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POLLEN AND SEED MORPHOLOGY OF CRAMBE SPECIES OF TURKEY.

Byline: A. S. Blbl, B. Tarikahya-Hacioglu, Y. Arslan and I. Subasi

ABSTRACT

Our study includes 5 taxa out of 10 naturally growing taxa in Turkey: C. orientalis L. subsp. orientalis, C. orientalis L. subsp. sulphurea (Stapf ex O.E.Schulz) Prina, C. tataria Sebek var. tataria, C. tataria Sebek var. aspera (M.Bieb.) Boiss. and C. maritima L. The morphological characters of the pollen and seeds from the studied species of Crambe were given. Data obtained by light microscope, stereomicroscope and scanning electron microscope were explained and summarized in Tables. As a result of our study, there are high similarities among taxa in means of pollen grains. Pollen shape is suprolate to prolate-spheroidal. Crambe orientalis subsp. orientalis had the biggest pollen grains with 25.90 m polar axis and 21.46 m equatorial axis lengths, while Crambe orientalis subsp. sulphurea had the smallest-sized pollen grains with 22.75 m polar axis and 21.07 m equatorial axis lengths. Seeds are brown to dark brown, elliptic to orbicular, with reticulate surface.

Crambe maritima had the biggest seeds with 2.8-4.1 x 2.3-3.4 mm, while Crambe orientalis subsp. sulphurea had the smallest-sized seeds with 2.0-2.6 x 1.7-2.4 mm. Since the group has stenopalynous pollens, our investigation supports the use of seed surface patterns as a diagnostic characteristic for species level instead of pollen morphology.

Key words: Crambe, light microscopy, pollen grains, scanning electron microscopy, seed coat morphology.

INTRODUCTION

Brassicaceae distributed mainly in the temperate regions of northern hemisphere (Hedge, 1976), comprises about 338 genera and 3709 species (Franzke et al.,2010). The genus Crambe (Brassicaeae) has approximately 35 taxa worldwide. Crambe species are large coarse-growing perennial or biennial herbs with large tap roots. With the globose, 1-seeded, indehiscent fruit structure it is easy to recognize the genus. In flora of Turkey, there are 6 species and 4 infraspecific taxa (Mutlu, 2012).

These taxa are Crambe alutacea Hand.-Mazz., C. grandiflora DC., C. hispanica L., C. maritima L., C. orientalis L. subsp. orientalis var. orientalis, C. orientalis L. subsp. orientalis var. dasycarpa O.E.Schulz, C. orientalis L. subsp. sulphurea (Stapf ex O.E.Schulz) Prina, C. tataria Sebek var. tataria, C. tataria Sebek var. aspera (M.Bieb.) Boiss., C. tataria Sebek var. parviflora (Hub.-Mor. and Reese) Hedge and Hub.-Mor. Our study includes 5 taxa out of 10: C. orientalis subsp. orientalis, C. orientalis subsp. sulphurea, C. tataria var. tataria, C. tataria var. aspera. and C. maritima.

Brassicaceae is a stenopalynous family, the pollen grains usually tricolpate, with a reticulate exine (Abdel-Khalik, 2002). Pollen morphology of Brassicaceae was investigated by several authors (Abdel-Khalik, 2002; Perveen et al., 2004; Bolurian, 2009; Pinar et al., 2009; Arora and Modi, 2011; Keshavarzi, et al. 2012; Mutlu and Erik, 2012). Pollen morphology can be used to discuss the systematic relationships among the genera of the Brassicaceae (Inceoglu and Karamustafa, 1977; Dogan and Inceoglu, 1990; Brochmann, 1992; Pinar et al., 2009). Seed coat morphology is known to be an important character for taxonomic and evolutionary studies (Koul et al., 2000; Abdel-Khalik, 2002; Tantawy et al., 2004; Karcz et al., 2005; Moazzeni et al., 2007; Pinar et al., 2007, 2009; Kaya et al., 2011; Mousavi and Sharifi-Rad, 2014). Our study provides information about the pollen and seed morphology of the available Turkish Crambe taxa by using light and electron microscopy.

