
+
-
+
(+)
+
-
-
-
-
-
-
-
-
-
Corrichtera annua
-
+
+
-
+
-
-
+
-
+
+
-
+
+
-
-
-
-
+
Diplotaxis acris
-
+
+
-
+
(+)
(+)
+
-
+
-
-
+
(+)
-
-
-
-
+
Diplotaxis harra
-
+
+
-
-
+
+
+
-
+
-
-
+
(+)
-
-
-
-
+
Eremobium lineare
-
+
+
-
-
+
-
+
-
+
(+)
-
-
-
-
-
-
-
-
Eruca sativa
-
+
+
-
-
+
+
+
-
+
-
-
-
-
-
-
-
-
-
Erucaria hispanica
-
+
+
-
+
-
-
+
-
+
-
-
-
-
-
-
-
-
-
Farsetiaa egyptiaca
-
+
+
-
+
-
-
+
-
+
-
-
+
-
+
-
-
-
+
Farsetia burtonae
-
+
+
-
+
(+)
-
+
-
+
+
-
+
-
+
+
-
-
+
Farsetia longisiliqua
-
+
+
-
+
(+)
-
+
-
+
-
-
+
-
+
+
-
-
+
Horwoodia dichsoniae
-
+
+
-
-
+
-
+
-
+
-
+
-
+
-
-
-
+
-
Isatis lusitanica
-
+
+
-
+
-
(+)
+
-
+
+
-
-
-
-
-
-
-
-
Labularia maritima
-
+
+
-
+
-
-
+
-
+
+
-
+
-
+
+
-
-
+
Lepidium sativam
-
+
+
-
-
+
-
+
-
+
-
-
-
-
-
-
-
-
-
Leptaleum filifolium
-
+
+
+
+
-
+
+
-
+
(+)
-
-
-
-
-
-
-
-
Malcolmia africana
-
+
+
-
-
+
-
+
-
+
(+)
-
+
-
+
+
+
-
+
Matthiola fruticulosa
-
+
+
-
-
+
-
+
-
+
(+)
-
+
-
+
-
+
-
+
Matthiola livida
-
+
+
-
+
-
-
+
-
+
(+)
-
+
-
+
-
+
-
+
Matthiola longipetala
-
+
+
-
+
-
-
+
-
+
(+)
-
+
-
+
-
+
-
+
Morettia conescens
-
+
+
-
+
(+)
+
+
-
+
-
-
+
-
+
-
+
-
+
Morettia parviflora
-
+
+
-
-
+
-
+
-
+
+
-
+
-
+
-
+
-
+
Moricandia sinaica
-
+
+
-
-
+
+
+
-
+
+
-
-
-
-
-
-
-
-
Notocera bicorne
-
+
+
-
+
-
-
+
-
+
+
-
+
+
+
+
-
-
+
Savignya parviflora
-
+
+
-
-
+
(+)
+
-
+
-
-
-
-
-
-
-
-
-
Schimpera arabica
-
+
+
-
-
+
-
+
-
+
+
-
+
+
-
-
-
+
-
Sisymbrium erysimoides
-
+
+
-
-
+
-
+
-
+
+
-
+
(+)
-
-
-
-
+
Sisymbrium irio
-
+
+
-
+
-
(+)
+
-
+
+
-
+
+
-
-
-
-
+
Sisymbrium orientale
-
+
+
-
+
-
+
+
-
+
-
-
+
+
-
-
-
-
+
Sinapis arvensis
-
+
+
-
+
-
-
+
-
+
-
-
+
(+)
-
-
-
-
+
Schouwia thebaica
-
+
+
-
+
-
-
+
-
+
(+)
-
-
-
-
-
-
-
-
Zilla spinosa
-
+
+
-
+
-
-
+
-
+
-
-
-
-
-
-
-
-
-
PI = Polygonal Isodiametric; PE = Polygonal elongated; Str = Straight anticlinal cell walls; Sin = Sinuous anticlinal cell walls; T = Thin cell walls; Th = Thick cell walls; B = Beaded; Sm = Smooth cuticle ; Stri = Striated cuticle; Ani = Anisocytic stomata ; Ano = Anomocytic stomata; TWT= Trichome thin cell wall; TWTh= Trichome thick cell wall ; UBr = Unbranched trichomes; Br = Branched trichomes ; BrT = T- shaped ; BrM = Many branched ; SW = Smooth cell walls ; WW = Warty cell walls; + = Present ; - = Absent ; (+) = Rarely present; ]+[ = Numerous.
