J. Life Earth Sci., Vol.
1(2): 1-7, 2006 ISSN 1990-4827
@2006, JLES, RU
POLLEN GERMINATION, VIABILITY AND TUBE GROWTH IN
FOURTEEN CULTIVATED AND WILD SPECIES OF CUCURBIT
GROWN IN BANGLADESH
M. Rashed Zaman
Department of Genetics and Breeding, University of Rajshahi, Rajshahi 6205, Bangladedsh
Abstract
Pollen germination, viability and tube growth in fourteen species of cucurbit were examined. All the male
flowers had three anthers each. The highest number of pollen grain (917.00 ? 52.69) per anther was recorded
in pumpkin (Cucurbita maxima Duch ex Poir.) whereas the lowest (479.00 ? 13.14) per anther was recorded
in snake gourd (Trichosanthes cucumerina L.). Viable pollen was highest (98.51%) in bitter gourd
(Momordica charantia L.) and lowest (86.87%) in pointed gourd (Trichosanthes dioica Roxb.). Pollen grains
of all species were starchy in nature. The highest percentage of pollen germination (97.19%) was recorded as
in bottle gourd (Lagenaria siceraria (Mol) Standl.) and lowest (77.98%) in pointed gourd. Rate of increasing
lengths of the tubes was very fast between first two hours and became very slow in last six hours. After 12
hrs. of germination the longest pollen tube (1649.20? ± 15.05) was found in pumpkin and the shortest
(766.08? ± 19.03) was found in pointed gourd.
Key words: Pollen germination; viability, tube-growth, cucurbit.
mvivsk: †PvÏwU cucurbit cÖRvwZi civM‡iYyi AsKz‡iv`Mg, mRxeZv I bvwjKv e„w× cix¶v Kiv n‡q‡Q| me¸wj cÖRvwZi
cyi“l dz‡ji cÖwZwU‡Z wZbwU K‡i civMavbx wQj| wgwó Kzgovi (Cucurbita maxima Duch ex Poir.) cÖwZ civMavbx‡Z
M‡o me‡P‡q †ekx civM‡iYy (917.52 ? 52.69) wQj Ges me‡P‡q Kg (479.00 ? 13.14) wQj wPwPsMv‡Z
(Trichosanthes cucumerina L.)| mwµq civM‡iYyi msL¨v me‡_‡K ‡ekx (98.51%) cvIqv hvq Kijv‡Z (Momordica
charantia L.) Ges me‡_‡K Kg (86.87%) wQj cU‡j (Trichosanthes dioica Roxb.)| cix®‹zZ me cÖRvwZi civM‡iYy
kK©iv cÖK…wZi| civM‡iYyi AsKz‡iv`M‡gi kZKiv nvi me©v‡c¶v †ekx ch©‡e¶xZ nq jvD-G (Lagenaria siceraria (Mol)
Standl.) hv cÖvq 97.19% Ges cU‡j me©v‡c¶v Kg wQj hv cÖvq 77.98%| cÖ_g `yB N›Uvq civMbvjx e„w×i nvi AZ¨vš—
†ekx wQj Ges †kq Qq N›Uv‡Z GB e„w×i nvi Lye Kg †`Lv hvq| AsKz‡iv`M‡gi 12 N›Uv ci wgwó Kzgov‡Z me‡P‡q †ekx
ˆ`‡N©¨i (1649.20? ? 15.05) Ges cU‡j me‡P‡q Kg ˆ`‡N©¨i (766.08? ? 19.03) civMbvjx cvIqv hvq|
Introduction seeds produced (Bertin, 1990; Waser and Price, 1991;
Holm, 1994; Melser et al., 1997; Mitchell, 1997;
Flower structure is important in relation to pollen Bosch and Waser, 1999; Dieringer and Cabrera,
distribution and reception and in many of the 2002). In addition, the amount of pollen deposited
hermaphroditic plant species flowers have evolved may also influence the quality of the progeny through
dramatically to suit particular pollination mechanisms the action of prefertilization (microgametophyte
(Ainsworth, 2000). In wind pollinated plants (where competition: Snow, 1986; Schlichting et al., 1987;
there is a strong correlation with dioecy; Renner and Winsor et al., 1987, 2000) and /or postfertilization
Ricklefs, 1995), sexu al specialization of male and mechanisms (selective abortion: Niesenbaum and
female flowers is common. Intraspecific variation in Casper, 1994; Rigney, 1995; Havens and Delph,
size of the pollen loads deposited on stigmas may 1996; Niesenbaum, 1999; Melser and Klinkhamer,
influence both the number and quality of the eventual 2001).