MATERIALS AND METHODS

Plant material: Specimens were collected in 2011 and 2012 for the General Directorate of Agricultural Research and Policy project "The genetic and morphological characteristics of Crambe L. species found in Anatolian natural flora and their facilities to be improvement materials". List of investigated species and localities were given in "Table 1". Our study includes 5 taxa out of ten naturally growing taxa in Turkey: C. orientalis subsp. orientalis, C. orientalis subsp. sulphurea, C. tataria var. tataria, C. tataria var. aspera and C. maritima.

Light microscopy (LM): According to Wodehouse method (1935), pollen samples were taken from plants and slides were prepared by using basic fuchsine mixture. At least 30 pollen samples for each species were examined in LMand micrographs were taken with Leica DM 750 digital imaging system. After the examination of each species, measurements were done for colpus length and thickness, pore length and thickness, pollen shape, exine thickness, intine thickness, polar axis length, equatorial axis length of pollen. The pollen terminology adopted by Faegri and Iversen (1975) and Brochman (1992), and shape classification follows Erdtman (1969) based on P/E ratio in "Table 2". At least 30 seed samples for each species were examined in stereomicroscope and micrographs were taken with Olympus SZ2-LGB digital imaging system.

After the examination of each species, measurements were done forseed shape, color, size, epidermal cell shape, anticlinal cell wall, periclinal cell wall. The seed terminology was used as Murley (1951), Koul et al. (2000) and Karcz et al. (2005).

Scanning Electron Microscopy (SEM): For SEM, pollen grains and seeds were directly mounted on stubs and covered with gold. The surface ornamentations of pollen grains and seeds were examined in detail with JeolTescan MAIA3 XMU model electron microscope.

RESULTS AND DISCUSSION

The morphological characters of the pollen and seeds from the studied species of Crambe were given below. Data obtained by light microscope, stereomicroscope and electron microscope were reviewed in "Tables 2 and 3", as well as in "Figures 1, 2 and 3".

Pollen Morphology

Crambe orientalis subsp. orientalis: The pollen grains are tectate or semitectate, radially symmetric, isopolar, tricolpate, prolate-spheroidal,outline is elliptic in the equatorial view and circular in the polar view "Figure 1A and B, Figure 2A and B". Polar axis is 25.90 m and equatorial axis 21.46 m.The colpus length is 17.07 m and the colpus width is 4.77 m. The mesocolpium length is 8.00 m. The apocolpium length is 5.94 m. Exine ornamentation is reticulate. The exine is 2.69 m thick and the ectexine is thicker than the endexine. Intine thickness is 0.53 m. The lumen width is 1.35 m, regular and polygonal or rarely irregular lumina. Murus width is 0.44 m, generally 4-5 sided and smooth muri "Table 2; Figure 1A and B, Figure2A and B".

Crambe orientalis subsp. sulphurea: The pollen grains are tectate or semitectate, radially symmetric, isopolar, tricolpate, prolate-spheroidal, outline is elliptic in the equatorial view and circular in the polar view "Figure1C and D, Figure2C". Polar axis is 22.75 m and equatorial axis 21.07 m. The colpus length is 16.62 m and the colpus width is 4.68 m. The mesocolpium length is 8.03 m. The apocolpium length is 5.64 m. Exine ornamentation is reticulate. The exine is 2.80 m thick and the ectexine is thicker than the endexine. Intine thickness is 0.54 m. The lumen width is 2.57 m, regular and polygonal or rarely irregular lumina. Murus width is 0.38 m, generally 4-5 sided and smooth muri "Table 2; Figure1C and D, Figure2C".