214
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Table 4: analysis of data: Means measurements of epidermal cells and stomata (N. = number, St. = stomata, E = epidermal cells). Levels not connected by same letter are significantly different. Species
N. stomata/
St. length
St. WIDTH
E. Length
E. Width
Anastatica hierochantica
30.32 KLM
31.50 BC
22.80 BCD
91.58 EFGHIJ
46.00 CDEFG
Brassica tourneforti
43.0 8C
19.90 L
11.30 R
87.20 HIJK
33.90 O PQR
Cardaria draba
39.04 EF
28.00 DE
17.20 LMN
83.00 JKLM
35.80 MNOPQ
Coronopus niloticus
28.92 MN
20.60 KL
17.00 LMNO
54.70 P
29.50 RST
Corrichtera annua
32.08 JKL
26.30 EFGH
22.40 BCDE
83.60 IJKLM
41.20 GHIJK
Diplotaxis acris
34.16 IJ
30.00 C
18.20 JKL
94.60 DEFGHI
45.10 DEFGH
Diplotaxis harra
37.84 FG
27.70 DE
17.30 LMN
99.00 DEFG
47.70 BCDE
Eremobium lineare
25.12 O
27.10 DEF
17.10 LMN
86.08 HIJKL
48.88 BCD
Eruca sativa
38.20 EFG
21.70 K
16.80 LMNO
89.76 FGHIJ
51.40 B
Erucaria hispanica
37.64 FGH
24.50 IJ
15.50 OPQ
85.60 HIJKL
46.30 CDEF
Farsetia aegyptiaca
25.52 O
31.10 B C
21.00 EFG H
90.08 FGH I J
38.00 KLMNO
Farsetia burlonae
38.36 EFG
24.70 H I J
21.30 DEFG
73.80 MNO
28.60 S T
Farsetia longisiliqua
21.84 P
24.20 I J
19.90 GHI
95.60 DEFGH
39.80 IJKLM
Horwoodia dichsoniae
22.12 P
24.20 I J
23.80 AB
137.10 B
59.60 A
Isatis lusitanica
34.68 HIJ
20.20 I J
19.60 IJ
81.50 JKLM
37.50 KLMNO
Labularia maritima
15.52 Q
25.70 FGH I
20.30 FG HI
85.74 HIJKL
28.78 S T
Leptaleum filifolium
30.02 KLM
24.98 GHI J
19.70 HIJ
147.20 B
26.80 T
Lepidium sativam
42.64 CD
20.40 K L
15.80 NOPQ
112.40 C
42.00 FGHIJK
Malcolmia africana
47.84 B
23.70 J
16.00 N O P
75.90 LMN
39.30 J KLMN
Matthiola fruticulosa
35.48 GHI
33.70 A
23.90 AB
103.60 CD
44.40 DEFGHI
Matthiola livida
32.68 IJK
28.10 D
24.90 A
99.20 DEF
50.40 BC
Matthiola longipetala
19.92 P
32.00 A B
2320 BC
160.22 A
47.90 BCD
Morettia conescens
40.96 CDE
23.50 J
17.90 KLM
70.20 N O
40.70 HIJKL
Morettia parviflora
22.04 P
20.10 K L
15.10 PQ
84.70 HJKLM
40.10 IJKLM
Moricandia sinaica
40.00 DEF
25.200GHIJ
21.60 DEF
89.64 FGH IJ
57.40 A
Notocera bicorne
26.92 NO
24.10 IJ
21.20 EFGH
64.40 O P
34.90 NOPQ
Savignya parviflora
39.16 EF
27.10 DEF
22.00 CDE
91.60 EFGHIJ
45.60 CDEFG
Schimpera arabica
54.20 A
20.80 K L
16.70 LMNO
70.40 N O
39.20 LKLMN
Sisymbrium erysimoides
56.68 A
21.00 KL
16.40 MNOP
78.10 KLMN
31.10 Q R S T
Sisymbrium irio
46.12 B
19.60 L
14.30 Q
74.30 MNO
26.80 T
Sisymbrium orientale
29.62 KLMN
25.60 FGHI
19.90 GHI
101.30 DE
43.00 EFGHI J
Sinapis arvensis
29.44 L M N
24.50 I J
20.10 FGHI
81.50 JKLM
31.36 PORST
Schouwia thebaica
29.80 KLMN
24.40 I J
19.10 IJK
83.80 IJKLM
