EFFECT OF MATURITY ON DRY MATTER ACCUMULATION AND QUALITY 
OF FORAGE FROM NATURAL GRASSLAND AND THREE INTRODUCED 
GRASSES IN THE ACCRA PLAINS, GHANA
A Thesis 
Presented to 
The Faculty of Graduate Studies 
of
The University of Guelph
by
FRANCIS KWASIVIE FIANU
In partial fulfilment of requirements 
for the degree of 
Doctor of Philosophy 
December, 1977
©  Francis Kwasivie Fianu, 1977
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,/C x ‘nS'i V-v
G  270381
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ABSTRACT
EFFECT OF MATURITY ON DRY MATTER ACCUMULATION AND QUALITY OF FORAGE
FROM NATURAL GRASSLAND AND THREE INTRODUCED GRASSES
IN THE ACCRA PLAINS, GHANA
Francis Kwasivie Fianu, Supervisor:
University of Guelph, 1977 Dr. J.E. Winch
Two studies were conducted in Legon, Ghana, in 1974 and- 1975
to characterize the dry matter accumulation and quality of a natural 
grassland sward (dominated by Sporobolus and Heteropogon) and of 
introduced giant star grass (Cynodon plectostachyus (k . Schum) Pilger), 
buffel grass (Cenchrus ciliaris L. cv. biloela) and pangola grass 
(Digitaria decumbens Stent.).
The natural grassland study was a split plot experiment with 
pretreatment slashing, grazing and burning in the main plots and.ten 
harvest dates during the major rainy season in the subplots. Each of 
the ten subplots was subdivided into three parts and harvested 
sequentially at the end of the rainy season, mid-dry season and at the 
end of the dry season.
Pretreatments did not affect the botanical composition, dry 
matter accumulation during the growing season or regrowth during the 
ensuing dry period. Sporobolus p.yramidalis Beauv. grew faster than 
Heteropogon contortus (L.) Beauv. at the early stages, dominated the 
sward and flowered at 3-6 weeks. Heteropogon initially grew slowly and 
flowered from week 6. By week 7 Heteropogon became the dominant species 
of the sward. Cenchrus sp., Bothriochloa sp. and Setaria sp. flowered 
within 4-fc weeks but gamba grass (Andropogon gayanus Kunth.) did not
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flower during the study. Dry matter accumulation in the natural grass­
land sward and its dominant species continued after flowering until the 
end of the rainy season.
Leaf production, and the In Vitro digestibility as well as 
nitrogen content of Sporobolus and Heteropogon were not affected by 
pretreatment. While Sporobolus maintained a high percentage of leaf 
throughout the growing season, leaf proportions dropped in Heteropogon 
at the mature stages.
Leaves were more digestible and contained more nitrogen than 
stems in both species. Heteropogon tended to be more digestible than 
Sporobolus. The two species were similar in leaf and whole plant 
nitrogen, but Sporobolus stems contained more nitrogen than those of 
Heteropogon.
In the study on introduced grasses, giant star and buffel were 
harvested at ten dates during the minor rainy season of 1974 (September 
16 - December 3l)j during the major rainy season of 1975» pangola grass 
was included in the experiment.
Pangola was sensitive to moisture stress during early growth 
and failed to grow during the minor rainy season of 1974* Buffel, on 
the other hand, was drought tolerant and grew even under light showers. 
Buffel flowered from week 3 in both seasons while giant star flowered 
only during the minor rainy season at week 6, and pangola flowered in 
week 6 during the major rainy season. Growth continued in all three 
grasses after flowering. In buffel, senescent leaves were retained on 
the plant whereas in the stoloniferous grasses, they were stripped off 
by rainfall.
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During the minor rainy season, giant star and buffel pro­
duced similar dry matter yields. In the major rainy season, however, 
buffel was superior in yield to the prostrate grasses which showed np 
consistent differences.
