不同基质对闭鞘姜生长发育和光合作用的影响

热带作物学报2021, 42(11): 32063211

Chinese Journal of Tropical Crops

Effects of Different Substrates on the Growth and Development of

Costus speciosus

LIU Xiaorong, WU Zhi, XU Yang, HAN Qingbin, WANG Dairong

Environmental Horticulture Institute, Guangdong Academy of Agricultural Sciences / Guangdong Key Lab of Ornamental Plant

Germplasm Innovation and Utilization, Guangzhou, Guangdong 510640, China

Abstract: In this study, the effects of six substrates consisting of red soil, peat, coir, and perlite in different proportions

on the growth of Costus speciosus were investigated. The physical and chemical properties of the six substrates were

measured after mixed. The sprouting rate, leaf number, stem diameter, plant height, plant width, diurnal variation of

photosynthetic characteristics, rhizome fresh weight (RFW) and rhizome dry weight (RDW) were measured. The results

revealed that the net photosynthetic rate (P

n

) in the peat + coir + perlite, 1∶2∶2 (S4), was the greatest. The P

n

curves

of the six substrates varied in single or double peaks, while the transpiration rate (T

r

) curves displayed a single peak. The

greatest plant height, RFW and RDW were also observed in S4. Based on the findings, S4 was considered a suitable

substrate for C. speciosus growth and dry matter accumulation.

Keywords: Costus speciosus; potting plant; soilless substrate; peat; coir; photosynthetic characteristics

DOI: 10.3969/.1000-2561.2021.11.020

不同基质对闭鞘姜生长发育和光合作用的影响

志，徐 扬，韩庆斌，王代容

All Rights Reserved.

刘晓荣，吴

广东省农业科学院环境园艺研究所/广东省园林花卉种质创新综合利用重点实验室，广东广州 510640

摘 要：采用随机区组设计，研究不同配比的红壤、泥炭、椰糠和珍珠岩6种基质配方对闭鞘姜生长的影响。测定6

种基质的物理和化学性质，观测萌芽率、叶片数、茎粗、株高、株幅、光合日变化、根茎鲜重和根茎干重。结果显示，

基质S4（泥炭+椰糠+珍珠岩=1∶2∶2）的植株净光合速率（P

n

）显著高于其他基质处理。在6种基质生长的植株叶片

净光合速率曲线呈单峰或双峰变化，而蒸腾速率曲线呈单峰变化。最大株高、最大根茎鲜重和根茎干重也出现在基质

S4种植的植株。从以上结果可知，基质S4比较适合闭鞘姜的生长和根状茎干物质积累。

关键词

：

闭鞘姜；盆栽植物；无土栽培基质；泥炭；椰糠；光合特性

中图分类号

：

S682.19 文献标识码

：

A

1 Introduction

Costus speciosus is a rhizomatous perennial

herb with pinkish white flowers on reddish bracts. It

has increased popularity in recent years due to its

medicinal and ornamental properties. Its traditional

potting substrate is soil, which is heavy and ine-

fficient for transport.

Substrate is a key factor that affects plant

growth in soilless cultivation. In addition to sup-

porting and fixing, substrate is important for trans-

ferring adequate oxygen, water, and nutrients from

the nutrient solution to plant roots. Peat has been

widely used in soilless cultivation over the last

century due to its excellent physical and chemical

properties, especially at the seedling stage

[1-3]

. Ho-

wever, as a non-renewable resource and increasing

收稿日期 2020-12-28；修回日期 2021-03-10

基金项目 广东省科技计划项目（No. 2015A020209082）；广东省农业科学院“新兴学科团队创意农业研究团队”项目。

作者简介 刘晓荣（1979—），女，副研究员,研究方向：观赏植物生理生态，E-mail：***************。

第11期 刘晓荣等: 不同基质对闭鞘姜生长发育和光合作用的影响 3207

price, peat has raised concerns among environ-

mental, scientific, and governmental agencies

[4-7]

,

which has resulted in policy changes and govern-

mental regulations of its use in several European

countries.

Coir is now widely used in the soilless cultiva-

tion across the world as an environmentally friendly

substrate which has abundant resources. It is light-

weight, good aeration, and a high water-holding

capacity that is more than eight times of its own

weight

[8]

. Previous studies found that coconut coir

is a good alternative to peat

[9-11]

. It is also cost effi-

cient for raising plant growth, which has been

widely used for growing various fruits, vegetables,

and flowers since the beginning of the century

[12-14]

.

