T. AYOUB
INTRODUCTION
In a previous paper (Ayoub, 1975), the importance of senna (Cassia acutifolia) as a
promising irrigated cash crop was put forward. The crop requires high temperatures
and an abundance of sunlight (Gupta, 1971), and these conditions prevail in the
more arid parts of the Sudan where soil salinity and alkalinity problems may be
important. A study was, therefore, undertaken on the effects of salt levels, salt
types, and soil alkalinity on senna germination, vegetative growth, yield, and
mineral composition. Some environmental factors which may effect salt tolerance
by senna, such as watering interval and air temperature, were also included in
these studies. The present paper reports some primary features of these studies.
MATERIALS AND METHODS
Germination studies
Germ Lnation experiments were carried out i n 10 -cm-diameter Petri dishes in a single randomized
design with 11 levels of NaCl solutions (0-40-0 mmhos cm"1) replicated four times. In each
sterilized Petri dish, 10 healthy surface-disinfected (HgCla) senna seeds were placed on a filter
paper bed, covered with another sheet of filter paper, and soaked with 10 ml of the specified
solution. This volume was sufficient to saturate the filter papers and to partially immerse the
seeds. The Petri dishes were covered and placed in the dark in an incubator maintained at
30 2 C. The germination coimt was taken every 24 h.
Interaction of temperature and salinity on germination
The germination tests were carried out in Petri dishes as described above. Salt levels used
were 0,50, 8-3, 16-3, 24-5, and 33-6 mmhos cm"1 NaCl at 25 C and each test was replicated four
ABSTRACT
Studies on the germination of Cassia acutifolia (senna) seeds indicated that they were highly
tolerant of salinities up to 160 mmhos cm"1, and 50 per cent reduction in germination occurred
at about 20-0 mmhos cm"1. Air temperature had a significant effect on senna germination at
substrate salinities between 10-0 and 25-0 mmhos cm"1.
Seedling growth was more sensitive to salinity and alkalinity than the germination stage.
Young plant survival and total pod yield were significantly reduced in soil salinities higher than
11-0 minhoa cm"1, and this was more pronounced when plants were irrigated at short intervals
with saline waters. The sensitivity of senna to higher levels of salinities was correlated with the
higher rate of chloride accumulation in the tissue resulting in specific chloride injury. However,
the possibility of sodium injury cannot be excluded.
485
times. Replicates were incubated at 12-15, 28-30, 3235, and 40-45 C and germination counts
were made on the 7th day.
ESP tolerance studies
Hudeiba Research Station Farm heavy clay soil (Vertisols) was air-dried, crushed, and sieved
through 2 mm mesh. Soils of different exchangeable sodium percentage (ESP) levels were produced following the method of Chang and Dregne (1955). Final ESP levels obtained were
approximately 5, 15, 24, 35, 45, and 55.
These soils were again air-dried, sieved, and 5'5 kg of each soil placed in 21-cm-diameter
plastic pots and randomized on a ventilated greenhouse bench. Each soil test was replicated
five times. Ten seeds of senna were sown in each pot and irrigated when necessary with lowsalt tap water (0-20 mmhos).
Germination counts were recorded and after a period of 30 d plants were harvested for dry
matter determinations.
Senna seed germination never exceeded 75 per cent and under field conditions the
maximum germination obtained was usually less than 60 per cent. This low germination percentage is mainly due to the presence of 'hard' seeds which do not imbibe
486
10090
80 70| 6 0 -
I 508
302010
4
5
6
Days from sowing
FIG. 1. Senna seed germination at different NaCl levels. The numbers on the graph are NaCl
solution conductivities in mmhos cm"1 at 25 C.
0
water even after 10 d soaking. Laboratory tests have indicated that these seeds
have an impermeable seed coat. The data shown in Fig. 1, presenting germination
percentage in relation to different salt levels of the substrate, are therefore based on
germinable seeds.
Control seeds completed germination on the 7th day. Seed germination was
satisfactory up to a salt level of about 16-0 mmhos cm"1 NaCl. Between salt levels
of 20-0 and 24-0 mmhos cm"1 germination was reduced 50 per cent and beyond the
salt level of 30-0 mmhos cm"1 germination was almost inhibited.