progeny (Herrera, 2002). Threshold effects, nonlinear Important role played by pollen in sexual plant
dose-response relationships, and maternal and reproduction has motivated a multitude of observational
paternal identity, act for determining the number of and experimental investigations regarding patterns
and consequences of variation in size of stigmatic because of the inadequate information regarding their
pollen loads. This contrasts sharply less number of area and production in Bangladesh. No work on
investigations that have so far documented patterns of pollen biology of this important group of plants has
variation in the size of pollen tube populations for been done in our country. So, an endeavor has been
naturally pollinated wild plants (Levin, 1990; Honig made to study the pollen biology of some wild and
et al., 1992; Aizen and Feinsinger, 1994; cultivated cucurbits grown in Bangladesh.
Niesenbaum, 1994; Plitmann and Levin, 1996;
Herrera, 1997; Quesada et al., 2001; Herrera, 2002). Materials and Methods
However, because of the increasing realization of the
importance of pollen in both fundamental and applied The 14 cucurbit species collected from natural
areas, there has been an explosion of knowledge on population, commercial and homestead gardens used
pollen biology (Mulcahy and Ottaviano 1983; for the present work and they are Pumpkin
Shivana and Johri 1985; Mulcahy et al. 1986; Giles (Cucurbita maxima Duch ex Poir.), Ash gourd
and Prakash 1987; Cresti et al. 1988; Iwanami et al. (Benincasa hispida (Thunb.) Cogn.), Bottle gourd
1988; Quesada et al., 1995; Delph et al., 1997; Corff (Lagenaria siceraria (Mol) Standl.), Snake gourd
et al., 1998). (Trichosanthes cucumerina L.), Ridge gourd (Luffa
acutangula (Roxb.) L.), Pointed gourd
Pollen germination and pollen tube growth are (Trichosanthes dioica Roxb.), Cucumber (Cucumis
prerequisites for fertilization and seed development. sativus L and C. anguina L.), Bitter gourd
Due to involvement of the pistillate tissue in the (Momordica charantia L.), Sweet gourd (M.
nature, physiological and biochemical investigations cochinchinensis Spreng.), Watermelon (Citrullus
on pollen germination and pollen tube growth in vivo lanatus Thunb.), Muskmelon (Cucumis melo L.),
are rather difficult. In vitro germination techniques Sponge gourd (Luffa cylindrica Roem.), and Ivy
have therefore been used extensively on a variety of gourd (Coccinia cordifolia (Voigt) L).
pollen systems. Such studies have provided
considerable information on the physiology and The fresh male flowers at blooming stage were
biochemistry of pollen germination and pollen tube collected for counting the number of anthers of each
growth (Shivana and Johri 1985; Heslop-Harrison flower and for estimating the pollen viability. The
1987; Steer and Steer 1989). Pollen germination and anthers were stained with 1% acetocarmine solution
tube growth are generally divided into four phases: and the stained pollen was considered as viable and
imbibition phase, lag phase, tube initiation phase, and non-stained as non-viable pollen. To determine the
rapid tube elongation phase (Linskens and Kroh, starchy and non-starchy pollen grain the anthers were
1970). The time taken for different phases varies stained with potassium iodide solution where non-
greatly, from species to species, depending on the starchy pollen did not take stain.