Although coir has a high water-holding capa-

city, it has poor aeration. Mixed and combined with

other coarser material could make up this short-

coming. Pan et al

[15]

demonstrated that Oncidium

grew best in a substrate combination of crushed

stone, bark, coconut shell and charcoal in a 2∶2∶

1∶1 ratio. A hanging ornamental plant was proved

that soil mixture (1 part cocopeat:1 part topsoil:1

part sand) was significantly better than cocopeat

only

[16]

. Bhardwaj

[17]

reported that the medium (coil

+ vermicompost + sand + pond soil) gave maximum

seed germination and seedling growth.

Although, the effects of different substrate

mixtures on flower growth and development have

been previously investigated, there were few reports

available on C. speciosus growth. The objective of

this study was to assess red soil, peat, coir, and per-

lite in different combinations on C. speciosus

growth and development, to develop a labor-effi-

cient and cost-saving substrate.

2 Materials and Methods

2.1 Plant and growth conditions

Rhizomes annually of C. speciosus were wild

germplasm obtained from native. One or two buds

were divided and individually grown in plastic pots

with a diameter of 10 cm and height of 8.0 cm. The

experiment was conducted in the greenhouse in En-

vironmental Horticulture Institute, Guangdong Aca-

demy of Agricultural Sciences, China (113°15 E,

23°08 N) from April, 2018 to October, 2018. The

temperature and relative humidity were recorded by

ZDR-20 data loggers (Hangzhou Zeda Instruments

Co. Ltd., Hangzhou, China). The minimum and ma-

ximum average temperature was 24.3 ℃ and 33.9 ℃,

respectively. Relative humidity was maintained at

the range of 70% to 80%.

2.2 Substrate treatment

Six substrates consisting of red soil, peat, coir,

and perlite in different proportions were used for the

experiment. The red soil was the native field soil.

Peat, coir and perlite were purchased from a hor-

ticultural supplier’s corporation (DGSTAR, Guang-

zhou, China). The mixtures by volume were as fol-

lows: S1 (red soil + perlite; 3∶1); S2 (peat + perlite;

3∶1); S3 (coir + perlite; 3∶1); S4 (peat + coir +

perlite; 1∶2∶2); S5 (peat + coir + perlite; 2∶1∶

2) and S6 (peat + coir + perlite; 2∶2∶1).

Coir was supplied in the form of compressed

bricks (30 cm × 30 cm × 12 cm), and peat was sup-

plied as compressed bails (300 L). Both substrates

were hydrated according to the manufacturer’s in-

structions. Initial substrate samples of each treat-

ment were collected. The potential of hydrogen (pH)

and electrical conductivity (EC) of extracted sub-

strate solutions were analyzed using the pour thr-

ough method

[18]

. The bulk density (BD), total po-

rosity, and aeration porosity of the media were

measured and analyzed

[19]

.

2.3 Experimental design

Plants were arranged in a randomized complete

block design, and each treatment replicated three

times, and in each replicate consisted of 10 plants.

Plants were fertilized using a 20 N-20 P-20 K com-

mercial water-soluble fertilizer (COMPO Expert

GmbH, Munster, Germany) and irrigated two or

three days with tap water. The EC and the pH value

of water are 0.23 mS·cm

–1

and 7.4 respectively.

2.4 Data collection

Data regarding all growth indices were col-

lected in late June before flowering time, including

the plant height, plant width, number of leaves, leaf

length, and leaf width of the third mature leaf from

the top of the plant. Rhizome fresh weight (RFW)

and rhizome dry weight (RDW) were measured in

October. Leaf gas exchange was measured using a

portable photosynthesis measuring system (LI-6400;

LICOR, Lincoln, NE, USA). Stomatal conductance,

intercellular carbon dioxide (CO

2

), net photosyn-

thetic rate (P

n

), and transpiration rate (T

r

) were re-

corded. Water use efficiency (WUE) was calculated

using the following equation: WUE = P

n

/T

r

.

All Rights Reserved.

3208

热带作物学报

第42卷

Diurnal photosynthetic variations were deter-

mined from 8∶30 to 16∶30 in three sunny days

using five plants per treatment, and from the top the

third leaf per plant was selected. Leaf length, leaf

width, chlorophyll content was determined using the

same leaves as those used for other growth parame-

ters above. 3 SPAD readings (Minolta Camera Co.,

Osaka, Japan) were taken on each leaf (inter area).