Crambe tataria var. tataria: The pollen grains are tectate or semitectate, radially symmetric, isopolar, tricolpate, subprolate, outline is elliptic in the equatorial view and circular in the polar view "Figure 1E and F, Figure 2D". Polar axis is 24.84 m and equatorial axis 21.76 m. The colpus length is 18.73 m and the colpus width is 4.73 m. The mesocolpium length is 7.24 m. The apocolpium length is 5.53 m. Exine ornamentation is reticulate. The exine is 2.52 m thick and the ectexine is thicker than the endexine. Intine thickness is 0.51 m. The lumen width is 1.03 m, regular and polygonal or rarely irregular lumina. Murus width is 0.32 m, generally 4-5 sided and smooth muri "Table 2; Figure 1E and F, Figure 2D".

Crambe tataria var. aspera: The pollen grains are tectate or semitectate, radially symmetric, isopolar, tricolpate, prolate-spheroidal, outline is elliptic in the equatorial view and circular in the polar view "Figure 1G and H, Figure 2E and F". Polar axis is 23.47 m and equatorial axis 20.81 m. The colpus length is 16.98 m and the colpus width is 4.50 m. The mesocolpium length is 7.96 m. The apocolpium length is 5.53 m. Exine ornamentation is reticulate. The exine is 2.48 m thick and the ectexine is thicker than the endexine. Intine thickness is 0.51 m. The lumen width is 1.34 m, regular and polygonal or rarely irregular lumina. Murus width is 0.42 m, generally 4-5 sided and smooth muri "Table 2; Figure 1G and H, Figure 2E and F".

Crambe maritima: The pollen grains are tectate or semitectate, radially symmetric, isopolar, tricolpate, subprolate, outline is elliptic in the equatorial view and circular in the polar view "Figure 1I and J, Figure 2G and H". Polar axis is 25.23 m and equatorial axis 21.14 m. The colpus length is 16.86 m and the colpus width is 4.73 m. The mesocolpium length is 8.34 m. The apocolpium length is 5.18 m. Exine ornamentation is reticulate. The exine is 2.68 m thick and the ectexine is thicker than the endexine. Intine thickness is 0.53 m. The lumen width is 1.36 m, regular and polygonal or rarely irregular lumina. Murus width is 0.33 m, generally 4-5 sided and smooth muri "Table 2; Figure 1I and J, Figure 2G and H".

Seed Morphology

Crambe orientalis subsp. orientalis: The seeds are 2.3-3.9 x 1.9-3.6 mm. Seed shape varies from elliptic, with smooth surface. Seed color varies from brown or dark brown. SEM showed reticulate surface pattern with regular to irregular polygonal, epidermal cell shape varies from isodiametric, raised, straight; smooth anticlinal cell walls and flat to concave: macro-reticulate periclinal cell walls "Figure 3A and B".

Crambe orientalis subsp. sulphurea: The seeds are 2.0-2.6 x 1.7-2.4 mm. Seed shape varies from orbicular, with smooth surface. Seed color varies from brown or dark brown. SEM showed reticulate surface pattern with irregular polygonal mostly hexagonal, epidermal cell shape varies from isodiametric, raised, straight; smooth anticlinal cell walls and flat to slightly concave periclinal cell walls "Figure 3C and D".

Crambe tataria var. tataria: The seeds are 2.2-3.4 x 2.1-3.3 mm. Seed shape varies from orbicular, with smooth surface. Seed color varies from brown or dark brown. SEM showed reticulate surface pattern with regular polygonal mostly hexagonal, epidermal cell shape varies from isodiametric, raised, straight; smooth anticlinal cell walls and flat to concave: macro-reticulate periclinal cell walls "Figure 3E and F".

Crambe tataria var. aspera: The seeds are 2.8-4.1 x 2.3-3.4 mm. Seed shape varies from orbicular, with smooth surface. Seed color varies from brown or dark brown. SEM showed reticulate surface pattern with regular polygonal mostly hexagonal, epidermal cell shape varies from isodiametric, raised, straight; smooth anticlinal cell walls and flat to concave: macro-reticulate periclinal cell walls "Figure 3G and H".

