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UNIVERSITI PUTRA MALAYSIA BIOLOGICAL STUDIES ON SONNERATIA CASEOLARIS OBTAINED FROM THE SELANGOR RIVER, PENINSULAR MALAYSIA LOO KHAI KIN. FS 2005 16

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Page 1: UNIVERSITI PUTRA MALAYSIA BIOLOGICAL STUDIES ON … filesedimen, dahan dan daun adalah 1.50 & 0.05 mg/g, 5.62 & 0.14 mglg and 22.79 + 1.08 mg/g. Kandungan nitrogen tanah didapati semakin

UNIVERSITI PUTRA MALAYSIA

BIOLOGICAL STUDIES ON SONNERATIA CASEOLARIS OBTAINED FROM THE SELANGOR RIVER, PENINSULAR MALAYSIA

LOO KHAI KIN.

FS 2005 16

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BIOLOGICAL STUDIES ON SONNERATZA CASEOLARZS OBTAINED FROM THE SELANGOR RIVER, PENINSULAR MALAYSIA

LOO KHAI KIN

Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia, in Fulfilment of the Requirements for the Master of Science

October 2005

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DEDICATION

This master thesis is especially dedicated to the following most patient persons in my life who made the impossible, possible.

My parents,

sisters and my wife.

Thank to God.

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Abstract of thesis presented to the Senate of Universiti Putra Malaysia in fulfilment of the requirements for the degree of Master of Science.

BIOLOGICAL STUDIES ON Sonneratia caseolaris OBTAINED FROM SELANGOR RIVER, PENINSULAR MALAYSIA

BY

LOO KHAI KIN

October 2005

Chairman : Misri Kusnan, PhD

Faculty : Science

Sonneratia caseolaris is a display tree for fireflies in Kuala Selangor. Fireflies has

become a tourist attraction to this area and contribute to the economy of the people

related to the industry. S. caseolaris trees are found in abundance in certain sections

along the Selangor river which are influenced by both freshwater from the

upperstream and seawater during high tide.

Analysis of the important elements in the plants showed that the nitrogen content in

the sediment, branches and leaves were 1.50 + 0.05 mglg, 5.62 + 0.14 mglg and

22.79 k 1.08 mglg, respectively. The soil nitrogen content was decreasing from the

upper stream to the seaward sites. The nitrogen content of the leaves of S. caseolaris

grown in the upper stream was higher than those of trees grown in the seaward sites.

The phosphorus content in the sediment, branches and leaves were 1.02 + 0.03 mglg,

0.258 r 0.01 mglg and 0.58 r 0.03 mglg, respectively. The phosphorus content in

the leaves of trees at the upper stream was higher than those of the seaward sites.

The potassium content in sediment, branches and leaves were 9.27 * 0.31 mglg,

14.69 r 0.58 mglg and 31.85 + 1.10 mglg, respectively. The distribution of

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potassium was more at the seaward sites and to a lesser extent at the upper stream.

However, the potassium concentration in leaves grown at the upper stream was

higher than the seaward sites. These results suggest the influence of water movement

in distributing nutrients to the S. caseolaris habitat along the Selangor river system.

S. caseolaris produced large numbers of fruits that may contain over a thousand

seeds per fruit of which around 90% are viable. Seed germination seems to be the

only way of propagation of this plant. Laboratory studies showed that high salinity

inhibited the germination seeds of S. caseolaris. The inhibition was proportional

with the salinity. Only about 2% of the seeds germinated in seawater. The present

study also showed that salinity affected the stem height of the seedlings. The stem

height was reduced at 13.0 and 26.0%0 salinity. The xylem, pith cells, parenchyma

cells of the cortex and air spaces of the stem seedlings were also smaller than the

stem of the seedlings treated at lower salinity (0 - 6.5%0). The shape of leaves grown

under high salinity (13.0 and 26.0%0) were more rounded and the leaves of the

seedlings grown under low salinity (0, 3.3 and 6.5%0) were longer. The leaf

mesophyll and the parenchyma cells of the seedlings exposed to high salinity (13.0

and 26.0%0) were also smaller than the seedlings exposed to low salinity (0 - 6.5%0).

