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UNIVERSITI PUTRA MALAYSIA
BAHARI BIN MOHD MASARUDDIN
FP 2012 24
COMBINING ABILITY, HERITABILITY, HETEROSIS, AND STABILITY ANALYSES OF WATERMELON HYBRIDS
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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
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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
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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.
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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
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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
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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.
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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.
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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
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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:
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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:
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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
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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