Crambe maritima: The seeds are 3.3-4.9 x 2.5-3.8 mm. Seed shape varies from elliptic, with smooth surface. Seed color varies from brown to dark brown. SEM showed reticulate surface pattern with regular 4, 5, 6 gonal, epidermal cell shape varies from isodiametric, raised, straight; smooth anticlinal cell walls and flat to slightly concave periclinal cell walls "Figure 3I and J".

Brassicaceae is a stenopalynous family (Erdtman, 1952), pollen grains 3-zonocolpate, generally prolote to subprolate oblate to oblate-spheroidal, or prolate-spheroidal with reticulate or granulate exine ornamentation (Abdel-Khalik, 2002).Our investigation supports Erdtman (1952) since there are high similarities among Crambe pollen grains. Crambe orientalis subsp. orientalis had the biggest pollen grains with 25.90 m polar axis and 21.46 m equatorial axis lengths, while Crambe orientalis subsp. sulphurea had the smallest-sized pollen grains with 22.75 m polar axis and 21.07 m equatorial axis lengths. Crambe maritima had the biggest seeds with 2.8-4.1 x 2.3-3.4 mm, while Crambe orientalis subsp. sulphurea had the smallest-sized seeds with 2.0-2.6 x 1.7-2.4 mm.

There are many researches on pollen morphology of Brassicaceae family. According to pollen studies of Inceoglu and Karamustafa (1977), Crambe tataria was observed with radially symmetric, isopolar, tricolpate, subprolate pollen grains and the outline shape was elliptic in the equatorial view and circular in the polar view. Polar axis was 25.80 m and equatorial axis was 21.60 m. The colpus length was 19.60 m and the colpus width was 5.00 m. Exine ornamentation was reticulate. The exine was 2.1 m thick and the ectexine was thicker than the endexine. Intine thickness was 0.6 m. The lumen width was 1.4 m. The measurements are similar to our results.

The pollen morphology of Crambe cordifolia Steven, istectate, radially symmetric, isopolar, tricolpate, prolate-spheroidal, outline is elliptic in the equatorial view and circular in the polar view. Polar axis is 20.5 m and equatorial axis 20.0 m. The colpus length is 13.75 m and colpus long sunken with acute ends. Exine ornamentation is reticulate. The exine is 2.5 m thick and the sexine is thicker than nexine (Perveen et al., 2004). The pollen size of the investigated species' in our research was larger in polar axis (25.90-22.75 m) and equatorial axis (21.46-21.07 m).

According to pollen morphology studies on selected taxa of Brassicaceae by (Pinar et al., 2009), the ornamentation of Hesperis L. isoblate-spheroidal, spheroidal and subprolate, radially symmetric, isopolar, tricolpate, reticulate, rarely tetracolpate, polar axis is 15.2-30.2 m and equatorial axis 12.5-27.1 m, outline is elliptic in the equatorial view and circular in the polar view. Crambe mainly differs from these taxa with the prolate-spheroidal to subprolateshape, ornamentation only tricolpate and polar axis is 22.75-25.90 m and equatorial axis 21.07-22.75 m.

Pollen grains of 39 species belonging to six tribes; Arabideae, Euclidieae, Hesperideae, Lunarieae, Matthioleae and Sisymrieae from Brassicaceae were studied by Abdel-Khalik (2002). In general the pollen morphology varies within a narrow range. The shape varies from prolate spheroidal, subprolate to prolate. The pollen morphologies of Brassica campestris L., Coronopus didymus (L.) Smith., Eruca sativa L., Farsetia hamiltonii Royle., Iberis amara L., Lepidium sativum L., Raphanus sativus L., Sisymbrium irio L. are tricolpate, oblate to suboblate, prolate-spheroidal to rarely oblate-spheroidal (Arora and Modi, 2011). Sexine is thinner or thicker than nexine. Tectum is reticulate but sometimes it may be granulate. Pollen morphology of the genus Clypeola (Brassicaceae), tricolpate rarely tetracolpate, prolate-spheroidal, subprolate to prolate, polar axis is 22.4-33.4 m and equatorial axis 19.2-23.9 m (Keshavarzi et al., 2012).

