SAGO PALM 22: 13−19 (2014)

Growth of sago Palm seedlings under different soil pH Conditions
at the experimental farm in Kendari, Indonesia
Preuk Chutimanukul1, Haruna Tanaka2, Hitoshi Naito3, Takayuki Inagaki1, Yulius B. Pasolon4,
Marselinus S. Padang4, Takashi Mishima1, Yoshihiko Nishimura5, Akemi Itaya1,
Tomohiro Uchiyama1, Ahmad Junaedi6 and Hiroshi Ehara1,*
1

Graduate School of Bioresources, Mie University, Tsu 514-8507, Japan
2

3

Japan International Cooperation Center, Shinjuku 163-0716, Japan

College of Life Sciences, Kurachiki University of Science and The Arts, Kurashiki 712-8505, Japan
4

5

6

Faculty of Agriculture, Halu Oleo University, Kendari 93232, Indonesia

Center for International Education and Research, Mie University, Tsu 514-8507, Japan

Faculty of Agriculture, Bogor Agricultural University, Kampus IPB Dramaga 16680, Indonesia
Corresponding author: H. Ehara (ehara@bio.mie-u.ac.jp)

Key words: acid soil, environmental stress, growth analysis, sago seedlings
Introduction

The sago palm (Metroxylon sagu Rottb.) is amonocotyledon plant that grows in swampy and peaty

soils where almost no other major crops can growwithout drainage or soil improvement (Sato et al.,
1979; Jong, 1995). The sago palm is one of the mostimportant bioresources, not only for sustainableagriculture, but also for rural development in swampy

areas of the tropics (Ehara, 2009). Its carbohydrates
can be further processed into various basic raw

materials for food, animal feed, and industrial uses
(Ehara et al., 2000). In Indonesia, the sago palm has

been culturally significant and has played an

important socioeconomic role for hundreds of years.

According to Oates (1999), by the early 14th century,

the sago palm was a major agricultural product in theregion from South Mindanao and Northern Borneo toSouth Sulawesi and the Maluku Islands. At themature stage, it produces a huge trunk that may reach
7-15 m in height and 1.2 m in average girth at thebase of the palm (Flach and Schuiling, 1989).

However, its size depends on several factors that

include the planted area and physiologicalcharacteristics. According to Anugoolprasert et al.
(2012b), the sago palm can grow in a widely different
soil pH range, from 4.3 to 7.0, under natural
conditions in tropical areas.

Plant growth is generally affected by relative

concentrations of hydrogen ions (H + ). A plant’s

ability to endure various pH levels depends largely on
its ability to take in and utilize nutrients at varying

concentrations in the soil solution. Most crops growwell in soil that is neutral, mildly acidic, or mildly
basic. Soils are naturally acidic or alkaline, so whencorrecting the pH of soil for healthy plant growth, it isimportant to better understand the growth response tosoil pH and to be sensible about the long-term effects
of different soil management on soil pH.
Blagodatskaya and Anderson (1998) reported that thevariations in soil pH are natural, due to the chemical
characteristics of different types of soils. Generally,

plant growth is limited by various factors relating toeach soil type; for example, acidic soils significantly

limit crop production worldwide, with approximately
50% of the world’s potentially arable soils being

acidic, illustrating an important constraint on

14

P. Chutimanukul et al.

agriculture (von Uexkull and Mutert, 1995). Someby Rembon et al. (2008). The germinated seeds grown

nutrient availability to plants. Land utilization often

in the experimental farm, Faculty of Agriculture, Halu

fertilizers can change the soil pH, and thus cause othernutrients to be more available, or can reduce certain
faces problems such as unsuitability for plant growthin Kendari were used as planting materials. The two
experimental plots were placed under natural sunlight
Oleo University. In each plot (14.44 m2) set in the 1or unwanted changes in soil pH. In the case where soil

ha sago palm pilot farm, 16 sago seedlings at the third

Calcium carbonate (CaCO 3 ) is one of the limingtransplanted. The compost with the trade name MOF-

pH is exceedingly low, lime or dolomite can be used
to improve the soil pH, bringing it to the desired level.