Leaf dry matter yield increased until week 8-9* Buffel and 
giant star produced more leaf dry matter than pangola grass. Leaf 
proportions in the plant declined steeply with maturity in buffel grass 
but slowly in the prostrate grasses. During the minor rainy season, 
whole plant In Vitro digestibility and nitrogen were similar in giant 
star and buffel during the minor rainy season, but buffel had the 
highest whole plant digestibility followed by pangola and giant star 
was the least digestible. The leaves were more digestible than stems, 
this difference being most striking in mature buffel grass.
Leaf nitrogen levels were higher than stem nitrogen levels 
in all the grasses but species differences in nitrogen content were not 
consistent. The nitrogen content of whole plants would probably be 
adequate for the maintenance requirements of a steer until week 7 during 
the minor rainy season and week 11 in the major rainy season. In Vitro 
digestibility was highly correlated with nitrogen content of leaves, 
stems and whole plants of all species except giant star stem.
It would appear that buffel grass should be harvested at 5 
weeks and giant star at 7 weeks during the minor rainy season. In the 
major rainy season buffel would be harvested at 9 weeks and giant star 
and pangola at 8 weeks for optimum combination of nutrient yield during 
the rainy season and regrowth during the ensuing dry period.
The natural grassland species and the introduced grasses were 
similar in digestibility at the early stages but the erect grasses -
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natural and introduced - declined more rapidly than the prostrate 
introduced ones.
For high animal performance both the natural and the intro­
duced species would have to be supplemented with concentrates.
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ACKNOWLEDGEMENTS
The author wishes to express his profound gratitude to his 
supervisor Professor John E. Winch and to the other members of his 
committee: Dr. E.N.W. Oppong, Professor of Animal Science and Dean of
Agriculture, University of Ghana, Legon; Professor J.W. Tanner,
Chairman of Crop Science Department, University of Guelph; Professor
D.J. Hume of Crop Science Department, Guelph; Professors D.G. Grieve 
and J. Buchanan-Smith of the Animal Science Department, Guelph, for 
their encouragement and to Professor Bruce Hunter, Crop Science 
Department, University of Guelph, for his administrative help while in 
Ghana.
The author is also indebted to Messrs. S.A. Gyadu, Adapoe 
and Amartey for their help in the supervision of the field work in 
Legon; to Messrs. Norman Abavon and Saforo for their help in applying 
the burning pretreatment; to .Dr. Samuel White of the Zoology Department, 
Legon, for' his help in drying some of the samples; to Dr. Bafi Yeboah, 
Messrs. D.A. Ayebo and J.M. Dzakuma of the Nungua Agricultural Research 
Station, Legon, for assisting with the grazing pretreatment; to 
Drs. Essie Blay, R.B. Dadson, E.O. Otchere and R.E. Larsen for taking 
on the author's lectures and student supervision while he was in Guelph.
Much gratitude is owed to Professor B.R. Christie, Crop 
Science Department, Guelph for his valuable suggestions in statistical 
analyses, to Mr. Ban Yu, Crop Science Department, Guelph, and Mr.
John Tofflemire and Dr. N.T. Ison, I.C.S., Guelph and Dr. Daniel Ennis, 
Food Science Department, Guelph for their help with computer programming. 
The establishment of the IVD lab in Legon emanated from this study and
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the author is grateful to Dr. J. Buchanan-Smith who proposed it;
Dr. J.E. Winch who strove to make it materialize; Mrs. Helen Major 
who started it; and to Dean E.N.W. Oppong, Dr. R.K. Assoku and Dr.
E.O. Otchere of the Animal Science Department, Legon who helped with 
the fistulations.
The study as well as the IVD laboratory was funded by CIDA 
under the Ghana Project; the author is grateful to CIDA and Professor 
J.C.M. Schute, Director of the Project.
The study leave granted by the University of Ghana, Legon 
to make this work possible is most gratefully acknowled.
Thanks are also due to the Universities of Ghana and Guelph 
for the use of their facilities.
