Introduction
More than 955 million hectares of saline-alkali
land are distributed all over the world. It accounts 2/3rd of the total cultivated lands, and its
area has still been increasing (Lin et al. 2012). Global annual population increase, coupled with decreasing
arable land and a growing condition of food crisis has attracted more and more
attention to the development and utilization of salinized land (Guo et al. 2015). Soil salinization occurs due to
excessive accumulation of salinity, which causes severe water loss to crops.
So, saline soils not only reduce the biological and economic
yield of crops strongly, but also poses a severe impact on product
quality (Guo et al. 2015). Introduction of new varieties of salt-tolerant crops in those
areas and to bring the crops under cultivation appears to be the most
economical and effective way to overcome the problem (Xu et al. 2019).
Peanut (Arachis hypogea L., Fam.: Fabaceae) is an important oil crop in the People’s Republic
of China and the crop is suitable for growing in neutral acid soil (Xu et al. 2019).
However, salt-alkali stress affects peanut yield. Previous studies have shown
that different peanut varieties have different salt stress tolerance. It has
been seen that with the increase of salt concentration, plant growth rate,
plant water content, leaf water potential, photosynthetic pigments, total
carbohydrates and other traits all decreased significantly (Hammad et al. 2010). Peanut with high oleic acid
content refers to peanut varieties with the oleic acid content of 75% or more
of the total fatty acids (Moore and Knauft 1989). It has a strong antioxidant capacity,
beneficial to human health, good storage durability, and the shelf life of
processed food is significantly prolonged. As a result, the market prospects
are optimistic. Therefore, if it is possible to cultivate
saline-alkali-tolerant peanut varieties with high oleic acid content, it will
be of great significance for the development of peanut production. The
cultivation prospect would then expand the peanut planting area, increase farmers'
income ensuring the safety of edible oil. In recent years, considerable
information on the mechanism of salt tolerance of ordinary peanuts has been
obtained. Tolerance to salinity stress in peanut is conferred by a higher
allocation of assimilation capacity to the kernel through maintaining total
sugar and chlorophyll-a contents close to unstressed
treatment (Mohammad et al. 2012). The salt sensitivity of peanut is the sum of the effect of
rooting zone NaCl on biomass and the effect of pegging zone NaCl on seed
development (Smitharani et al. 2014). WRKY and Na+/H+ genes might be
responsible for imparting tolerance to salinity stress in peanut (Bera et al. 2013). But the information on high oleic acid peanut varieties was not
available, just on the production and quality of high oleic acid peanuts under
salt and alkali stress ((Su et al. 2017; Su et al. 2018; Xu et al. 2019). For the first time and through a two-year field salt-tolerance
stress study, Wang put forwarded the identification standard of salt-tolerance
of high-oleic peanuts based on relative yield (Su et al. 2017). Chi Xiaoyuan
found different levels of reduction on the oil content, oleic acid content, oil
sub-proportion of high-oleic acid peanut varieties (lines) planted in
saline-alkali land (Xu et al. 2019). In the present experiment, four high-oleic acid varieties
(lines) with different responses to saline-alkali obtained through a two-year
field salt-alkali stress test were screened. For the first time, the
physiological indexes of saline-alkaline combined with yield and yield-related
traits to discuss the resistance mechanism of high-oleic peanut varieties,
which will provide a reference for the future selection of salt-tolerant for
high-oleic peanut varieties.
Materials and Methods
Experiment 1
The field test was carried out between
2016 and 2017 in Baicheng Jilin Province (122.83° E
45.62 °N) which includes
saline-alkali and cinnamon soil. The experimental materials were 27 high oleic
acid varieties (lines) of peanuts which were collected from Shandong Peanut
Research Institute. The test plots were uncultivated fallow saline-alkali land
with vegetation cover mainly by reeds and alkali grasses. After de-weeding and
ploughing the land the plots were made ready for plantation. The divided zone
did contain one ridge (2.0 × 0.6 m) per zone and 1 row per ridge. Each row has
25 holes, and each hole was sown with 1 capsule of peanut. Both the small and
the large peanut groups were designed with random blocks and 4 replicates (the
yield data was calculated after combining 4 replicates). The peanut lines were
planted on May 27 and the crops were harvested on September 29.
