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Abstract
Previous studies have shown that salicylic acid (SA) plays a role in the response of plants to salt and osmotic stresses. Therefore, an experiment
was conducted to investigate the impact of exogenous salicylic acid on the growth, physiology and antioxidant activity of carrot (Daucus carota L.
cv. Nantes) grown under combined stress of salinity and boron toxicity. The treatments consisted of salt (control, NaCl, and Na2SO4), boron ( B: 0
and +B: 25 mg B kg 1) and salicylic acid ( SA: 0 and +SA: 0.5 mmol SA kg 1). The diameter of the storage root was increased by NaCl salinity
in the absence of B toxicity, however, it was increased by Na2SO4 salinity under B toxicity. For the storage root yield, NaCl salinity was more toxic
than Na2SO4 salinity. With its role in plant growth regulation, SA application positively affected the storage root dry weight, S concentration,
carotenoids and anthocyanin content and increased the total antioxidant activity (AA) of the shoot and storage root. SA application regulated
proline and toxic ion (B, Cl) accumulation in the storage root and shoot. This study reports the long term effects of SA under stress conditions and
reveals that SA was not as effective as in alleviating abiotic stress as reported in the literature conducted with short-term studies. That means longterm effects of SA would be significantly different from its short-term effects.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Antioxidants; Ascorbate peroxidase; B toxicity; Catalase; Salicylic acid; Salinity
1. Introduction
Salinity and sodicity can reduce plant growth and alter ionic
relations by ionic and osmotic effects (Tarakcioglu and Inal,
2002), and oxidative stress (Borsani et al., 2001). There is an
abundance of information concerning the effects of NaCl,
however, relatively little is known about the responses of plants
to Na2SO4, although there are indications that NaCl is more
phytotoxic than Na2SO4 (Rogers et al., 1998). Boron is
frequently associated with other salts and generally found in
high concentrations in irrigation water of arid regions where
saline soils and saline irrigation water can be prevalent (BenGal and Shani, 2002). Reductions in boron accumulation in the
presence of high sulfate-salinity have been reported in the stems
* Corresponding author. Tel.: +90 312 5961392; fax: +90 312 3178465.
E-mail address: inal@agri.ankara.edu.tr (A. Inal).
1
Future address: Suleyman Demirel University, Faculty of Agriculture,
Department of Soil Sci. & Plant Nutr., Isparta, Turkey.
0304-4238/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2007.03.012
121
122
Boron
(B)
Salt
(S)
Shoot
DW
Storage root
DW
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO
Control
Nacl
Na2SO4
14.3
12.0
14.1
12.7
11.5
12.8
15.9
12.3
41.4
12.9
10.6
14.5
25.8
20.5
25.6
22.6
18.4
30.4
28.2
21.6
27.9
27.9
20.9
30.9
10.0
6.35
9.78
8.95
5.93
9.98
10.2
6.13
10.6
9.18
6.38
9.80
22.2
24.5
22.2
22.6
23.9
23.5
22.0
24.4
22.0
21.9
21.6
23.4
ns
ns
ns
ns
12.9
13.4
23.9
26.2
8.50
8.71
23.2
22.6
ns
**
ns
ns
15.1
12.1
14.3
12.8
11.0
13.7
26.9B
21.0C
26.8B
25.2B
19.6C
30.6A
10.1
6.24
10.2
9.06
6.15
9.89
22.1C
24.5A
22.1C
22.3C
22.7BC
23.5B
ns
**
ns
**
F-test signicance
Mean for SA
SA
+SA
F-test signicance
Mean for B S
B
+B
F-test signicance
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Values in columns followed by different letters, where available according to Ftest significance, are significantly different at p < 0.05. ns: Non-significant and
**
p < 0.01.
