UNIVERSITI PUTRA MALAYSIA

BAHARI BIN MOHD MASARUDDIN

FP 2012 24

COMBINING ABILITY, HERITABILITY, HETEROSIS, AND STABILITY ANALYSES OF WATERMELON HYBRIDS

© COPYRIG

HT UPM

COMBINING ABILITY, HERITABILITY, HETEROSIS, AND STABILITY ANALYSES OF WATERMELON HYBRIDS

By

BAHARI BIN MOHD MASARUDDIN

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

March 2012

© COPYRIG

HT UPM

ii

Abstract of thesis presented to the senate of University Putra Malaysia in fulfilment of the requirement for the degree of Master of Science

COMBINING ABILITY, HERITABILITY, HETEROSIS, AND

STABILITY ANALYSES OF WATERMELON HYBRIDS

By

BAHARI BIN MOHD MASARUDDIN

March 2012

Chairman: Associate Professor Mohd Rafii bin Yusop, PhD

Faculty: Agriculture

An evaluation on watermelon hybrids derived from a complete 4 x 4 diallel crosses

were conducted at two locations, MARDI Bukit Tangga Research Station, Kedah and

MARDI Seberang Perai Research Station, Penang with two planting seasons at each

location. Objectives of the study were to identify the superior and highly stable F1

watermelon hybrid for yield and fruit quality, to estimate the combining ability of

inbred lines in a complete diallel cross, to determine the genetic control and heritability

of important traits in the watermelon population and to quantify and determine the

level of heterosis revealed by the F1 hybrids. Results from evaluation across

environments of the watermelon genotypes showed that inbred line CS-19 had longest

vine length, thickest rind and heaviest fruit weight among all genotypes, whereas

inbred line BL-14 was earliest in flowering and inbred CH-8 was best for the earliest in

days to fruit maturity. Meanwhile hybrid CH-8 x CS-19 had the highest fruit yield per

plant compared to other genotypes. Analysis of combining ability among the four

watermelon inbred lines showed the presence of both additive and non-additive gene

effects in each environment as well as across environments. However, the additive

© COPYRIG

HT UPM

iii

gene effects are more important than non-additive gene effects since the mean squares

for general combining ability (GCA) were greater than the mean squares for specific

combining ability (SCA). Inbred line CS-19 was the best general combiner ability for

vine length, fruit yield, fruit weight and number of fruits, while inbred line BL-14 was

the best general combiner for days to flower and days to fruit maturity. Inbred line

6372-4 was best general combiner for flesh colour, while inbred line CH-8 was the

best general combiner for total soluble solid contents. The hybrids that showed

significant SCA effect were BL-14 x 6372-4 for rind thickness, fruit yield and fruit

weight, CS-19 x CH-8 for days to flower, flesh colour and fruit weight; CS-19 x 6372-

4 only for fruit weight and BL-14 x CH-8 only for flesh colour. The environment has a

significant influence on the magnitude of correlation coefficients for some of the traits

studied. In hybrid populations, fruit yield was highly correlated with vine length, rind

thickness and fruit weight at all environments. In general, the hybrids exhibited low to

moderate heterosis values for most of the traits. However none of the hybrids had

heterosis over better-parent for days to flower, days to fruit maturity and flesh colour

traits. Hybrid 6372-4 x BL-14 had moderate heterosis over mid and better-parent for

fruit yield. The estimates of broad- and narrow-sense heritability values were generally

low at each environment as well as across environments. Days to fruit maturity showed

moderate broad-sense (30%) and narrow-sense heritability (33%) estimates compared

to other traits. Through combined used of the five stability statistics, hybrid BL-14 x

6372-4 was identified to be highly stable for vine length, days to flower, days to fruit

maturity, total soluble solids content and fruit yield, while hybrid CS-19 x 6372-4 was

highly stable for fruit weight. These superior hybrids will be recommended for

commercial cultivation after multi-location trials and large scale evaluations have been

done in the future.

© COPYRIG

HT UPM

iv

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

KEUPAYAAN BERGABUNG, KEBOLEHWARISAN, HETEROSIS, DAN ANALISIS KESTABILAN KE ATAS TEMBIKAI HIBRID

Oleh

BAHARI BIN MOHD MASARUDDIN

Mac 2012

Pengerusi: Profesor Madya Mohd Rafii bin Yusop, PhD

Fakulti: Pertanian

Satu penilaian ke atas hibrid tembikai yang dihasilkan daripada kacukan diallel 4 x 4

lengkap dijalankan dalam dua musim penanaman di dua lokasi iaitu Stesen

Penyelidikan MARDI Bukit Tangga dan Stesen Penyelidikan MARDI Seberang Prai.

