POLLEN FLORA OF PAKISTAN -XLII. BRASSICACEAE ANJUM ...

Pak. J. Bot., 36(4): 683-700, 2004. 

POLLEN FLORA OF PAKISTAN -XLII. BRASSICACEAE 

ANJUM PERVEEN, M. QAISER AND REHANA KHAN * 

Department of Botany, University of Karachi, Karachi-75270, Pakistan 

* Federal Urdu University of Science, Arts & Technology, 

Gulshan-e-Iqbal, University Road, Karachi, Pakistan 

Abstract 

Pollen morphology of 77 species belonging to 36 genera of the family Brassicaceae from 

Pakistan has been examined by light and scanning electron microscope. Pollen grains are usually 

radially symmetrical, isopolar sub-prolate to prolate, or prolate-spheroidal rarely oblate-spheroidal, 

tricolpate rarely 4-8 colpate. Sexine thinner or thicker than nexine. Tectum fine to coarsely 

reticulate with more or less regular pattern of muri or reticulate – rugulate. On the basis of tectal 

surface four distinct pollen types are recognized viz., Arabis bijuga-type, Farsetia ramosissimatype, 

Draba lanceolata –type and Erysimum melicentae – type. 

Introduction 

Brassicaceae is a cosmopolitan family of about 350 genera and 3000 species, 

occurring mainly in north temperate zone particularly in Mediterranean region 

(Mabberely, 1987). In Pakistan it is distributed by 92 genera and 250 species (Jafri, 

1973). Cruciferae are annual, biennial and perennial herbs. Corolla is cruciform (4 petals 

arranged in cross manner. Some common members of Brassicaceae are species of 

Brassica oleracea (Broccoli, Brussels sprout, Cabbage, Cauliflower, Kale etc.). Oil is 

also extracted from the seed of Brassica species. Many Crucifers are cultivated as 

ornamentals like Aubrieta deltoidea (aubrietia), Erysimum cheire (wallflower), Arabis 

sp., (rock cress). Chiguriaeva (1973) examined the pollen morphology of the family 

Brassicaceae in relation to taxonomy. Javied & Naqshi (1975) also utilized pollen data in 

the classification of family Brassicaceae. Jonsell (1986) during monographic studies of 

Brassicaceae also examined pollen morphology of this family. Lahham & Al-Essawi 

(1987) examined the pollen morphology of the family Brassicaceae from Jordan. Pollen 

morphology of the family Brassicaceae has been examined by Erdtman (1963), Sharma 

& Nair (1973), Carter et al., (1975). Moore & Webb (1978). The most comprehensive 

studies of Brassicaceae pollen is that of Rollins & Banerjee (1979). However, there are 

no reports on pollen of the family Brassicaceae from Pakistan. Present investigations are 

based on the pollen morphology of 77 taxa representing 36 genera of the family 

Brassicaceae by light and scanning electron microscope. 

Materials and Methods 

Pollen samples were obtained from Karachi University Herbarium (KUH) or 

collected from the field. The list of voucher specimens is deposited in KUH. The pollen 

grains were prepared for light (LM) and scanning microscopy (SEM) by the standard 

methods described by Erdtman (1952). For light microscopy, the pollen grains were 

mounted in unstained glycerin jelly and observations were made with a Nikon Type-2 

microscope, under E40, 0.65 and oil immersion (E100, 1.25), using 10x eye piece. For

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ANJUM PERVEEN ET AL., 

SEM studies, pollen grains suspended in a drop of water were directly transferred with a 

fine pipette to a metallic stub using double sided cello tape and coated with gold in a 

sputtering chamber (Ion-sputter JFC-1100). Coating was restricted to 150 A. The S.E.M 

examination was carried out on a Jeol microscope JSM-2. The measurements are based 

on 15-20 readings from each specimen. Pollen diameter, polar axis (P) and equatorial 

diameter (E), aperture size, apocolpium, mesocolpium and exine thickness were 

measured (Tables 1-4). 

The terminology used is in accordance with Erdtman (1952), Kremp (1965), Faegri 

& Iversen (1964) and Walker & Doyle (1975). 

General pollen characters of the family Brassicaceae 

Pollen grains usually radially symmetrical, isopolar, sub-prolate or prolate to prolatespheroidal, 

rarely oblate-spheroidal, tricolpate often 4-8 colpate. Sexine thinner or thicker 

than nexine. Tectum fine to coarse reticulate or reticulate-rugulate. On the basis of tectum 

types, four distinct pollen types are recognized viz., Arabis bijuga-type, Farsetia 

ramosissima-type, Draba lanceolata – type and Erysimum melicentae - type. 