123
Table 2
MDA (nmol g 1 FW) content, membrane permeability (EC,%), ascorbic acid (Asc, mmol kg 1 FW) contents, total antioxidant activity (AA, mmol kg 1 FW) of
shoots and storage roots, and proline (mmol kg 1 FW) content, APX and CAT enzyme activity (mmol g 1 DW min 1), and H2O2 content (mmol kg 1 FW) of carrots
shoots
Salicylic
acid
Boron
(B)
F-test
cance
Salt
(S)
MDA
Shoot
Storage root
Shoot
Storage root
Shoot
Storage root
Shoot
Storage root
Proline
APX
CAT
H2O2
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
8.74
8.89
9.65
9.58
8.26
10.8
8.58
9.81
10.1
9.97
9.03
10.8
6.52
6.71
7.49
7.00
6.90
6.70
6.90
6.84
7.32
8.06
7.10
7.30
6.02
6.17
6.08
6.14
6.32
6.38
5.72
6.90
5.78
6.27
6.57
6.92
26.2
26.6
27.2
25.1
25.6
25.9
26.2.
26.4
27.1
25.4
25.7
25.2
20.6
21.4
25.9
22.9
25.9
27.8
23.7
25.1
25.3
25.8
28.2
28.7
13.8A
11.5B
11.9AB
11.0B
12.1AB
12.2AB
12.0AB
12.0AB
11.5B
12.9AB
11.8AB
12.2AB
39.0
34.5
43.1
35.7
35.2
48.5
39.2
36.9
43.5
39.2
35.7
48.3
33.3
32.7
35.7
31.3
34.4
35.8
34.6
37.7
34.0
34.0
40.8
36.2
0.17
0.21
0.32
0.21
0.29
0.33
0.23
0.22
0.38
0.26
0.34
0.32
0.59
1.18
1.03
1.26
1.43
1.37
0.77
1.13
1.01
1.39
1.43
0.98
0.82
1.08
1.15
0.89
1.32
1.22
0.92
1.44
1.03
1.10
1.07
1.11
9.48
12.0
12.2
9.95
11.6
11.5
10.3
11.4
12.7
10.7
10.4
9.33
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
9.33
9.71
6.88
7.25
6.19
6.36
26.1
26.0
24.1
26.1
12.1
12.1
39.3
40.5
33.8
36.2
0.25
0.29
1.14
1.12
1.08
1.11
11.1
10.8
ns
ns
ns
ns
ns
ns
**
**
ns
ns
ns
8.66B
9.35B
9.86AB
9.77AB
8.64B
10.8A
6.71
6.77
7.40
7.53
7.00
7.00
5.87
6.53
5.93
6.21
6.44
6.65
26.2
26.5
27.1
25.2
25.7
25.5
22.2
23.2
25.6
24.4
27.1
28.2
12.9
11.7
11.7
11.9
12.0
12.2
39.1C
35.7DE
43.3B
37.5CD
35.4E
48.4A
33.9
35.2
34.8
32.6
37.6
36.0
0.20B
0.21B
0.35A
0.24B
0.31A
0.32A
0.68
1.16
1.02
1.33
1.43
1.17
0.87
1.26
1.09
0.99
1.19
1.16
9.91
11.7
12.4
10.3
11.0
10.4
ns
ns
ns
ns
ns
**
ns
**
ns
ns
ns
signifi-
EC
Asc
AA
Shoot
Mean for SA
SA
+SA
F-test
cance
signifi-
Mean for B S
B
+B
F-test
cance
signifi-
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Values in columns followed by different letters, where available according to F-test significance, are significantly different at p < 0.05. ns: Non-significant.
*
p < 0.05.
**
p < 0.01.