materials used to improve or amend the soil, which is

a common solution to acidity. However, the soils arecharacterized by high calcium carbonate (CaCO 3)

leaf stage (three leaves that have leaflets expanded) or

4 months after germination in a polyvinyl bag were821 (Rembon et al., 2008) made from the sago pith

residue after starch extraction (123 g C, 10 g N, 6 g
P2O5, 20 g K2O, 30 g CaO, 10 g MgO/kg) was applied

content (more than 5%), which increases the amountin all the plots at the rate of 40 kg with physiologicaland Somponges, 1987). Hence, the amount of lime or

The two treatment plots were set up as follows: (1)of CaCO3 in the soil, and as a result, the availabilityof most nutrients is considerably decreased (Chein

dolomite required to correct an acidic pH will varyfrom soil to soil. It is generally known that excessive
use of lime or dolomite causes saline soils, and saline

neutral fertilizers, 300 g urea, and 210 g
superphosphate of lime (SP36 containing 36% P2O5).
control: native acid soil with fertilizers applied as

described above; (2) calcium application plot: 3kg

calcium carbonate (CaCO 3 ) was also applied tosoils contain soluble salts in quantities that affectincrease soil pH to above 6 prior to transplanting. The

the ion composition of crops at maturity (Sharma,

experiment was conducted beginning on December 7,

plant growth at various stages and create yield

differences between crops, along with differences in1997).

In this study, we analyzed the growth of sago palm

seedlings grown in the native acid soil and in soil

treated with calcium to increase the soil pH at theexperimental farm in Southeast Sulawesi, Indonesia.
Although the growth of the sago palm is generally
considered to decelerate under a low pH condition,such growth depression is usually observed in peatsoil compared with that in mineral soil, as reported bySato et al. (1979). The purpose of the current study is
to clarify the effect of different pH levels in soils thatare amended with CaCO 3 and that have the same
parent material on the growth of the sago palm.
Materials and Methods

The seeds of Manno type sago palm that werecollected from Sentani, Jayapura, Province of Papua,Indonesia, on July 11, 2010, were transferred to

Kendari, Province of Southeast Sulawesi, as explained

soil profile was characterized based on USDA
classification methods (Pasolon et al., 2009). The
2011, and 3 plants each among 16 plants in theexperimental plot were taken as samples at the

beginning of the experiment as well as on June 24,
2012, and October 2, 2012.

The plant growth parameters such as plant height(from soil surface to the top of the standing palm) andplant length (from soil surface to the tip of the longest
leaf: total length of shoot) were measured, and the leaf

number per plant, leaflet number per leaf, and deadleaf number were counted at the plant sampling in

June and October 2012. The dry matter weight ofleaflets, rachises, petioles, bases of shoots (leaf sheathpart holding the other leaves’ sheath inside) and roots,
and the leaflet area was measured, and analyses of thegrowth from December 2011 to June 2012 and from
June 2012 to October 2012 were performed. The

leaflets of the most active leaves (the uppermost or

second expanded leaf from the top) were used formeasuring the Soil and Plant Analyzer Development

Growth of Sago Palm Seedlings under Different Soil pH Conditions at the Experimental Farm in Kendari, Indonesia

15(SPAD) value, indicating chlorophyll content, using

length in both the control and calcium application

porometer (Decagon Devices, Inc., Pullman, USA) at

plant, and dry matter weight of each part for 6 months

the Chlorophyll Meter SPAD-502 (Minolta Co., Ltd.,

Japan), and the stomatal conductance using a leaf
the sampling in October 2012. The statisticalafter transplanting were almost the same as in thewas a significant difference in leaf number per plant;

Statistical Analysis System (SAS) for Windows v9.0.

however, the difference was not so large.
Additionally, in the results of growth analysis for 6

Results and Discussion

months, there were no remarkable differences

The soil type of the experimental site was alluvial

between the control and calcium application plotsfans, which originally formed from the surrounding(Table 3).

sedimented yellow podzolic soils. The texture of the

Growth parameters at 10 months after transplanting

virgin soil was loam, which contained 44% sand, 44%
silt, and 12% clay, and its soil pH was 3.5 (KCl) toare shown in Table 4. There were no apparent