The increase of salinity had resulted in the increase of the vapor pressure deficit,

which on the other hand, reduced transpiration and stomata conductance.

This study also showed that suspended solids and inundation levels may influence

the growth of the seedlings. The stem height, total and average leaf area, length and

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width of the leaf and root fresh weight and internal structure of the seedlings were

varied with the levels of suspended solids and the inundation level.

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Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai memenuhi keperluan ijazah untuk Master Sains.

BEBERAPA KAJIAN BIOLOGI TENTANG Sonneratia caseolaris YANG DIDAPATI DARI SUNGAI SELANGOR, SEMENANJUNG MALAYSIA

oleh

LOO KHAI KIN

Oktober 2005

Pengerusi : Misri Kusnan, PhD

Fakulti : Sains

Sonneratia caseolaris adalah pokok kediaman untuk kelip-kelip di Kuala Selangor.

Kelip-kelip telah menarik pelancong ke tempat ini dan memberi sumbangan kepada

ekonomi kepada penduduk ang berkaitan dengan industri ini. Pokok S. caseolaris

boleh dijumpai dengan banyak di beberapa kawasan sepanjang Sungai Selangor

yang dipengaruhi oleh air tawar dari hulu sungai dan air masin dari hilir sungai.

Analisis elemen penting dalam pokok menunjukkan kandungan nitrogen dalam

sedimen, dahan dan daun adalah 1.50 & 0.05 mg/g, 5.62 & 0.14 mglg and 22.79 +

1.08 mg/g. Kandungan nitrogen tanah didapati semakin berkurangan dari hulu ke

hilir sungai. Kandungan nitrogen dalam daun S. caseolaris yang tumbuh di hulu

sungai adalah lebih tinggi dari di hilir sungai. Kandungan fosforus dalam sedimen,

dahan dan daun adalah 1.02 2 0.03 mglg, 0.258 & 0.01 mg/g and 0.58 & 0.03 mglg.

Kandungan fosforus dalam dam pokok di hulu sungai adalah lebih tinggi dari hilir

sungai. Kandungan kalium dalarn sedimen, dahan dan daun adalah 9.27 ? 0.31 mglg,

14.69 k 0.58 mglg and 31.85 + 1.10 mg/g. Penyebaran kalium dalam sedimen adalah

lebih di kawasan hilir sungai. Walaubagaimanapun, kandungan kalium dalam daun

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bertumbuh di hulu sungai adalah lebih dari di hilir sungai. Keputusan ini

mencadangkan adanya pengaruh pergerakan air dalam penyebaran nutrien di

kawasan pertumbuhan S. caseolaris di sepanjang Sungai Selangor.

S. caseolaris mengeluarkan banyak buah yang mengandungi sehingga seribu biji

dalarn satu buah dan 90% adalah biji boleh bercambah. Percambahan adalah satu-

satunya cara bagi pokok ini membiak. Kajian makmal menunjukkan saliniti yang

tinggi akan menyekat percambahan biji S. caseolaris. Sekatan ini berhubungkait

dengan saliniti. Hanya 2% biji bercambah dalam air laut. Kajian ini juga

menunjukkan saliniti mempengaruhi tinggi pokok. Pokok adalah rendah dalam

saliniti 13.0 dan 26.0%0. Xilem, sel pith, sel parenkima dalam korteks dan ruang

kosong dalam batang pokok ini adalah lebih kecil dari pokok hidup dalam 0 - 6.5%0.

Bentuk daun bertumbuh dalam saliniti tinggi (13.0 dan 26.0%0) adalah lebih bulat,

manakala bertumbuh dalam saliniti rendah (0-6.5%0) adalah lebih panjang. Mesofil

dan parenkima dalam daun bertumbuh dalam saliniti tinggi (13.0 dan 26.0%0) adalah

lebih kecil dari salinti rendah (0 - 6.5%0). Penambahan saliniti akan meninggikan

tekanan wap defisit tetapi mengurangkan fotosintesis, transpirasi dan rintangan

stomata.