Experiment 2
Materials: The materials were 4 high oleic acid varieties (lines) that were
selected from the Experiment 1, after two years of field salt-alkali stress
tests. The peanut varieties (lines) were: Small 16S5, Huayu
669 and large Huayu 967, 16L2. The control variety
was Fuhua 12. Huayu 669 and
Huayu 967 performed best in experiment one, 16S5 and
16L2 performed the worst. Fuhua 12 is the main local
variety.
Test design: The experiment was carried out in pot planting and for which two kinds
of soils were used. These are saline-alkali soil and cinnamon soil (control), the former soil was collected from the uncultivated
saline-alkali land spot in Baicheng (Jilin province)
and the latter soil category was collected from 0–30 cm surface soil (cinnamon soil) in peanut breeding field in Fuxin
(Liaoning Province). The cinnamon soil had a salt content of 0.114% and a pH of
6.5. On the other hand, the total salt content of saline-alkali soil was 0.241%
having a pH value as high as 10.1. Each variety of the two types of soil was
set up with 3 replicates, each containing 2 pots and 6 seedlings per pot
following a random block design.
Methods: The pot culture experiment was conducted in the dry shed of the peanut
breeding test base (Fuxin) of the Institute of Sandy Land Management and
Utilization of Liaoning from 29 May to 9 October 2017. The pot used for the
experiment was 21 cm high with an outer diameter of 35 cm. The saline-alkali
soil was transported from the salt-alkali field of Experiment 1. After mixing,
stirring, and sieving the test soil was made ready for carrying out the
experiment. Each experimental pot was filled with 20 kg of soil. Ten peanut
seeds were sown in each pot and when the seedling grew, 6 seedlings were kept
for running the experiment by removing the others. Watering to the test pots
was done at every 7–10 days according to the soil moisture.
The volume of water in each pot was the same and when the temperature in the
shed is high, the shading net was used. Ten peanut plants with uniform growth
were selected during the harvest of the cinnamon soil, and all plants in the
saline-alkali soil were harvested following the measurement of their yield and
yield-related traits.
The economic yield was the dry weight yield of the pods per plant but
the biological yield was determined by the drying method. The samples were
washed with water and then dried with filter paper. Different organs of the
experimental peanut plants such as roots, stems and petioles, leaves and pods
were separated and dried in an oven at 105℃ for 30 min and then dried to a constant weight at 70℃ (Tian et al. 2019).
Relative
yield: The relative yield was calculated
following the formula: Relative yield = yield under
stress/yield of the variety under non-stress × 100%. The grading standard for
peanut salt-tolerant identification was done via relative yield determination
as mentioned above. The scales were: relative yield> 85% is high tolerance,
60%<relative yield≤85% is medium tolerance, 45%<relative
yield≤60% is low tolerance and relative yield≤45% is susceptible
(Su et al. 2018).
Photosynthetic
index: It was measured on the functional
leaves (the main stem inverted three leaves) at the flowering stage of the
experimental plants. A sunny day without wind was selected for the measurement,
and the data on photosynthesis was recorded with the help of a LI-6400
photosynthesis instrument (LI-COR of American), repeating 5 times for each
plant.
Chlorophyll
determination: The chlorophyll concentration
(relative amount of chlorophyll at two wavelengths) was estimated by measuring
soil plant analysis development (SPAD) values. The SPAD-502 Plus chlorophyll
meter of Nissan was used to determine the SPAD value of peanut functional
leaves on 20 July, 25 August, and 27 September 2017, and the measurement was
repeated for 5 times.
Chlorophyll stability index: Chlorophyll
Stability to salt stress was assessed after Seydi et al. (2007). The Chlorophyll Stability Index
(CSI) was modified as the percentage of chlorophyll of the salt-stressed sample
relative to its content in the control sample as follows:
CSI=
Salt
tolerance index (STI): To determine this parameter, the
procedure of Seydi et al. (2007) was adopted using the following formula:
STI=
Malondialdehyde (MAD) determination: The functional leaves of peanuts were sampled at plump pods mature
stage, quickly placed in liquid nitrogen, and measured by Soluble Bao
Malondialdehyde (MDA) content detection kit (BC0025). At each treatment of each
variety (system) the data measurement was repeated for 3 times.
Relative value calculation:
Relative value =
Data Analysis
DPS 7.05 software was used for data
collation and analysis, and multiple comparisons were used for the analysis of
variance.