The pollen morphology of the genus Arabis (Brassicaceae) istricolpate, but some of them are 2- and 4-colpate, prolate-spheroidal, subprolate, suboblate, prolate and oblate, polar axis and equatorial axis are 11.76-38.22 m (Mutlu and Erik, 2012). Cardaria draba L. Desv are tricolpate, prolate, tectum was reticulate (Mousavi and Sharifi-Rad, 2014). The major difference of Crambe from all these taxa mentioned above is the size and prolate-spheroidal, subprolate shape of pollen grains.

The observed seed morphology revealed that the shape of seeds varied from orbicular or elliptic with smooth surface in Crambe taxa. Seed color varied from brown or dark brown. SEM showed reticulate surface pattern with regular polygonal mostly hexagonal, epidermal cell shape varied from isodiametric, raised, straight; smooth anticlinal cell walls and flat to concave: macro-reticulate periclinal cell walls. The analysis of the mean L (length) and W (width) values showed that the largest seeds occur in Crambe maritime 3.3-4.9 x 2.5-3.8 mm, while the smallest L and W values were recorded for Crambe orientalis subsp. sulphurea 2.0-2.6 x 1.7-2.4 mm "Table 3". Crambe seeds had differences from the other investigated Brassicaeae members in literature.

For example Eruca sativa v. longirostris, E. sativa v. oblongifolia had 1.8 x 1.5 mm, sub-globose seeds (Tantawy et al., 2004), while Crambe species had 2-4.9x1.7-3.8 mm, orbicular to elliptic shaped seeds. Erucaria hispanica and Raphanus raphanistrum had sub-globose to ovate seed shape (Tantawy et al., 2004). The seeds of Raphanus sativus and R. raphanistrum were 0.81.0 mm, shiny brown, glabrous textured and kidney shaped (Kasem et al., 2011). Brassica juncea and Sinapis arvensis had round plump or shriveled seed shape and the seed size was 1.2-2 x 1.1-1.7 mm, seed coat color was tan, brown, dark brown, or yellow (Wang et.al., 2013). zdogru et al. (2016), detected four main seed surface types (papillate, reticulate, reticulate-papillate and undulate) in the genus Ricotia. Ricotia tenuifolia, R. aucheri, R. davisiana and R. varians had reticulate seed surface like Crambe species investigated in our research.

The seed shape of R.aucheri was ovoid to reniform, R. isatoides was reniform, R. tenuifolia was reniform to elliptic-ovoid, R. carnosula, R. davisiana, R. cretica and R. lunaria was orbicular, R. sinuate was orbicular or oblong and R. varians was ovoid. Crambe L differs from these taxa with orbicular to elliptic seed shape.

In conclusion, our investigation supports the use of seed surface patterns as a diagnostic character for species level instead of pollen morphology. The other seed characteristics like seed size, seed shape, seed wing, and seed color that might be helpful to distinguish some taxa (Tantawy et al., 2004; Pinar et al., 2007, 2009; Kaya et al., 2011, Bona, 2013).

Table 1. List of investigated taxa and localities.

SPECIES###LOCALITY

Crambe orientalis subsp.###Kahramanmara: City center, 2 nd km on road to Ahir Mountain, near city forest, 943 m

orientalis###B.Tarikahya 2788

Crambe orientalis subsp.###Kahramanmara: Malatya-Elbistan road, ca.35 km to Elbistan, Yukariyalak-2 turnout, near

sulphurea###field, 1456 m, B.Tarikahya 2799

Crambe tataria var.