type of reserve food material in the pollen and the Brewbaker and Kwack’s media as suggested by
external factors. Brewbaker and Kwack (1963) and Roberts’ media as
Cucurbits form an important group of vegetable per schedule of Roberts et al. (1983) were used to
crops comprising both cultivated and wild species. study in vitro germination rate of pollen grains by
This group includes mostly seed propagated ones, sitting drop culture method. The culture was
besides few vegetatively propagated ones like maintained in a humidity chamber to prevent
pointed gourd and also few perennials like ivy gourd. evaporation. Ten humidity chambers were prepared
From nutritional point of view bitter gourd is rich for in the same procedure with ten pair of petri plates for
vitamin C, pumpkin contains high carotenoid each experiment. Cultures along with humidity
pigments, sweet gourd is high in protein and pointed chambers were incubated under desired temperature
gourd is fairly high in calcium (Bhuiya et al., 1977; (22 ? 2?C is optimal) for 1 to 1.5 hrs. Respond to
Gopalan et al., 1982). A few cultivars of squashes germination of the cultured pollen grains was
and pumpkins are relatively high in energy and expressed in percentage.
carbohydrates (Seshadri, 1993). Cucurbits are of The prepared slides were observed under a
tremendous economic importance as food plants and microscope and the time required for germination of
it has been difficult to estimate or quantify them the pollen from each chamber was recorded with the
help of a stopwatch. The tube lengths of in vitro germination and none of the pollen grain was found
germinating pollen grains were recorded after 1-, 2-, to germinate in that medium. The highest percentage
6- and 12 hours of setting the experiment. of pollen germination was recorded as 97.19% in
bottle gourd whereas the lowest was 77.98% in
Results pointed gourd. The time required for germinating
All the male flowers of the 14 species had three pollen tube was maximum (101.60 mins.) for snake
anthers each. Anther size varied proportionately gourd and it was minimum (13.20 mins.) for
based on the size of flowers. Average value for cucumber.
highest number of pollen grain per flower was found Tube lengths of in vitro germinating pollen
to be 917.00 ? 52.69 in pumpkin and the lowest value grains were recorded after 1, 2, 6 and 12 hrs of
was 479.00 ? 13.14) in snake gourd. The germination. The rate of increasing lengths of the
microphotographs showing pollen morphology of tubes was very fast between first two hours and
different cucurbit species are presented in Fig. 1. became very slow in last six hours. After 12 hrs of
germination the longest pollen tube (1649.20? ±
Highest number of viable pollen (98.51%) was 15.05) was found in pumpkin and the shortest
observed in bitter gourd while the lowest (86.87%) in (766.08? ± 19.03) was found in pointed gourd.
pointed gourd. Pollen grains of all the 14 species
were found to be starchy in nature as it was Statistically the differences between different
determined by potassium iodide solution. Pollen values for pollen grains per anther, germination rates,
grains were found to germinate frequently in times required to germinate pollen tube and lengths
Brewbaker and Kwack’s medium. But Roberts’ of pollen tube (after 12 h) among 14 cucurbits were
medium was found not suitable for in vitro pollen found to be highly significant (P<0.001).
Table 1. Pollen characters in different cucurbit species.
Pollen In vitro germinated Pollen Time required to
Mean No. of pollen/
Cucurbits viability pollen grain (five germination germinate pollen
anther ? S.E.
(%) focuses) (%) tube (min.)
Pumpkin 917.00abc* ? 52.69 96.84 348.80cde ? 15.51 95.46 51.80b ? 1.42
Ash gourd 776.67cd ? 11.94 96.79 496.60a ? 18.17 94.30 13.40g ? 0.83
Bottle gourd 656.00de ? 53.13 97.22 388.60abcd ? 14.91 97.19 27.00d ? 1.09
Snake gourd 479.00f ? 13.14 91.08 173.20g ? 3.73 82.55 101.60a ? 1.28
Ridge gourd 553.33ef ? 13.51 97.93 297.40def ? 58.45 85.51 19.80ef ? 0.44
Pointed gourd 581.67ef ? 30.45 86.87 165.80g ? 4.56 77.98 51.60b ? 1.69
Cucumber 761.33cd ? 25.25 96.29 410.00abcd ? 25.59 95.53 13.20g ? 0.44
Cucumber (short) 890.00abc ? 38.18 97.82 373.60bcd ? 35.76 94.63 18.80efg ? 0.44
Bitter gourd 945.00ab ? 16.68 98.51 149.00g ? 6.19 91.19 21.40e ? 1.04
Sweet gourd 956.67a ? 15.33 98.22 245.40efg ? 11.66 92.05 27.20d ? 2.63
Watermelon 850.33abc ? 13.24 94.17 478.00ab ? 36.24 95.03 44.20c ? 0.82
Muskmelon 778.33cd ? 14.53 97.72 422.22abc ? 39.76 88.29 15.20fg ? 0.99
Sponge gourd 669.33de ? 44.71 89.03 367.00bcd ? 8.95 88.01 16.40efg ? 0.61
Ivy gourd 794.33bcd ? 13.46 91.91 228.80fg ? 11.99 80.11 13.40g ? 0.61
*Means followed by same letter are statistically similar as per Duncun’s multiple range test
Pumpkin (X1000) Ash gourd (X250) Bottle gourd (X1000)
Snake gourd (X1000) Ridge gourd (X400) Pointed gourd (X250)
Cucumber (X400) Cucumber (short) (X1000) Bitter gourd (X250)
Sweet gourd (X250) Watermelon (X250) Muskmelon (X100)
Sponge gourd (X1000) Ivy gourd (X250)
Fig. 1. Microphotographs showing morphology of the pollen indifferent cucurbit species.
