Salt Stress Inhibits Germination and Early Seedling Growth in Cabbage (Brassica oleracea capitata L.)
Kyeong Bo Lee,
Kwang Yong Jung,
Deog Bae Lee,
Mi Suk Han
Eui Shik Rha
Salinity induced inhibition in germination and early stages of cabbage (Brassica oleracea capitata L.) [two varieties (autumn cabbage and spring cabbage)] were measured in response to increasing NaCl concentration. The salinity (NaCl) concentrations in solution were 0 (control), 4.7, 9.4 and 14.1 dS m- 1. Different concentrations of salt stress had considerable effect on germination, germination rate (1/t50, where t50 is the time to 50% of germination), root and shoot lengths, root, shoot and plant fresh weight of cabbage. Final germination in cabbage (autumn cabbage and spring cabbage) showed significant inhibition with increasing salt stress up to 14.1 dS m- 1 NaCl. The required time for germination increased with increasing concentration of salt. The seedling growth was strongly inhibited by all salt levels, particularly at 14.1 dS m- 1. Furthermore Root growth was more affected then shoots growth by salt stress. Fresh weights of root, shoot and plant were also severely affected by different salinity treatments. Linear regression revealed a significant negative relationship between salinity and final germination, germination rate, root and shoot lengths and fresh weights of roots, shoots and plants.
Soil salinity is one of the most important constraints that limit crop production in arid and semi arid regions (Neumann, 1995). Irrigated land is particularly at risk with approximately one third being significantly affected by salinity. Despite its relatively small area, irrigated land is estimated to produce one third of the world s food (Munns, 2002). The growth of plants is ultimately reduced by salinity stress although plant species differ in their tolerance to salinity (Munns and Termaat, 1986). One approach to reducing the deleterious effects of soil salinity on crop production is the development of salt tolerant cultivars (Epstein et al., 1980). Although some crops are moderately tolerant of saline conditions, many crops are adversely affected by even low levels of salt (Greenway and Munns, 1980). The salt tolerance of crops has generally been expressed as the yield decreased expected for a given level of soluble salts in the root medium as compared with yield under nonsaline conditions (Maas and Hoffman, 1977).
There have been numerous reviews of the effects of salinity on plant physiological processes and subsequent effects on yield (Greenway and Munns, 1980; Munns, 1993; Shannon et al., 1994; Neumann, 1995). In spite of this extensive literature there is still a controversy with regard to the mechanisms of salt tolerance in plants (Neumann, 1995). Salinity impairs seed germination, reduces nodule formation, retards plant development and reduces crop yield (Greenway and Munns, 1980). Salt and water stresses could reduce germination either by limiting water absorption by the seeds (Dodd and Donovan, 1999). Germination in saline seedbeds may be restricted by low soil moisture and osmotic potential or by toxic concentrations of specific ions (Roundy, 1987). Salinity stress can affect seed germination through osmotic effects (Welbaum et al., 1990). Successful seedling establishment depends on the frequency and the amount of precipitation as well as on the ability of the seed species to germinate and grow while soil moisture and osmotic potentials decrease (Roundy, 1985).
The present study was undertaken to evaluate the effect of NaCl salinity tolerance in two varieties of cabbage differing in salt tolerance during seed germination and early seedling growth and also to find the relation between salinity and growth.
MATERIALS AND METHODS
Seeds of cabbage (Brassica oleracea capitata L.) [two varieties (autumn cabbage cv Gaeul baechu and spring cabbage cv Bom baechu)] were obtained from Jeollabuk-do Agricultural and Extension Services, Iksan, Korea, were used in this experiment.
Experiment was established to evaluate the effect of different NaCl concentrations on germination, germination rate (1/t50, where t50 is the time to 50% of germination), root and shoot lengths, fresh weights of roots, shoots and plants of the seedlings. Plastic Petri dishes (87 mm diameter, 15 mm height) with a tight-fitting lid were used for the experiment. The solution consisted of 0.0 (control), 4.7, 9.4 and 14.1 dS m- 1. Ten seeds for each of the four NaCl treatments as well as control were used. Seeds were hand sorted to eliminate broken and small seeds. Seed were allowed to germinate in laboratory condition on filter paper (Whatman No. 2) in Petri dishes soaked with 5 mL of the respective salt concentration. Petri dishes were sealed with parafilm to prevent evaporation of water and minimizing the changes in concentration of the solutions.