###Kirehir: Kirehir-Kirikkale road, 85 km to Kirikkale, road side, 1180 m, B.Tarikahya 2808

Tataria

Crambe tataria var.

###Ankara: Ankara-Konya road, ca. 60 km to .Kohisar, TOF, 1135 m, B.Tarikahya 2775

Aspera

Crambe maritima###Sinop: Trkeli-Abana road, Haciveli district, near road, sandy dunes, s.l., B.Tarikahya 2813

Table 2. The palynological measurements Crambe taxa. P: polar axis, E: equatorial axis, I: thickness of intine, t: Polar, Clg: length of colpus, Clt: latitude of colpus, M: median, V: variation, S: standart deviation.

###TAXON NAME

###C. orientalis subsp.###C. orientalis subsp.###C. tataria var. tataria###C. tataria var. aspera###C. maritima

###orientalis###sulphurea

Pollen shape###Subprolate###Prolate-Spheroidal###Subprolate###Prolate-Spheroidal###Subprolate

P/E###1.21###1.08###1.14###1.13###1.19

Polar###M / var

axis###25.89(31.31-37.60)###22.75(24.00-28.30)###24.84(24.46-28.91)###23.47(23.50-28.20)###25.23(24.00-29.45)

(m)###S###1.72###1.04###1.06###1.04###1.24

Equatorial M / var###21.46(26.07-33.72)###21.07(21.98-26.23)###21.76(21.07-25.97)###20.80(20.17-25.25)###21.14(20.89-25.47)

axis(m)###S###1.83###1.11###1.39###1.35###1.47

###M / var###2.69(1.23-2.01)###2.80(1.10-1.67)###2.53(0.90-1.70)###2.47(1.00-1.76)###2.64(1.10-1.67)

Exine (m)

###S###0.21###0.28###0.24###0.19###0.14

###M / var###0.53(1.20-2.32)###0.58(0.78-1.79)###0.51(0.70-1.67)###0.52(0.87-1.95)###0.53(0.87-1.88)

I (m)

###S###0.33###0.25###0.24###0.17###0.26

###M / var###0.40(0.25-0.75)###0.35(0.25-0.60)###0.40(0.25-0.65)###0.30(0.25-0.85)###0.35(0.20-0.60)

i (m)

###S###0.13###0.10###0.11###0.15###0.11

###M / var###1.35(1.33-2.44)###2.57(1.67-2.86)###1.03(1.00-1.86)###1.34(1.12-1.86)###1.36(1.10-2.47)

Lumen width

###S###0.28###0.23###0.25###0.21###0.32

###M / var###0.44(1.82-3.10)###0.38(1.86-3.23)###0.32(1.47-2.43)###0.43(0.25-0.60)###0.33(1.57-3.06)

Murus width

###S###0.33###0.38###0.24###0.32###0.31

Apocolpium M / var###5.94(5.55-7.86)###5.63(5.00-7.13)###5.53(5.55-7.56)###5.53(4.90-7.14)###5.18(5.00-6.87)

(m)###S###0.53###0.59###0.51###0.61###0.51

Mesocolpium M / var###8.00(6.78-9.81)###8.03(5.64-8.09)###7.24(6.21-8.17)###7.96(5.20-8.00)###8.34(5.95-7.97)

(m)###S###0.83###0.57###0.45###0.65###0.44

###M / var###17.07(9.90-12.00)###16.62(7.38-9.50)###18.73(7.27-10.41)###16.98(8.20-9.35)###16.86(6.34-9.42)

Clg(m)

###S###0.77###0.58###0.69###0.34###0.77

###M / var###4.77(5.14-7.73)###4.68(4.33-6.76)###4.73(4.59-6.52)###4.50(5.01-6.77)###4.73(4.11-6.59)

Clt(m)

###S###0.55###0.68###0.52###0.53###0.55

###M / var###7.71(5.14-7.73)###7.58(4.33-6.76)###6.95(4.59-6.52)###7.75(5.01-6.77)###7.72 (4.11-6.59)

T

###S###0.55###0.68###0.52###0.63###0.55

Table 3. The seed properties of Crambe taxa.