Kajian ini juga menunjukkan pepejal terampai dan tahap rendaman mungkin

mempengaruhi pertumbuhan anak pokok. Tinggi, jumlah dan purata luas daun,

panjan dan lebar daun, berat basah akar dan struktur dalaman dalam anak pokok

vii

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adalah berlainan dalam kepekatan pepejal terabung dan tahap rendaman yang

berlainan.

... V l l l

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ACKNOWLEDMENTS

I am most indebted and grateful to my supervisors Dr. Misri Kusnan, Prof. Dr.

Saberi and Assoc. Prof. Dr. Japar Sidik Bujang who had guided me along the way

towards the completion of this master degree. They had given me a lot of

constructive comments, advice and ideas. With their support and motivation, my

research was conducted efficiently.

Also, I would like to extend my sincere gratitute to Assoc. Prof. Dr. Ahmad Ismail,

Dr. Abdul Rahim and Dr. Hishamuddin who had supported me towards the

completion of this master thesis and gave me a lot of ideas on how to write good

scientific papers. Their time, kindiness and constant encouragement are very much

appreciated. Thanks to the Malaysia Nature Society (MNS) for supporting me in my

field work.

My thanks are also extended to Abu Hena, Dr. Yap, Nik, Mr. Gopal, Mr. Hidir and

all my friends who indirectly contributed so much towards this achievement.

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J certi* that an Examination Cornittee met on 6 October 2005 to conduct the final e~am~ation of L.oo Klai Kin on his master thesis entitled "Riologi~al Aspects of Snnnerc??lo r r ~ s e o l a ~ s in Kuda Selmgor" in accordance with 1Jniveniti Pertaaim Malaysia (Iiigher Degree) Act 1980 md Universiti Pertmian Malaysia (Higher Degree) Rega!atttien~ 1981. T!x Committee r~.ommends that the candidate be awarded the relevm? degree. Members of the Exmhat1on Co11~v&ee are as felle9s:

Abdul T a t i f f Mahamad, Phll Eefesser F2clAty Science md Techelegies Universiti Kebangsaan Malaysia (Member)

Professor/ Deputy Dean School of Graduate Studies Universiti P&a Mdaysia

27 DEC 2005

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This thesis submitted to the Senate of Universiti Putra Malaysia and has been accepted as fulfilment of the requirement for the degree of Master of Science. The members of the Supervisory Committee are as follows:

Misri Kusnan, PhD Lecturer Faculty Science and Environmental Studies Universiti Putra Malaysia (Chairman)

Saberi Othman, PhD Professor Faculty Science and Technologies Universiti Pendidikan Sultan Idris (Member)

Japar Sidik Bujang, PhD Associate Professor Faculty Science Universiti Putra Malaysia (Member)

AINI IDERIS, PhD Professor/Dean School of Graduate Studies Universiti Putra Malaysia

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DECLARATION

I hereby declare that the thesis is based on my original work except for quotations and citations which have been duly acknowledged. I also declare that it not been previously or concurrently submitted for any other degree at UPM or other institutions.

LOO KHAI KIN

xii

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TABLE OF CONTENTS

Page

DEDICATION ABSTRACT ABSTRAK ACKNOWLEDGEMENTS APPROVAL DECLARATION LIST OF TABLES LIST OF FIGURES LIST OF ABBREVIATIONS

CHAPTER

1 INTRODUCTION

LITERATURE REVIEW 2.1 Sonneratia caseolaris 2.2 Development of seeds 2.3 Nutrients

2.3.1 Nitrogen 2.3.2 Phosphorus 2.3.3 Potassium

2.4 Salinity 2.4.1 Salinity effects to seed germination 2.4.2 The effect of salinty to plants

2.5 Total suspended solid

3 Study Area

4 Nutrients in sediments and plants 4.1 Introduction 4.2 Materials and methods

4.2.1 Sample Collection 4.2.2 Preparation of samples for analysis 4.2.3 Sulphuric acid-Hydrogen Peroxide Digestion 4.2.4 Nitrogen analysis 4.2.5 Phosphorus analysis 4.2.6 Potassium analysis 4.2.7 Classification of the station and statistic analysis