Seed germination was evaluated after every 12 h. After 36 h seeds had started to germinate (seeds were considered to be germinated with the emergence of the radical). The germinating seeds were counted at daily intervals. The lengths of roots and shoots of the germinated seeds, which were more than 2 mm in length, were measured and recorded after 20 days of sowing. In all treatments a continuous increase in the number of germinating seeds as well as in the lengths of roots and shoots was observed during the subsequent days of germination.
The experiment was established by using a randomized complete block design with three replications. Analysis of variance was performed using MS-Excel and differences between the means were compared through Least Significant Difference (LSD) test (p<0.05) (Li, 1964). Linear regression equations were developed by using Minitab version 14.0 statistical software package.
Germination percentage of cabbage (spring cabbage and autumn cabbage) was strongly
inhibited by all salinity treatments. The inhibition being strongest particularly
at the higher level of salt treatment compared to control. Highest percentage
of germination was observed in autumn cabbage while lowest germination was investigated
in spring cabbage (Fig. 1A). The final germination rate of
seeds of these plant species under various conditions of salinity was expressed
as a 1/t50 of the germination of seeds of the same population as
Effect of different treatments of NaCl stress on germination
(A) and germination rate (B) of cabbage (autumn cabbage and spring cabbage)
Seed germination delayed as the salinity level increased. The germination
response of both varieties under investigation showed marked differences in
the timing of initiation and completion of germination. The germination started
within 36 h and was complete on the 6th day. Figure 1B showed
that autumn cabbage comparatively took more time to complete germination.
Experiment was prolonged to investigate the effect of salinity (NaCl) on seedling
vigor of germinating seeds of cabbage (autumn cabbage and spring cabbage). It
was investigated that an increased salinity level caused delayed emergence of
root and shoot as compared to controls. The increase in length of root and shoot
was continuously observed in frequent hours of germination in both cabbage varieties
in the control as well as salt treatments. The average length (Fig.
2) of root and shoot of the seedlings of the both cabbage varieties raised
in increasing levels of salt solutions shows that both cabbage varieties showed
a strong inhibition, but the magnitude of decrease in length was more prominent
in root as compared to shoot in all NaCl salt treatments in both. Highest reduction
of root and shoot length was in spring cabbage as compared to autumn cabbage
Effect of different treatments of NaCl stress on root length
(A) and shoot length (B) of cabbage (autumn cabbage and spring cabbage)
Increase in salt concentration caused a significant reduction in fresh weights.
Result presented in Fig. 3 revealed highly significant differences
in cabbage (spring cabbage and autumn cabbage) for root, shoot and plant fresh
weight. Figure 3 also shows that fresh weight of root, shoot
and plant of cabbage (autumn cabbage and spring cabbage) was strongly affected
by all salinity treatments. Root, shoot and plant fresh weight was significantly
inhibited in both varieties at all salinity levels (4.7-14.1 dS m- 1
NaCl), whereas fresh shoot weight was reduced more as compared to fresh root
weight. Reduction in fresh shoot weight was more in spring cabbage then autumn
cabbage (Fig. 3B). Fresh root weight of autumn cabbage was
strongly inhibited by all salt treatments as compared to spring cabbage (Fig.
3B). Maximum reduction in plant weight was investigated in autumn cabbage
Linear regression equations were developed to find the relationship between
salinity and final germination, germination rate, root and shoot lengths and
fresh weights of roots, shoots and plants (Table 1). Linear
regression revealed a significant negative relationship between salinity and
final germination, germination rate, root and shoot lengths and fresh weights
of roots, shoots and plants.
Effect of different treatments of NaCl stress on fresh
weights of roots (A), shoots (B) and plants (C) of cabbage (autumn cabbage
and spring cabbage)
Relationship between salinity and germination percentage,
germination rate, root length, shoot length and fresh weight of plants,
shoots and roots.
| x denotes the parameters in linear equations
Linear regression also revealed a strong (R2 = 0.98 p<0.001)
negative significant relationship between salinity and germination rate. There
was also a weak (R2 = 0.61 p = 0.02) negative significant relationship
between Salinity and shoot length (Table 1).
Seed germination is important growth stage often subject to high mortality
rates. Salinity inhibits the germination and germination rate of cabbage (autumn
cabbage and spring cabbage) as the salt treatment increased (Fig.
1). It is assumed that in addition to toxic effects of certain ions, higher
concentration of salt reduces the water potential in the medium, which hinders
water absorption by germinating seeds and thus reduces germination (Maas and
Nieman, 1978). Salt-induced inhibition of seed germination could be attributed
to osmotic stress or to specific ion toxicity (Huang and Redmann, 1995). These
results are similar in line with Francois et al. (1984). They found that
soil salinity up to 50 mM did not significantly inhibit germination of Sorghum
bicolor seeds, but salt levels greater than 100 mM delayed germination.