###TAXON NAME

###C. orientalis subsp. C. orientalis subsp.

###C. tataria var. tataria C. tataria var. aspera###C. maritima

###orientalis###sulphurea

Seed color###Brown or dark brown Brown or dark brown###Brown or dark brown###Brown or dark brown###Dark brown

Seed shape###Elliptic###Orbicular###Orbicular###Orbicular###Elliptic

Seed size(mm)###2.3-3.9x1.9-3.6###2.0-2.6x1.7-2.4###2.2-3.4x2.1-3.3###2.8-4.1x2.3-3.4###3.3-4.9x2.5-3.8

###Surface###Reticulate###Reticulate###Reticulate###Reticulate

###Reticulate

###pattern

Epidermal cell Regular to irregular###Regular polygonal###Regular polygonal.

###Irregular polygonal###Regular 4.5.6 gonal

###shape###polygonal###mostly hexagonal###mostly hexagonal

Anticlinal cell###Raised.straight;###Raised.straight;###Raised.straight;###Raised.straight;###Raised.straight;

###wall###smooth###Smooth###smooth###smooth###smooth

Periclinal cell###Flat to concave:###Flat to slightly###Flat to concave:###Flat to concave:###Flat to slightly

###wall###macro-reticulate###concave###macro-reticulate###macro-reticulate###concave

Ornamentation###Reticulate###Reticulate###Reticulate###Reticulate###Reticulate

Acknowledgments: This investigation was financed by General Directorate of Agricultural Research and Policy, Republic of Turkey, Ministry of Food, Agriculture and Livestock under [Grant number TBAD/13/A01/P01/007].

REFERENCES

Abdel-Khalik, KNE.(2002). Biosystematic studies on Brassicaceae (Cruciferae) in Egypt. Ph.D. Dissertation. Wageningen University.

Arora, A. and A. Modi (2011). Pollen Morphology of Some Desertic Crucifers. Indian J. Fundamental and Applied Life Sciences 1: 11-15.

Bolurian, S. (2009). A systematic study of certain species of the Alyssum belonging to the Mustard family (Brassicaceae) in Iran. M.A. thesis. AL-Zahra University.

Bona, M. (2013). Seed-coat microsculpturing of Turkish Lepidium (Brassicaceae) and its systematic application.Turkish J. Botany 37: 662-668.

Brochmann, C.(1992). Pollen and seed morphology of nordic Draba (Brassicaceae): phylogenetic and ecological implications. Nordic J. Botany 1: 657-673.

Dogan, C., and O. Inceoglu (1990). Pollen morphology of some Isatis L. taxa in Turkey. Turkish J. Botany14: 12-31.

Erdtman, G. (1952). Pollen morphology and plant taxonomy. Angiosperms. Chronica Botanica Co., Waltham Mass.

Ergtman, G. (1969). Handbook of Palynology: Morphology-Taxonomy-Ecology, An Introduction to the Study of Pollen Grains and Spores. Copenhagen: Munksgaard.

Faegri, K., and J. Iversen (1975). Text Book of Pollen Analysis.New York: Hafner Press.

Franzke, A., M.A. Lysak, I.A. Al-Shehbaz, M.A. Koch and K. Mummenhoff (2010). Cabbage family affairs: the evolutionary history of Brassicaceae. Trends in Plant Science 839: 1-9.

Hedge, I.C. (1976). A systematic and geographical survey of the old world Cruciferae. In: Vaughan, J.G., MacLeod, A.J.andB.M.G., Jones eds. The Biology and Chemistry of the Cruciferae. London: Academic Press.