4.2 (H2O). The soil pH in the control and calcium

differences in plant height and plant length between

application plots was 4.4 (KCl) to 5.4(H2O) and 6.5(KCl) to 6.9 (H2O) respectively, and recognized asacid (control) and neutral (calcium application plot)(Table 1). Calcium application treatment increased
Olsen P2O5, exchangeable Ca content, and CEC anddecreased Al

length, leaflet number per leaf, dead leaf number per
control and calcium application plots (Table 2). Theredifference of the data was determined using the3+plots (Table 2). The growth in plant height, plantcontent in the soil. Then K2O (HCl25%), Morgan K 2 O, exchangeable Mg, and K

the control and calcium application plots. The leaf
number per plant was 16.9 and 16.1 in the control andcalcium application plots, respectively, and they wereat almost the same level. Considering the result thatthe leaf number and dead leaf number were at thesame level in the two plots, the different soil pH
levels in the case of the same soil parent material had

contents in the soil tended to be lower in the calcium

no apparent effect on leaf emergence and senescence.

application plot than in the control.

These results were in agreement with a former report

The sago seedlings grew to around 100 cm in plantby Anugoolprasert et al. (2012a), in which a different

Table 1. Property of the virgin soil and soils in experimental plots.
Plot
Virgin soil
Control
Calcium
application

Olsen
P 2O 5(mg/kg)Bray IP 2O 5(mg/kg)Morgan
K 2O(mg/kg)

60
170

-

5.0
140.9

27
103

90

53

122.3

46

P 2O 5
K 2O(HCl 25%) (HCl 25%)
(mg/kg)(mg/kg)

pH
(HCl)

pH
(KCl)

Total
N
(mg/kg)

4.2
5.4

3.5
4.4

400
900

20
380

6.9

6.5

600

410

Exchangeable cation (cmol c/kg)

Plot

CEC

Al3+

Ca

Mg

K

Na

(cmolc/kg)(cmolc/kg)

Virgin soil

0.73

0.20

0.05

0.13

2.34

0.84

Control

1.40

0.55

0.20

0.12

2.66

0.02

Calcium
application

6.05

0.28

0.09

0.16

4.17

0.00

P. Chutimanukul et al.

16

Table 2. Growth parameters of the control and calcium application plots at 6 months after transplanting (June 2012).
C

Plot
C
Control
Calcium application

Plant height(cm)

Plant length
(cm)

Leaf
number/plantLeafletnumber/leaf

87.3 ± 2.9 a
82.6 ± 3.1 a

100.3 ± 2.8 a
96.4 ± 2.4 a

11.9 ± 0.2 a
10.6 ± 0.3 b

13.1 ± 0.3 a
13.1 ± 0.4 a

1Dead leaf
number1
0.5 ± 0.2 a
1.3 ± 0.3 a

C

3

Plot
C
Control
Calcium application

Leaflet(g plant-1 )Rachis(g plant-1 )Petiole(g plant-1 )Base(g plant-1 )Root(g plant-1 )40.0 ± 2.3 a
33.5 ± 0.3 a

3.6 ± 0.2 a
3.0 ± 0.1 a

69.7 ± 5.1 a
57.6 ± 2.2 a

9.4 ± 1.1 a
5.0 ± 0.9 a

27.1 ± 2.5 a
28.3 ± 1.6 a

Whole(g plant-1 )3
149.7 ± 10.3 a
127.3 ± 3.0 a

Each value represents the mean ± SE (n=3). Different letters in the table indicate significant difference at the 0.05 probability
level, according to the T-test.

Table 3. Parameters of growth analysis from December 2011 to June 2012.
Plot

RGR
(mg g -1 d-1)

NAR
(mg cm -2 d-1)

LAR
(cm 2 g-1)

SLA
(cm 2 g-1)

LWR
(g g -1)

Control P
Calcium application

15.56 ± 0.34 a
14.44 ± 0.12 a

0.543 ± 0.023 a
0.470 ± 0.024 a

28.9 ± 0.6 a
31.5 ± 1.5 a

113.2 ± 1.8 a
116.3 ± 4.8 a

0.255 ± 0.002 a
0.271 ± 0.003 a

CC
Each value represents the mean ± SE (n=3). Different letters in the table indicate significant difference at the
0.05 probability level, according to the T-test.