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To the memory of the late 
Reverend Samuel Yameke Brew 
of the Methodist Mission, Nyakrom, Ghana
iii
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TABLE OF CONTENTS
GENERAL INTRODUCTION ........................................  1
GENERAL LITERATURE REVIEW .................................... 4
GENERAL MATERIALS AND METHODS ................................  11
PAPER 1: DRY MATTER ACCUMULATION IN NATURAL GRASSLAND
IN THE ACCRA PLAINS ................................  16
PAPER 2: EFFECT OF MATURITY ON QUALITY OF SPOROBOLUS
PYRAMIDALIS AND HETEROPOGON CONTORTUS IN THE 
ACCRA PLAINS ......................................  34
PAPER 3: DRY MATTER ACCUMULATION IN GIANT STAR, BUFFEL
AND PANGOLA GRASSES IN THE ACCRA PLAINS ................ 55
PAPER 4: EFFECT OF MATURITY ON QUALITY OF GIANT STAR,
BUFFEL AND PANGOLA GRASSES IN THE ACCRA PLAINS ......... 76
GENERAL DISCUSSION ..........................................  106
GENERAL CONCLUSIONS ..........   114
LITERATURE CITED ............................................  116
APPENDICES .................................................  128
Page
iv
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LIST OP TABLES
0.1 Schedule of activities in the study of natural grassland .... 12
0.2 Schedule of activities in the study of introduced grasses ... 14
1.1 Analyses of variance of yield of dry matter from natural 
grassland during the major rainy seasons of 1974 and 1975 ••• 22
1.2 Regrowth yield from natural grassland during the dry
period (July-August) 1974 ........................   28
1.3 Regrowth yield from natural grassland during the dry
period (July-August) 1975 ....................    29
1.4 Seasonal total yield from' natural grassland during 1974
(upper) and 1975 (lower) ...........................    32
2.1 Yield of leaves of Sporobolus and Heteropogon during the
major rainy season of 1974 ...............................  39
2.2 Yield of leaves of Sporobolus and Heteropogon during the
major rainy season of 1975 ...................     40
2.3 Percent In Vitro digestible dry matter of whole plants 
of Sporobolus and Heteropogon during the major rainy
seasons of 1974 (upper) and 1975 (lower) .............    44
2.4 Regressions of percent In Vitro digestibility on maturity 
in Sporobolus and Heteropogon during the major rainy
seasons of 1974 and 1975 .................................  46
2.5 Percent nitrogen in whole plants of Sporobolus and 
Het eropogon during the major rainy seasons of 1974
(upper) and 1975 (lower) ...................    50
2.6 Regressions of percent nitrogen on maturity in Sporobolus 
and Heteropogon during the major rainy seasons of 1974
and 1975 ................................................ 51
3.1 Regrowth yields from giant star and buffel grasses during
the dry period (December 1974 "to April 1975)  ..............  66
3.2 Regrowth yields from giant star, buffel and pangola
grasses during the dry period (August-September) 1975 .......  67
3 .3 Seasonal total yields from giant star and buffel grasses
during 1974 ............................................. 70
Table Page
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LIST OF TABLES (cont'd)
Table
3.4 Seasonal total yields from giant star, buffel and 
pangola grasses during 1975 • • • •  .....................  •
4.1 Yield of leaves of giant star and buffel grasses
during the minor rainy season of 1974 .................
4.2 Yield of leaves of giant star, buffel and pangola 
grasses during the major rainy season of 1975 ..........
4*3 Percent In Vitro dry matter digestibility of whole plants 
of giant star and buffel grasses during the minor rainy 
season of 1974 (upper) and of giant star, buffel and 
pangola grasses during the major rainy season of 1975 
(lower)  ............................ ..............
4 .4 Regressions of percent In Vitro dry matter digestibility 
on maturity of giant star and buffel grasses during the 
minor rainy season of 1974 and of giant star, buffel and 
pangola grasses during the major rainy season of 1975 ••••
4»5 Percent nitrogen of whole plants of giant star and buffel 
grasses during the minor rainy season of 1974 (lower) and 
of giant star, buffel and pangola grasses during the 
major rainy season of 1975 (lower) ....................
4*6 Regressions of percent nitrogen on maturity of giant star 
and buffel grasses during the minor rainy season of 1974 
and of giant star, buffel and pangola grasses during the 
major rainy season of 1975 .... .......................