124
Table 3
Storage root total carotenoids (mg 100 g 1 FW) content, shoot and storage root anthocyanin (Acy, mg 100 g
and carotenoids, mg g 1 FW), and stomatal resistance (s cm 1) of carrot shoots
Salicylic
acid
Boron
(B)
Salt
(S)
Storage roots
total carotenoids
Acy
Shoot
Storage root
Chlorophyll
Carotenoids
Stomatal resistance
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
13.8
14.7
15.3
16.1
16.7
14.7
16.5
16.3
20.1
18.8
17.8
15.5
179
207
221
210
185
196
228
189
217
2.5
175
195
13.0
11.8
14.5
9.44
12.5
13.4
16.1
21.4
17.1
14.5
20.7
18.4
1.81
1.63
1.40
1.72
1.57
1.79
2.02
1.75
1.78
1.38
1.47
1.70
0.36
0.32
0.28
0.35
0.32
0.39
0.40
0.36
0.39
0.30
0.29
0.36
3.96
5.11
5.90
3.77
6.95
5.70
4.75
7.55
6.15
3.82
7.40
6.83
ns
ns
ns
ns
ns
ns
15.2
17.5
200
202
ns
12.4
18.0
1.65
1.68
ns
0.34
0.35
ns
5.23
6.08
15.1B
15.5AB
17.7A
17.5AB
17.3AB
15.1B
204
198
219
207
180
195
1.46
16.6
15.8
12.0
16.6
15.9
1.92A
1.69BC
1.59BC
1.55BC
1.52C
1.74AB
0.38A
0.34ABC
0.33BC
0.32C
0.30C
0.37AB
4.35
6.33
6.02
3.79
7.18
6.27
**
ns
ns
**
**
ns
F-test significance
Shoot
Mean for SA
SA
+SA
**
F-test significance
**
Mean for B S
B
+B
F-test significance
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Values in columns followed by different letters, where available according to F-test significance, are significantly different at p < 0.05. ns: Non-significant.
*
p < 0.05.
**
p < 0.01.
Salicylic
acid
125
Salt
(S)
B
Shoot
Storage root
Shoot
Storage root
Shoot
Storage root
Shoot
Storage root
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
50.0
40.7
51.0
303
161
293
52.7
45.3
50.2
289
148
303
14.3
11.1
17.0
88.8
66.3
95.3
13.9
10.2
19.3
71.9
53.7
89.0
1.46
6.37
7.86
1.44
4.88
6.34
1.56
6.50
7.78
2.03
5.15
6.68
0.75
2.39
3.47
0.83
1.97
2.09
0.73
2.20
3.46
0.81
1.71
2.08
12.3
49.7
10.1
12.4
44.5
11.6
9.90
49.4
9.90
12.0
44.5
8.80
2.40
12.4
1.50
2.50
9.50
1.60
1.70
10.7
0.70
2.30
10.2
1.70
2.46
2.46
4.09
2.88
2.47
3.27
2.69
2.42
3.85
2.85
2.47
3.16
0.36
0.44
0.87
0.50
0.30
0.75
0.39
0.68
0.88
0.49
0.42
0.80
ns
ns
ns
ns
ns
ns
ns
ns
150
148
48.8
43.0
4.72
4.95
1.92
1.83
23.4
22.4
4.98
4.55
2.94
2.91
0.54
0.61
ns
**
ns
ns
ns
**
51.4C
43.0C
50.6C
296A
155B
298A
14.1D
10.7D
18.1D
80.4B
60.0C
92.2A
1.51D
6.43B
7.82A
1.73D
5.01C
6.51B
0.74D
2.30B
3.47A
0.82D
1.84C
2.08BC
11.1CD
49.6A
10.0D
12.2C
44.5B
10.2D
2.05C
11.6A
1.10D
2.40C
9.85B
1.65CD
2.58CD
2.44D
3.97A
2.86C
2.47D
3.21B
0.38D
0.56C
0.88A
0.49C
0.36D
0.77B
**
**
**
**
**
**
**
**
F-test significance
Na
Cl
Mean for SA
SA
+SA
F-test significance
Mean for B S
B
+B
F-test Significance
Control
NaCl
Na2SO4
Control
NaCl
Na2SO4
Values in columns followed by different letters, where available according to F-test significance, are significantly different at p < 0.05. ns: Non-significant.
*
p < 0.05.
**
p < 0.01.
126
127
Alalsavar, C., Al-Farsi, M., Quantick, P.C., Shahidi, F., Wiktorowicz, R., 2005.
Effect of chill storage and modified atmosphere packaging (MAP) on
antioxidant activity, anthocyanins, carotenoids, phenolics and sensory
quality of ready-to-eat shredded orange and purple carrots. Food Chem.
89, 6976.
Al-Hakimi, A.M.A., Hamada, A.M., 2001. Counteraction of salinity stress on
wheat plants by grain soaking in ascorbic acid, thiamin or sodium salicylate.