Table 4. Growth parameters of the control and calcium application plots at 10 months after transplanting (October 2012).
Plot

Plant height(cm)

Plant length
(cm)

Leaf
number/plantLeafletnumber/leaf

Dead leaf
numberControl
Calcium application

124.4 ± 4.0 a
112.5 ± 4.6 a

134.0 ± 1.0 a
123.5 ± 3.8 a

16.9 ± 0.2 a
16.1 ± 0.4 a

34.0 ± 0.7 a
30.5 ± 1.2 a

3.1 ± 0.0 a
3.0 ± 0.2 a

Plot

Leaflet(g plant-1 )Rachis(g plant-1 )Petiole(g plant-1 )Base(g plant-1 )Root(g plant -1 )Whole
(g plant-1 )Control
Calcium application

127.0 ± 6.6 a
101.0 ± 10.4 b

14.0 ± 1.3 a
10.6 ± 1.7 a

199.7 ± 19.6 a
123.7 ± 12.8 b

38.1 ± 3.8 a
26.2 ± 3.3 b

34.3 ± 1.0 a
37.8 ± 1.9 a

413.5 ± 31.8 a
299.2 ± 30.2 b

6.6 a
1Each value represents the mean ± SE (n=3). Different letters in the table indicate significant difference at the 0.05 probabi lity
6.6 a1level, according to the T-test.

pH condition of culture solution, such as pH 3.6, 4.5,

weights between the two plots, though the leaflet,months during summer in a greenhouse at Mie

calcium application plot with a higher soil pH

and 5.7, had no effect on the number of emerged

leaves, live green leaves, or dead leaves for 4.5

University, Japan.

The difference in dry matter weight tended to bedifferent from plant part to part (Table 4). There wereno significant differences in rachis and root dry

petiole, base, and total dry weights were larger in thecontrol, with a lower soil pH condition, than in thecondition. Considering that there were no significant
differences in the leaf number per plant and leaflet

number per leaf in the two experimental plots and thatthe leaf dry weight was larger in the control than inGrowth of Sago Palm Seedlings under Different Soil pH Conditions at the Experimental Farm in Kendari, Indonesia

17

the calcium application plot, the size of a single leaflet

growth rates of individuals might be attributed to theto observe the difference in the size of single leaflet,

Considering that there was no difference in themight be larger in a lower soil pH condition than in ahigher soil pH condition. It took longer than 6 months

which might be related to the leaf emergence rate, one
leaf per month in general (Jong, 1995). It wasconsidered that a comparatively long period would beneeded for the emergence of a leaf to reflect the effectof the experimental procedure.

Table 5 shows the results of growth analysis for 4

months from 6 months after transplanting. There wasa significant difference in relative growth rate (RGR),
and the value was larger in the control than in the

calcium application plot. RGR can be divided into net

assimilation rate (NAR), which is a good index toevaluate photosynthetic activity, and leaf area ratio(LAR) to investigate the morphogenesis of anindividual plant. The NAR of the control plants waslarge compared with that of the plants in the calcium
application plot. The components of the LAR, specificleaf area (SLA), indicating leaf thickness, and leaf
weight ratio (LWR), which is the ratio of leaf weight
to whole plant weight, were the same in the two

experimental plots. Although there was a significant

difference in NAR, there was no difference in theP

LAR. Based on these results, the difference in theCdifference in assimilation rate, not to morphologicalcharacteristics.

LAR, dry matter might increase, keeping a goodbalance with the increase in the single leaflet area andtotal leaf area. Two parameters relating to the NAR,

SPAD value and stomatal conductance, are shown in
Table 6. There was no significant difference in the

SPAD value indicating chlorophyll content per unit
leaf area; however, the stomatal conductance washigher in the control than in the calcium application

plot. Omori et al. (2000) reported that stomataldensity is related to thickness of leaf. Ehara (2009)
stated that higher stomatal density would be observedin sago palms grown in soil that contains higher

exchangeable Ca. However, there was no significant
difference in the SLA indicating leaf thickness

between the two experimental plots, and theexchangeable Ca content in the soil was much higher

in the calcium application plots. Considering theseresults, the difference in stomatal conductance in thecurrent experiment might be attributed to stomatal
aperture, which might account for the difference ingrowth rate through assimilation rate, that is,

photosynthesis ability. Such differences in stomatal

Table 5. Parameters of growth analysis from June 2012 to October 2012.
Plot

RGR
(mg g-1 d-1)