4*7 Correlation coefficients between In Vitro dry matter
digestibility and nitrogen in leaves and stems of giant 
star and buffel grasses during the minor rainy season of 
1974 and of giant star, buffel and pangola grasses during 
the major rainy season of 1975 ........................
4.8 Yield of In Vitro digestible dry matter of v/hole plants 
of giant star and buffel grasses during the minor rainy 
season of 1974 (upper) and of giant star, buffel and 
pangola grasses during the major rainy season of 1975 
(lower) .............................................
4.9 Yield of nitrogen of whole plants of giant star and buffel 
grasses during the minor rainy season of 1974 (upper) and 
of giant star, buffel and pangola grasses during the major 
rainy season of 1975 (lower)..........................
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LIST OP FIGURES
1.1 Dry matter accumulation during the major rainy seasons 
of 1974 and 1975 natural grasslands in the Accra
Plains ......................................      23
1.2 Accumulation of dry matter of species components in 
natural grassland during the major rainy season of
1974 in the Accra Plains ................................  25
1.3 Accumulation of dry matter of species components in 
natural grassland during the major rainy season of
1975 in the Accra Plains ................................  26
2.1 Percent In Vitro dry matter digestibility of leaves and 
stems of Sporobolus and Heteropogon during the major
rainy season of 1974 ..................................  42
2.2 Percent In Vitro dry matter digestibility of leaves and 
stems of Sporobolus and Heteropogon during the major
rainy season of 1975 ..................................  43
2.3 Percent nitrogen content of leaves and stems of 
Sporobolus and Heteropogon during the major rainy
season of 1974 .......................................  48
2.4 Percent nitrogen content of leaves and stems of 
Sporobolus and Heteropogon during the major rainy
season of 1975 ................    49
3*1 Accumulation of dry matter in giant star, buffel and
pangola grasses during the minor rainy season of 1974 
and the major rainy season of 1975 .....................  62
4.1 In Vitro dry matter digestibility of leaves and stems 
of buffel and giant star grasses during the minor rainy 
season of 1974 .......................................  87
4.2 In Vitro dry matter digestibility of leaves and stems 
of giant star, buffel and pangola grasses during the
major rainy season of 1975 ...........................  88
4.3 Nitrogen content of leaves and stems of giant star and
buffel grasses during the minor rainy season of 1974 ...... 93
4«4 Nitrogen content of leaves and stems of giant star,
buffel and pangola grasses during the major rainy 
season of 1975 .......................................  94
Figure Page
vii'
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LIST OF APPENDICES
I Analyses of variance of weekly yields from natural
grassland during the major rainy season of 197,4.....
II Analyses of variance of weekly yields from natural
grassland during the major rainy season of 1975 ...... -*-30
III Analyses of variance of dry matter of species 
components of natural grassland during 1974
(upper) and 1975 (lower) ........................... 131
IV Weekly yields of species components of natural
grassland during the major rainy season of 1974 ......  132
V Weekly yields of species components of natural
grassland during the major rainy season of 1975 ......  133
VI Analyses of variance of weekly regrowth yields
from natural grassland during 1974 ..................  134
VII Analyses of variance of weekly regrowth yields
from natural grassland during 1975  .............. 135
VIII Analyses of variance of regrowth yields during
the dry periods of 1974 (upper) and 1975 (lower) .....  136
IX Yields of regrowth from natural grassland during
the dry period of 1974 ......................    137
X Yields of regrowth from natural grassland during
the dry period of 1975 ...........................   138
XI Analyses of variance of seasonal total yields
from natural grassland during 1974 ..................  139
XII Analyses of variance of seasonal total yields from
natural grassland during 1975 ....................... 140
XIII Analyses of variance of weekly yields of leaves-of
Sporobolus and Heteropogon during the major rainy 