4.3 Result and discussion

5 Morphology of fruits and seeds of Sonneratia caseolaris 5.1 Introduction

. . 11

iii vi ix X

xii ... X l l l

xiv xix

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5.2 Materials and methods 5.3 Result and discussion

6 Effect of salinity on germination 6.1 Introduction 6.2 Materials and methods 6.3 Result and discussion

Effect of salinity on seedlings growth 7.1 Lntroduction 7.2 Materials and methods 7.3 Result and discussion

7.3.1 Effect of salinity on the seedling's growth 7.3.2 Effect of salinity on physiological growth of the seedlings.

Effect of salinity on anatomical structure 8.1 Lntroduction 8.2 Materials and methods 8.3 Result and discussion

Effect of total suspended solid on seedlings growth 9.1 Introduction 9.2 Materials and Methods 9.3 Result and discussion

9.3.1 Effect on growth of the seedlings 9.3.2 Effect of inundation and total suspended solid on transpiration rate,

stornatal conductance and vapor pressure deficit of the seedlings 136

Effect of inundation and total suspended solid on anatomical structure 10.1 Introduction 141 10.2 Materials and Methods

10.2.1 Simulation experiment 142 10.2.2 Process of plant tissue 142 10.2.3 Slide Observation 145 10.2.4 Statistical analysis 145

10.3 Result and discussion 145

GENERAL DISCUSSION 12 SUMMARY

REFERENCES APPENDICES BIODATA OF THE AUTHORS

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LIST OF TABLE

Table

3.1 Current extent of mangrove forest reserves and stateland mangroves in Malaysia

Some physico-chemical characteristics (salinity and pH) of water and plant densities at the sampling site.

The fruit weight, fruit thickness, fruit diameter, seed weight, number of seed and percentage of seed germinated in distilled water of S. caseolaris.

Correlation of fruit weight, fruit thickness, fruit diameter, seed weight, number of seed and percentage of seed germinated in distilled water of S. caseolaris.

Stem height with time in S. caseolaris exposed to different salinity (%o) levels.

Measurement of components of seedlings exposed to different salinity levels.

Average root length, shoot fresh and dry weight, root fresh and dry weight of the seedlings of S. caseolaris after the end of the high and low suspended solid treatment.

Area of components @m2) of seedlings exposed to high and 149-150 low suspended solid and different inundation level.

. . . Xl l l

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LIST OF FIGURES

Figure

2.1

2.2

2.3

3.1

4.1

4.2

4.3

Page

7

8

8

45

56

57

5 8

Flower of Sonneratia caseolaris.

Pneumatophores of Sonneratia caseolaris.

Fruit of Sonneratia caseolaris.

Sampling site.

Nitrogen, phosphorus and potassium concentration in soil.

Nitrogen, phosphorus and potassium concentration in leaves.

Nitrogen, phosphorus and potassium concentration in branches.

The mature fruits of S. caseolaris.

Percentage of seed germinated in distilled water with time.

Percentage of germinated and ungerminated seed after 20 days exposed to different salinity.

Percentage of germinating S. caseolaris seeds exposed under different salinity.

Percentage of germinating inhibited S. caseolaris seeds after treatment in distilled water.

The percentage of germination in distilled water of 70 ungerminate S. caseolaris seeds exposed to different salinity.

Seedlings of S. caseolaris after 8 weeks exposed to different salinity.

The number of leaves present in the seedlings exposed to different salinity levels with time.

Leaf survivorship in seedlings exposed to different salinity levels with time.

Cumulative leaf fall number of the seedlings exposed to different salinity with time.

Cumulative leaf number of seedlings exposed to different salinity levels with time.

xiv

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Total leaf area per seedling exposed to various salinity with time.

Leaf area of seedlings exposed to different salinity levels with time.

Leaf of S. caseolaris exposed to different salinity levels.

Leaf length of seedlings exposed to different salinity levels with time.

Mean leaf width of seedlings exposed to different salinity levels with time.

Maximum leaf width of seedlings exposed to different salinity levels with time.

Effect of different salinity levels on photosynthetic rate with time.