Results
Yield of field test for 2016 and 2017 in Baicheng
From Table 1, pod yield ranged from
1000.05 to 5208.00 kg/ha and the kernel yield
ranged from 709.95 to 3249.75 kg/ha. Huayu669 had the highest
yield in small peanut group having pod yield 2950.05 kg/ha, kernel yield
2032.50 kg/ha and surviving plants 19.00. The values were higher than the
varieties (lines). Huayu967 had the highest yield in big peanut group having
pod yield 5208.00 kg/ha, kernel yield 3249.75 kg/ha and surviving plants 21.00,
all of which were higher than ten varieties (lines).
Based on the information presented in Table 2–3 and according to the principle of low, the tolerant
performances by different varieties were Huayu 669 medium, Huayu 967 high and 16S5 low and 16L2 performed as susceptible.
Yield and yield-related indicators of pot culture
Number of surviving plants: There should
be 36 plants of each entry at last, with the prolongation of salt-alkali
stress, some plants gradually died. The plants of Huayu669 were more survivable
than other varieties during the whole growing period (Table 4). Huayu967 ranked
second but in 16L2 and 16S5 and Fuhua12 many plants died before 3 August 2017. After growing for 89 days and on 25 August 2017 there
were no more plants died for Huayu967.
Economic and biological yields of different varieties during mature period under
salt-alkali stress: Table 5
showed the relative value of yield and yield-related indices for all the tested
varieties. It is seen that Huayu 967 and Huayu 669 occupied in the top two for each index. On the
other hand, 16L2, 16S5, Fuhua 12 ranked 3rd, 4th and 5th positions,
respectively. According to the grading standard of peanut salt-tolerant
identification, Huayu 967 and Huayu
669 showed medium tolerant, 16L2 was low tolerant, and 16S5 and Fuhua 12 were susceptible (Table 5).
Comparison of
yield-related traits of various varieties under
saline-alkali stress: From Table 6,
the relative values of yield-related traits have decreased in different degrees
under salt-alkali stress. Each index of Huayu 967 and
Huayu 669 ranged from 62–90% and 60–98%, respectively and could be said
that the indices were mostly higher than 60%. The relative values of total
branches, fruiting branches and total pods for Huayu
669 reached more than 90%, with the smallest decrease. However, the relative
values of plump pods for Huayu 669 and Huayu 967 were 63 and 64%, respectively which is higher
than other varieties.
Comparison of
physiological indexes of various varieties under
salt-alkali stress of pot culture
Table 1: Surviving plants and yield of small peanut and large
peanut (second column) varieties (lines) in 2016
Varieties
(lines) |
Plants
per block |
Pod
yield (kg/ha) |
Kernel
yield (kg/ha) |
Varieties
(lines) |
Plants
per block |
Pod
yield (kg/ha) |
Kernel
yield (kg/ha) |
15S1 |
13.30abc |
2179.05 |
1420.65 |
15L1 |
14.00AB |
1954.05 |
1238.85 |
15S3 |
8.70c |
1303.95 |
925.80 |
Huayu967 |
21.00A |
5208.00 |
3249.75 |
15S8 |
15.00abc |
1546.05 |
1066.80 |
16L2 |
5.30B |
1000.05 |
709.95 |
15S9 |
11.00c |
1429.05 |
951.75 |
15L4 |
8.70B |
1195.95 |
819.30 |
16S5 |
9.70c |
1333.05 |
867.75 |
15L8 |
11.70B |
1828.95 |
1130.25 |
15S13 |
15.00abc |
1378.95 |
1031.55 |
15L9 |
10.30B |
1458.00 |
1073.10 |
15S24 |
20.30a |
2688.00 |
1771.35 |
15L10 |
1.00B |
1945.95 |
1224.00 |
Huayu669 |
19.00ab |
2950.05 |
2032.50 |
15L11 |
14.00AB |
1683.00 |
1031.70 |
15S28 |
13.30abc |
1624.95 |
1078.95 |
15L15 |
11.00B |
