universiti putra malaysia weak neck problem in …psasir.upm.edu.my/id/eprint/27009/1/fp 2011...

UNIVERSITI PUTRA MALAYSIA

WEAK NECK PROBLEM IN MUSA SP. CV. RASTALI POPULATIONS IN RELATION TO MAGNESIUM,

BORON AND SILICON AVAILABILITY

EKA TARWACA SUSILA PUTRA

FP 2011 34

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WEAK NECK PROBLEM IN MUSA SP. CV.

RASTALI POPULATIONS IN RELATION TO

MAGNESIUM, BORON AND SILICON

AVAILABILITY


DOCTOR OF PHILOSOPHY

UNIVERSITI PUTRA MALAYSIA

2011

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WEAK NECK PROBLEM IN MUSA SP. CV. RASTALI

POPULATIONS IN RELATION TO MAGNESIUM,

BORON AND SILICON AVAILABILITY

BY


Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia,

in Fulfillment of the Requirements for the Degree of Doctor of Philosophy

April 2011

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DEDICATION

I would like to dedicate my thesis to:

My beloved parents,

Sapardi

Marsih

For giving me do’a, support, sacrifices and encouragement

and

My dearest wife and sons,

Atin

Khadafi

Ilham

for their do’a, motivation, inspiration and love

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Abstract of thesis presented to the Senate of Universiti Putra Malaysia in

fulfillment of the requirement for the degree of Doctor of Philosophy

WEAK NECK PROBLEM IN MUSA SP. CV. RASTALI POPULATIONS

IN RELATION TO MAGNESIUM, BORON AND SILICON

AVAILABILITY

By


April 2011

Chairman : Assoc. Prof. Zakaria bin Wahab, PhD

Faculty : Agriculture

Weak neck is the most significant physiological damage in Musa sp. cv. Rastali

accessions. It has the ability to reduce fruit quality and selling value. A holistic

approach is needed to solve the weak neck problem. The objectives of the

research were: (1) to determine the genetic diversity of the Rastali population

based on morphological and molecular markers, (2) to identify the specific allele

that is associated with weak neck in the Rastali populations in comparison to other

cultivars, and (3) to elucidate the relationship between fertilizer recommendation

containing Mg, B and Si and the weak neck problem of the Rastali population.

A total of 78 accessions of Rastali (74 accessions of Rastali from Peninsular

Malaysia and 4 accessions from Central Java, Indonesia), and six different

cultivars with no occurrence of weak neck were selected for comparison. Out of

these, only 29 accessions were able to be morphologically characterized because

many of them were without reproductive organ. A total of eight qualitative

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reproductive descriptors were used in this experiment. These 78 accessions were

used for the molecular study. Genomic DNA were extracted from leaf samples

using GeneAll® Plant DNA extraction kit and quantified using a

spectrophotometer. Polymerase chain reaction (PCR) was performed in a total

volume of 15µL. A total of 10 SSR primer pairs were used in this study.

Meanwhile, forward primers were labelled with FAM fluorescent label. The

detection of the DNA fragments was made on ABI Prism® 310 automated

sequencer and the allele sizes were also obtained.

The field experiment was done at Universiti Putra Malaysia under field conditions

arranged in split-split plot design with five blocks. The main factor was NPK

fertilizer which was recommended by Department of Agriculture Malaysia (DOA)

and United Plantation Berhad (UPB). The sub-factors were applications and non-

application (as control) of kieserite (Magnesium), boric acid (Boron) and sodium

silicate (Silicon). The sub-sub factors were the Musa sp. cv. Rastali accessions,

consisting of four groups, which were chosen for their different morphological

characters. In particular, the Musa sp. cv. Berangan accession was selected as the

control treatment.

The findings showed that there is genetic diversity among Rastali accessions.

Two male reproductive morphological descriptors (namely, male neuter flower

along the rachis and male bud shape) were found to be the most important

morphological characters which have caused the variation among the Rastali

accessions. Existence of variations among Rastali was further supported by

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genetic analysis using 10 SSR markers. The SSRs markers were able to separate

the accessions of Rastali. A significant 39% of genetic diversity was found

among 78 accessions of Rastali. Out of the 78 Rastali accessions, eight were

excluded from the main cluster, and showed higher heterogeneity compared to

those in the main group (the coefficients of similarity ranging from 0.13 to 0.24).