Inceoglu, O., and F. Karamustafa (1977). The pollen morphology of plants in Ankara region II. Cruciferae. Communications Series C2: Botanique21: 111-118.

Karcz, J., KsiazczykT. And J. Maluszynska (2005). Seed coat patterns in rapid-cycling Brassica forms, Acta Biologica Cracoviensia Series Botanica 47: 159-165.

Kasem, W.T., Ghareeb, A. and E.Marwa (2011). Seed morphology and seed coat sculpturing of 32 taxa of family Brassicaceae. J. American Science, 7(2): 166-178.

Kaya, A., nal, M., zgke, F., DoganB.and E. Martin (2011). Fruit and seed morphology of six species previously placed in Malcolmia (Brassicaceae) in Turkey and their taxonomic value. Turkish J. Botany 35: 653-662.

Keshavarzi, M, Abassian, S. and M. Sheidai (2012). Pollen Morphology of the genus Clypeola (Brassicaceae) in Iran. Phytologia Balcanica 18: 17-24.

Koul, KK, Ranjna, N. and S.N.Raina (2000). Seed coat microsculpturing in Brassica and allied genera (subtribes Brassiccinea, Raphaninae, Moricandiinae). Annals of Botany 86: 385-397.

Moazzeni, H, Zarre, S., Al-Shehbaz I.A. and K. Mummenhoff (2007). Seed coat microsculpturing and its systematic application in Isatis (Brassicaceae) and allied genera in Iran. Flora 202: 447-454.

Mousavi, S.M. and J. Sharifi-Rad (2014). Anatomical, palynological and micromorphological study of seed, trichome and stomata of Cardaria draba L. Desv (Brassicaceae) in Sistan, Iran. International J. Biosciences 5: 63-69.

Murley, M.R. (1951). Seeds of the Cruciferae of North Eastern America. American Midland Naturalist 46: 1-81.

Mutlu, B. (2012). Crambe. In: A. Gner, S. Aslan, T. Ekim, M. Vural, and M.T. Baba eds. Trkiye Bitkileri Listesi (Damarli Bitkiler). Istanbul: Nezahat Gkyigit Botanik Bahesi ve Flora Arastirmalari Dernegi Press.

Mutlu, B. and S.Erik (2012). Pollen morphology and its taxonomic significance of the genus Arabis (Brassicaceae) in Turkey. Plant Systematics and Evolution 298: 1931-1946.

zdogru, B., Akaydin, G., Erik, S. and K.Mummenhoff (2016). Seed morphology of Ricotia (Brassicaceae) and its phylogenetic and systematic implication. Flora-Morphology, Distribution, Functional Ecology of Plants222: 60-67.

Perveen, A., QaiserM. And R. Khan (2004). Pollen Flora of Pakistan - XLII. Brassicaceae. Pakistan J. Botany 36: 683-700.

Pinar, N.M., Adiguzel N. and F. Geven (2007). Seed coat macrosculpturing in some Turkish Aethionema R. Br. (Brassicaceae). Pakistan J. Botany 39: 1025-1036.

Pinar, N.M., A. Duran, eter, T. and G.N. Tug (2009). Pollen and Seed Morphology of the Genus Hesperis L. (Brassicaceae) in Turkey.Turkish J. Botany 33: 83-96.

Tantawy, M.E., Khalifa, S.F., Hassan, S.A. and G.T. Al-Rabiai (2004). Seed exomorphic characters of some Brassicaceae (LM and SEM Study). International J. Agriculture and Biology 6: 821- 830.

Wang, Y.H., Wei, W., Kang, D.M and K.P. Ma (2013). Seed coat microsculpturing is related to genomic components in wild Brassica juncea and Sinapis arvensis. PloS one, 8(12): e83634.

Wodehouse RP. (1935). Pollen Grains, their structure, identification and significance in science and medicine. New York: McGraw Hill Press.
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Publication:Journal of Animal and Plant Sciences
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