Objektif kajian ini adalah untuk mengenalpasti tembikai hibrid F1 yang stabil serta

berhasil dan berkualiti tinggi, menganggar keupayaan bergabung titisan tulen tersebut

di dalam kacukan diallel lengkap untuk menentu pengawalan genetik dan

kebolehwarisan bagi beberapa ciri yang penting dalam populasi tembikai tersebut dan

untuk menggangarkan tahap heterosis dalam hibrid F1. Hasil penilaian ke atas hibrid

tembikai di pelbagai persekitaran menunjukkan titisan tulen CS-19 mempunyai jalar

yang terpanjang, kulit yang paling tebal dan buah yang terberat berbanding di kalangan

semua genotip. Manakala titisan tulen BL-14 memberikan bilangan hari berbunga yang

terawal dan titisan tulen CH-8 pula menunjukkan bilangan hari buah matang yang

tersingkat. Sementara itu hibrid CH-8 x CS-19 menghasilkan hasil buah per pokok

yang tertinggi berbanding dengan genotip yang lain. Analisis keupayaan bergabung

© COPYRIG

HT UPM

v

dikalangan empat titisan tulen tembikai menunjukkan wujudnya kesan kedua-dua gen

aditif dan bukan-aditif di setiap persekitaran dan juga merentasi persekitaran.

Walaubagaimanapun kesan gen aditif adalah lebih penting berbandingan kesan gen

bukan aditif berdasarkan min kuasadua untuk keupayaan bergabung am (GCA) adalah

lebih besar berbanding mean kuasadua untuk keupayaan bergabung khusus (SCA).

Titisan tulen CS-19 memberikan keupayaan am terbaik untuk panjang jalar, hasil buah,

berat buah dan bilangan buah, manakala titisan tulen BL-14 untuk sifat bilangan hari

berbunga dan bilangan hari buah matang. Sementara itu titisan tulen 6372-4 adalah

terbaik untuk warna isi, dan titisan tulen CH-8 adalah keupayaan bergabung am terbaik

untuk jumlah pepejal terlarut. Hibrid yang menunjukkan kesan SCA yang bererti

adalah BL-14 x 6372-4 untuk hasil buah dan berat buah, CS-19 x CH-8 untuk bilangan

hari berbunga, berat buah dan warna isi, manakala CS-19 x 6372-4 untuk berat buah

dan BL-14 x CH-8 untuk warna isi. Persekitaran mengakibatkan pengaruh yang bererti

ke atas magnitud pekali korelasi untuk sebahagian ciri yang dikaji. Di dalam populasi

hibrid ini, hasil buah adalah sangat berkorelasi dengan panjang jalar, ketebalan kulit

dan berat buah di semua persekitaran. Secara umumnya hibrid tersebut memberikan

nilai heterosis yang rendah hingga sederhana bagi kebanyakan ciri-ciri tersebut.

Sementara itu tiada hibrid yang mempunyai nilai heterosis berdasarkan nilai induk

terbaik untuk bilangan hari berbunga betina, bilangan hari buah matang dan warna isi.

Walaubagaimanapun didapati hibrid 6372-4 x BL-14 mempunyai nilai sederhana bagi

heterosis berdasarkan purata dan induk terbaik untuk ciri hasil buah. Nilai

kebolehwarisan luas dan sempit, secara umumnya adalah rendah di setiap persekitaran

dan juga merentasi persekitaran. Ciri bilangan hari buah matang menunjukkan

kebolehwarisan luas (30%) dan sempit (33%) yang sederhana berbanding dengan ciri-

ciri yang lain. Menerusi penggunaan kombinasi lima kaedah statistik kestabilan, hibrid

© COPYRIG

HT UPM

vi

BL-14 x 6372-4 dikenalpasti sangat stabil untuk panjang jalar, bilangan hari berbunga,

bilangan hari buah matang, jumlah pepejal terlarut dan hasil buah, manakala hibrid

CS-19 x 6372-4 pula untuk berat buah. Hibrid yang unggul ini adalah disyorkan untuk

ditanam secara komersial setelah penilaian berskala besar dijalankan di masa akan

datang.

© COPYRIG

HT UPM

vii

ACKNOWLEDGEMENTS

In the name of Allah the Most Gracious and Merciful

This study was made possible by the generous financial and administrative support of

the Malaysian Agriculture Research and Development Institute (MARDI), University

Putra Malaysia (UPM) and the Ministry of Agriculture and Agro-based Industries

(MOA). In the process of completing this research and my Master of Science degree, I

have benefited from the expertise and guidance of my chairman of my supervisory

committee, Associate Professor Dr Mohd Rafii Yusop, who has been an invaluable

adviser and counselor. His encouragement and moral support made it all possible for

me to complete my work here in UPM. I am also grateful for comments and

suggestions from my committee member, Professor Dr Ghizan Saleh. I would also like

to express my gratitude to all the staff members of MARDI Bukit Tangga Station and

MARDI Seberang Prai Station for helping me conducting the research.

Finally, last but not least, thanks to my parents (Mohd Masaruddin), my lovely wife

(Salina Husain), children (Nur Farhana, Muhamad Ikhram Nabil, Nurfarina Nabilah

and Muhamad Danish Hakim) and the rest of the family and friends who have assisted

me throughout the duration of this research.