In the selection of the overall allele at each Rastali accession, one allele with the

size of 238 bp, was found to be always present in all the accessions of Rastali, but

was absent in the other six cultivars. This allele was located at locus A-25 and

was considered the most common allele. Meanwhile, the allele which was located

on locus A-13 and with the size of 277bp was only found in the other six cultivars

but absent in all the Rastali accessions.

The findings also showed that combined application of magnesium, boron and

silicon could improve the Mg, B and Si contents in the leaves and fruit of the

Rastali accessions and Berangan, in combination with the NPK fertilizer doses

from DOA or UPB. These conditions increased the stomatal length and width and

physiological activities of the Rastali accessions and Berangan at low level of

NPK fertilizer (UPB), in combination with Mg, B and Si applications. In

particular, the fingers of the R08 (in combination with the NPK fertilizer doses

from UPB) and R62 (in combination with the NPK fertilizer doses from DOA)

which had received Mg, B and Si were found to be lighter and smaller than those

without any Mg, B and Si applications. Meanwhile, the bunches of the R08, R62

and Berangan which had received Mg, B and Si were found to be lighter and

smaller than those without any Mg, B and Si applications, in combination with the

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NPK fertilizer doses from DOA. These characters have been shown to have the

potential in reducing the intensity of the weak neck problem in the Rastali

accessions, especially for the R08 and R62.

Although the applications of magnesium, boron and silicon enhanced stomatal

length and width and physiological activities of the Rastali accessions and

Berangan at low level of NPK fertilizer (UPB), they decreased polygalacturonase

(PG) and pectinmethylesterase (PME) activities in the Rastali accessions, except

the PG on Day 4 after harvesting for the R08 and R62. Nevertheless, the PG and

PME activities in Rastali were still higher compared to those of Berangan. The

applications of Mg, B and Si also increased the lignin and cellulose content in the

fruit neck zone of the R34 (in combination with the NPK fertilizer doses from

DOA) and R08 (in combination with the NPK fertilizer doses from UPB). The

decline in the PG and PME activities, as well as the increase in the lignin and

cellulose content in the R08 (in combination with the NPK fertilizer doses from

UPB) and R34 (in combination with the NPK fertilizer doses from DOA) which

had received Mg, B and Si were able to increase the relative thickness of the cell

walls during fruit growth. Application of Mg, B and Si resulted in small-sized

cell but relatively thicker cell wall. Meanwhile, the applications of Mg, B and Si

decreased the ethylene activity in the R08 (in combination with the NPK fertilizer

doses from DOA or UPB), R34 and R12 (in combination with the NPK fertilizer

doses from UPB).

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The decrease in ethylene, PG and PME activities delayed the accumulation of

soluble soilids content (SSC) in the R08 (in combination with the NPK fertilizer

doses from DOA) and chlorophyll degradation in the R12, which had obtained

Mg, B and Si. The slowing down of chlorophyll degradation and the lowering of

SSC resulted in maintained finger firmness of the fruit at a high level. Thus, the

fruit has been found to remain strong and hard, especially in the abscission layer,

although it has been ripened. The R08 and R12 which had obtained Mg, B and Si

were able to produce fruit with stronger abscission layer (higher firmness), in

combination with the NPK fertilizer doses from DOA. The stronger abscission

layer is not easy to crack and it is also not prone to weak neck. Therefore, the

applications of Mg, B and Si in combination with the NPK fertilizer doses from

DOA have the potential to reduce the weak neck intensity in the R08, although

totally eliminating the weak neck problem as in Berangan has yet to be achieved.

Based on the data, it can therefore be concluded that (1) Rastali cultivar has weak

neck problem, (2) there are early indications that the 238bp and 277bp alleles

have the possibilities to be associated with weak neck, and (3) the combined

applications of Mg, B and Si was able to reduce the weak neck intensity from

80.00% to 24.33% in the Rastali accessions and Berangan, in combination with

the NPK fertilizer doses from DOA.

