UNIVERSITI PUTRA MALAYSIA - core.ac.uk · menguji sama ada rumput guinea mempunyai kesan alelopati pada kekacang menunjukkan bahawa panjang tumbuhan stylo menurun secara bererti (P

UNIVERSITI PUTRA MALAYSIA

COMPATIBILITY OF FOUR FORAGE LEGUMES WITH GUINEA GRASS (Panicum maximum Jacq.)

MOHAMMED BABA

FP 2013 38

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COMPATIBILITY OF FOUR FORAGE LEGUMES WITH GUINEA GRASS

(Panicum maximum Jacq.)

By

MOHAMMED BABA

Thesis submitted to the School of Graduate Studies,

Universiti Putra Malaysia, in Fulfilment of the Requirements for the

Degree of Doctor of Philosophy

December 2013

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COPYRIGHT

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photographs and all other artwork, is copyright material of University Putra Malaysia

unless otherwise stated. Use may be made of any material contained within the thesis for

non-commercial purposes from the copyright holder. Commercial use of material may

only be made with the express, prior, written permission of University Putra Malaysia.

Copyright © Universiti Putra Malaysia

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DEDICATION

To my loving parents, wife and daughters

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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

COMPATIBILITY OF FOUR FORAGE LEGUMES WITH GUINEA GRASS

(Panicum maximum) JACQ

By

MOHAMMED BABA

December 2013

Chairman : Associate Professor Mohd Ridzwan Abd Halim, PhD

Faculty : Agriculture

Livestock production in the tropics is constrained by decline in quantity and quality of

the predominant tropical grasses during the growing season. Incorporation of forage

legumes into grassland agriculture has been used as a panacea to mitigate the problem.

However, in tropical regions, not much of success has been achieved due to strong

competitive ability of the grass which reduces the content of the companion legume. It is

with this background that four experiments were conducted to evaluate the compatibility

of grass-legume mixtures and their suitability for plantation livestock integration.

An investigation into the nature of competition between Panicum maximum (guinea

grass) and each of the following legumes of contrasting growth habits; Stylosanthes

guianensis (stylo), Centrosema pubescens (centro) Macroptilium bracteatum (burgundy)

and Arachis pintoi (arachis) showed that dry matter yield in guinea-centro mixtures (2:2

and 3:1) species combination tended to be higher than the corresponding grass

monoculture (differences were not significant). Mean relative yield total values were

higher in guinea-stylo (1.41) and guinea-centro (1.4) the values seemed to be higher at

2:2 species combination except in the case of guinea-burgundy. The climbing/trailing

legume (centro) appeared to be more compatible with guinea grass especially at 2:2

species combination compared to the erect (stylo), erect/climbing (burgundy) and

prostrate (arachis) legumes.

Compatibility of forage mixtures evaluated at 50:50 species combination in relation to

dry matter yield and chemical composition indicates that guinea-arachis produced

significantly greater (P<0.01) cumulative total dry matter yield compared to other

mixtures. However, the yield was not significantly different from that of guinea

monoculture. The erect legume (stylo) produced significantly higher (P<0.01)

cumulative total dry matter yield than arachis in mixture with guinea. Similarly, relative

yield of stylo tended to be higher than those of arachis and burgundy. Thus as a

compromise between dry matter yield and pasture quality, grass-stylo mixture was

recommended.

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The productivity of four mixtures guinea-arachis, guinea-burgundy, guinea-centro and

guinea-stylo evaluated under 3 shade levels 0 (1701.9µmol m-2

s-1

), 50 (583.3 µmol m-2

s-1

) and 70% (520.2 µmol m-2

s-1

) indicates that guinea-centro tended to produce higher

yield among mixtures though differences were not significant. Centro in mixture with

guinea recorded significantly higher yield (P<0.05) than arachis and burgundy.

Interaction between legume species and shade levels on dry matter yield showed that the

dry matter yields of centro and stylo were significantly greater than those of arachis and

burgundy under 0% shade, however, centro recorded significantly higher yield than all

legumes at 50% shade, a similar trend was observed at 70% shade but the difference was

only significant when centro was compared with arachis. The dry matter yields of grass

and legumes decreased with increasing shade level. Guinea-centro mixture was

recommended based on higher dry matter yield of centro compared to other legumes

especially at 50% shade.