1699.95 |
1099.95 |
Huayu20 |
11.70bc |
1467.00 |
971.10 |
15L16 |
10.30B |
1741.95 |
1263.00 |
Baiyuanhua
1 |
15.70abc |
1513.05 |
1099.95 |
15L17 |
10.70B |
1738.05 |
1070.55 |
Baiyuanhua
2 |
16.70abc |
1458.00 |
1032.30 |
15L18 |
9.30B |
1483.05 |
909.15 |
|
|
|
|
Huayu33 |
8.00B |
1413.00 |
897.30 |
Note:
Lower case letter represents significance at 0.05,
uppercase letter represents probability level significance at 0.01
Table
2: Yield performance of pods and kernels
for 2016 and 2017 (kg ha-1)
varieties
(lines) |
2017 Saline-alkali soil |
2017 Cinnamon soil |
2016 Saline-alkali soil |
2016 Cinnamon soil |
Relative yield of pod (%) |
Relative yield of kernel (%) |
||||||
|
Pod
yield |
Kernel
yield |
Pod
yield |
Kernel
yield |
Pod
yield |
Kernel
yield |
Pod
yield |
Kernel
yield |
2017 |
2016 |
2017 |
2016 |
Huyu669 |
2783.40AB |
2076.45AB |
4073.25bc |
3091.40 |
2950.05 |
2032.50 |
4315.95 |
3185.25 |
68.33 |
68.35 |
67.17 |
63.81 |
Huyu967 |
3083.40AB |
2301.90AB |
3600.00bc |
2678.40 |
5208.00 |
3249.75 |
4288.05 |
3126.00 |
85.65 |
121.45 |
85.94 |
103.96 |
16S5 |
1528.76ABC |
1091.78ABC |
2980.05c |
2246.90 |
1333.05 |
867.75 |
2662.95 |
1927.95 |
51.3 |
50.06 |
48.61 |
45.01 |
16L2 |
1833.30ABC |
1370.10ABC |
3939.90bc |
2892.00 |
1000.05 |
709.95 |
2737.95 |
2050.80 |
46.53 |
36.53 |
47.38 |
34.62 |
Note:
Lower case letter represents significance at 0.05,
uppercase letter represents probability level significance at 0.01
Table
3: Judging tolerance based on relative
yield
Varieties (lines) |
According to the relative yield of pod |
According to the relative yield of kernel |
According to the principle of low |
||
2017 |
2016 |
2017 |
2016 |
||
Huayu 669 |
medium tolerant |
medium tolerant |
medium tolerant |
medium tolerant |
medium tolerant |
Huayu 967 |
high tolerant |
high tolerant |
high tolerant |
high tolerant |
high tolerant |
16S5 |
low tolerant |
low tolerant |
low tolerant |
low tolerant |
low tolerant |
16L2 |
low tolerant |
susceptible |
low tolerant |
susceptible |
susceptible |
Table
4: Number of surviving plants
|
July
20 |
August
3 |
August
19 |
August
25 |
September
11 |
October
9 |
Huayu
967 |
22
|
18
|
17
|
14
|
14
|
14 |
16L2 |
19
|
14
|
12
|
9
|
8
|
6 |
16S5 |
21
|
14
|
11
|
8
|
5
|
5 |
Huayu
669 |
28
|
27
|
26
|
26
|
26
|
19 |
Fuhua12 |
11
|
8
|
8
|
8
|
7
|
7 |
Table
5: Relative value of yield and
yield-related indicators of each entry under saline-alkali stress (%)
|
Root
dry weight |
Stem
dry weight |
Leaf
dry weight |
Salt
tolerance index |
Economic
yield |
Fuhua12 |
47 |
37 |
43 |
41 |
32 |
Huayu
967 |
93 |
71 |
82 |
78 |
80 |
16L2 |
66 |
53 |
55 |
55 |
46 |
16S5 |
62 |
51 |
57 |
50 |
35 |
Huayu
669 |
96 |
62 |
67 |
66 |
70 |
Comparison of
net photosynthetic rate during flowering stage: From Fig. 1, under saline-alkali stress, the net photosynthetic rate of all
varieties (lines) was less than 10 μmol·CO2·m-2·s-1, and the order was: Huayu669﹥Huayu967﹥16L2﹥Fuhua12﹥16S5. The two medium tolerant varieties of Huayu
669 and Huayu 967 showed up to 5.29 and 4.91 μmol·CO2·m-2·s-1,
respectively and the values were larger than the normal cinnamon i.e., 3.98 and
4.54 μmol·CO2·m-2·s-1,
respectively. The net photosynthetic rate of other three varieties (lines)
under saline-alkali soil was lower than that of
cinnamon soil. The smallest was recorded for Fuhua12
(3.02 μmol·CO2·m-2·s-1) which is
1.41 μmol·CO2·m-2·s-1 lower than the
value of normal cinnamon soil and the percentage of decrease was 31.9%.