Interaction of salinity and temperature on senna germination
Fig. 2 shows the effect of salinity at different ah" temperatures on senna germination. Substrate salinity, ah- temperature, and their interaction highly significantly
affected senna germination. Up to a salinity of 6-0 mmhos cm"1 there was no difference between temperatures from 28 to 45 C hi germination percentage. Air temperature had its greatest effect on germination at salinities between 10-0 and 25-0
mmhoa cm"1. Fifty per cent reduction in germination occurred at salinities of about
10-0, 170, and 22-0 mmhos cm"1 when air temperatures were 40-45,32-35, and 2830 C, respectively. Senna did not germinate at temperatures below 15 C.
Effect of ESP levels on senna germination and seedling groioth
The data ha Table 1 shows the effect of different ESP levels on senna seed germination and seedling growth. Seed germination was satisfactory up to an ESP level of
I 40
5-0
10-0
350
-O,
35. Beyond that level some seeds germinated, but most of the seedlings died shortly
after that. Seedlings growing in ESP levels of 25 or more were stunted and had
bleached-yellowish (chlorotic) secondary leaves. The effect of ESP was, therefore, more pronounced at the seedling growth stage than during seed germination.
1. Effect of exchangeable sodium percentage (ESP) on senna seed germination
and seedling growth
TABLE
ESP
Relative
germination (%)
Relative
growth (%)
5
15
25
35
45
55
100-0
96-3
88-9
88-9
48-2
33-3
100-0
601
45-8
32-8
25-0
100
300
15-0
20-0
25-0
NsCI levd (mmhos cm"1)
F i o . 2. Effect of NaCl levels a t different air t e m p e r a t u r e s on senna seed germination:
28-30 C;
, 32-35 C; A
A, 40-45 C.
o-o
487
TABLE
00
00
2. Effect of equal conductances of different salt types on senna growth, survival, and mineral composition
Relative
growth
Sftlt treatments
Plant
survival
Type
mmhos cm"1
ITIM
No salt
NaCl
0-2
2-5
50
100
20-0
1-4
22-0
43-9
87-8
175-6
100
96
66
29
17
100
100
88
49
19
4-6
4-4
6-9
19-9
38-0
73-9
67-5
60-9
57-0
42-7
CaCla
2-5
5-0
10-0
20-0
14-6
29-3
58-0
117-2
100
41
23
12
100
86
54
14
4-5
7-1
31
8-2
64-5
54-5
52-4
53-2
Na 2 SO 4
2-5
5-0
10-0
20-0
16-5
33-0
66-0
132-0
79
89
61
23
94
97
85
25
12-3
14-1
13-6
16-7
10-0
20-0
73-2
146-4
46
20
64
25
19-9
30-1
CaCla+NaCl
Ca
Cl
60-9
44-4
51-0
41-9
38-7
69-9
87-3
95-8
103-8
124-8
4-9
18-2
34-2
106-6
121-2
105-3
116-0
120-0
100-3
87-3
108-3
112-3
143-7
190
04-0
94-0
186-0
64-7
63-4
65-2
58-1
105-3
83-9
60-0
53-5
05-9
60-4
91-3
87-3
7-5
7-7
9-5
49-9
52-4
46-1
69-9
128-7
141-2
106-6
107-2
Na
Mg
489
There was a progressive reduction in growth with increasing salinity of the substrate. This effect was more apparent in the plants exposed to chloride salinity.
Fifty per cent reductions in growth were found at 4-0 mmhos cm"1 CaCl2, at 7-0
mmhos cm"1 NaCl, and at about 13-0 mmhos cm"1 Na2 SO4. Data on leaf shedding
and plant survival confirmed the results reported for plant growth.
Mineral composition of shoots
Mineral contents of plant shoots as mEq per 100 g dry wt. are also shown in
Table 2. The control plants contained very little Na and Cl, but their K contents
were high. Increasing NaCl in the substrate increased Cl and Na and decreased K
contents of shoots, Cl being accumulated at much higher rates. Sodium accumulated
in larger amounts in the shoots from NaCl treatments than from Na2S(>4, and Cl
Irrigation interval
7-5
10-1
Watering
effect
16-1
27-8
1-4
5-3
3-3
1-5
5-6
3-6
23-6
130
25-3
81-3
53-3
32-0
93-3
62-7
327-7
187-2
0-3
0-7
0-5
4-3
5-4
7d
14 d
Salt effect
S.e. means salt effect =
7d
14 d
Salt effect
S.e. means salt effect =
No. of
661-3
691-3
254-7
330-0
458-0
510-7
67-20, watering effect
pods/pot
228-3
176-7
202-5
= 42-49.