Effect of different salinity levels on stomata1 conductance in seedlings with time.

Effect of different salinity levels on transpiration rate in seedlings with time.

Effect of different salinity levels on vapor pressure deficit in seedlings with time.

The comparative cell sizes of different leaf vascular bundle of seedlings exposed to salinity levels.

The comparative cell sizes of lower part of leaf midrib of seedlings exposed to salinity levels.

The comparative of cell sizes of leaf blade of seedlings exposed to salinity levels.

The comparative of cell sizes of stem vascular bundle of seedling exposed to salinity levels.

The comparative of cell sizes of pith cells of seedling exposed to salinity levels.

The comparative of cell sizes of stem cortex of seedling exposed to salinity levels.

Experimental setup to simulate high tide and low tide.

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The lnt-ln0 stem height with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

The average increment of stem height with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Increment of stem height in S. caseolaris after 8 exposed to low suspended solid and high suspended solid and different inundation level

Seedlings exposed exposed to low suspended solid and high suspended solid and different inundation level after 8 weeks.

Number of leaves with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level

Number of leaves of S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level

Total leaves production with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Total leaves production of S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level.

Accumulative leaves fall with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Total leaves senesced of S. caseolaris after 8 week exposed to low suspended solid and high suspended solid and different inundation level.

Total leaves area with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Total leaves area S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level

Mean leaf area with time in S. caseolaris exposed to low suspended solid and high suspended solid and different

xvi

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inundation level

Mean leaf area with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Leaf length with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Leaf length S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level.

Average leaf width with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Average leaf width S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level.

Maximum leaf width with time in S. caseolaris exposed to low suspended solid and high suspended solid and different inundation level.

Maximum leaf width S. caseolaris after 8 weeks exposed to low suspended solid and high suspended solid and different inundation level.

Transpiration rate with time in S. caseolaris exposed to low and high suspended solid and different inundation level.

Stomata conductance with time in S. caseolaris exposed to low 139 and high suspended solid and different inundation level.

Vapor pressure deficit with time in S. caseolaris exposed to low and high suspended solid and different inundation level.

The comparative cell sizes of different leaf vascular bundle of seedlings exposed to low and high suspended solid and different inundation level.

The comparative cell sizes of different leaf blade of seedlings exposed to low and high suspended solid and different inundation level.

The comparative cell sizes of different parenchyma cells in leaf midrib of seedlings exposed to low and high suspended

xvii

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solid and different inundation level.

The comparative cell sizes of different stem vascular bundle of seedlings exposed to low and high suspended solid and different inundation level.

The comparative cell sizes of different stem cortex of seedlings exposed to low and high suspended solid and different inundation level.

The comparative cell sizes of different root vascular bundle of seedlings exposed to low and high suspended solid and different inundation level.

xviii

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mM

NaCl

pmol

co2

LIST OF ABBREVIATIONS

Hectare

Percent

Part per thousand

Kilogram

Kilometer

Liter

Meter

Milligram

Millimeter

Degree Celsius

MilliMolar

Sodium chlorite

MicroMolar

Carbon dioxide

second

xix

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CHAPTER 1

INTRODUCTION

Mangrove plants are salt tolerant plants of tropical and subtropical intertidal regions

of the world, which are under regular or occasional inundation by water of riverine,

estuaries or oceanic origin (Hamilton and Snedaker, 1984; Lacerda, 1998).

Mangrove plants are well known to be resistant to high salinity and waterlogged

conditions. They are also able to tolerate high concentration of nutrients and heavy

metals (Chen et al., 1995).

Lacerda et al. (1993) reported that there is a total area of 4,062,335 ha mangroves in

America, whereas, there are 3,257,700 ha mangroves in Africa (Diop, 1993) and

6,877,600 ha in Asia (Saenger et al., 1983). The highest species diversity occurs in

Asia with a total of more than 70 species recorded, followed by 20-30 species in

Africa. Among 70 species of true mangrove plant, 65 of them contribute

significantly to the structure of mangrove forests. Approximately 40 of these species

found in Southeast Asia, about 15 species are found in African and 10 species in

America (Fields, 1995). Kitamura et al. (1997) had recorded 27 species of true

mangroves in Bali and Lombok, Indonesia. In addition there were 1 species of

mangrove fern, 1 species of palm, 2 species of mangrove shrub and 19 species

coastal plants associated with the mangrove. Saberi (2000) had recorded seventeen

species of true mangrove and seven species of sub-mangrove plants in the transition

zone to dry land forest in Rhizophora mucronata dominated area in Sungai Pulai,

Kukup and Sepang mangrove forest.