Biol. Plant. 44, 253261.
Alpaslan, M., Gunes, A., 2001. Interactive effects of boron and salinity stress on
the growth, membrane permeability and mineral composition of tomato and
cucumber plants. Plant Soil 236, 123128.
Apostol, K.G., Zwiazek, J.J., MacKinnon, M.D., 2002. NaCl and Na2SO4 alter
responses of jack pine (Pinus banksiana) seedlings to boron. Plant Soil 240,
321329.
Asada, K., 1992. Ascorbate peroxidase-a hydrogen peroxide scavenging
enzyme in plants. Physiol. Plant. 55, 235241.
Asrar, Z.A., Khavari-Nejad, R.A., Heidari, H., 2005. Excess manganese effects
on pigments of Menta spicata at flowering stage. Arch. Agron. Soil Sci. 51,
101107.
Bates, L.S., Waldren, R.P., Teare, J.D., 1973. Rapid determination of proline for
water stress studies. Plant Soil 39, 205207.
Ben-Gal, A., Shani, U., 2002. Yield, transpiration and growth of tomatoes under
combined excess boron and salinity stress. Plant Soil 247, 211221.
Borsani, O., Valpuesta, V., Botella, M.A., 2001. Evidence for a role of salicylic
acid in the oxidative damage generated by NaCl and osmotic stress in
Arabidopsis seedlings. Plant Physiol. 126, 10241030.
., 2003. Molecular events following perception of ultraBrosche, M., Strid, A
violetB radiation by plants. Physiol. Plant. 117, 110.
Brugnoli, E., Bjorkman, O., 1992. Growth of cotton under continuous salinity:
influence on allocation pattern, stomatal and light energy. Planta 187, 335
347.
Change, B., Maehly, A.C., 1955. Assay of catalases and peroxidase. Meth.
Enzymol. 2, 764775.
Cheng, I.F., Zhao, C.P., Amolins, A., Galazka, M., Doneski, L., 1996. A
hypothesis for the in vivo antioxidant activity of salicylic acid. Biometals
9, 285290.
Dat, J.F., Lopez-Delgado, H., Foyer, C.H., Scott, I.M., 1998. Parallel changes in
H2O2 and catalase during thermotolerance induced by salicylic acid or heat
acclimation in mustard seedlings. Plant Physiol. 116, 13511357.
Dat, J., Vandenabeele, S., Vranova, E., Van Montagu, M., Inze, D., Van
Breusegem, F., 2000. Dual action of the active oxygen species during plant
stress responses. Cell Mol. Life Sci. 57, 779795.
Edreva, A., Gurel, A., Gesheva, E., Hakerlerler, E., 2002. Reddening of cotton
(Gosspiyum hirsutum L ). Leaves Biol. Plant. 45, 303306.
El-Motaium, R., Hu, H., Brown, P., 1994. The relative tolerance of six Prunus
rootstocks to boron and salinity. J. Am. Soc. Hort. Sci. 119, 1169
1175.
Everard, J.D., Gucci, R., Kann, S.C., Flore, J.A., Loescher, W.H., 1994. Gas
exchange and carbon partitioning in the leaves of celery (Apium graveolens)
at various levels of root zone salinity. Plant Physiol. 106, 281292.
Fadzilla, N.M., Gill, V., Finch, R.P., Burdon, R.H., 1996. Chilling, oxidative
stress and antioxidant responses in shoot cultures of rice. Planta 199, 552
556.
Gibberd, M.R., Turner, N.C., Storey, R., 2002. Influence of saline irrigation on
growth, ion accumulation and partitioning, and leaf gas exchange of carrot
(Daucus carota L.). Ann. Bot. 90, 715724.
Gossett, D.R., Banks, S.W., Millhollon, E.P., Cran Lucas, M., 1996. Antioxidant
response to NaCl stress in a control and a NaCl-tolerant cotton cell line
grown in the presence of paraquat, buthionine sulfoximine and exogenous
glutathione. Plant Physiol. 112, 803809.