seasons of 1974 (uPPer) and 1975 (lower) ............. 141
XIV Analyses of variance of yield of leaves of
Sporobolus and Heteropogon during the major rainy 
seasons of 1974 (upper) and 1975 (lower) ............. I42
Appendix Page
viii
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ix
XV Analyses of variance of weekly percent In Vitro dry 
matter digestibility of leaves (upper) and stems (lower) 
of Sporobblus and Heteropogon during the major rainy
season of 1974 ....................................... ^ 3
XVI Analyses of variance of weekly percent In Vitro dry 
matter digestibility of leaves (upper) and stems (lower) 
of Sporobolus and Heteropogon during the major rainy
season of 1975 .......................    144'
XVII Analyses of variance of percent In Vitro dry matter 
digestibility of leaves (upper) and stems (lower) of 
Sporobolus and Heteropogon during the major rainy
season of 1974 .......... •...........................  145
XVIII Analyses of variance of percent In Vitro dry matter 
digestibility of leaves (upper) and stems (lower) of 
Sporobolus and Heteropogon during the major rainy
season of 1975 .......... ........................ . 146
XIX Analyses of variance of percent In Vitro dry matter 
digestibility of whole plants of Sporobolus and 
Heteropogon during the major rainy seasons of 1974
(upper) and 1975 (lower) .............................  147
XX Percent In Vitro digestible dry matter of leaves, stems 
and 'whole plants of Sporobolus and Heteropogon during
the major rainy season of 1974 ........................ 148
XXI Percent In Vitro digestible dry matter of leaves, stems 
and whole plants of Sporobolus and Heteropogon during
the major rainy season of 1975 ......................... 149
XXII Analyses of variance of percent In Vitro dry matter 
digestibility (leaves vs. stems) in Sporobolus (upper) 
and Heteropogon (lower) during the major rainy season of
1974 ................................................ 150
XXIII Analyses of variance of percent In Vitro dry matter 
digestibility (leaves vs. stems) in Sporobolus (.upper) 
and Heteropogon (lower) during the major rainy season
of 1975 .............................................  151
XXIV Analyses of variance of weekly percent In Vitro dry 
matter digestibility in whole plants of Sporobolus and 
Heteropogon during the major rainy season of 1974 (upper)
and 1975 (lower) ..................................... ^^2
LIST OP APPENDICES (cont'd)
Appendix Page
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XXXV Analyses of variance of weekly percent nitrogen 
of leaves (upper) and stems (lower) of 
Sporobolus and Heteropogon during the major rainy
season of 1974 ...................................  153
XXVI Analyses of variance of weekly percent nitrogen 
of leaves (upper) and stems (lower) of Sporobolus 
and Heteropogon during the major rainy season of
1975 ............................................  154
XXVII Analyses of variance of percent nitrogen in leaf, 
stem and whole plant of Sporobolus and Heteropogon 
during the major rainy seasons of 1974 (upper)
and 1975 (lower) .................................  155
XXVIII Percent nitrogen in leaves,"stems;and whole plant 
of Sporobolus and Heteropogon during the major
rainy season of 1974 .............................  156
XXIX Percent nitrogen in leaves,.stemsand whole plant 
of Sporobolus and Heteropogon during the major
rainy season of 1975 .............................  157
XXX Analyses of variance of percent nitrogen of 
leaves vs. stems in Sporobolus (upper) and 
Heteropogon (lower) during the major rainy
season of 1974 ................................... 158
XXXI Analyses of variance of percent nitrogen of leaves 
vs. stems in Sporobolus (upper) and Heteropogon
(lower) during the major rainy season of 1975 .......  159
XXXII Analyses of variance of weekly percent nitrogen 
of whole plants of Sporobolus and Heteropogon 
during the major rainy seasons of 1974 (upper)
and 1975 (lower) .................................  160
XXXIII Analyses of variance of weekly yields from 
giant star, buffel and pangola grasses during the 
minor rainy season of 1974 (uPPer) and the major
rainy season of 1975 (lower) ......................  161
XXXIV Analyses of variance of yields from giant star, 
buffel and pangola grasses during the minor 
rainy season of 1974 and the major rainy season
of 1975 .........................................  162