Some of the results obtained in this study are similar to those of some other
workers who showed that in general, increased salinity results in decrease in
germinability and delayed rate of germination (El-Sharkawi and Springuel, 1979).
In 1985 Ayers and westcot investigated that salinity delay germination of several
species but does not appreciably reduce the final germination percentage.
The root and shoot length are the most important parameters for salt stress
because roots are in direct contact with soil and absorb water from soil and
shoot supply it to the rest of the plant. For this reason, root and shoot length
provides an important clue to the response of plants to salt stress (Jamil and
Rha, 2004). Figure 2 showed that lengths of roots and shoots
of cabbage (autumn cabbage and spring cabbage) raised in increasing levels of
salt solutions showed a strong inhibition. It was also observed that the degree
of reduction increased with the increasing concentration of salt. Inhibition
of plant growth by salinity may be due to the inhibitory effect of ions. High
salinity may inhibit root and shoot elongation due to slowing down the water
uptake by seeds (Werner and Finkelstein, 1995) may be another reason for this
decrease. Reduction of plant growth under saline conditions is a common phenomenon
in plants (Ashraf and Harris, 2004), but such reduction occurs differently in
different plant organs. For example in present study, decrease in length of
root was more prominent as compared to shoot in all NaCl salt treatments. (Fig.
2). Similar kind of results was earlier reported by Jamil et al.
(2005). They observed that Salt stress inhibited the growth of shoot more than
root in Brassica species. Demir and Arif (2003) also obtained similar
results. They observed that the root growth of safflower was more adversely
affected compared to shoot growth by soil salinity. Our results were also similar
with the findings of Hussain and Rehman (1995, 1997). They found that the roots
of seedlings were more sensitive than the shoots.
Fresh weights of roots, shoots and plants were significantly inhibited in cabbage
(autumn cabbage and spring cabbage) at all salinity levels (Fig.
3). Some researchers could argue that because dry weights were not much
affected compared to the fresh weights, growth reduction would be attributable
to osmotic effects. Jeannette et al. (2002) reported that faster rate
of germination allowed the emerging seedlings to accumulate more biomass relative
to the control but conversely, total fresh weight of root and shoot of cultivated
accessions was significantly reduced with increased salt stress. These results
are also similar in line with Shannon and Grieve (2000) indicated that salinity
reduced fresh weight of all nine vegetables with increasing salt concentration.
Linear regression revealed a significant negative relationship between salinity
and final germination, germination rate, lengths of roots and shoots and fresh
weights of roots, shoots and plants (Table 1). A negative
relation of salinity on germination has been reported in several studies (Boorman,
1968; Khan and Ungar, 1984; Greenway and Munns, 1980). Salinity reduced total
plant biomass by negatively affecting root, stem and leaf mass (Greenway and
Munns, 1980; Ashraf and Harris, 2004).
The present research was conducted by the research fund of Rural Development Administration (RDA), Korea in 2006.
Ashraf, M. and P.J.C. Harris, 2004. Potential biochemical indicators of salinity tolerance in plants. Plant Sci., 166: 3-16.
CrossRef | Direct Link |
Ayers, R.S. and D.W. Westcot, 1985. Water quality for agriculture. FAO Irrigation and Drainage Paper No. 29, (Rev.), FAO, Rome, pp: 174
Boorman, L.A., 1968. Some aspects of the reproductive biology of Limonium vulgare Mill. and Limonium humile Mill. Ann. Bot., 32: 803-824.
Dodd, G.L. and L.A. Donovan, 1999. Water potential and ionic effects on germination and seedling growth of two cold desert shrubs. Am. J. Bot., 86: 1146-1153.
PubMed | Direct Link |
El-sharkawi, H.M. and I.V. Springuel, 1979. Germination of some crop plant seeds under salinity stress. Seed Sci. Technol., 15: 151-162.
Epstein, E., J.D. Norlyn, D.W. Rush, R.W. Kinsbury, D.B. Kelly, G.A. Cunningham and A.F. Wrona, 1980. Saline culture of crops: A genetic approach. Sci., 210: 399-404.
CrossRef | Direct Link |
Francois, L., E.T. Dononvan and E.V. Mass, 1984. Salinity effects on seed yield, growth and germination of grain sorghum. Agron. J., 76: 741-744.
