IJERD – International Journal of Environmental and Rural Development (2023) 14-2

Research article
erd

Growth and Mineral Uptake of Moringa oleifera Lam. in
Low-Permeability Soils at Different Salinity Levels
TAKASHI KUME*
Graduate School of Agriculture, Ehime University, Matsuyama, Japan
Email: kume@ehime-u.ac.jp

CHULEEMAS BOONTHAI IWAI
Department of Agriculture, Khon Kaen University, Khon Kaen, Thailand

TETSUYA SHIMAMURA
Graduate School of Agriculture, Ehime University, Matsuyama, JapanSHINSUKE HARUTA
Graduate School of Agriculture, Ehime University, Matsuyama, Japan
Received 24 July 2023

Accepted 1 October 2023(*Corresponding Author)

Abstract This study investigated the uptake of minerals by moringa (Moringa oleifera Lam.)and examined the salt tolerance of moringa under different salinity treatments (0, 4, 8, and16 dS/m). The effect of root growth on soil permeability at different salinity levels was also
examined. Moringa showed significant negative effects of salinity on growth parameters at 8
dS/m. Significant growth inhibition was observed for moringa at 16 dS/m. The higher C/F
ratios, calculated as (assimilated organ mass)/ (non-assimilated organ mass), in moringa in
the 16 dS/m treatment may be due to the inhibition of nitrogen uptake by the roots, causing
photosynthesis in the leaves to produce assimilates to sustain the body of the tree. In eachmoringa organ, the Na concentration increased as salinity increased. Significant differencesin concentration were observed for Ca and K in leaves and Fe in stems at different salinity
levels. The correlation analysis showed that only Mg and P in the branches and Fe and Mg
in the stems were significantly negatively correlated with Na concentration, suggesting that
increases in Na concentration cause limited inhibition of mineral uptake by moringa. A
significant positive correlation was found between Na and P in the roots. It was suggested
that moringa roots may have promoted growth by increasing P uptake in response to increased
Na. There was a positive correlation between the length of moringa main roots and saturated
hydraulic conductivity. The saturated hydraulic conductivity of the soil without moringa
cultivation was 1.1 × 10-6 cm/s, and moringa root growth increased the saturated hydraulicconductivity by two orders of magnitude (10-4 cm/s) at 0 and 4 dS/m and by one order ofmagnitude (10-5 cm/s) at 8 and 16 dS/m.
Keywords moringa, soil salinity, root growth, mineral uptake, soil permeability

INTRODUCTION
Soil salinity is a major threat to land globally, and 833 million ha of soil is affected by salt (FAO,
2021). Coastal paddy fields in the Tohoku region were salinized by the tsunami caused by the Great
East Japan Earthquake. In addition to natural disasters, coastal areas are also said to be widely
affected by salt damage due to sea level rise caused by climate change. In addition, it has been
calculated that the saline front will move up to 20 km upstream of the Mekong Delta in the 2030s
due to climate change (Khang et al., 2008).
Saline soils need to be amended by improving soil permeability to discharge salts from soil
layers. It is known that tree roots, such as Alnus Mill. roots, enhance soil permeability (Vergani andGraf, 2015). However, other studies suggest that the effect of trees and roots on soil permeabilityⒸ ISERD

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depends on soil type, soil and land use history, and vegetation cover type (Bonell et al., 2010).
Therefore, the effect of each plant on the improvement of soil permeability should be investigated.
Moringa (Moringa oleifera Lam.) is highly salt tolerant, and all parts of moringa can be used
for multiple purposes (Horn et al., 2022). Moringa seedlings survived up to 8 dS/m with a slight
reduction in biomass chlorophyll a, crude protein, and mineral concentrations with increasing salinity(Nouman et al., 2012). Two-month-old moringa transplants might be NaCl tolerant up to 8 dS/m(Elhag and Abdalla, 2014). Farooq et al. (2022) investigated the impact of various salinity levels on
the root attributes of moringa and revealed higher salinity levels (7.5 dS/m and 11.5 dS/m)significantly minimized the root surface area compared to 1.5 dS/m and 3.5 dS/m. These results showthat salt tolerance has been studied, but a comprehensive assessment of mineral concentrations ofmoringa under saline conditions has not been completed. Research on root growth parameters has
been conducted; however, the effect of moringa root growth on the improvement of soils has notbeen examined.
OBJECTIVE
This study investigates the growth and mineral uptake of moringa and the salt tolerance of moringa
at different salinity levels. Root growth and its effect on soil permeability in soils with different
salinity levels were also examined.
METHODOLOGY
A cultivation experiment was conducted in a greenhouse for two months. Root boxes (width: 60 cm,
height: 40 cm, depth: 10 cm) made of acrylic plates were used for the cultivation experiments. The
root boxes were filled with a 20-cm-deep plow layer and a 10-cm-deep plow layer, as shown in Table
1. The soil texture of paddy soil was CL (Clay Loam) and the soil hydraulic conductivity of twolayers of soil as shown in Table 1 was 1.1 × 10-6 cm/s.
Table 1 Physical properties of soils in the cultivation experiment