2.5 Statistical analysis

The data were analyzed using statistical soft-

ware (SAS version 8.1; SAS Institute, Cary, NC). It

was used one-way PROC ANOVA to evaluate vari-

ance of substrate pH, EC, density, total porosity,

aeration porosity, hold-water porosity and gas-water

porosity ratio, number of leaves, stem base diameter,

plant height, leaf length and width, RFW and RDW,

stomatal conductance, intercellular CO

2

, P

n

, T

r

and

WUE and leaf SPAD. Mean separation used least

significant difference (LSD) at P = 0.01 or 0.05.

3 Results

3.1 Substrate physical and chemical proper-

ties

The physical characteristics of the six sub-

All Rights Reserved.

Substrate

S1

pH

5.58±0.39

B

EC

/(mS·cm)

–1

strates were provided in Tab. 1. S1 and S2 had the

lower pH values (5.58 and 4.84, respectively) sig-

nificantly different from each other. No significant

differences were detected among S3, S4, S5, and S6

with regard to pH values. S1 had the lowest EC

value, although there was no significant difference

between S1 and S3. There was the highest bulk

density (0.972 g·L

–1

) And lowest water holding ca-

pacity (54.82%) in S1. S4 had the greatest total po-

rosity (84.67%) and water holding porosity (76.88%),

but had a lower bulk density. No significant differ-

ences were detected in the aeration porosity or gas-

water porosity ratios among the six media treatments.

3.2 Effects of different substrates on vegeta-

tive parameters

The six substrates did not significantly affect

the sprouting rate or leaf length (Tab. 2). The great-

est number of leaves was observed in S4 (25.3).

Although the greatest stem base diameter was ob-

served in S6, no significant differences were detected

among S3, S4, and S6. Plant height was greater in S3

and S4 than in S5. The smallest leaf width was ob-

served in S1 (4.47 cm), but no significant differences

were detected among the other five substrates. The

Tab. 1 Physical and chemical characteristics of six substrate types

Bulk density

/(g·L

–1

)

Total porosity

/%

Aeration

porosity/%

Hold-water

porosity/%

Gas-water

porosity ratio

25.33±3.21

C

0.972±0.029

A

65.67±0.02

C

10.85±0.11 54.82±0.12

b

0.20±0.11

S2 4.84±0.45

C

58.00±1.15

A

0.249±0.006

B

71.00±0.04

BC

7.46±0.04 63.54±0.07

ab

0.12±0.03

S3 6.56±0.21

A

26.33±6.51

C

0.141±0.009

C

72.67±0.06

BC

14.05±0.10 58.62±0.06

b

0.24±0.05

76.88±0.03

a

0.10±0.03 S4 6.72±0.26

A

56.00±7.51

A

0.149±0.015

C

84.67±0.01

A

7.78±0.05

S5 6.77±0.04

A

64.00±2.00

A

0.147±0.026

C

75.33±0.03

B

11.36±0.13 63.97±0.13

ab

0.18±0.13

S6 6.37±0.19

A

40.67±5.68

B

0.168±0.020

C

69.00±0.01

BC

15.92±0.16 64.37±0.07

ab

0.25±0.09

Note: The data in the table are mean ± SD. S1: Red soil + perlite, 3∶1; S2: Peat + perlite, 3∶1; S3: Coir+ perlite, 3∶1; S4: Peat + coir

+ perlite, 1∶2∶2; S5: Peat + coir + perlite, 2∶1∶2; S6: Peat + coir + perlite, 2∶2∶1. pH: Potential of hydrogen; EC: Electrical conduc-

tivity. Different uppercase letters in each column indicate extrmely significant difference (P<0.01). Different lowercase letters in each column

indicate significant difference (P<0.05) based on Duncan’s test.