LIST OP APPENDICES (oont'd)
Appendix Page
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LIST OP APPENDICES (cont'd)
Appendix
XXXV
XXXVI
XXXVII
XXXVIII
XXXIX
XL
XLI
XLII
XLIII
Page
Accumulation of dry matter of giant star, buffel and
pangola grasses during the minor rainy season of
1974 (upper) and the major rainy season of 1975 (lower) .. 163
Analyses of variance of regrowth yields from giant 
star, buffel and pangola grasses during the dry periods
of December 1974 - May 1975 (upper) and- August -
September 1975 (lower)  ........  ..t.......... 164
Analyses of variance of weekly regrowth yields from
giant star, buffel and pangola grasses across regrowth
dates during 1974 (upper) and 1975 (lower)  .......    I65
Analyses of variance of seasonal total yields from 
giant star, buffel and pangola grasses during 1974 
(upper) and 1975 (lower) .............................  166
Analyses of variance of yield of leaves of giant star 
and buffel grasses during the minor rainy season of
1974 and of giant star, buffel and pangola grasses in
the major rainy season of 1975 ........................  167
Analyses of variance of percent In Vitro dry matter 
digestibility of leaves, stems and whole plants of 
giant star, buffel and pangola grasses during the minor 
rainy season of 1974 and the major rainy season of
1975 ................................................ 168
Analyses of variance of weekly percent In Vitro dry 
matter digestibility of leaves, stems and whole plants 
of giant star, buffel and pangola grasses during the 
minor rainy season of 1974 .....................   169
Analyses of variance of weekly percent In Vitro dry 
matter digestibility of leaves, stems and whole plants 
of giant star, buffel and pangola grasses during the 
major rainy season of 1975 ............................ 170
Analyses of variance of weekly percent In Vitro dry 
matter digestibility (leaves vs. stems) of giant star, 
buffel and pangola grasses during the minor rainy 
season of 1974 (upper) and the major rainy season of
1975 (lower)  .............     171
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xii
LIST OP APPENDICES (cont'd)
XLIV Percent In Vitro dry matter digestibility of leaves, 
stems and whole plants of giant star and buffel 
grasses during the minor rainy season of 1974 ..........  172
Appendix Page
XLV Percent In Vitro dry matter digestibility of leaves,
stems and whole plants of giant star, buffel and 
pangola grasses during the major rainy season of 1975 •••• 1^3
XLVI Analyses of variance of percent nitrogen in leaves,
stems and whole plants of giant star, buffel and 
pangola grasses during the minor rainy season of 1974 
and the major rainy season of 1975 ................. 174
XLVII Analyses of variance of weekly percent nitrogen content
of leaves, stems and whole plants of giant star, buffel 
and pangola grasses during the minor rainy season of
1974 ................................................ 175
XLVIII Analyses of variance of weekly percent nitrogen content
of leaves, stems and whole plants of giant star, buffel 
and pangola during the major rainy season of 1975 176
XLIX Analyses of variance of weekly percent nitrogen (leaves
vs. stems) of giant star, buffel and pangola grasses 
during the minor rainy season of 1974 (upper) and the 
major rainy season of 1975 (lower) ................. 177
L Percent nitrogen of leaves, stems and whole plants of
giant star and buffel grasses during the minor rainy 
season of 1974 ....................................... 178
LI Percent nitrogen of leaves, stems and whole plants of
giant star, buffel and pangola grasses during the 
major rainy season of 1975  ......................... 179
LII Analyses of variance of yield of In Vitro digestible
dry matter and nitrogen of whole plants of giant star,
buffel and pangola grasses during the minor rainy
season of 1974 and the major rainy season of 1975 .......  180
LIII Analyses of variance of weekly yields of In Vitro
digestible dry matter (leaves vs. stems) of giant star, 
buffel and pangola grasses during the minor rainy 
season of 1974 (upper) and the major rainy season of
1975 (lower) ...........................    181
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xiii
LIV Yield of In Vitro digestible dry matter of leaves,
stems and whole plants of giant star and buffel 
grasses during the minor rainy season of 1974 ’.........