Greenway, H. and R. Munns, 1980. Mechanisms of salt tolerance in nonhalophytes. Ann. Rev. Plant Physiol., 31: 149-190.
Huang, J. and R.E. Redmann, 1995. Salt tolerance of Hordeum and Brassica species during germination and early seedling growth. Can. J. Plant Sci., 75: 815-819.
Hussain, M.K. and O.U. Rehman, 1995. Breeding sunflower for salt tolerance: Association of shoot growth and mature plant traits for salt tolerance in cultivated sunflower (Helianthus annuus L.). Helia, 18: 69-76.
Hussain, M.K. and O.U. Rehman, 1997. Evaluation of sunflower (Helianthus annuus L.) germplasm for salt tolerance at the shoot stage. Helia, 20: 69-78.
Jamil, M. and E.S. Rha, 2004. The effect of salinity (NaCl) on the germination and seedling of suger beet (Beta vulgaris L.) and cabbage (Brassica oleracea capitata L.). Korean J. Plant Res., 7: 226-232.
Jamil, M., C.C. Lee, S.U. Rehman, D.B. Lee, M. Ashraf and E.S. Raha, 2005. Salinity (NaCl) tolerance of Brassica species of germination and early seedling growth. EJEAFChe, 4: 970-976.
Direct Link |
Jeannette, S., R. Carig and J.P. Lynch, 2002. Salinity tolerance of Phaseolus species during germination and early seedling growth. Crop Sci., 42: 1584-1594.
Kaya, M.D., A. Ipek and A. Ozturk, 2003. Effects of different soil salinity levels on germination and seedling growth of safflower (Carthamus tinctorius L.) Turk. J. Agric. For., 27: 221-227.
Direct Link |
Khan, M.A. and I.A. Ungar, 1984. The effect of salinity and temperature on the germination of polymorphic seeds and growth of Atriplex triangularis wild. Am. J. Bot., 71: 481-489.
Li, C.C., 1964. Introduction to Experimental Statistics. McGraw Hill Book Company, New York, USA.
Maas, E.V. and R.H. Nieman, 1978. Physiology of plant tolerance to salinity. In: Crop Tolerance and suboptimal land conditions. Chapter, 13: 277-299.
Maas, E.V., 1990. Crop Salt Tolerance. In: Agricultural Salinity Assessment and Management, Tanji, K.K. (Ed.). American Society of Civil Engineers, New York, pp: 262-304.
Munns, R. and A. Termaat, 1986. Whole plant responses to salinity. Aust. J. Plant Physiol., 13: 143-160.
CrossRef | Direct Link |
Munns, R., 1993. Physiological processes limiting plant growth in saline soils: Some dogmas and hypotheses. Plant Cell Environ., 16: 15-24.
CrossRef | Direct Link |
Neumann, P.M., 1995. Inhibition of Root Growth by Salinity Stress: Toxicity or an Adaptive Biophysical Response. In: Structure and Function of Roots, Baluska, F., M. Ciamporova, O. Gasparikova and P.W. Barlow (Eds.). Kluwer Academic Publishers, Dordrecht, The Netherlands, pp: 299-304.
Roundy, B.A., 1985. Root penetration and shoot elongation of tall wheatgrass and basin wild rye in relation to salinity. Can. J. Plant Sci., 65: 335-343.
Roundy, B.A., 1987. Seedbed salinity and the establishment of range plants. Proceedings of the Symposium on Seed and Seedbed Ecology of Rangeland Plants, April 20-24, USDA-ARS, Washington, DC., pp: 68-71.
Shannon, M.C., C.M. Grieve and L.E. Francois, 1994. Whole-Plant Response to Salinity. In: Plant-Environment Interactions, Wilkinson, R.E. (Ed.). Marcel Dekker, New York, pp: 199-244.
Shannon, M.C., C.M. Grieve, S.M. Lesch and J.H. Draper, 2000. Analysis of salt tolerance in nine leafy vegetables irrigated with saline drainage water. J. Am. Soc. Hortic. Sci., 125: 658-664.
Direct Link |
Welbaum, G.E., T. Tissaoui and K.J. Bradford, 1990. Water relations of seed development and germination in muskmelon (Cucumis melo L.). 111. Sensitivity of germination to water potential and abscisic asid during development. Plant Physiol., 92: 1029-1037.
PubMed | Direct Link |
Werner, J.E. and R.R. Finkelstein, 1995. Arabidopsis mutants with reduced response to NaCl and osmotic stress. Physiol. Plant., 93: 659-666.
CrossRef | Direct Link |