Plow layer
Plow sole layer

Dry bulk
density(Mg/m3)
1.2
1.7

Particle density(Mg/m3)A mixing ratio of experimental soil

2.53
2.59

Paddy soil (CL)Gravel

100%
75%

0%
25%

The soils were subjected to four salinity treatments: one without NaCl (0 dS/m) and three with
NaCl added to the soil to adjust the soil salinity to ECe = 4, 8, and 16 dS/m. Indian moringa seeds
were used, and a total of 24 sample trees were grown, with six moringa tree replications in eachsalinity treatment. 10 mm irrigation water was supplied when the soil surface was dried, and no
fertilizer was used.
Growth parameters (tree height and dry matter (DM) weight of leaves, branches, stems, androots) were measured. Dry matter weights were determined after samples were oven-dried at 80 °C
for 48 hours. The C/F ratio was calculated as the (assimilated organ mass) ÷ (non-assimilated organmass), and the aerial C/F ratio was calculated as (assimilated organ mass) ÷ (aboveground nonassimilated organ mass); these ratios were used as indices to describe the structure of material
production systems (Nomoto and Yokoi, 1981).
The mineral concentrations in each organ of the harvested moringa were determined by ICP
optical emission spectrometry (Varian, Vista-MPX). A soil permeability experiment was performed
after harvesting the aboveground portions of the moringa plants, with the roots remaining in the rootbox. The root box was placed in a bucket filled with water and it was capillary saturated, then water
was ponded to a depth of 10 cm, and changes in water level over time were recorded. The measured
data were substituted into Darcy's equation to obtain the saturated soil hydraulic conductivity.
ANOVA was performed at the 5% significance level for the height, dry matter, and mineralⒸ ISERD

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concentration of each part of moringa grown under each salinity condition. When the ANOVA
showed significant differences, multiple comparisons (Tukey test) were used to test between samples.
RESULTS AND DISCUSSION
Effect of Salinity Treatments on Growth Parameters
Significant differences were found in tree height, root dry matter weight, total dry matter weight, andthe C/F ratio under different salinity levels (Table 2). There were no significant differences between
0 dS/m and 4 dS/m in any of the parameters. All the parameters except the CF/ratio at 8 dS/m hadsignificantly smaller values than those at 0 and 4 dS/m. All the parameters except the C/F ratio at 16
dS/m had significantly smaller values than those at the other salinity levels. The C/F ratio at 16 dS/m
(0.346) was significantly larger than that at 4 dS/m (0.164). Noumen et al. (2012) showed thatmoringa was able to survive under 8 dS/m conditions with little reduction in nutritional value. Elhag
and Abdalla (2014) showed that moringa is salt tolerant up to 8 dS/m, with significantly lower values
for tree height and leaf dry weight at 16 dS/m. Farooq et al. (2022) also concluded from theirexperiment results that germination rate, root surface area, and aboveground growth are largely
unaffected at salinity levels up to 3.5 dS/m. Our results were similar to those of previous studies.
Table 2 Effects of salinity treatments on growth parameters
Different letters indicate significant difference at p < 0.05 (DM: dry matter)
Salinity Tree height
treatment(cm)

Leaf DM(g)

Branch DM(g)Stem DM(g)

Root DM(g)

Total DM(g)