Key to the pollen types 

1. + Tectum reticulate-rugulate ..…………….…………… Farsetia ramosissima-type 

- Tectum fine-coarse reticulate ……….…………………………………………... 2 

2. + Tectum finely reticulate ……………………………………... Arabis bijuga-type 

- Tectum not as above ………………………………….………………………… 3 

3. + Tectum medium reticulate .…………………………. Erysimum melicentae - type 

Tectum coarsely reticulate…………………………...….. Draba lanceolata –type 

Pollen type: Arabis bijuga - type (Fig. 1 A-C) 

Pollen class: Tricolpate 

P/E ratio: 108-140 

Shape: Prolate-spheroidal to sub-prolate rarely prolate. 

Apertures: Colpus long sunken with acute ends. 

Exine: Sexine thicker than nexine. 

Ornamentation: Finely reticulate more or less regular pattern of muri. 

Measurements: Size: Polar axis P=16 (25.5) 32.5 μm and equatorial diameter E=15 

(21.5) 28 μm, trilobed, with apertures on the angles of the outline of the grain in polar 

view, colpi 10 (15.5) 20 μm long. Exine 1.0 (1.5) 2.5 μm thick, sexine as thick as nexine. 

Tectum finely reticulate. 

Species included: Arabidopsis mollissima (C.A. Mey) Busch, A. traxacifolia (T.And.) 

Jafri, Arabidopsis wallichii (Hook.f. Thoms.) Busch, A. stricta (Camb.) Busch., Arabis 

amlpexicaule Edgew., A. bijuga Watt., A. pterosperma Edgew, Bassica juncea (L.) 

Czern., B. napus L.ssp. napus, B. nigra (L.) Koch., B. oleracea var. botrytis L., B. 

oleracea var. capitata, B. rapa ssp. rapa, B. rapa ssp. campestris, Cardamine impatiens 

L., Chorispora sibirica (L.) DC., Douepia tortuosa Camb., Drabopsis verna Koch., 

Euclidium syriacum (L.) R.Br., Malcolmia scorpioides (Bunge) Boiss.

POLLEN FLORA OF PAKISTAN -XLII BRASSICACEAE 

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Key to the species and group 

1. + Pollen grains prolate-spheroidal ……..……………… Douepia tortuosa - subtype 

(Arabis pterosperma, Doupia tortuosa, Bassica juncea, Brassica napus ssp. 

napus, B. nigra, B. rapa ssp. rapa) 

- Pollen grains sub-prolate to prolate …………………………………………….. 2 

2. + Pollen grains subprolate ………………………. Arabidopsis mollissima- subtype 

(Arabidopsis mollissima, A. traxacifolia, Arabis amlpexicaule, A. bijuga, 

Brassica oleracea var. capitata, B. rapa ssp. campestris, Cardamine impatiens, 

Chorispora sibirica, Drabopsis verna, Euclidium syriacum Malcolmia 

scorpioides) 

- Pollen grains prolate …..…………….…………… Arabidopsis wallichii-subtype 

(Arabidopsis wallichii, A. stricta, Brassica oleracea var. botrytis L.) 

Pollen type: Erysimum melicentae (Fig. 1D-F. Fig. 2 A & B). 


P/E ratio: 110-151 

Shape: Sub-prolate to prolate rarely prolate-spheroidal 


Exine: Sexine thicker or thinner than nexine. 

Ornamentation: Medium reticulate with more or less regular pattern of muri. 


(21.5) 32 μm, (trilobed, with apertures on the angles of the outline of the grain in polar 

view, colpi 10 (15.5) 17.5 μm long. Exine 1.0 (1.96) 2.5 μm thick, sexine as thick as 

nexine. Tectum medium reticulate. 

Species included: Alliaria petiolata (M.Bieb.) Cav. Grande, Arabis saxicola Edgew, 

Alyssum desertorum Staph, Arabidopsis pumila (Steph) Busch, Cardaria drapa (L.) 

Desv,. Cithareloma lehmannii Bunge, Coronocarpus didymus (L.) Smith, Diceratella 

canescens (Boiss.) Boiss., Goldbachia laevigata (M.Bieb.) DC. Eremobium aegypticum 

(Spreng) Boiss., Leptaleum filifolium (Willd.) DC., Malcolima strigosa Boiss. 