Table 2. Pollen tube growth in different cucurbit species using Brewbaker and Kwack’s medium.
Mean length (? ) ? S.E. after germination time of
Cucurbits
1 hour 2 hours 6 hours 12 hours
Pumpkin 308.56 ? 38.07 649.04 ? 27.75 1448.84 ? 18.72 1649.20a ? 15.05
Ash gourd 106.40 ? 15.05 498.44 ? 17.80 989.52 ? 24.26 1223.60f ? 15.05
Bottle gourd 329.89 ? 27.75 766.08 ? 19.03 1287.44 ? 23.31 1447.04bcd ? 43.61
Snake gourd 95.76 ? 9.52 585.20 ? 30.09 745.60 ? 60.75 820.08g ? 32.96
Ridge gourd 170.24 ? 9.52 787.36 ? 27.75 1181.04 ? 27.75 1520.32bc ? 35.12
Pointed gourd 106.40 ? 15.05 446.88 ? 24.26 744.80 ? 30.09 766.08g ? 19.03
Cucumber 266.00 ? 15.05 872.48 ? 19.03 1255.52 ? 19.03 1276.80ef ? 33.65
Cucumber (short) 287.28 ? 19.03 872.48 ? 32.27 1244.88 ? 38.66 1500.24bc ? 17.80
Bitter gourd 202.16 ? 9.52 680.96 ? 68.34 1393.84 ? 23.31 1415.12cde ? 35.61
Sweet gourd 127.68 ? 19.03 829.92 ? 44.13 1351.28 ? 32.27 1564.08ab ? 19.03
Watermelon 436.24 ? 17.80 1074.64 ? 17.80 1340.64 ? 34.97 1415.12cde ? 28.55
Muskmelon 659.68 ? 38.66 1127.84 ? 17.80 1393.84 ? 40.93 1372.56de ? 27.75
Sponge gourd 148.96 ? 9.52 819.28 ? 32.27 1170.40 ? 26.06 1415.12cde ? 44.13
Ivy gourd 74.48 ? 11.65 521.36 ? 23.31 893.76 ? 23.31 1181.04f ? 27.75
*Means followed by same letter are statistically similar as per Duncun’s multiple range test
Discussion decrease in viability with the stage of development of
the male gametophyte and found binucleate pollen to
Flowering in cucurbits normally starts at 40-45 survive long time than trinucleate. Pacini et al. (1997)
days old plants, although it varies due to changes of found pollen viability to be decreased with time. The
weather. In typical monoecious sex form, as 14 rapid decrease in pollen viability of Cucurbita pepo
species used in the present study, the numbers of is related to the fact that the flowers of both sexes of
male flowers are sometimes produced in greater this monoecious plant are accessible to insects (bees
proportion than the pistillate flowers in the same and bumble bees) for only 6 hrs and only the pollen is
plant. Thus intraspecific variation in the size of partially dehydrated (Nepi and Pacini, 1993, 2001;
pollen loads deposited on stigmas may influence both Winsor and Stephenson, 1995). After that period the
the number and quality of the eventual progeny flower closes and wilts. There is no adaptive
(Herrera, 2004). Pollen production in different genera advantage to be gained by the pollen for remaining
is variable. In watermelon and pump kin, pollen viable for longer time than the pollinators, which
production is good while in crops like muskmelon the have access to the flower. Polyploid pollen grains of
pollen production is scanty (Seshadri, 1993). In the Cucumis melo showed a lower germination
present experiment the number of pollen per flower percentage in vitro and a slower germination rate
was found higher in sweet gourd followed by bitter than haploid pollen (Susin and Alvarez, 1997).