0 dS/m

53.5±7.9ab 2.020±0.35 0.977±0.20 2.470±0.62

9.45±2.20ab

14.9±2.6ab

0.164±0.02a 0.711±0.10

4 dS/m

63.5±7.1a 2.390±0.49 1.110±0.28 2.960±0.73

10.20±1.40a

16.7±2.1b

0.164±0.01a 0.650±0.10

8 dS/m

52.5±8.5ab 1.890±0.70 1.030±0.40 2.760±1.20

4.15±0.88b

9.78±2.5a

0.199±0.04ab 0.555±0.04

16 dS/m27.5±3.9b 0.812±0.20 0.307±0.12 0.650±0.24

1.42±0.40c

3.18±0.1c

0.346±0.09b 0.925±0.20

C/F ratio

AerialC/F ratio

Mean ± standard deviation (Data adopted from Kume et al., 2023)

Fig. 1 Composition of dry matter weight by the organ of moringa grownat different salinity levels
The compositions of moringa organs at 0 dS/m and 4 dS/m were similar, while the percentagesof moringa organ dry matter weights at 8 dS/m were approximately the same as those at 0 dS/m, with
a 7% increase in stem dry matter weight and a 10% decrease in root dry matter weight (Fig. 1).
Moringa in the 16 dS/m treatment had 10% greater leaf dry matter weight, 2% greater branch dry
matter weight, 5% greater stem dry matter weight, and 17% lower root dry matter weight thanmoringa in the 0 dS/m treatment, with a significant difference in the C/F ratio (0.164 at 0 and 4 dS/m
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IJERD – International Journal of Environmental and Rural Development (2023) 14-2

compared to 0.346 at 16 dS/m). The aerial C/F ratios were not significantly different among thesalinity levels. Carbon is acquired by aboveground plant parts, and nitrogen is acquired by
belowground plant parts (Hirose, 1988). Individuals showing abnormally high C/F ratios are thought
to die due to an imbalance between assimilated and non-assimilated organs (Higo, 1987). Moringain the 16 dS/m soil, where growth inhibition was observed, showed a significantly higher C/F ratio
than moringa in the other treatments, and that photosynthesis in the leaves produces assimilates tosustain the body of the tree.
Mineral Uptake by Moringa
The Na concentrations in the organs were significantly different among the different salinity levels(Table 3). In each moringa organ, the Na concentration increased as salinity increased. Tukey’s test
showed no significant differences in Na concentration between the 0 dS/m treatment and 4 dS/mtreatment for all organs. In leaves, branches, and stems, the Na concentrations at 8 dS/m and 16 dS/m
were significantly different from those at 0 dS/m. For roots, the Na concentration at 8 dS/m wassignificantly different from that at 0 dS/m and 4 dS/m, and the Na concentration at 16 dS/m wassignificantly different from those at the other salinity levels.
Table 3 Mean values of mineral concentrations in each organ of moringa grown
at various salinity levels
Different letters indicate significant difference at p < 0.05 (DM: dry matter)

LeavesSalinityNa
treatment mg/g DM
0 dS/m 1.110±0.21a

K
Ca
Fe
P
Mg
Cu
Zn
mg/g DM mg/g DM
mg/g DM mg/g DM mg/g DM μg/g DM μg/g DM
24.9±1.4a 8.69±0.89ab 0.166±0.05 7.72±0.86 3.18±0.25 11.3±2.7 50.1±17.0

4 dS/m 0.958±0.13a

21.7±1.9a 10.10±1.20a 0.122±0.09 6.24±0.70 2.58±0.35 10.3±0.9 44.8±8.8

8 dS/m 2.410±0.30b 18.5±0.7ab 7.90±0.36ab 0.183±0.05 5.79±0.91 2.66±0.40 13.8±4.2 82.7±25.0
16 dS/m 1.890±0.35ab 14.2±2.1b 6.10±0.50b 0.120±0.02 5.42±0.87 2.61±0.36 8.03±0.7 34.9±4.3
Stems