Grattan, S.R., Grieve, C.M., Poss, J.A., 2005. Does salinity reduce borons toxic
effect in broccoli? [Abstract]. Am. Soc. Hort. Sci. 40,1075. American
Society of Horticulture Science Meeting.
Grieve, C.M., Poss, J.A., 2000. Wheat response to interactive effects of boron
and salinity. J. Plant Nutr. 23, 12171226.
Gueta-Dahan, Y., Yaniv, Z., Zilinskas, B.A., Ben-Hayyim, G., 1997. Salt and
oxidative stress: similar and specific responses and their relation to salt
tolerance in Citrus. Planta 203, 460469.
128
Gunes, A., Inal, A., Alpaslan, M., Cicek, N., Guneri, E., Eraslan, F., Guzelordu,
T., 2005. Effects of exogenously applied salicylic acid on the induction of
multiple stress tolerance and mineral nutrition in maize (Zea mays L.). Arch.
Agron. Soil Sci. 51, 687695.
Harborne, J.B., 1977. Flavonoid sulphates: a new class of natural product of
ecological significance in plants. In: Reinhold, L., Harborne, J.B., Swain, T.
(Eds.), Progress in Phytochemistry, vol. 4. Pergamon, New York, USA, pp.
189208.
Harper, J.R., Balke, N.E., 1981. Characterization of the inhibition of K+
absorption in oats roots by salicylic acid. Plant Physiol. 68, 13491353.
He, Y.L., Liu, Y.L., Chen, Q., Bian, A.H., 2002. Thermotolerance related to
antioxidation induced by salicylic acid and heat hardening in tall fescue
seedlings. J. Plant Physiol. Mol. Biol. 28, 8995.
Hodges, D.M., DeLong, J.M., Forney, C.F., Prange, R.K., 1999. Improving the
thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds. Planta 207, 604611.
Ismail, A.M., 2003/2004. Response of maize and sorghum to excess boron and
salinity. Biol. Plant. 47, 313316.
Johnson, J.M., Ulrich, A., 1959. Analytical methods for use in plant analysis. II,
Bull. No. 766, California Agric. Exp. Sta. Berkeley, CA.
ktem, H.A., 2003. Antioxidant responses of tolerant
Karabal, E., Yucel, M., O
and sensitive barley cultivars to boron toxicity. Plant Sci. 164, 925933.
Keles, Y., Oncel, I., Yenice, N., 2004. Relationship between boron content and
antioxidant compounds in Citrus leaves taken from field with different
water source. Plant Soil 265, 345353.
Kennedy, B.F., De Filippis, L.F., 1999. Physiological and antioxidative
responses to NaCl of the salt tolerant Grevillea ilicifolia and salt sensitive
Grevillea arenaria. J. Plant Physiol. 155, 746754.
Khan, W., Prithiviraj, B., Smith, D.L., 2003. Photosynthetic responses of corn
and soybean to foliar application of salicylates. J. Plant Physiol. 160, 485
492.
Klessig, D.F., Malamy, J., 1994. The salicylic acid signal in plants. Plant Mol.
Biol. 26, 14391458.
Kraus, T.E., Fletcher, R.A., 1994. Paclobutrazol protects wheat seedlings from
heat and paraquat injury. Is detoxification of active oxygen species
involved? Plant Cell Physiol. 35, 4552.
Lichtenthaler, H.K., Wellburn, W.R., 1983. Determination of total carotenoids
and chlorophylls a and b of leaf extracts in different solvents. Biochem. Soc.
Trans. 11, 591592.
Lin, C.C., Kao, C.H., 2000. Effect of NaCl stress on H2O2 metabolsim in rice
leaves. Plant Growth Reg. 30, 151155.
Metwally, A., Finkemeier, I., Georgi, M., Dietz, K.-J., 2003. Salicylic acid
alleviates the cadmium toxicity in barley seedlings. Plant Physiol. 132, 272
281.
Molassiotis, A., Sotiropoulos, T., Tanou, G., Diamantidis, G., Therios, I., 2006.
Boron-induced oxidative damage and antioxidant and nucleolytic responses
in shoot tips culture of the apple rootstock EM9 (Malus domestica Borkh).
Environ. Exp. Bot. 56, 5462.