SPAD values of various varieties (lines) at different growth stages: SPAD value correlated positively with
the chlorophyll content in the leaf. The SPAD value of the leaf reflects the
level of the chlorophyll content. Table 7 showed CSI of all varieties in the
three periods (the flowering, the poding and the
plump pod mature stage) were consistent. As shown in Fig. 2, under salt-alkali stress, the highest chlorophyll
content in each period was recorded for Huayu 967,
which were 37.75, 44.47 and 41.7 for 3 stages,
respectively. Followed by Huayu669 and 16L2, the smallest were Fuhua 12 and 16S5, the chlorophyll content of 16S5 was only
38 at the flowering stage. The CSI of Huayu 967, Huayu 669, 16L2 in each period were greater than 75%, and
the susceptible varieties (lines) were between 55 and 74%.
Table
6: Relative value of yield-related traits
(%)
|
Main
stem height |
Lateral
branch length |
Total
branches |
Fruiting
branches |
Total
pods |
Plump
pods |
Fuhua12
|
53 |
49 |
69 |
63 |
56 |
37 |
Huayu
967 |
90 |
62 |
73 |
81 |
75 |
64 |
16L2
|
81 |
45 |
71 |
85 |
61 |
59 |
16S5
|
56 |
47 |
72 |
74 |
71 |
48 |
Huayu
669 |
62 |
60 |
98 |
98 |
94 |
63 |
Table
7:
CSI of each entry during different growth periods (%)
|
Flowering stage |
Poding stage |
Plump pods mature stage |
Fuhua12 |
72 |
74 |
73 |
Huayu
967 |
85 |
89 |
96 |
16L2 |
75 |
77 |
82 |
16S5 |
59 |
65 |
66 |
Huayu
669 |
75 |
83 |
86 |
Table
8: MAD Content of the third leaf from top
during maturing period (nmol g-1)
|
Fuhua12 |
Huayu967 |
16L2 |
16S5 |
Huayu669 |
Saline-alkali
soil |
55.69 |
41.81 |
640 |
36.65 |
49.71 |
cinnamon
soil |
52.25 |
34.40 |
23.60 |
25.67 |
48.93 |
relative
value |
1.07 |
1.22 |
2.56 |
1.43 |
1.02 |
Table
9: Correlation analysis of yield,
agronomic trait and physiological indicators under saline-alk
ali stress
|
Fresh pod weight per plant |
Dry pod weight per plant |
Dry root weight per plant |
Dry stem weight per plant |
Dry leaf dry weight |
Biological yield |
Main stem height |
Lateral branch length |
Total No. of branches |
Fruiting branches |
Total pods |
Plump pods |
Photosynthetic rate |
SPAD |
MDA |
Dry
pod weight per plant |
0.72 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Dry
root weight per plant |
-0.62 |
-0.11 |
|
|
|
|
|
|
|
|
|
|
|
|
|
Dry
stem weight per plant |
-0.14 |
-0.38 |
0.33 |
|
|
|
|
|
|
|
|
|
|
|
|
Dry
leaf dry weight |
0.07 |
-0.48 |
-0.07 |
0.88* |
|
|
|
|
|
|
|
|
|
|
|
Biological
yield |
-0.07 |
-0.45 |
0.18 |
0.97** |
0.96** |
|
|
|
|
|
|
|
|
|
|
Main
stem height |
-0.06 |
-0.03 |
0.21 |
0.75 |
0.49 |
0.63 |
|
|
|
|
|
|
|
|
|
Lateral
branch length |
0.03 |
-0.09 |
-0.003 |
0.73 |
0.57 |
0.65 |
0.98** |
|
|
|
|
|
|
|
|
Total
No. of branches |
0.66 |
0.15 |
-0.99** |
-0.36 |
0.06 |
-0.19 |
-0.29 |
-0.09 |
|
|
|
|
|
|
|
Fruiting
branches |
0.91* |
0.50 |
-0.73 |
-0.30 |
0.05 |
-0.15 |
-0.40 |
-0.28 |
0.80 |
|
|
|
|
|
|
Total
pods |
0.73 |
0.61 |
-0.71 |
-0.76 |
-0.48 |
-0.66 |
-0.60 |
-0.50 |
0.76 |
0.83 |
|
|
|
|
|
Plump
pods |
0.88 |
0.50 |
-0.79 |
-0.44 |
-0.07 |
-0.29 |
-0.47 |
-0.34 |
0.88 |
0.99** |
0.90* |
|
|
|
|
Photosynthetic
rate |
0.73 |
0.15 |
-0.96** |
-0.23 |
0.20 |
-0.05 |
-0.25 |
-0.05 |
0.98** |
0.86 |
0.72 |
0.89* |
|
|
|
SPAD |
0.81 |
0.20 |
-0.83 |
-0.09 |
0.33 |
0.93 |
-0.27 |
-0.10 |
0.88* |
0.93* |
0.67 |
0.91* |
0.95* |
|
|
MDA |
0.20 |
0.06 |
-0.09 |
0.67 |
0.53 |
0.60 |
0.95* |
0.98** |
0.01 |
-0.12 |
-0.36 |
-0.19 |
0.05 |
0.02 |
|
Oleic
acid content |
0.71 |