BranchesRoots

0 dS/m 0.574±0.08a

24.2±2.3

11.10±1.4 0.538±0.08a 7.21±0.86 3.21±0.50 9.82±0.9 55.8±6.6

4 dS/m 1.570±0.16a

18.9±1.3

9.95±1.10 0.479±0.01a 6.58±0.60 3.12±0.31 10.3±1.4 57.8±13.0

8 dS/m 3.930±0.68b

19.5±2.8

10.90±1.10 0.115±0.05b 5.02±0.10 2.14±0.21 11.3±1.7 69.3±20.0

16 dS/m 4.670±0.62b

25.5±7.0

10.10±0.67 0.142±0.02b 6.38±1.20 1.94±0.21 14.1±2.2 73.3±13.0

0 dS/m 0.499±0.10a

30.2±1.8

10.50±1.20 0.171±0.04 9.13±0.35 1.73±0.28 13.2±1.7 80.0±18.0

4 dS/m 1.240±0.15a

25.9±1.3

11.80±1.20 0.109±0.01 8.66±0.43 1.68±0.21 9.11±2.5 56.4±21.0

8 dS/m 3.870±0.80b

31.6±3.78.33±0.52

16 dS/m 4.680±0.62b

24.5±3.0

10.60±2.60 0.172±0.08 8.04±1.30 1.23±0.12 8.83±2.5 48.6±8.9

0 dS/m 0.768±0.11a

13.7±1.1

6.23±0.95

0.716±0.03 4.38±0.32 3.25±0.24 14.7±1.2 91.3±17.0

4 dS/m 1.550±0.12a

12.4±0.75.30±0.59

0.560±0.01 5.00±0.27 2.31±0.15 12.6±1.6 63.2±14.0

8 dS/m 5.650±0.75b

13.8±1.3

5.36±0.74

0.714±0.07 5.02±0.47 2.38±0.13 12.4±0.1 75.9±11.0

16 dS/m 9.070±0.84c

13.3±2.0

5.58±0.48

1.040±0.20 6.68±1.00 2.89±0.33 15.5±1.8 78.9±15.00.114±0.04 6.66±0.80 1.07±0.10 6.02±0.7 41.0±4.6

Apart from Na, significant differences in concentration were observed for Ca and K in leaves
and Fe in stems. For these values, the concentration decreased with increasing salinity. Cu, Mg, P,
and Zn did not show significant differences among organs or salinity levels. In leaves, the Na
concentration increased and the K concentration decreased with increasing soil salinity. Mg and P in
the branches and Fe and Mg in the stems showed a decreasing trend in response to increasing Na(Table 4). The P concentration increased with increasing Na concentration in the roots. There was nocorrelation between Na and other minerals. The P concentration was significantly correlated withvarious ion concentrations in all organs. The P concentration was significantly positively correlatedwith Ca, Cu, Fe, and K in leaves; with Ca, Cu, Fe, Na, and Zn in branches; with Fe, K, and Mg inⒸ ISERD

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IJERD – International Journal of Environmental and Rural Development (2023) 14-2

stems; and with K, Mg, and Na in roots. In branches, the Ca concentration was significantly positively
correlated with Cu, Fe, Mg, P, and Zn, and the Cu concentration was significantly positively
correlated with Fe, Mg, P, and Zn.
Table 4 Correlation matrix of mineral concentrations in each organ

LeavesBranches

Stems

Ca
Cu
Fe
K
Mg
Na
P
Zn
Ca
Cu
Fe
K
Mg
Na
P
Zn
Ca
Cu
Fe
K
Mg
Na
P
Zn

Ca
-

Cu
0.28
*

Fe
0.19
0.85
-

K
0.57
0.10
0.10
-

*
*
Fe
0.78
0.75
-

*

Ca
*
*

*
*
Cu
0.71
*

*

*

*

*
*
Ca
-

*
*
Cu
-0.31
-

*
*
Fe
0.24
-0.22
-

*
*

*
*

*
*
*
*
Cu
0.13
-

Ca
Fe
Ca
-0.07
Cu
0.37
Fe
Roots
K
*
*
Mg
*
Na
P
Zn
*
* Indicates a significant correlation coefficient at p < 0.05.