Allelopathy study conducted using laboratory bioassay and pot trial to test whether

guinea grass had allelopathic effect on companion legumes indicates that shoot length in

stylo was significantly (P<0.05) reduced at 1% leaf extract concentration. Dry matter

yield and root length in burgundy were significantly decreased by root powder while the

yields of centro and stylo were enhanced by leaf powder. In conclusion, the

climbing/trailing legume (centro) was observed to be the most compatible with guinea.

This is based on the facts that in study 1, guinea-centro mixture showed tendency

towards higher dry matter yield than the corresponding monoculture of grass (guinea). In

study 2, however, the guinea-centro mixture was not part of the treatments owing to lack

of germination of the centro seeds. In study 3, centro produced significantly higher dry

matter yield than arachis and burgundy and surpassed other legumes at 50 % shade.

Centro was also not negatively affected by the allelopathic effect of guinea grass as

observed in study 4.

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Abstrak tesis yang dikemukakan kepada Senate Universiti Putra Malaysia sebagai

memenuhi keperluan untuk ijazah Doktor Falsafah

KESERASIAN CAMPURAN EMPAT KEKACANG FORAJ DENGAN RUMPUT

GUINEA (Panicum maximum) JACQ

Oleh

MOHAMMED BABA

Disember 2013

Pengerusi : Profesor Madya Mohd Ridzwan Abd Halim, Ph.D

Fakulti : Pertanian

Pengeluaran ternakan di kawasan tropika dikekang oleh penurunan dalam kuantiti dan

kualiti rumput tropika utama semasa musim yang semakin meningkat. Penubuhan

kekacang makanan ternakan ke padang rumput pertanian telah digunakan sebagai

penawar untuk menyelesaikan masalah tersebut. Walau bagaimanapun, di kawasan-

kawasan tropika, tidak banyak kejayaan yang telah dicapai kerana kemampuannya

kompetitif kuat rumput yang mengurangkan kandungan kekacang teman. Ia adalah

dengan latar belakang ini bahawa empat percubaan telah dijalankan untuk menilai

keserasian campuran rumput kekacang dan kesesuaiannya untuk integrasi ternakan

perladangan.

Siasatan ke atas sifat persaingan antara Panicum maximum (rumput guinea) dengan

setiap satu daripada kekacang berikut yang berlainan tabiat pertumbuhan; Stylosanthes

guianensis (Stylo), Centrosema pubescens (centro) Macroptilium bracteatum (burgundy

bean) dan Arachis pintoi (arachis) menunjukkan bahawa hasil kering campuran guinea-

entro (2:2 dan 3:1) gabungan spesies cenderung untuk menjadi lebih tinggi daripada

monokultur rumput (perbezaan ini tidak ketara). Hasil relatif jumlah adalah lebih tinggi

dalam campuran guinea-stylo (1.41 ) dan guinea-centro (1.4) pada nisbah 2:2 kecuali

untuk guinea-burgundy. Kekacang yang memanjat (centro) kelihatan lebih serasi dengan

rumput guinea terutamanya pada kombinasi 2:2 berbanding yang tegak ( stylo),

tegak/memanjat (burgundy) dan yang melata (Arachis).

Keserasian campuran foraj dinilai pada nisbah 50:50 gabungan spesies dari segi hasil

kering dan komposisi kimia menunjukkan bahawa guinea-arachis lebih tinggi hasil

(P<0.01) terkumpul berbanding campuran lain. Walau bagaimanapun, hasil ini tidak

ketara berbeza dengan guinea monokultur. Kekacang menegak (stylo) mempunyai hasil

lebih tinggi (P< 0.01) daripada Arachis dalam campuran dengan guinea. Begitu juga,

hasil relatif stylo cenderung untuk menjadi lebih tinggi daripada arachis dan burgundy.

Oleh itu sebagai satu kompromi di antara hasil bahan kering dan kualiti, campuran

rumput Stylo disyorkan.