Mukherjee, S.P., Choudhuri, M.A., 1983. Implications of water stress-induced
changes in the levels of endogenous ascorbic acid and hydrogen peroxide in
Vigna seedlings. Physiol. Plant. 58, 166170.
Nakano, Y., Asada, K., 1981. Hydrogen peroxide is scavenged by ascorbatespecific peroxidase in spinach chloroplasts. Plant Cell Physiol. 22, 867880.
Navarro, J.M., Garrido, C., Martinez, V., Carvajal, M., 2003. Water relations
and xylem transport of nutrients in pepper plants grown under two different
salts stress regimes. Plant Growth Reg. 41, 237245.
Page, A.L., Miller, R.H., Keeney, D.R., 1982. Methods of Soil Analysis. Part 2:
Chemical and Microbiological Properties, second ed. SSSA, Madison,
Wisconsin, USA.
Poss, J.A., Grattan, S.R., Grieve, C.M., Shannon, M.C., 1999. Characterization
of leaf boron injury in salt-stressed Eucalyptus by image analysis. Plant Soil
206, 237245.
Prieto, P., Pineda, M., Aguilar, M., 1999. Spectrophotometric quantitation of
antioxidant capacity through the formation of a phosphomolybdenum
complex: specific application to the determination of vitamin E. Anal.
Biochem. 269, 337341.
Reay, F.P., Fletcher, R.H., Thomas, V.J.G., 1998. Chlorophylls, carotenoids and
anthocyanin concentrations in the skin of Gala apples during maturation
and the influence of foliar applications of nitrogen and magnesium. J. Sci.
Food Agric. 76, 6371.
Rogers, M.E., Grieve, C.M., Shannon, M.C., 1998. The response of lucerne
(Medicago sativa L.) to sodium sulphate and chloride salinity. Plant Soil
202, 271280.
Sairam, R.K., Srivastava, G.C., Agarwal, S., Meena, R.C., 2005. Differences in
antioxidant activity in response to salinity stress in tolerant and susceptible
wheat genotypes. Biol. Plant. 49, 8591.
Senaratna, T., McKay, C.E., McKersie, B.D., Fletcher, R.A., 1988. Uniconazole
induced chilling tolerance in tomato and its relationship to antioxidant
content. J. Plant Physiol. 133, 5661.
Shakirova, F.M., Sakhabutdinova, A.R., Bezrukova, M.R., Fatkhutdinova, R.A.,
Fatkhutdinova, D.R., 2003. Changes in the hormonal status of wheat
seedlings induced by salicylic acid and salinity. Plant Sci. 164, 317322.
Smirnoff, N., 1993. The role of active oxygen in the response of plants to water
deficit and desiccation. New Phytol. 125, 2758.
Tarakcioglu, C., Inal, A., 2002. Changes induced by salinity, demarcating
specific ion ratio (Na/Cl) and osmolality on ion and proline accumulation,
nitrate reductase activity, and growth performance of lettuce. J. Plant Nutr.
25, 2741.
Tissa, S., Darren, D., Eric, B., Kinsley, D., 2000. Acetyl salicylic acid (aspirin)
and salicylic acid induce multiple stress tolerance in bean and tomato plants.
Plant Growth Reg. 30, 157161.
Turan, M., Aydin, A., 2005. Effects of different salt sources on growth,
inorganic ions and proline accumulation in corn (Zea mays L.). Eur. J.
Hort. Sci. 70, 149155.
Wolf, B., 1971. The determination of boron in soil extracts, plant materials,
composts, manures, water and nutrient solutions. Commun. Soil Sci. Plant
Anal. 2, 363374.
Yadav, H.D., Yadav, O.P., Dhankar, O.P., Oswal, M.C., 1989. Effect of chloride
salinity and boron on germination, growth, and mineral composition of
chickpea (Cicer arietinum L.). Annals Arid Zone 28, 6367.
Yan, B., Dai, Q., Liu, X., Huang, S., Wang, Z., 1996. Flooding-induced
membrane damage, lipid oxidation and activated oxygen generation in
corn leaves. Plant Soil 179, 261268.