0.73 |
-0.59 |
-0.8 |
-0.6 |
-0.74 |
-0.57 |
-0.5 |
0.62 |
0.76 |
0.98** |
0.83 |
0.57 |
0.53 |
-0.37 |
Note: * significance at 0.05, **
probability level significance at 0.01
Comparison of
malondialdehyde content of various varieties (lines) under salt-alkali stress: It can be seen from
Table 8 that under saline-alkali stress, the malondialdehyde concentration of
all the varieties (lines) was higher than the value of cinnamon soil. The line
16S5 had the lowest malondialdehyde concentration under saline-alkali stress,
followed by Huayu 967, Huayu
669 and 16L2 had the highest malondialdehyde content. The relative values of Huayu 669, Fuhua 12, Huayu 967 were low, and the concentration of malondialdehyde increased by
78, 44 and 7.4 nmol g-1,
respectively under saline-alkali stress. However, 16L2 and 16S5 increased much
more, reaching 36.77 and 198 nmol g-1, respectively.
Correlation
analysis of various indicators
Fig.
1: Photosynthetic rate of each entry
during flowering period
Fig. 2: SPAD value of each entry during different growth and
development periods under saline-alkali stress
Table 9 showing the biological yield
under salt-alkali stress is extremely positively correlated with stem and leaf
dry weight per plant. The height of the main stem was
significantly correlated with the length of lateral branches. Oleic acid
content was significantly positively correlated with the total pods. Poding branches correlated significantly and positively
with the fresh weight of pods per plant (Lauter and Meiri
1990), and highly significantly positive correlation was shown with the number
of plump pods. The net photosynthetic rate correlated
positively and the level of significance was high with the total number of branches and the number of plump pods. There was a significant positive correlation between the concentrations of malondialdehyde
some biological traits. The concentration of malondialdehyde was
significantly positively correlated with the height of the main stem and the
length of the lateral branches, respectively.
Discussion
The seed yield of 210 high yielding
peanut germplasm accessions, under saline condition, ranged from 0 to 2030 kg ha-1 (Singh et al. 2016). In the present investigation,
the kernel yields ranged from 867.75 to 3249.75 kg ha-1. The yield per plant
of Huayu 25 under non-saline soil was 12.69 ± 1.32a
(g) and under saline-alkaline soil the value was 4.48 ± 0.38c (Tian et al. 2019), wherein
the STI value was 35%. This had shown no big difference with susceptible
varieties but lower than the salt-tolerant varieties of experiment 2 (Huayu 967 and Huayu
669). So, it is understood that the
salt-tolerant varieties (lines) in this research performed well.
According to the grading standard of peanut salt-tolerant
identification, the high-oleic acid peanut varieties (lines) selected in
Experiment 2 under pot salt-alkali stress showed the salt-tolerant ability as
follows: Huayu 967 and Huayu
669 were medium tolerant, 16L2 low tolerant, 16S5 and Fuhua
12 susceptible. Compared with the same varieties (lines) in the field, the
tolerance of Huayu 669 remained unchanged but Huayu 967 decreased from high tolerance to medium tolerance
and 16S5 decreased from low tolerance to susceptible. It could be speculated
that it may be because the saline-alkali soil used for pot experiment was the
salt spot in the saline-alkali plot, and the intensity of saline-alkali stress
was greater than the field experiment.