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Abstrak Tesis yang dikemukakan kepada senat Universiti Putra Malaysia sebagai

memenuhi keperluan untuk ijazah Doktor Falsafah

KEPERLUAN MAGNESIUM, BORON DAN SILIKON DAN

HUBUNGANNYA DALAM MASALAH LERAI BUAH

PADA POPULASI MUSA SP. CV. RASTALI

Oleh


April 2011

Pengerusi : Prof. Madya Zakaria bin Wahab, PhD

Fakulti : Pertanian

Lerai buah merupakan kerosakan fisiologi yang paling banyak didapati pada Musa

sp. cv. Rastali. Ia mampu mengurangkan kualiti dan harga pasaran buah.

Pendekatan holistik diperlukan dalam menyelesaikan masalah lerai buah.

Objektif kajian ini adalah untuk: (1) menentukan kepelbagaian genetik populasi

Rastali berdasarkan morfologi dan penanda molekul (2) mengenalpasti alel khusus

yang terdapat pada populasi Rastali berbanding kultivar yang lain dan (3)

menentukan hubungan di antara baja yang disyorkan yang mengandungi

magnesium (Mg), boron (B), dan silikon (Si) terhadap lerai buah pada populasi

Rastali.

Sejumlah 78 aksesi Rastali (74 aksesi Rastali dari Semenanjung Malaysia dan 4

aksesi dari Jawa Tengah, Indonesia) dan enam kultivar berbeza yang tidak

mempunyai lerai buah dipilih untuk perbandingan. Daripada jumlah ini, hanya

29 aksesi sahaja yang dapat dicirikan morfologinya kerana ketiadaan organ

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reproduktif pada kebanyakan kultivar yang mengalami lerai buah. Sebanyak

lapan nilai kualitatif untuk ciri-ciri reproduktif digunakan dalam eksperimen ini.

Untuk kajian molekular, 78 aksesi telah digunakan. DNA diekstrak daripada daun

menggunakan GeneAll®

Plant DNA extraction kit dan kuantitinya diukur dengan

menggunakan spektrofotometer. Reaksi Rantaian Polimerase dihasilkan dalam

jumlah 15µL. Sepuluh primer SSR digunakan untuk kajian ini. Primer kehadapan

dilabel menggunakan FAM fluorescent. Serpihan bahagian DNA dikenalpasti

menggunakan ABI Prism®

310 automated sequencer dan saiz alel diperolehi.

Kajian lapangan telah dijalankan di Universiti Putra Malaysia, yang mana ia

disusun dalam plot yang dipisah-pisahkan dengan lima blok. Faktor utama adalah

baja NPK, iaitu yang disyorkan oleh Jabatan Pertanian Malaysia (DOA) dan

United Plantation Berhad (UPB). Faktor-faktor sampingan adalah diaplikasikan

dan tidak diaplikasikan (sebagai pengawal) kieserite (Mg), asid borik (B) dan

natrium silikat (Si). Faktor sampingan-sampingan bagi aksesi Rastali,

mengandungi empat kumpulan yang dipilih untuk ciri-ciri morfologi yang

berbeza. Musa sp. cv. Berangan dipilih sebagai rawatan kawalan.

Hasil kajian menunjukkan terdapat kepelbagaian genetik pada aksesi Rastali. Dua

morfologi reproduktif jantan (iaitu bunga jantan mandul sepanjang rakis dan

bentuk kudup jantan) merupakan ciri morfologi yang paling penting yang

menyebabkan kepelbagaian di antara aksesi Rastali. Perbezaan yang wujud di

antara Rastali kemudian disokong oleh analisis genetik menggunakan 10 penanda

SSR mampu mengasingkan aksesi Rastali. Kepelbagaian genetik yang signifikan

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dijumpai di antara 78 aksesi Rastali. Daripada 78 aksesi Rastali, 8 aksesi

terkeluar daripada kumpulan utama dan menunjukkan heteroginiti yang tinggi

berbanding yang terdapat di kumpulan utama (skala persamaan koefisien bermula

0.13 hingga 0.24). Keputusan ini juga menunjukkan bahawa dalam pemilihan

daripada keseluruhan aksesi Rastali, satu alel yang bersaiz 238bp ditemui sentiasa

ada dalam keseluruhan aksesi Rastali, tetapi tiada dalam enam kultivar yang lain.