690
650
67-0
47-0
60-3
53-7
65-9
66-ft
11
1-7
1-4
1-4
1-8
1-6
60
5-2
25-0
28-3
26-7
250
26-0
25-5
43-3
41-6
7d
14 d
Salt effect
S.e. means salt effect =
7d
14 d
Salt effect
S.e. means salt effect =
TABLE
490
accumulated in greater amounts from CaCl2 treatments than from NaCl. Increasing
substrate content of CaCl2 increased shoot Cl, Ca, and Mg contents with slight decrease in K contents.
Effect of frequency of irrigation ivith salt water on senna yield and yield components
Significant reductions in dry matter yields and plant heights occurred at soil
salinities of 10-0 mmhos cm""1 and above. When soil salinity was raised from 7-5
6-0-
Logy =4-9889-00129*
r - -0-895 /><0001
ve pod yield
5-0-
3 3-02
eo
3 2-0-
l-0<
ft-0
55
110
165
220
Leaf bUde O (mEq/100 g dry wt.)
275
F I G . 3. Relation between senna leaf chloride content and log per cent relative pod yield.
Results of senna pod yield and yield components from different NaCl levels are
shown in Table 3.
' Pod yield was significantly affected by salt levels, frequency of irrigation, and
their interaction. Watering every 7 d with lower-salt waters resulted in appreciably
higher yields than watering every 14 d. Contrary to this, frequent irrigation with
high-salt waters injured plants and resulted in lower pod yields than irrigation
every 14 d with the same water.
The number of pods per pot in the various treatments were comparable to pod
yield while increasing salt level resulted in a small but significant (P < 0-05)
depression in pod weight.
Plant survival was significantly reduced by salt levels higher than 10-0 mmhos
cm" 1 in the soil. This effect was more noticeable at the shorter watering interval
than at the longer watering interval, indicating that the effect of NaCl salinity on
senna was not solely an osmotic phenomenon. Injured plants contained considerable
amounts of Cl which was closely related to the degree of visible injury.
Pod yield was highly correlated with pod number (r = 0-9947), less with plant
survival (r = 0-7221), and least with pod weight (r = 0-3818).
491
to 10-1 mmhos cm" reductions of 65 and 35 per cent were found in dry matter and
plant height, respectively. Increasing the watering interval from 7 to 14 d reduced
both parameters but not significantly.
The relationship between pod yield and Cl contents of senna leaves is demonstrated in Fig. 3. Significant yield reduction occurred when the Cl content of the
blade was above 28 mEq per 100 g dry wt. Fifty per cent yield reduction occurred
at Cl content of about 62 mEq, and at Cl content of greater than 110 mEq pod yield
was very low. The Na contents also varied with NaCl levels, but these variations
seemed to be of minor magnitude.
DISCUSSION
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492
ACKNOWLEDGEMENTS
The author is grateful to the Director General, A.R.C, Sudan for permission to
publish this paper and to Dr. R. H. Nieman of U.S. Salinity Laboratory, Riverside,
for his valuable comments and encouragement.
LITERATURE CITED
AYOTTB, ALI T., 1975. J. exp. Bot. 26, 891-96.
BERNSTEIN, L., 1970. Agric. Inform. Bull. No. 283, U.S. Dept. Agrio.
and FRANCOIS, L. E., 1975. Agron. J. 67, 185-90.
CHANG, C. E., and DBEGNE, H. E., 1955. Proc. Soil Set. Soc. Am. 19,
CHAPMAN, H. D., and PRATT, P. F., 1961. In Methods of analysis for
29-35.
soils, plants and waters.
FBANCOIS,
More frequent irrigation with saline water induced, earlier and more severe injury
than less frequent irrigation. Bernstein and Francois (1975) found that more frequent sprinkling with saline waters resulted in greater osmotic shock than less
frequent watering. Moreover, it was noted in this study that water use by the crop
was reduced greatly under high salinity. It is, therefore, inadvisable to give extra
water to senna under saline conditions.
In conclusion, the following points can be mentioned. Senna is moderately tolerant to salinity. It seems to be more salt tolerant at germination and later growth
stages, the seedling stage being the most sensitive. Avoiding adverse environmental
conditions like high air temperature (Francois and Goodin, 1972), low relative
humidity (O'Leary, 1975), and bad cultural practices (Sandoval and Benz, 1973)
at that critical stage may improve the salinity tolerance by senna and its survival
in the field.