Produktiviti empat campuran guinea-Arachis, guinea-burgundy, guinea-centro dan

guinea-stylo dinilai di bawah 3 aras naungan 0 ( 1701.9 μmol m-2

s-1

), 50 ( 583.3 μmol

m-2

s-1

) dan 70 % ( 520.2 μmol m-2

s-1

) menunjukkan bahawa guinea-centro cenderung

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untuk mengeluarkan hasil yang lebih tinggi di kalangan campuran walaupun perbezaan

ini tidak ketara . Centro dalam campuran dengan guinea mencatatkan hasil lebih tinggi

(P< 0.05) daripada Arachis dan burgundy. Interaksi antara spesies kekacang dan aras

naungan pada pengeluaran bahan kering menunjukkan bahawa hasil bahan kering centro

dan stylo nyata lebih besar daripada arachis dan burgundy di bawah 0% teduh,

bagaimanapun, centro mencatatkan hasil lebih tinggi daripada semua kekacang pada

50% naungan, trend yang sama diperhatikan di naungan 70%, tetapi perbezaan itu hanya

ketara apabila centro telah dibandingkan dengan arachis. Hasil bahan kering rumput dan

kekacang menurun dengan peningkatan tahap teduh. Campuran guinea- centro disyorkan

berdasarkan hasil bahan kering tinggi centro berbanding kekacang lain terutama pada

50% teduh.

Kajian allelopathy dijalankan menggunakan bioesei makmal dan percubaan pasu untuk

menguji sama ada rumput guinea mempunyai kesan alelopati pada kekacang

menunjukkan bahawa panjang tumbuhan stylo menurun secara bererti (P<0.05) dengan

campuran 1% ekstrak daun. Hasil bahan kering dan panjang akar dalam burgundy telah

menurun dengan ketara oleh serbuk akar manakala hasil centro dan stylo telah

dipertingkatkan dengan serbuk daun. Kesimpulannya, kekacang menegak/memanjat

(centro) diperhatikan menjadi yang paling serasi dengan guinea. Ini adalah berdasarkan

kepada fakta bahawa dalam kajian 1, campuran guinea-centro menunjukkan

kecenderungan ke arah hasil bahan kering tinggi daripada monokultur rumput (guinea).

Dalam kajian 2, bagaimanapun, campuran guinea-centro tidak merupakan sebahagian

daripada rawatan kerana kekurangan percambahan benih centro. Dalam kajian 3, Centro

mempunyai hasil bahan kering yang tinggi sedikit dari Arachis dan burgundy dan

mengatasi kekacang lain pada 50% teduh. Centro juga tidak terjejas oleh kesan alelopati

rumput guinea sebagaimana yang berlaku dalam kajian 4.

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ACKNOWLEDGEMENTS

All praise is due to Allah Subhanahu wataala for enabling me to accomplish this

monumental task in life. I would like to express my immense gratitude to Assoc. Prof.

Dr. Mohd Ridzwan Abd Halim. Chairman of the Supervisory Committee for his

consistent guidance, very useful suggestions and constructive criticisms throughout the

study period. Iam highly indebted to Prof. Dr. Abd. Razak Alimon, Abdul Shukor

Juraimi and Tuan Hj. Idris Abu Bakar, members of my Supervisory Committee for their

guidance comments and suggestions.

My sincere appreciation goes to Mr Suhaimi and Mr Mazlan Bangi both of Botany and

Physiology laboratory, Department of Crop Science respectively. Mr Mahdi Ebrahimi of

Physiology Department, Faculty of Veterinary Medicine who assisted with the in vitro

work. Mrs Nassrin Sultana and Miss Noor Anisa who have also made contributions

towards the success of this study. Words will not be adequate to express my gratitude to

Mr Fabian Tengoua, Mr shamsuddin Rufai and Mr Ibrahim Wasiu all of Institute for

Tropical Agriculture, they have been with me all through the field work.

I am highly indebted to Bayero University Kano, Nigeria, for granting me the

permission to carry out the study. Worthy of note also is the contribution of Universiti

Putra Malaysia in the areas of research facilities, enabling environment and funds.

I remain eternally grateful to my parents for their prayers and useful advices, my wife

and daughters for their patience, understanding and moral support while the programme

lasted, my two younger brothers, Kabir Muhammad and Nasir Muhammad for their

support and encouragement and my dear friend Musa Danlami Usman.

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I certify that a Thesis Examination Committee has met on 16th December 2013 to

conduct the final examination of Mohammed Baba on his Doctor of Philosophy thesis

entitled “Compatibility of four forage legumes with guinea grass (Panicum maximum

Jacq.)” in accordance with the Universities and University Colleges 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:

Siti Aisha Hassan, PhD

Associate Professor

Faculty of Agriculture

Universiti Putra Malaysia

(Chairman)

Yahya Awang, PhD

Associate Professor



(Internal Examiner)

Hawa Jaafar, PhD

Associate Professor



(Internal Examiner)

Jim Bidlack, PhD

Professor

University of Central Oklahoma

United State of America

(External Examiner)

NORITAH OMAR, PhD

Associate Professor Deputy Dean

School of Graduate Studies


Date: 17 February 2014

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

accepted as fulfillment of the requirements for the degree of Doctor of Philosophy. The

members of the Supervisory Committee are as follows:

Mohd Ridzwan Abd. Halim, Ph.D.