Alel ini berada pada lokus A-25 dan boleh dipertimbangkan sebagai alel yang

paling biasa. Sementara itu, alel yang bersaiz 277 bp yang berada pada lokus A-

13, hanya ditemui pada enam kultivar yang lain tetapi tiada pada semua aksesi

Rastali.

Hasil penyelidikan ini juga menunjukkan gabungan aplikasi Mg, B dan Si dapat

meningkatkan kandungan Mg, B dan Si di dalam daun dan buah pada aksesi

Rastali dan Berangan, dengan kombinasi baja NPK dos daripada DOA dan UPB.

Keadaan ini meningkatkan panjang dan lebar stomata dan aktiviti fisiologi aksesi

Rastali dan Berangan pada tahap rendah baja NPK (UPB), dengan kombinasi

aplikasi Mg, B dan Si. Secara khusus, buah R08 (dalam kombinasi dengan baja

NPK dos dari UPB) dan R62 (dalam kombinasi dengan baja NPK dos dari DOA)

yang telah menerima Mg, B dan Si didapati lebih ringan dan lebih kecil

berbanding dengan yang tidak diaplikasikan dengan Mg, B, and Si. Sementara

itu, tandan bagi R08, R62 dan Berangan yang telah mendapat Mg, B dan Si

didapati ringan dan kecil berbanding dengan yang tidak diaplikasikan dengan Mg,

B and Si, dengan kombinasi baja NPK dos daripada DOA. Ciri ini mempunyai

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potensi untuk mengurangkan intensiti masalah lerai buah pada aksesi Rastali,

terutamanya untuk R08 dan R62.

Walaupun aplikasi Mg, B dan Si dapat meningkatkan panjang dan lebar stomata

serta aktiviti fisiologi pada aksesi Rastali dan Berangan pada kadar rendah baja

NPK (UPB), ia akan mengurangkan aktiviti poligalakturonase (PG) dan

pectinmethylesterase (PME) dalam aksesi Rastali, kecuali PG pada hari ke 4

selepas dituai untuk R08 dan R62. Bagaimanapun, aktiviti PG dan PME dalam

Rastali masih lebih tinggi berbanding dengan Berangan. Aplikasi Mg, B, dan Si

juga dapat meningkatkan kandungan lignin dan selulosa di zon leher buah R34

(dalam kombinasi dengan baja NPK dos daripada DOA ) dan R08 (dalam

kombinasi dengan baja NPK dos daripada UPB). Penurunan aktiviti PG dan

PME, serta peningkatan lignin dan selulosa dalam R08 (dalam kombinasi dengan

baja NPK dos daripada UPB) dan R34 (dalam kombinasi dengan baja NPK dos

daripada DOA ) yang menerima Mg, B dan Si mampu meningkatkan ketebalan

relatif dinding sel semasa pertumbuhan buah. Aplikasi Mg, B dan Si

menghasilkan saiz sel yang kecil tetapi dinding sel yang lebih menebal.

Sementara itu, aplikasi Mg, B dan Si menurunkan aktiviti etilena pada R08 (dalam

kombinasi dengan baja NPK dos daripada DOA atau UPB), R34 dan R12 (dalam

kombinasi dengan baja NPK dos daripada UPB).

Penurunan aktiviti etilena, PG dan PME akan menangguhkan pengumpulan

kandungan pepejal boleh larut (SSC) pada R08 (dalam kombinasi dengan baja

NPK dos daripada DOA) dan degradasi klorofil pada R12, yang telah menerima

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Mg, B dan Si. Perlambatan degradasi klorofil dan penurunan SSC akan

mengekalkan ketegasan buah pada peringkat tinggi. Oleh itu, buah akan didapati

tetap kuat dan keras, terutamanya pada lapisan luruhan, walaupun buah telah

masak. R08 dan R12 yang mendapat Mg, B dan Si mampu menghasilkan buah

dengan lapisan luruhan yang lebih kuat (ketegasan lebih tinggi), dalam kombinasi

dengan baja NPK dos daripada DOA. Lapisan luruhan yang kuat tidak mudah

retak dan tidak mudah terdedah kepada lerai buah. Oleh sebab itu, aplikasi Mg, B

dan Si dengan kombinasi baja NPK dos daripada DOA mempunyai potensi untuk

menurunkan intensiti lerai buah pada R08, walaupun secara keseluruhannya

penyelesaian masalah lerai buah seperti pada Berangan masih belum dapat

dicapai.