NAR
(mg cm -2 d-1)

LAR
(cm 2 g-1)

SLA
(cm 2 g-1)

LWR
(g g -1)

Control
Calcium application

10.18 ± 0.75 a8.30 ± 0.97 b

0.388 ± 0.036 a
0.297 ± 0.037 b

26.7 ± 0.5 a
28.2 ± 0.2 a

91.3 ± 1.1 a
92.8 ± 0.9 a

0.293 ± 0.005 a
0.303 ± 0.001 a

Each value represents the mean ± SE (n=3). Different letters in the table indicate significant difference at the
0.05 probability level, according to the T-test.

Table 6. Stomatal conductance and SPAD value at 10 months after transplanting in October 2012.
Plot
Control
Calcium application

Stomatal conductance(mmol m -2 s-1)
62.4 ± 5.3 a
43.5 ± 5.2 b

SPAD
67.1 ± 2.0 a
67.3 ± 2.4 a

Each value represents the mean ± SE (n=3). Different letters in the table indicate significant
difference at the 0.05 probability level, according to the T-test.

C18

P. Chutimanukul et al.

conductance and NAR would be reflected inwere no significant differences in plant size andexperimental plots. Anugoolprasert et al. (2012b)

interaction. This experiment indicated that the growthdifference in soil properties, such as the K2O and/or

exchangeable K content in soil, of the tworeported that the sago palm adapts to a range of pH

values from 4.3 to 7.0 even in soils that have different

texture in southern Thailand. They supposed that thesago palm grown under any conditions of soil pH
might exhibit an avoidance mechanism to restrict the

distribution of any excess of undesirable nutrients inthe plant, which would account for the mineralcontents in the plant under a wide range of soil pH
conditions. According to Anugoolprasert et al.
(2012a), sago palm seedlings can maintain nutrient

uptake under a wide range of low pH conditions (pH
5.7 to pH 3.6) in a culture solution.

Peaty soil is highly to weakly acidic and contains

low levels of calcium, potassium, and phosphorus
compared with mineral soil (Tie et al., 1991;
Yamaguchi et al., 1994). Although the sago palm

grows in mineral soil and in deep to shallow peat soil,

the growth of the sago palm will be better in mineralsoil or shallow peat soil (Sato et al., 1979). Kakuda etal. (2000) supposed that the reason the growth of thesago palm in peat soil will decelerate compared withgrowth rate between the two experimental plots, as

shown in Table 2, which might be related to microbial

and biomass production of sago palm seedlings at 10

months after planting tended to be better in acidic soil
than in neutral soil conditions, as shown in Table 4.

This phenomenon may be due to nutrient competition

between Ca and Mg with K. Table 1 indicates thatlime application reduced the availability K2O and theexchangeable K cation. In the current study, thegrowth of sago palm seedlings at soil pH 5.4 (H2O)[4.4 (KCl)] was not small in comparison with that at
soil pH 6.9 (H2O) [6.5 (KCl)] on an experimental
farm.

It is still unclear whether the sago palm will show apreferable growth at a lower soil pH condition. Basedon the results of the current study, the experimental
procedures at field level show that the growth of thesago palm will not decelerate under acidic conditions
compared with that under neutral conditions in soils
that have the same parent material.
Acknowledgements

This article is listed as contribution No. 49 from thethat in mineral soil might be the small amount ofLaboratory of Crop Production and Ecology, Miewill be one of important factors limiting sago palm

Project launched by Secretariat of Agriculture,

nitrogen supply per unit of cubic capacity in peat soil.
Considering the former studies, nitrogen provision

growth. On the other hand, Lina et al. (2009) reported
that nitrogen did not significantly improve the growthparameters, such as cumulative increase in height,
monthly growth rate, base diameter, number of leaves

per palm, and number of leaflets per palm, in the early

growth stages in the field experiment because
nitrogen was provided by nitrogen fixation bacterium.

Shrestha et al. (2007) reported that beneficial

University. This study was in part supported by

Global-Scale Issue International Research Network
Forestry and Fisheries Research Council, Ministry ofAgriculture, Forestry and Fisheries, to whom we
express our gratitude.
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