LV Yield of In Vitro digestible dry matter of leaves,
stems and whole plants of giant star, buffel and
pangola grasses during the major rainy season of 1975 ••• 1®3
LVI Analyses of variance of weekly yields of nitrogen
(leaves vs. stems) of giant star, buffel and pangola
grasses during the minor rainy season of 1974 (upper)
and the major rainy season of 1975 (lower) 184
LVII Yield of nitrogen of leaves, stems and whole plants of
giant star and buffel grasses during the minor rainy 
season of 1974 • ••••............................. .. 185
LVIII Yields of nitrogen of leaves, stems and whole plants 
of giant star, buffel and pangola grasses during the 
major rainy season of 1975 .................*....... .
LIX Weekly distribution of rainfall during 1974 ............ 1®7
LX Weekly distribution of rainfall during 1975 ...........
LXI Weekly distribution of rainfall during the minor
rainy season of 1974 and the ensuing dry period ....... 189
LXII Weekly distribution of rainfall during the major
rainy season of 1975 and the ensuing dry period.......  190
LIST OP APPENDICES (cont'd)
Appendix Page
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GENERAL INTRODUCTION
Livestock raising in Ghana is primarily associated with the 
savanna areas which account for about two—thirds of the country's land 
area. About 70$ of the sheep and goats and almost all the cattle in 
the country subsist on the natural grazing in these savannas. These 
grasslands have been designated the Guinea Savanna Zone and the 
Coastal Grasslands and Thickets (Taylor 1952). The Accra Plains are 
located in the Coastal Grassland and Thickets Belt.
The Accra Plains contain about 240,000 hectares of savanna 
grassland located about 10 km west of the city of Accra extending to 
the Volta Estuary in the East and from the Akwapim Hills in the North, 
to the Atlantic Ocean on the South. The area is an undulating plain 
less than 80 m above sea level except for a few isolated rocky hills 
up to 300 m above sea level (Brammer 1967). The soils which vary from 
clay to sand are deficient in nitrogen and phosphorus (Nye and 
Bertheux 1957, Brammer 1956, 1962, 1967)-
Precipitation in the Accra Plains varies from 65O mm to 
1000 mm per year with peaks in June, during the major rainy season, and 
September during the minor rainy season (Walker 1962). This low rain­
fall regime limits forage quality and yields, particularly during the 
dry season (Blair 1963)*
For this reason losses of up to llfo of liveweight during the 
dry season is one of the major problems confronting the livestock indus­
try of Ghana (Rose Innes 1961). Such losses are not uncommon in the 
tropics and in some areas the losses during the dry season are as high 
as wet season gains (French 1939) and result in the delay of over two
1
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2years in the maturity of cattle. The contention that quantity and 
quality of forage impose, a severe limitation on livestock growth is 
supported by findings from work underway at the University of Ghana's 
Kade Agricultural Research Station. In that study, yearling Wungua 
Blackhead Sheep, raised on cover crops under plantation tree crops, 
gained weight equivalent to that of three year olds of the same breed 
raised on savanna grassland (Oppong, private communication).
In attempts to find higher yielding and better quality 
fodder, many species of forage have been introduced into Ghana since 
the late 1920's (Gold Coast Department of Agriculture Annual Reports 
1931-1943? Kannegieter 1961, 1965> Rose-Innes 1966). Among the species 
thought to be adapted to Ghanaian conditions on the basis of their 
vigour are buffel grass (Cench.rus ciliaris L.), giant star grass 
(Cynodon plectostachyus Pilger) and pangola grass (Digitaria decumbens 
Stent.) (Evans 1961, Thompson 196l> Asare 1972).
Information is scarce on the dry matter accumulation of 
native and introduced forage grasses, as they grow during the rainy 
season in Ghana. Dry matter accumulation in natural grassland in the ’ 
Accra Plains was reported by Lansbury, Rose Innes and Mabey (1965). 
Yields increased to 3000 kg/ha during the major rainy season and the 
herbage contained 1.6% N which fell to 0 .42 - 0*54% hy the beginning 
of the dry season. Similar deterioration in nitrogen content was 
reported by Sen and Mabey (1965)• These studies did not involve 
digestibility or leaf production of the individual species, and there 
have been no such studies on the introduced species.