Berdasarkan data tersebut, dapatlah disimpulkan bahawa (1) kultivar Rastali

mempunyai masalah lerai buah, (2) terdapat penunjuk awal bahawa alel 238bp

dan 277bp berkemungkinan berkaitan kepada masalah lerai buah, (3) kombinasi

aplikasi Mg, B dan Si mampu mengurangan intensiti masalah lerai buah dari

80.00% kepada 24.33% pada aksesi Rastali dan Berangan, dalam kombinasi

dengan baja NPK dos daripada DOA.

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ACKNOWLEDGEMENTS

Bismillahi Allahu Akbar and Alhamdulillahirabbil’alamin, All praise to Allah

S.W.T. whose countless blessing enabled me to accomplish this study. My

sholawat and salam is addressed to His righteous messenger, prophet Muhammad

S.A.W.

First of all I would like to express my most sincere gratitude to Assoc. Prof. Dr.

Zakaria Wahab for his suggestions, important advice, unflagging interest,

concerns, inspiration and encouragement during the period of this study. His

broad knowledge and logical way of thinking have been of great value to me. His

understanding, assistance, encouragement and guidance have provided a good

basis for my thesis. I gratefully acknowledge my indebtedness to my supervisory

committee members, Prof. Dr. Ghizan Saleh and Dr. Nur Ashikin Psyquay

Abdullah, who provide many valuable comments and suggestions, which

significantly improved the thesis.

I take this opportunity to express my thanks to all the farmers in Peninsular

Malaysia and Central Java, Indonesia for permission given to collect the samples

from their fields. The appreciation is also due to the Faculty of Agriculture,

Universitas Gadjah Mada for its approval to pursue my PhD program at Faculty of

Agriculture, Universiti Putra Malaysia. I owe my sincere gratitude to Prof. Dr.

Didik Indradewa and Dr. Endang Sulistyaningsih for giving me support and

motivation during my study. Special thanks are due to my mentors Mr. Priyono

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Suryanto (PhD candidate) and Mr. Abdul Rohman (PhD candidate), for giving me

constant encouragement, motivation, inspiration and support. I would like to

thank Dr Sri Rahayu and Dr. Tata Wijayanta, who teach me the hard life. Special

thanks to Dr. Usman Jakfar and ikhwahfillah for their support and kindness.

My keen appreciation goes to Tuan Haji Mohd Shahril Ab. Rahman for his

valuable assistance in the field. Sincere thanks are extended to Encik Daud

Mustam and Tuan Haji Suhaimi Aman for their assistance and kindness to allow

me to use the SEM in their laboratory. A special thank to Encik Mazlan Bangi for

his assistance and kindness to allow me to use Photosynthetic Analyzer,

Porometer and SPAD 502. Not to forget Tuan Haji Mohd Khairi Kandar and

Encik Yusoff Suki for their assistance to determine the nutrient content using

AAS and AA in their laboratory. Sincere thanks to Encik Shamsudin Bojang for

his kindness to allow me to use the CPD in his laboratory. I would like to thank

Puan Salmi Yaacob, Puan Salmah Kassim and Encik Mohd Helmi Hamisan, for

providing chemicals, valuable information and assistance during my laboratory

work. I would like to thank my special friends, Rahmatollah Behmaram, Ali

Rajbarfard, Mahmood Reza, Majid Faroughi, Alireza Biabani, Zahra Nouri,

Elnaz, Nahid Babaei and Pedram Kashiani. They gave me friendship and provide

useful discussions to improve my thesis.

I would like to express my profound appreciation to my parents, Sapardi and

Marsih, for their do’a, support, sacrifices and encouragement. I would also like to

express my grateful appreciation to my father and mother-in-law, Sugiran and

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Sukemi, for their do’a and support. Last but not least, I would like to thank to my

beloved wife, Suprihatin Wijayanti, S.P., and my little heroes, Muammar Iqbal

Khadafi Tarwaca and Muammar Ilham Hanafi Tarwaca for their sacrifices,

patience and moral support that enable me to successfully carry out the PhD

study.