Tab. 2 Effects of six substrates on vegetative parameters of C. speciosus

Substrate

S1

Sprouting

rate/%

100

Number of

leaves

Stem base

diameter/mm

Plant height

/cm

Leaf length

/cm

15.62±12.12

a

16.43±12.08

a

15.98±11.43

a

Leaf width

/cm

RFW

/g

RDW

/g

20.81

c

23.11

bc

39.98

ab

44.80

a

22.24

bc

27.95

abc

23.30±3.56

bc

8.47±1.27

c

81.51±13.85

ab

4.47±0.51

b

146.03

c

4.89±0.75

ab

163.17

bc

4.97±0.47

ab

231.85

a

S2 99.0 22.80±3.09

bc

9.43±1.51

bc

87.39±15.72

ab

S3 100 21.60±3.65

bc

10.50±2.78

ab

93.23±29.44

a

S4 99.0 25.30±1.92

a

10.60±1.36

ab

92.59±11.41

a

S5 98.0 21.10±3.33

c

8.82±1.85

c

75.64±20.94

b

16.43±3.27

a

4.98±0.79

ab

207.72

ab

15.67±2.89

a

4.68±0.88

ab

163.05

bc

S6 97.0 21.80±2.75

bc

11.41±2.44

a

83.06±9.98

ab

16.68±3.42

a

5.12±0.83

a

193.62

abc

Note: The data in the table are mean ± SD. S1: Red soil + perlite, 3

∶

1; S2: Peat + perlite, 3

∶

1; S3: Coir+ perlite, 3

∶

1; S4: Peat + coir + perlite,

1

∶

2

∶

2; S5: Peat + coir + perlite, 2

∶

1

∶

2; S6: Peat + coir + perlite, 2

∶

2

∶

1. RFW: Rhizome fresh weight; RDW: Rhizome dry weight. Different

lowercase letters in each column indicate significant difference (P<0.05) based on Duncan’s test.

第11期 刘晓荣等: 不同基质对闭鞘姜生长发育和光合作用的影响 3209

RFW (231.85 g) and RDW (44.80 g) of S4 were

greater than those of S1, S2. The lowest RFW

(146.03 g) and RDW (20.81 g) were observed in S1.

3.3 Effects of different substrates on photo-

synthetic physiological characteristics

No significant differences were detected in

stomatal conductance and T

r

among the six substrates

(Tab. 3), but intercellular CO

2

concentration, P

n

, and

WUE were significant. The intercellular CO

2

con-

centration of S1 was greater than S2, and S6. The P

n

of S4 was significantly greater than S1, S2, S5, and

S6. The WUE of S3 was greater than S1, S5 and S6.

Substrate

S1

stomatal conductance

/(mmol·m

–2

·s

–1

)

Intercellular CO

2

concentration/(mg·L

–1

)

3.4 Diurnal changes of leaf photosynthetic

parameters

The diurnal variation curve of leaf P

n

in S1 dis-

played two single peaks (Fig. 1). The first peak was

appeared at 10：30 (11.48 μmol·m

–2

s

–1

), and the second

peak was at 14：30 (13.35 μmol·m

–2

s

–1

). The diurnal

variation curves of leaf P

n

in the other five substrates

were similar and displayed one peak at 12：30. The

average diurnal P

n

of the six substrates were 7.77,

8.50, 9.43, 12.16, 9.71 and 9.00 μmol·m

–2

s

–1

, respec-

tively.

Tab. 3 Correlation coefficients of P

n

and environmental factors of C. speciosus leaves in six substrates.

P

n

/

(μmol·m

–2

·s

–1

)

T

r

/

(mmol·m

–2

·s

–1

)

WUE/

(μmol·CO

2

·mmol

-1

)

207.03±11.61

a

305.06±7.80

a

8.54±2.20

bc

6.19±2.90

a

1.67±0.90

c

S2 174.71±22.90

a

185.73±8.10

b

11.82±4.70

bc

3.98±0.70

a

3.08±1.34

abc

S3 191.35±27.20

a

155.55±10.20

b

13.75±5.60

ab

4.34±3.80

a

3.75±1.74

a

S4 180.14±15.20

a

213.42±8.90

ab

15.98±5.66

a

7.80±8.70

a

2.79±1.96

ab

S5 113.58±5.40

a

252.08±7.60

ab

5.90±2.20

c

3.87±1.50

a

1.81±0.99

bc

S6 98.50±7.60

a

170.40±11.10

b

8.37±1.90

bc

5.89±6.60

a

2.20±0.93

bc

Note: The data in the table are mean ± SD. S1: Red soil + perlite, 3∶1; S2: Peat + perlite, 3∶1; S3: Coir+ perlite, 3∶1; S4: Peat + coir

+ perlite, 1∶2∶2; S5: Peat + coir + perlite, 2∶1∶2; S6: Peat + coir + perlite, 2∶2∶1. P

n

: Net photosynthetic rate; T

r

: Transpiration rate;

All Rights Reserved.

WUE: Water-use efficiency. Different lowercase letters in each column indicate significant difference (P<0.05) based on Duncan’s test.

Fig. 1 Diurnal changes of net photosynthetic rate (P

n

)

on C. speciosus leaves in six substrates

The diurnal variation of T

r

of all six substrates

displayed one peak (Fig. 2), but the times were dif-

ferent. The peak in S1 appeared at 12：30, while the

peaks in S2, S3, and S4 reached their maximum at

14：30. In S5 and S6, the peak appeared at 10：30.

The maximum leaf T

r

was observed in S6 (8.17

mmolm

–2

s

–1

), while the minimum was observed in

S4 (5.22 mmolm

-2

s

-1

). The average diurnal T

r

of

the six substrates were 3.53, 3.28, 3.72, 2.99, 3.50,

and 3.70 mmol·m

–2

s

–1

, respectively.