1. + Pollen grains prolate-spheroidal ……………………………. Goldbachia laevigata 

- Pollen grains sub-prolate to prolate …………………………...…………………. 2 

2. + Pollen grains prolate .…………………………...… Alyssum desertorum-sub-type 

(Alyssum desertorum, Arabis saxicola, Arbidopsis pumila, Cardaria drapa) 

- Pollen grains sub-prolate …………………………… Alliaria petiolata –subtype 

(Alliaria petiolata, Cithareloma lehmannii, Coronocarpus didymus, Diceratella 

canescens, Goldbachia laevigata, Eremobium aegypticum, Leptaleum filifolium, 

Malcolima strigosa) 

Pollen type: Draba lanceolata- type (Fig. 2 C-F; Fig. 3A-F; Fig. 4A-F) 


P/E ratio: 106-168 

Shape: Prolate to subprolate rarely prolate-spheroidal 

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Fig. 1. Scanning Electron micrographs of pollen grains. Arabis bijuga: A, Equatroial view, B, Exine pattern. A. 

pterosperma: C, Polar view. Leptaleum filifolium: D, Equatorial view. Cithareloma lehmannii: E, polar view, 

F, Exine pattern. Scale bar = A, C, D & E = 10; B, F = 1 μm


Fig. 2. Scanning Electron micrographs of pollen grains. Leptaleum filifolium: A, Exine pattern. Eremobium 

aegypticum: B, pollen grains. Isatis minima: C, Equatroial view, D, Exine pattern. Capsella burspastoris: E., 

Polar view, F, Exine pattern. Scale bar =B, C & E = 10; A, D & F = 1 μm 

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Fig. 3. Scanning Electron micrographs of pollen grain. Boreava orientalis: A, Equatorial view. Euclidium 

tenuissimum: B, Equatorial view, C, Exine pattern. Farsetia hamiltonii: D, Exine pattern, E, Equatorial view. 

Alyssum desertorum: F, Exine pattern. Scale bar = A , B & E = 10; C, D & F = 1 μm


Fig. 4. Scanning Electron micrographs of pollen grains. Draba stenocarpa: A, Exine pattern, B, Equatorial 

view. Draba trinervia: C, Polar view. Christolea himalayensis: D, Equatorial view, E, Exine pattern. Barbrea 

intermidia: F, Equatorial view. Scale bar = B –D & F= 10; A & E = 1 μm 

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Ornamentation: Coarsely reticulate. 


(18.5) 22.1 μm, (trilobed, with apertures on the angles of the outline of the grain in polar 

view, colpi 10 (21.5) 32.5 μm long. Exine 1.5 (2) 2.5 μm thick, sexine as thick as nexine. 

Tectum coarsely reticulate. 

Species included: Alyssum desertorum Staph, Boreava orientalis Jaub. & Spach, 

Capsella bursa-pastoris (L.) Medik., Chorispora bungeana Fish & Mey, Christolea 

himalayensis (Camb.) Jafri, Chorispora tenella (Pallas) DC., Draba trinervis Schultz, D. 

altaica (C.A. Mey) Bunge, D. lanceolata Royle, D. melanopus Kom., D. stenocarpa 

Hook. F. & Thoms. Euclidium tenuissimum (Pallas) B. Fedtsch., Erysimum meilicentae 

Dunn., E. griffithianum Boiss, Farsetia hamiltonii Royle, Diceratella floccose (Boiss.) 

Boiss., Farsetia heliophila Bunge ex Coss., Isatis stocksii Rech.f Boiss, I. minima 

Bunge, Lepidium virginicum L, Matthiola macranica Rech f., Mathiola flavida Boiss., 

Malcolmia cabulica (Boiss.) Hook. F. Thoms, Malcolmia africana (L.) R.Br., Nasturtium 

officinale R. Br., Microsisymbrium flaccidum Schulz. 


1. + Pollen grains prolate-spheroidal ..……………….…………Isatis stocksii-subtype 

(Isatis stocksii, Isatis minima) 

- Pollen grains not as above ………………………………………………………. 2 

2. + Pollen grains prolate …………………………. Capsella bursa-pastoris. -subtype 

(Capsella bursa-pastoris, Boreava orientalis, Chorispora tenella, Malcolmia 

cabulica) 

- Pollen grains sub-prolate ………………………… Matthiola macranica-subtype 

(Chorispora bungeana, Christolea himalayensis, Draba trinervis, D. altaica, D. 

lanceolata, D. melanopus, D. stenocarpa, Euclidium tenuissimum, Erysimum 

meilicentae, E. griffithianum, Farsetia hamiltonii, Diceratella floccose, Farsetia 

heliophila, Lepidium virginicum, Matthiola macranica, Matthiola flavida, 

Malcolmia africana, Nasturtium officinale, Microsisymbrium flaccidum) 

Pollen type: Farsetia ramosissima - type (Fig. 5 A-G) 


P/E ratio: 102-113 

Shape: Sub-prolate to prolate-spheroidal rarely prolate. 