gourd, pumpkin and cucumber (short). The lowest
number of pollen was recorded in the snake gourd. The duration of pollen viability has been also
studied by Stanley and Linskens (1974) and found
The pollen viability in the present study was not to be correlated with the type of pollination.
found to differ from species to species. Highest However, Ottaviano and Mulcahy (1989) stated that
percentage of pollen viability (98.51%) was recorded pollen must be programmed to have high viability in
for bitter gourd whereas the lowest (86.87%) was the habitat of the species. The duration of pollen
recorded for pointed gourd in Brewbaker and viability varies greatly between species, and is related
Kwack’s medium. Roberts’ medium was not found to to the type of pollination. In general, plants with
be suitable for in vitro pollen germination of entomophilous pollination have pollen with longer
cucurbits and no germination occurred in any of the viability than those with anemophilous pollination
14 species. Stanley and Linskens (1974) correlated (Bassani et al., 1994). Pollen carried on the wind is
dispersed as soon as the anther open, and does not and Aconitum columbianum (Ranunculaceae). Am J Bot
have to await for arrival of the pollinating insect. 86, 871-879.
Brewbaker JL, Kwack BH. 1963. The essential role of calcium
Highest percentage of in vitro pollen germination ion in pollen germination and pollen tube growth. Am J
(97.19%) was recorded for bottle gourd and the Bot 50, 747-758.
lowest (77.98%) for pointed gourd. Germination time
Cai, G., Moscatelli, N. and Cresti, M. 1997. Cytoskeletal
of pollen tube also found to differ among the cucurbit organization and pollen tube growth. Trends Plant Sci., 2:
species. The shortest time taken by cucumber was 86-91.
only 13.20 ± 0.44 against 101.60 ± 1.28 minutes by
Corff JL, Argen J, Schemske DW. 1998. Floral display,
snake gourd. Rapid growth of pollen tube depends on pollinator discrimination, and female reproductiv e
constant fusion of vesicles forming the plasmalemma, success in two monoecious Begonia species. Ecology
and continuous secretion of cell wall material 79(5),1610–1619.
(Blevins and Lukaszewski, 1998). Jackson (1991) Cresti M, Gori P, Pacini E (eds). 1988. Sexual reproduction in
stated that capture of the secreted pollen proteins for higher plants. Springer, Berlin Heidelberg.
membrane and wall building, proceeds through Cresti M, Blackmore S, van Went JL. 1992. Atlas of Sexual
borate complexes with sugar residues. The pollen reproduction in Flowering Plants, Springer. Berlin.
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Delph LF, Johannsson MH, Stephenson AG. 1997. How
forms after germination of the pollen on stigma of a environmental factors affect pollen performance: ecological
receiving flower. The function of the pollen tube is to and evolutionary perspectives. Ecology 78(6), 1632–1639
transmit gametes from pollen grain to the ovary, and
Dieringer G, Cabrera L. 2002. The interaction between
this can occur over long distances (several pollinator size and the bristle staminode of Penstemon
centimetres in some cases) also (Cresti et al., 1992). digitalis (Scrophulariaceae). Am J Bot 89, 991-997.
Because of its fundamental importance to the process
Giles K L, Prakash J (eds). 1987. Pollen cytology and
of fertilization in higher plants, the pollen tube has development. Int Rev Cytol 107. Academic Press,
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aim of understanding the cell biology involved and Gopalan C, Rama-Sastri BV Balasubramanium SC. 1982.
regulating it through biotechnology (Cai et al., 1997; Nutritive Value of Indian Foods. Indian Council of
Stephenson et al., 2003). Medical Research, National Institute of Nutrition,
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Havens K, Delph LF. 1996. Differential seed maturation
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