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Mangroves play an important role in protection and stabilization of the coastal zone,

nursery ground for a variety of economically important molluscs, crustaceans and

fish and are sources of important products to coastal human populations in the form

of timber, firewood and charcoal, chemicals, medicine and waterways for

aquaculture (Lacerda, 1998) and sanctuary of wildlife. Mangroves have been used as

venues for scientific research and educational activities (Department of Primary

Industries, 1990). The mangrove ecosystem at Kuala Selangor is an example with

such value in the country. Mangrove area at Kuala Selangor is unique for its

fireflies, Pteroptyx tener, that became tourist attraction. They were observed in great

numbers in selected Sonneratia caseolaris trees, which formed their important

habitat.

As for socio-economic benefit of the tourism industry, it was reported that each

boatman could earn about RM 360 to RM 640 per month in Kampung Kuantan. Of

those visiting this area 68% were locals and 32% were foreigners. Of the foreigners,

the highest numbers came from Japan, UK and other ASEAN countries (notably

Singapore). Two thirds of all visitors came to know of the fireflies though word of

mouth (PE Research Sdn. Bhd., 2001).

In Kuala Selangor the largest populations of Sonneratia caseolaris are found

between Kampung Lanun and Batu Lapan about 24 kilometers along the Selangor

river. Sonneratia caseolaris vegetation spread out in low salinity along the river.

Metroxylon sago, Nypa fruticans, and Acanthus ilicifolius are also spread out in low

salinity and associated with S. caseolaris.

Page 25: UNIVERSITI PUTRA MALAYSIA BIOLOGICAL STUDIES ON … filesedimen, dahan dan daun adalah 1.50 & 0.05 mg/g, 5.62 & 0.14 mglg and 22.79 + 1.08 mg/g. Kandungan nitrogen tanah didapati semakin

However, the mangrove area worldwide, including in Kuala Selangor, Malaysia are

decreasing naturally or anthropogenically. There are a number of factors leading to

the degradation of mangrove ecosystem, such as climate (Galloway, 1982), water

temperature (Galloway, 1982), tides (Chapman, 1976; Fields, 1995), wave action

(Fields, 1995), salinity (Chapman, 1976; Fields, 1995), soil composition (Chapman,

1976; Fields, 1999, accretion rate (Chapman, 1976), soil aeration (Chapman, 1976)

and animal biota (Chapman, 1976; Fields, 1995). With the growing population and

changing technology, there is increasing pressure to develop mangrove areas for

residential, commercial, agricultural and recreational purposes. For example, during

the Cultural Revolution of the 1960s large areas of mangroves were destroyed along

the coast of southern China and replaced by paddy fields for growing rice (Field,

1995). The reclaimed land proved unsuitable and today much of it lies derelict. In

Thailand and the Philippines large areas of mangroves have been converted to fish

and shrimp ponds for commercial production. In Kalimantan, mangroves have been

chipped for paper production. In other countries e.g. Thailand, mangroves have been

destroyed for mining, port construction and replacement agriculture.

Dam building and sewage dumping are example of activities that can cause the

increase of salinity seawards (Mastaller, 1997). Salinity play an important role on

the growth of mangroves (Chapman, 1976; Field, 1995; Mastaller, 1997). Plants

vary greatly in their tolerance to salinity (Hesse, 1971). The harmful effects of

excess salts upon plant growth is partly due to a specific effect of certain ions and

partly to the increase in osmotic pressure around the plant roots which inhibits water

uptake (Hesse, 1971). These will affect the seed germination, growth and

physiological parameter of the plants especially respiration, water balance and

3