Ornamentation: Reticulate – rugulate. 

Measurements: Size: Polar axis P = 14 (23.5) 32.5 μm and equatorial diameter E=12 

(18) 24 um, trilobed, with apertures on the angles of the outline of the grain in polar view, 

colpi 8.75 (16.5) 25.5 μm long. Exine 1.5 (2) 2.5 μm thick, sexine as thick as nexine. 

Tectum reticulate-rugulate. 

Species included: Barbaraea vulgaris, Conringia planisiliqua Fisch & Mey., Crambe 

cardifolia Steven, Descurainia sophia (L.) Webb. & Berth., Diplotaxis griffithii (Hook. f.


Fig. 5. Scanning Electron micrographs of pollen grains. Diplotaxis harra: A, Polar view, D. grifithii: B, 

Equatorial view, C, Exine pattern. Crambe cordifolia: D, Polar view. Conringia planisliqua: E, Equatorial view, 

F, Exine pattern, G, Polar view. Scale bar = A, B, D, E & G = 10; C & F = 1 μm 

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& Thoms.) Boiss., Diptychocarpus strictus (Fish.M.Beib.) Trautv, Diplotaxis harra 

(Forssk.) Boiss, Erysimum crassicaule (Boiss.) Boiss., Erysimum melicentae Dunn., E. 

repandum L., Farsetia ramosissima Hochst. ex Boiss., Malcolmia behboudiana Rech.f., 

Moricandia sinaica (Boiss.) Boiss. Lepidium aucheri Boiss., Isatis costata C.A.Mey 


1. + Pollen grains oblate-spheroidal………………………..……………..Isatis costata 

- Pollen grains prolate-spheroidal or prolate to sub-prolate ……………………… 2 

2. + Pollen grains prolate …………………………….. Erysimum crassicaule-subtype 

(Erysimum crassicaule,E. repandum, Barbaraea vulgaris) 

- Pollen grains sub-prolate …..…………..…………… Diplotaxis griffithii-subtype 

(Conringia planisiliqua, Crambe cardifolia, Descurainia sophia, Diplotaxis 

griffthii, Diptychocarpus strictus, Diplotaxis harra, Erysimum crassicaule, 

Erysimum melicentae, E. repandum, Farsetia ramosissima, Malcolmia 

behboudiana, Moricandia sinaica, Lepidium aucheri) 

Discussion and Conclusions 

Brassicaceae is a stenopalynous family (Erdtman, 1952). Pollen grains are generally 

prolate to sub-prolate or prolate – spheroidal rarely oblate – spheroidal, 3 - colpate often 

4-8 –colpate as in Erysimum stocksiana. Tectum reticulate, reticulum varies from fine – 

coarse. Appel & Al-Shehbaz (2003) also reported tricolpate reticulate pollen in the family 

Brassicaceae. However, they reported sparsely spinulose and punctate tectum in the 

genus Heliophila and 10-colpate pollen in few genera of Brassicaceae. Erdtman (1963) 

divided the species of Brassicaceae into two pollen types on the bases of exine thickness. 

Moore & Webb (1987) classified the family in tricolpate with reticulate pollen. Khalik 

(2002) divided the family into three pollen types on the basis of lumina size. The generic 

delimitation based on pollen morphology is difficult. However, on the basis of tectal 

surface four distinct pollen types are recognized viz., Arabis bijuga – type, Erysimum 

melicentae - type, Farsetia ramosissima - type, Draba lanceolate - type. The Arabis 

bijuga – type is easily recognized by having finely reticulate pollen, 20 species are 

included in this pollen type viz., Arabidopsis mollissima (C.A. Mey) Busch, A. 

traxacifolia (T.And.) Jafri, Arabidopsis wallichii (Hook. f. Thoms.) Busch, A. stricta 

(Camb.) Busch., Arabis amlpexicaule Edgew., A. bijuga Watt., Arabis pterosperma 

Edgew, Brassica juncea (L.) Czern., B. napus L., ssp., napus, B. nigra (L.) Koch., B. 