Associate Professor



(Chairman)

Abd. Razak Alimon, Ph.D.

Professor



(Member)

Abdul Shukor Juraimi, Ph.D.

Professor



(Member)

___________________________

BUJANG BIN KIM HUAT, PhD.

Professor and Dean

School of Graduate Studies


Date:

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DECLARATION

Declaration by Graduate Student

I hereby confirm that:

this thesis is my original work

quotations, illustrations and citations have been duly referenced

the thesis has not been submitted previously or concurrently for any other degree at

any institutions

intellectual property from the thesis and copyright of thesis are fully-owned by

Universiti Putra Malaysia as according to the Universiti Putra Malaysia (Research)

Rules 2012;

written permission must be owned from supervisor and deputy vice –chancellor

(Research and innovation) before thesis is published in book form

there is no plagiarism or data falsification/fabrication in the thesis and scholarly

integrity was upheld as according to Rule 59 in Rules 2003 (Revision 2012-2013).

The thesis has undergone plagiarism detection software

Signature: ------------------------------- Date: 16 December, 2013

Name and Matric No: MOHAMMED BABA (GS26109)

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Declaration by Members of Supervisory Committee

This is to confirm that:

the research conducted and the writing of this thesis was under our supervision,

Supervision responsibilities as stated in Rule 41 in Rules 2003 (Revision 2012-

2013) were adhered to.

Signature ------------------------------- Signature -------------------------------

MOHD RIDZWAN ABD HALIM, PhD ABD RAZAK ALIMON, PhD

(Chairman) (Member)

Signature -------------------------------

ABDUL SHUKOR JURAIMI, PhD

(Member)