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I certify that a Thesis Examination Committee has met on 27 April 2011 to

conduct the final examination of Eka Tarwaca Susila Putra on his thesis entitled

"Weak Neck Problem in Musa Sp. Cv. Rastali Populations in Relation to

Magnesium, Boron and Silicon Availability" in accordance with the Universities

and University College Act 1971 and the Constitution of the Universiti Putra

Malaysia [P.U.(A) 106] 15 March 1998. The Committee recommends that the

student be awarded the Doctor of Philosophy.

Members of the Thesis Examination Committee were as follows:

Mohd Ridzwan bin Abd Halim, PhD

Associate Professor

Faculty of Agriculture

Universiti Putra Malaysia

(Chairman)

Siti Aishah binti Hassan, PhD

Associate Professor



(Internal Examiner)

Mohd Rafii bin Yusop, PhD

Associate Professor



(Internal Examiner)

Thomas C. Kaspar, PhD

Professor

United States Department of Agriculture - Agricultural Research Service

United States of America

(External Examiner)

_______________________________

NORITAH OMAR, PhD

Associate Professor and Deputy Dean

School of Graduate Studies


Date: 26 July 2011

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This thesis was submitted to the Senate of Universiti Putra Malaysia and has been

accepted as fulfillment of the requirement for the degree of Doctor of Philosophy.

The members of the Supervisory Committee were as follows:

Zakaria bin Wahab, PhD

Associate Professor



(Chairman)

Ghizan bin Saleh, PhD

Professor



(Member)

Nur Ashikin Psyquay Abdullah, PhD

Senior Lecturer



(Member)

________________________________

HASANAH MOHD GHAZALI, PhD

Professor and Dean

School of Graduate Studies


Date:

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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 has not

been previously or concurrently submitted for any other degree at UPM or other

institution.