© COPYRIG

HT UPM

viii

I certify that an Examination Committee has met on 12 March, 2012 to conduct the

final examination of Bahari bin Mohd Masaruddin on his Master of Science thesis

entitled “Combining Ability, Heritability, Heterosis and Stability of Watermelon

Hybrids” in accordance with the Universities and University College Act 1971 and

Constitution of the Universiti Putra Malaysia [P.U. (A) 106] 15 March 1998. The

Committee recommends that the student be awarded the Master of Science.

Members of the Thesis Examination Committee were as follows:

Mahmud Tengku Muda Mohamed, PhD Professor Department of Crop Science Faculty of Agriculture Universiti Putra Malaysia (Chairman) Halimi Mohd Saud, PhD Associate Professor Department of Agriculture Technology Faculty of Agriculture Universiti Putra Malaysia (Internal Examiner) Faridah Qamaruz Zaman, PhD Associate Professor Department of Biology Faculty of Science Universiti Putra Malaysia (Internal Examiner) Mariam Abd Latif, PhD Associate Professor Datin School of Food Science and Nutrition Universiti Malaysia Sabah (External Examiner)

SEOW HENG FONG, PhD Professor and Deputy Dean School of Graduate Studies Universiti Putra Malaysia

Date: 23 April 2012

© COPYRIG

HT UPM

ix

This thesis was 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 were as follows:

Mohd Rafii Yusop, PhD Associate Professor Department of Crop Science Faculty of Agriculture Universiti Putra Malaysia (Chairman)

Ghizan Saleh, PhD Professor Department of Crop Science Faculty of Agriculture Universiti Putra Malaysia (Member)

BUJANG BIN KIM HUAT, PhD Professor and Dean

School of Graduate Studies Universiti Putra Malaysia Date:

© COPYRIG

HT UPM

x

DECLARATION

I declare that the thesis is my original work except for quotations and citations which have been duly acknowledged. I also declare that it has not been previously, and is not concurrently, submitted for any other degree at University Putra Malaysia or at any other institution.

BAHARI MOHD MASARUDIN

Date:

© COPYRIG

HT UPM

xi

TABLE OF CONTENTS PAGE ABSTRACT ii ABSTRAK iv ACKNOWLEDGEMENT vii APPROVAL viii DECLARATION x LIST OF TABLES xiii LIST OF FIGURES xviii LIST OF APPENDICES xix LIST OF ABBREVIATIONS xx

CHAPTER

1 INTRODUCTION 1

2 LITERATURE REVIEW 5 2.1 Origin and history of watermelon 5 2.2 Taxonomy of watermelon 5 2.3 Germplasm of watermelon 6 2.4 Morphology and botany of watermelon 7 2.5 Fruit quality characteristics of watermelon 9 2.6 Environmental conditions for growing watermelon 10 2.7 Production of watermelon 11 2.8 Uses and nutritional content of watermelon 12 2.9 Watermelon breeding in Malaysia 14 2.10 Diallel analysis 15 2.11 Combining ability 16 2.12 Genetic variance and heritability 18 2.13 Heterosis 20 2.14 F1 hybrids 21 2.15 Correlations among traits 23 2.16 Genotype x environment interaction and stability statistics 23

3 MATERIAL AND METHODS 27 3.1 Locations of experiment 27 3.2 Plant materials 27 3.2.1 Parental lines 27 3.2.2 Development of F1 hybrids by diallel cross 28 3.3 Experimental design 29 3.4 Field planting 29 3.5 Data collection 30 3.6 Data analysis 31 3.6.1 Analysis of variance 32 3.6.2 Combining ability 34 3.6.3 GCA:SCA ratio 35 3.6.4 Heritability 36 3.6.5 Heterosis 37 3.6.6 Correlations among traits 38

© COPYRIG

HT UPM

xii

3.6.7 Stability analysis 38

4 RESULTS AND DISCUSSION 43 4.1 Performance of F1 progenies 43 4.1.1 Analysis of variance in individual and across environment 43 4.1.2 Environment means 46 4.1.3 Genotype means in individual and across environments 49 4.1.4 Relationships among traits 62 4.2 Combining ability analysis 67 4.2.1 Analysis of variance of GCA and SCA 67 4.2.2 GCA and SCA variance components 88 4.3 Heritability 90 4.3.1 Broad-sense heritability 90 4.3.2 Narrow-sense heritability 91 4.4 Heterosis 92 4.4.1 Heterotic response in individual environment 93 4.4.2 Heterotic response across environments 101 4.5 G x E and stability analysis 106 4.5.1 Mean against CV 106 4.5.2 Variety superiority measure (Pi) 113 4.5.3 Stability variance (σ²i) 116 4.5.4. Rank sum method 120 4.5.5 Non parametric ranking (Si

3) 128 4.5.6 Overall performance of genotypes based on various ranking methods 138

5 GENERAL DISCUSSION 147 5.1 Performance of watermelon genotypes 147

5.2 General and specific combining ability effects 148 5.3 Heritability 151

5.4 Heterosis 152 5.5 G x E interaction and stability analysis 153

6 CONCLUSION 155

REFFERENCES 158

APPENDICES 171

BIODATA OF STUDENT 177

LIST OF PUBLICATIONS 178