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TABLE OF CONTENTS

Pages

ABSTRACT iii

ABSTRAK vi

AKNOWLEDGEMENTS ix

APPROVAL x

DECLARATION xii

LIST OF TABLES xviii

LIST OF FIGURES xxi

LIST OF PLATES xxiii

CHAPTER

1. INTRODUCTION 1

2. LITERATURE REVIEW 3

2.1.Plant Competition 3

2.1.1. Physiological Factors Affecting Plant competition 3

2.1.2. Morphological Factors Affecting Plant competition 5

2.2. Competition Model of DeWit 1960 6

2.3. Influence of Grass-Legume Mixtures on Nutritive Value 8

2.4. Influence of Grass-Legume Mixtures on Dry Matter Yield 8

2.5. Allelopathic Interactions in Grass-Legume Mixtures. 9

2.6. Nitrogen Fixation by Legumes 10

2.6.1. Factors Affecting Nitrogen Fixation 10

2.7. Influence of Fertilizer on Grass-Legume mixture 13

2.7.1. Effect of Nitrogen Fertilizer 13

2.7.2. Effect of Phosphorus Fertilizer 13

2.8. Sustainable Grass-Legume Pasture 14

2.8.1. Selection of Appropriate Forage Legume 14

2.8.2. Use of Fertilizer 14

2.8.3. Grazing Pressure 14

2.8.4. Acceptability of forage legumes by farmers 15

2.9. Contributions of Grass-legume mixtures to Animal Production 15

2.10. Silvopasture 15

2.11. Benefits of Silvopastures 15

2.12. Habitats of Plantation Crops 16

2.13. Forages under Plantation 16

2.14. Effect of Shade 17

2.14.1. Water Availability in Pastures 17

2.14.2. Photosynthesis 18

2.14.3. Growth and Morphology of Forage 18

2.14.4. Nutritive Value 20

2.14.5. Nodulation 21

2.15. Botanical and Agronomic Characteristics of Four Tropical 21

Legumes and Grass

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2.15.1. Arachis pintoi 21

2.15.2. Centrosema pubescens 23

2.15.3. Stylosanthes guianensis 24

2.15.4. Macroptilium bracteatum 25

2.15.5. Panicum maximum 27

3. NATURE OF COMPETITION BETWEEN GUINEA GRASS AND

FOUR LEGUME FORAGES 3.1. Introduction 29

3.2. Materials and Methods 30

3.2.1. Location of the Experiment 30

3.2.2. Experimental Design 30

3.2.3. Field Culture and Experimental Materials 30

3.2.4. Environmental Condition 31

3.2.5. Harvesting and Plant Measurements 32

3.2.6. Competition Indices Computed 33

3.2.7. Data Analysis 33

3.3. Results 34

3.3.1. Dry Matter Yield 34

3.3.2. Relative Yield 37

3.3.3. Relative Crowding Coefficient 41

3.3.4. Aggresivity Index 43

3.4. Discussion 44


3.4.2. Relative Yield 45

3.4.3. Relative Crowding Coefficient 46

3.4.4. Aggresivity Index 46

3.5. Conclusion 46

4. EVALUATION OF GUINEA-LEGUME MIXTURES FOR

COMPATIBILTYIN RELATION TO DRY MATTER YIELD

CHEMICAL COMPOSITIONAND IN VITRO DRY MATTER

DIGESTIBILITY

4.1. Introduction 48



4.2.2. Experimental Designs 49

4.2.3. Experimental Materials and Field Culture 49


4.2.5. Harvesting and Plant Measurements 50

4.2.6. Analysis of Chemical Composition 50

4.2.7. Analysis of Mineral Concentrations 51

4.2.8. Cumulative Gas Production and In vitro Dry Matter 52

Digestibility

4.2.9. Determination of Volatile Fatty Acids in Rumen Liquor 52


4.3. Results 53

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4.3.2. Percentage Relative Yield of Legumes in Mixtures 58

4.3.3. Percentage Grass and Legumes in Mixtures 59

and Leaf to Stem Ratio

4.3.4. Chemical Composition 60

4.3.5. Mineral Composition. 61

4.3.6. Cumulative Gas Production and In vitro Dry Matter 63

Digestibility

4.3.7. Volatile Fatty Acids Production 64

4.3.8. Correlation Coefficient 65

4.4. Discussion 65


4.4.2. Chemical Composition 66

4.4.3. Mineral Concentration. 68

4.4.4. Gas Production and In vitro Dry Matter Digestibility 69

4.4.5. Volatile Fatty Acids (VFA) Production 70

4.5. Conclusion 71

5. PRODUCTIVITY OF GUINEA-LEGUME MIXTURES UNDER

DIFFERENT SHADE LEVELS





5.2.3. Experimental Materials and Field Culture 73


5.2.5. Measurements of Physiological parameters 74

5.2.6. Harvesting and other Measurements 75

5.2.7. Analysis of Chemical Composition 75


5.3. Results 76

5.3.1. Light Intensity Measurement 76

5.3.2. Cumulative Dry Matter Yield 76

5.3.3. Mean Percentage Grass and Legumes in mixtures 76

5.3.4. Morphological Characteristics of Mixtures 78

5.3.5. Effect of light Intensity on LAI and Light Extinction 81

Coefficient.

5.3.6. Physiological Characteristics of Mixtures 82

5.3.7. Chemical composition 85

5.3.8. Correlation Coefficient 86

5.4. Discussion 87

5.4.1. Cumulative Dry Matter Yield 87

5.4.2. Mean Percentage Grass and Legumes in Mixtures 88

5.4.3. Morphological Characteristics of Mixtures 88

5.4.4. LAI and Light Extinction Coefficient. 89

5.4.5. Physiological Characteristics of Mixtures 89

5.4.6. Chemical Composition of Mixtures. 91

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5.5. Conclusion 92

6. ALLELOPATHIC POTENTIAL OF PANICUM MAXIMUM



6.2.1. Location of the experiment 93



6.2.4. Experimental Procedure 94

6.2.5. Data analysis 95

6.3. Results 95

6.3.1. Effect of Leaf Extract 96

6.3.2. Effect of Root Extract 98

6.3.3. Effect of Leaf and Root Powder 99

6.4. Discussion 102

6.4.1. Effects of Leaf and Root Extract 102

6.4.2. Effects of Leaf and Root Powder 103

6.5. Conclusion 104

7. GENERAL DISCUSSION 105

BIBILIOGRAPHY 109

APPENDICES 133

BIODATA OF STUDENT 141

LIST OF PUBLICATIONS 142

Documents

UNIVERSITI PUTRA MALAYSIA - core.ac.uk · menguji sama ada rumput guinea mempunyai kesan alelopati pada kekacang menunjukkan bahawa panjang tumbuhan stylo menurun secara bererti (P