Previous studies showed that the saline-alkali stress significantly
inhibited the growth and development of peanut plants, where the main stem
height, lateral branch length was
significantly reduced (Zhang et al. 2016). Main stem height, lateral branch length, and yield are used as
indicators to evaluate peanut salt-tolerance, and the total number of branches
needs to be studied further (Wang et al. 2013). In experiment 2, Huayu 967 and Huayu 669, which had more tolerance, also showed reduction
in main stem and lateral branch, but less than the intolerant varieties. The
relative value of fruiting branch number of the susceptible varieties was as
low as 63%, while they were as high as 81 and 98% of Huayu
967 and Huayu 669. And the fruiting branch number
under salt-alkali stress was significantly positively correlated with the yield
per plant, a very significant positive correlation with the plump pods.
Salt-alkali stress can destroy the order and structure of thylakoids in
the chloroplast (Zheng and Zhang 1998). The
chlorophyll content in the leaves of Huayu 967, was higher than other varieties (lines) at various
growth stages. The CSI of medium-tolerant and low-tolerant varieties (lines) in
each period was greater than 75%. So, CSI can be used as an indicator for
the high oleic acid peanut regarding salt tolerance. The net photosynthetic
rate of Huayu 669 and Huayu
967 did not decrease under salt-alkali stress but had a certain promotion
effect, which was consistent with previous studies (Ren et al. 2017). But a 31.9% reduction of
photosynthetic rate took place in susceptible variety Fuhua
12. Correlation analysis showed that the net photosynthetic rate under
salt-alkali stress correlated strongly, significantly and positively with the total number of branches and plump pods. The chlorophyll
content was significantly positively correlated with the total number of
branches, the number of fruiting branches, plump pods and the net
photosynthetic rate.
When plants are exposed to salt-alkali stress for a long time, the
plant membrane system gets destroyed, resulting an
increase in the malondialdehyde concentration. This increase
affects and destroys the normal metabolism of cells,
inhibiting plant growth and even withered to death (Qiao et al. 2013). The concentration of
malondialdehyde can reflect the degree of plant senescence and suffer from the
adversity. Under salt stress, the leaves and roots of salt-alkali tolerant
varieties have lower malondialdehyde concentration (Qiao
et al. 2013). In Experiment 2, Huayu 967 had a lower malondialdehyde concentration and
less increase under salt-alkali stress, which confirmed its higher salt-alkali
tolerance.
Results of this study revealed that compared with non-saline-alkali
stress, varieties with strong saline-alkali tolerance (Huayu
669 and Huayu 967) can reduce the main stem height
and lateral branch length from the morphological point of view. These changes
in the crop maintain more fruiting branches. From the physiological and biochemical point of view, less chlorophyll was destroyed, and the
net photosynthetic rate did not decrease but increased. The process had helped
accumulating enough organic matter and ensured a
sufficient source for the source-sink fluency and economic crop output below
ground.
Conclusion
Three research findings of the two best
performing high-oleic acid varieties (Huayu 967 and Huayu 669) were identified: (1) from the perspective of field yield, more
than 2950.05 kg ha-1 can be
selected as salt-tolerant varieties; (2) from the morphological point of view, the relative value of main stem
height, lateral branch length, total branches, fruiting branches, total pods
and plump pods should be higher than 60% and (3) from the physiological and biochemical point of view, the
STI was not less than 66; maintaining more chlorophyll content, the CSI should
be greater than 75%; strong net photosynthetic capacity to ensure the
accumulation of organic compounds and the malondialdehyde concentration should
be less. So, for the selection of high oleic acid peanut saline-alkali
tolerance, we could refer to the above values. Compared with the field test, pot culture has the following advantages:
Firstly, the use of pot can better control the consistency of the growth
environment of each test material under saline-alkali soil and cinnamon soil. The number of surviving plants can
intuitively reflect the degree of tolerance, to more
accurately determine the salt-tolerance of various varieties. Secondly, the pot experiment can reduce
the test period and test points. The saline-alkali soil in Experiment 2 has
been evenly mixed by stirring to ensure uniform salt-alkali stress and can
replace the multi-point field test for many years. Next, a transcriptome
analysis of four varieties will be carried out to explore the salt-tolerant
genes.
Acknowledgements
The first author acknowledges the
financial grant from Dean Fund of Liaoning Academy of Agricultural Sciences
(2019-QN-09); Foundation items: China Agricultural Research System (CARS-13);
Liaoning Provincial Department of Science and Technology Key R&D Program (2017201004); Shandong Taishan
Industry Leading Talent Project (LJNY201808); Shandong Province Key R & D
Program Special (2018GNC110027).
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