36.10 LMNOP
Zilla spinosa
39.52 EF
26.50 DEFG
23.70 AB
88.10 GHIJK
31.90 PQRS
215
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Figures 1- 6 (L.M. photographs) Leaf and stem epidermises:1) Anastatica hierochantica: Leaf epidermis shows cells with thin, straight anticlinal cell walls, stomata types. 2) Matthiola longipetala: Leaf epidermal cells with thin, sinuous anticlinal cell walls. 3) Diplotaxis harra: Leaf epidermis cells with thick, sinuous anticlinal cell walls. 4) Sisymbrium orientale: Leaf epidermal cells with thin sinuous beaded anticlinal cell walls 5) Diplotaxis harra: Stem epidermal cells with thick straight beaded anticlinal cell walls, 6) Schimpera arabica: Stem epidermal cells with thick straight anticlinal cell walls. Figs.1,2,x300; 3,x200; 4,x500; 5,x540; 6,x200.
216
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Vol 20, No. 9;Sep 2013
Figures 7 - 12 (L.M. photographs), nonglandular unicellular trichomes: 7) Brassica tourneforti: Stem epidermis shows unbranched with thick warty cell walls. 8) Cardaria draba: Leaf epidermis shows unbranched with thick smooth cell walls. 9) Horwoodia dichsoniae: Stem epidermis shows unbranched with thin warty cell walls.10 and 11) Malcolmia Africana: Leaf epidermis shows Y- shaped and multi-branched with thick smooth cell walls. 12) Morettia canescens: Stem epidermis shows multi-branched with thick warty cell walls. Figs. 7,x160; 8,x250;9,x250; 10, x125; 11, x250; 12,x320.
217
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Vol 20, No. 9;Sep 2013
Figures. 13 – 16 (SEM photographs) Nonglandular unicellular trichomes: 13 - 14) Anastatica hierochantica: Leaf epidermis shows many branches with thick warty cell walls; 15) Diplotaxis harra: Leaf epidermis shows unbranched with thick warty cell walls; 16) Morettia canescens: Leaf epidermis shows T- shaped branched with thick warty cell walls.
218
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Vol 20, No. 9;Sep 2013
60 50
Number of stomata /unit of area
Stomata
40 30 20 10 0
Species Figure (17): Number of stomata / unit of area. (0.23 mm2)
219
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Vol 20, No. 9;Sep 2013
40 35
stomata Length
Stomata Width
Mesuerment
30 25 20 15 10 5 0
Species
Figure 18: Stomata measurements (length and width).
220
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Vol 20, No. 9;Sep 2013
180 160
Epidermal cells Length Epidermal cells Width
140
100 80 60
(µm)
Measurements
120
40 20 0
Species Figure 19: Epidermal cells measurements (length and width).
221
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