_______________________________


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TABLE OF CONTENTS

Page

DEDICATION ii

ABSTRACT iii

ABSTRAK viii

ACKNOWLEDGEMENTS xiii

APPROVAL xvii

DECLARATION xviii

TABLE OF CONTENTS xix

LIST OF TABLES xxiii

LIST OF FIGURES xxvii

LIST OF ABBREVIATIONS xxix

CHAPTER

1. INTRODUCTION 1

2. LITERATURE REVIEW 6

2.1 Musa sp. cv. Rastali 6

2.2 Fruit Growth and Development 7

2.2.1 The Role of Stomata in the Growth and Development of Fruit 8

2.2.2 The Roles of Photosynthesis and Transpiration in the

Growth and Development of Fruit 9

2.3 Weak Neck Problem in Rastali 11

2.4 Genetic Diversity among Rastali Accessions and the Relationship

with Weak Neck Problem 15

2.5 Crop Nutrition Management and Possibilities to Overcome

Weak Neck Problem in Rastali 20

2.5.1 The Role of Magnesium 23

2.5.2 The Role of Boron 25

2.5.3 The Role of Silicon 28

2.6 Physiochemical Changes in Rastali Accessions during Ripening in

Relation to Weak Neck Problem 32

2.6.1 Physiochemical Changes during Ripening 32

2.6.2 The Role of Polygalacturonase and Pectin Methylesterase

in Fruit Cell Degradation 33

2.6.3 The Role of Ethylene in Fruit Cell Degradation and Ripening 34

3. CELL ULTRASTRUCTURE AND PEEL NUTRIENT CONTENT OF

NECK ZONE IN SIX CULTIVARS OF MUSA SPP. FRUIT DURING

RIPENING 38

3.1 Introduction 38

3.2 Materials and Methods 41

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3.2.1 Location 41

3.2.2 Plant Materials 41

3.2.3 Cell Morphology 43

3.2.4 Nutrient Content Analysis in Fruit Peel 43

3.2.5 Data Analysis 44

3.3 Results 44

3.3.1 Cell Morphology Before and After Ripening 44

3.3.2 Nutrient Content in the Fruit Peel 47

3.4 Discussion 50

3.5 Conclusion 52

4. MORPHOLOGICAL VARIATION AND GEOGRAPHICAL

DISTRIBUTION OF MUSA SP. CV. RASTALI IN PENINSULAR

MALAYSIA 53

4.1 Introduction 53



4.2.2 Nutrient Content Analysis 55

4.2.3 Morphological Descriptors 56


4.3 Results 57

4.3.1 Climatic Conditions 58

4.3.2 Nutrient Contents 58

4.3.3 Principal Component Analysis 59

4.3.4 Cluster Analysis 65

4.3.5 Geographical Distribution of Rastali Accessions 65

4.4 Discussion 68

4.5 Conclusion 72

5. GENETIC VARIATION AMONG MUSA SP. CV. RASTALI

ACCESSIONS AND ITS ASSOCIATION WITH A SINGLE

MICROSATELLITE ALLELE 73

5.1 Introduction 73



5.2.2 Molecular Method 76


5.3 Results 77

5.4 Discussion 86

5.5 Conclusion 88

6. PHYSIOLOGICAL ACTIVITIES, STOMATAL MORPHOLOGY,

NUTRIENT CONTENT, GROWTH AND YIELD OF MUSA SP. CV.

RASTALI IN RELATION TO MAGNESIUM, BORON AND SILICON

AVAILABILITY 89

6.1 Introduction 89



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6.2.2 Methodology 96

6.2.3 Observations 97

6.2.3.1 Physiological Activities 97

6.2.3.2 Stomatal Morphology 98

6.2.3.3 Nutrient Content 101

6.2.3.4 Growth 102

6.2.3.5 Yield 102


6.3 Results 104

6.3.1 The Physiological Activities at 5 Months after Planting 104

6.3.2 Physiological Activities at 7 Months after Planting 109

6.3.3 Stomatal Morphology at Flowering 116

6.3.4 Stomatal Conductance at Flowering 117

6.3.5 Transpiration Rate at Flowering 118

6.3.6 Chlorophyll a and b Content and Relative Chlorophyll Content

at Flowering 128

6.3.7 Photosynthetic Rate at Flowering 129

6.3.8 Stomatal Conductance and Transpiration Rate at Fruit

Growth Stage 130

6.3.9 Nutrient Content, Growth and Yield 135

6.4 Discussion 151

6.4.1 Physiological Activities at 5 and 7 Months after Planting 151

6.4.2 Stomatal Morphology, Conductance and Transpiration Rate

at Flowering 155

6.4.3 Relative Chlorophyll Content, Chlorophyll Contents and

Photosynthetic Rate at Flowering 161

6.4.4 Stomatal Conductance and Transpiration Rate at Fruit

Growth Stage 165

6.4.5 Nutrient Content, Growth and Yield 166

6.5 Conclusion 171

7. CELL DEVELOPMENT, ENZYME AND HORMONE

ACTIVITIES AND WEAK NECK IN MUSA SP. CV. RASTALI IN

RELATION TO MAGNESIUM, BORON AND SILICON

AVAILABILITY 173

7.1 Introduction 173



7.2.2 Methodology 181

7.2.3 Observations 181

7.2.3.1 Cell Growth and Degradation 181

7.2.3.2 Enzyme Activities 182

7.2.3.3 Fiber Content 183

7.2.3.4 Ethylene Activity 184

7.2.3.5 Fruit Firmness 185

7.2.3.6 Soluble Solids Content (SSC) 185

7.2.3.7 Peel Colour 186

7.2.3.8 Weak Neck Intensity 187

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7.3 Results 188

7.3.1 Cell Characteristics during Fruit Growth and Ripening 188

7.3.2 Polygalacturonase dan Pectin Methylesterase Activities during

Fruit Ripening 205

7.3.3 Fiber Content in the Fruit Neck Zone 208

7.3.4 Ethylene Production 213

7.3.5 Soluble Solids Content 214

7.3.6 Finger and Neck Zone Firmness 218

7.3.7 Fruit Peel Colour 223

7.3.8 Weak Neck during Fruit Ripening 226

7.4 Discussion 229

7.5 Conclusion 240

8. SUMMARY, GENERAL CONCLUSION AND

RECOMMENDATIONS 242

REFERENCES 249

APPENDIX 275

BIODATA OF STUDENT 285

LIST OF PUBLICATION 286

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