oleracea var. botrytis L., B. oleracea var. capitata, B. rapa ssp. rapa, B. rapa ssp., 

campestris. Cardamine impatiens L., Chorispora. sibirica (L.) DC., Douepia tortuosa 

Camb., Drabopsis verna, Euclidium syriacum (L.) R.Br., Malcolmia. scorpioides 

(Bunge) Boiss. This pollen type is further divided into three subtypes on the basis of 

pollen shape classes (see key to the species or species group). Erysimum melicentae – 

type is delimited by its medium reticulate tectum, 12 species have medium reticulate 

tectum such as Alyssum desertorum Staph, Alliaria petiolata (M.Bieb.) Cav. Grande, 

Arabis saxicola Edgew, Arbidopsis pumila (Steph) Busch, Cardaria drapa, Cithareloma 

lehminniii, Coronocarpus didymus (L.) Smith, Diceratella canescens (Boiss.) Boiss., 

Goldbachia laevigata (M.Bieb.) DC., Eremobium aegypticum (Spreng) Boiss., Erysimum 

melicentae Dunn., E. repandum L., E. griffithianum Boiss., Leptaleum filifolium (Willd.)


DC., Malcolima. strigosa Boiss. This type also divided into two subtype based on pollen 

shape class (see key to the species and species group). Draba lanceolata- type is the 

largest among all the four pollen types (28 species). This pollen type is characterized by 

its coarsely reticulate tectum. viz., Alyssum desertorum Staph, Boreava orientalis Jaub. & 

Spach, Capsella bursa-pastoris (L.) Medik., Chorispora bungeana Fish & Mey, 

Christolea himalayensis (Camb.) Jafri, Chorispora tenella (Pallas) DC., Draba trinervis 

Schultz, D. altaica (C.A. Mey) Bunge, D. lanceolata Royle, D. melanopus Kom., D. 

stenocarpa Hook. F. & Thoms., Euclidium tenuissimum (Pallas) B. Fedtsch, Erysimum 

melicentae Dunn., E. griffithianum Boiss., Farsetia hamiltonii Royle, Diceratella 

floccose (Boiss.) Boiss., Farsetia heliophila Bunge ex Coss., Isatis stocksii Rech.f 

Boiss, I. minima Bunge, Lepidium virginicum L, Matthiola macranica Rech f., Matthiola 

flavida Boiss., Malcolmia africana (L.) R.Br., Nasturtium officinale R. Br., 

Microsisymbrium flaccidum Schulz,. It is also divided into three subtype (see key to the 

species). Farsetia ramosissima - type is easily distinguished by its reticulate – rugulate 

tectum. Jonsell (1987) reported similar pollen in the genus Farsetia 13 species are 

included in this pollen type like, Conringia planisiliqua Fisch & Mey., Crambe 

cardifolia Steven, Descurainia sophia (L.) Webb. & Berth., Diplotaxis grifftihii (Hook.) 

& Thoms. Boiss., Diptychocarpus strictus (Fish. M. Beib.) Trautv, Diplotaxis harra 

(Forssk.) Boiss, Erysimum crassicaule (Boiss.) Boiss. Erysimum melicentae Dunn., E. 

repandum L., Farsetia ramosissima Hochst. ex Boiss, Malcolmia behboudiana, Rech.f., 

Moricandia sinaica (Boiss.) Boiss. Lepidium aucheri Boiss. Like other three types this 

pollen type is also classified into two subtypes 

Pollen morphologically the family is more closely related to Tamricaceae where both 

families have tricolpate pollen with reticulate tectum (Qaiser & Perveen, 2004). Ertdman 

(1952) reported that the family is more close to family Capparaceae. However, 

Capparaceae is eurypalynous family, in this family tricolporate pollen are common 

(Perveen & Qaiser, 2001). Pollen morphology confirms the homogeneous nature of the 

family. 

References 

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(Ed.): K. Kubitzki. V. Flowering plants Dicotyledons, Malvales, Capparales and Non-Betalain 

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Carter, A.L., S.T. Willams and T. Mcneilly. 1975. Scanning electron microscope studies of pollen 

behaviour on immature and mature brusseles sprout (Brassica oleracea var. gemmifera) 

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Chiguriaeva, A.A. 1973. Pollen morphology of Cruciferae in pollen and spore morphology of 

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Kremp, G.O.W. 1965. Encyclopaedia of Pollen Morphology. Univ. Arizona Press, Tuscon, U.S.A. 

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(Received for publication 12 November 2003)

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