The diurnal variation of WUE in S1, S3, and

Fig. 2 Diurnal changes of transportation rate (T

r

) on C.

speciosus leaves in six substrates

S6 exhibited a linear rise-fall pattern (Fig. 3). The

peaks of S1 and S3 appeared at 10：30 (4.70 μmol

CO

2

mmol

–1

H

2

O) and 12：30 (5.0 μmol CO

2

mmol

–1

H

2

O) respectively. From 8：30 to 12：30, S6 rose in

a straight line, slowly decreased at 14：30, and

subsequently rose to 5.22 μmol CO

2

mmol

–1

H

2

O.

In contrast to S1, S2, and S6, the diurnal variation

of WUE in S2, S4, and S5 exhibited a linear

fall-rise pattern. The S2 exhibited a linear down-

ward trend from 8：30 to 16：30. From 8：30 to

3210

热带作物学报

第42卷

Fig. 3 Diurnal changes of water use efficiency (WUE) on

C. speciosus leaves in six substrates

14：30, S4 exhibited a downward trend and rose after

14：30 to 6.52 μmol CO

2

·mmol

–1

H

2

O. The S5

reached its minimum level (1.57 μmol CO

2

·mmol

–1

H

2

O) at 10：30, and subsequently increased.

3.5 Effects of different substrates on foliar

SPAD readings

The SPAD readings of S1 were the highest

(44.1), while S3 was the lowest (34.7) (Fig. 4). The

order of SPAD readings among the six substrates

was as follows: S1>S2>S4>S6>S5>S3. No signifi-

cant differences were detected among S1, S2, and

S4. The SPAD readings of S1 was significantly

higher than S3 (P<0.05), which was about 1.27

times.

Fig. 4 Effects of six substrates on foliar SPAD readings

4 Discussion

Soil and peat were the most commonly used

growing substrates in the container production of

annual and perennial ornamental plants

[20]

. However

the density of soil was heavy, difficult to move, and

contains many potentially harmful micro-organisms.

Peat was uneconomical or unrecyclable, making

growers look for alternatives.

In this study, the greatest of S4 over the other

combinations probably related to its characteristics

including higher total porosity and hold-water po-

rosity. The number of leaves, RFW, and RDW of S4

significantly increased compared with other five

substrates. Although sprouting rate and leaf lengths

were not significantly different.

The total porosity and maximum water holding

capacity are important factors for plant growth.

However, porosity and bulk density are interacted

each other. Bulk porosity is low and the air content

is reduced. The air porosity of the substrate is large;

therefore it is more suitable for plant growth. Mid-

dle density was more suitable at the seedling stage;

similar findings were also reported by Chen

[21]

.

The results revealed that the stem base diame-

ter, RFW and RDW were lower in S1 the soil-based

substrate potentially due to its large bulk density

(0.972 g·L

–1

), matching the findings that the density

range of substrate was 0.19~0.70 g·L

–1

for most

potting commercial horticultural crops

[22]

.

Different substrates affected P

n

, T

r

and pore

conductance of two gerbera

[23]

. This study revealed

that the P

n

differed among the six substrates and the

greatest value observed in S4. Intercellular CO

2

and

WUE also differed among the six substrates, in the

following orders S1>S5>S4>S2>S6>S3 and S3>

S2>S4>S6>S5>S1, respectively. However, like P

n

,

the diurnal variation curves of leaf photosynthesis

were similar and exhibited one peak, except S1.

Interestingly, the T

r

of the six substrates displayed

single peak, but the times were different.

The maximum value of P

n

was in S4, which

promoted plant leaf growth and increased rhizome

accumulation. The results confirmed previously

reported findings, in which P

n

directly reflected

plant light energy and the ability to accumulate

photosynthetic products

[24]

.

SPAD-502 meter has been provided a rapid and

nondestructive measurement of leaf chlorophyll

content. Several studies demonstrated that SPAD

readings were significantly related to extracted

chlorophyll

[25-28]

. In the study, the greenest leaves

were observed in S1. Although no significant dif-

ferences were detected between S1 and S4, the de-

gree of leaf greenness reflected plant growth and

physiological health. In future studies if combined

with fertilizer management, the leaf chlorophyll

content would be improved. Therefore, S4 would be

an excellent substrate for C. speciosus growth and

development.

All Rights Reserved.

第11期 刘晓荣等: 不同基质对闭鞘姜生长发育和光合作用的影响 3211

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