Mg
0.15
0.15
0.19
0.33
-

Na
-0.20
0.05
0.05
-0.06
0.06
-

P
0.46
0.44
0.67
0.67
0.23
-0.02
-

Na
-0.11
-0.35
-0.05
-0.21
-0.47
*

*

Mg
0.66
0.80
0.59
-0.10
*
*
*
Mg
0.67
-0.22
0.58
-0.01
*
*

P
0.54
0.55
0.42
0.00
0.48
-0.45
*
P
0.19
0.27
0.42
0.68
0.56
-0.25
-

K
0.42
0.07
-0.12
*

Mg
0.50
0.28
0.12
0.58
-

*

*

Na
-0.11
0.17
0.32
0.14
0.08
*

K
-0.36
-0.08
-0.18
-

K
-0.17
0.56
0.05
-

Na
0.11
0.31
-0.57
-0.05
-0.42
-

P
0.20
0.09
-0.02
0.65
0.45
0.63
-

Zn
0.16
0.96
0.83
0.05
0.10
0.22
0.35
Zn
0.57
0.89
0.52
0.06
0.78
-0.30
0.43
Zn
-0.32
0.91
-0.21
0.31
-0.15
0.19
0.09
Zn
0.06
0.74
0.40
-0.13
0.16
-0.01
-0.25
-

In the study by Elhag and Abdalla (2014), leaf Ca and K in moringa also decreased withincreasing Na concentration. Increases in Na have been found to decrease K in triticale, barley, andrice (Flowers and Hjibagheri, 2001; Naidoo, 2007). There were no significant differences in theconcentrations of P, Cu, Mg, and Zn. Matsumaru (1990) found no clear relationship between Fe, Mn,
and Zn concentrations and increasing salinity in a cucumber salt tolerance experiment. The resultsof the correlation analysis in this study also showed that only Mg and P in the branches and Fe andMg in the stems were significantly negatively correlated with Na concentration, suggesting that
increases in Na concentration cause limited inhabitation of mineral uptake by moringa. A significant
positive correlation was found between Na and P in the roots. The same results were obtained by
Elhag and Abdalla (2014) and Nouman (2012). It was suggested that moringa roots may havefunctioned to promote growth by increasing P uptake in response to increased Na.
Root Growth and Soil Permeability
There was a positive correlation between the length of moringa main roots and saturated hydraulicconductivity in the salinity-treated soils (Fig. 2). The relationship between the circumference of themain root and saturated hydraulic conductivity was highly positive (Fig. 3). The saturated hydraulicⒸ ISERD

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IJERD – International Journal of Environmental and Rural Development (2023) 14-2

conductivity of the soil without moringa was 1.1 × 10-6 cm/s, and moringa root growth increased thesaturated hydraulic conductivity by two orders of magnitude (10-4 cm/s) at 0 and 4 dS/m and by one
order of magnitude (10-5 cm/s) at 8 and 16 dS/m. Sophora japonica has tap roots, which growrelatively deep and vertically; these roots loosen the soil more deeply and greatly increase soil
infiltration rates (Zhang et al., 2019). Alunus incana roots were shown to improve soil permeability(Vergani and Graf, 2015). The length and circumference of the main root were greatest at 0 dS/m
and 4 dS/m. The circumference was 13.1 cm and 13.9 cm at 0 and 4 dS/m, respectively, and 6.8 cm
at 16 dS/m, approximately half of that at 4 dS/m. There was a strong positive correlation between
root dry weight and the length and circumference of the main root in each treatment (length: r = 0.99,
p = 0.0006; circumference: r = 0.98, p = 0.001).

Fig. 2 Relationship between root length and saturated hydraulic conductivity

Fig. 3 Relationship between root circumference and saturated hydraulic conductivity
CONCLUSION
Moringa has high salt tolerance and can be grown in paddy soils up to 8 dS/m with almost no growth
inhibition. As soil salinity increased, Na concentrations in each moringa organ also increased. The
results of the correlation analysis in this study also showed that only Mg and P in the branches andFe and Mg in the stems were significantly negatively correlated with Na concentration, suggestingthat increases in Na concentration cause limited inhabitation of mineral uptake by moringa. There
was a strong positive correlation between the length of moringa main roots and saturated hydraulicconductivity, the results showed that moringa root growth increased the soil permeability.
ACKNOWLEDGEMENTSⒸ ISERD

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IJERD – International Journal of Environmental and Rural Development (2023) 14-2

This research was financially supported by The Mitsui & Co. Environment Fund and JSPS
KAKENHI 21H03692.
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