The objective of the work undertaken herein was to assess the 
dry matter accumulation, leaf dry matter production, nitrogen content
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3and digestibility of the major components of a natural grassland during 
the major rainy seasons of 1974 and 1975i ant* °f introduced pangola 
(Digitaria decumbens Stent.), giant star (Cynodon plectostachyus 
(K. Schum.) Pilger) and buffel grass (Cenchrus ciliaris L.) during the 
minor rainy season of 1974 and the major rainy season of 1975* 
both natural grassland and introduced species, the amount of dry matter 
accumulated after the rainy season harvests was estimated at the end 
of the rainy season, the mid-dry season and the end of the dry season. 
The study was conducted at the University of Ghana, Legon, in the 
Accra Plains.
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GENERAL LITERATURE REVIEW
The vegetation of the Accra Plains consists of clumps of 
broken thickets (Loxton 1955) and has been described by Rose Innes 
(1962) as "peppercorn tree savanna". The woody species include Grewia 
carpinifolia. Fagara xanthoxyloides, Gardenia ternifolia, Clausena 
anisata, Dichrostachys glomerata and several others. The dominant 
grasses vary from Andropogon gayanus and Hyparrhenia spp. in less 
disturbed upland areas, to Vetiveria fulvibarbis in the rolling plains. 
Short perennials such as Heteropogon contortus, Sporobolus pyramidalis, 
Cenchrus ciliaris, Bothriochloa bladhii and Brachiaria falcifera as 
well as such annuals as Eragrostis spp. and Aristida spp. replace 
Vetiveria where there is excessive grazing, trampling, burning and 
other forms of disturbance to the vegetation. Panicum maximum occurs 
in pure stands on the more humid verge of this zone and stretches into 
the contiguous moist forest belt to the north (Brand and Brammer 1956, 
Brammer 19&71 Rose Innes 1962).
Little is known about the effect of maturity on dry matter 
accumulation and quality of these species, during the rainy season or 
during the dry season. Lansbury et_ al. (1965) studied two grassland 
communities on the Accra Plains, one on clay soil dominated by Vetiveria 
fulvibarbis (Trin.) Stapf., Andropogon canaliculatus Schumach., and 
Heteropogon contortus (L.) Beauv. ex Roem et Schult; the other was on 
sandy soil and the major species were Schizachyrium schweinfurthii 
(Hack.) Stapf., A. canaliculatus, V. fulvibarbis, Setaria sphacelata 
(Schumach.) Stepf. et Hubbard, Sporobolus pyramidal is Beauv. and
H. contortus. Harvesting the plots at stages of development varying
4
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5from 2 to 32 weeks of growth during the major and minor rainy seasons 
(May to November), they found that total nitrogen was over 1.6% at the 
start of the growing season but fell to 0 .42$ in the sandy soil and 
0 .54$ in clay soil. Dry matter yields in the sandy communities tended 
to be higher than in the clay soil. Sen and Mabey (1965) also repor­
ted similar deterioration in quality with maturity in the grasslands 
of the Accra Plains.
Little information is available concerning the growth and 
development of cultivated grasses introduced into Ghana to help solve 
the deficiencies in natural grassland species. However in Rhodesia, 
Brockington (1961) studied the growth of twelve cultivated species 
including buffel grass and giant star grass. He reported that buffel 
commenced growth before the rains, and continued growing during early 
flowering and seed setting until seed shedding. In contrast with the 
bunch type grasses such as buffel grass, giant star grass had a con­
tinuous cycle of initiation and replacement of aerial shoots. After 
flowering, vegetative growth continued in both giant star and buffel 
but no new shoots developed. Similar findings have been made by 
Virguez (1965) on pangola grass in Brazil, and by Taerum (1970a,b,c) 
on buffel in Kenya.
During early vegetative growth, the grasses are succulent, 
leafy, high in nitrogen content and low in fibre (French 1957 > Sen and 
Mabey 1965» Arias and Butterworth 1965). As growth proceeds, the cells 
undergo secondary thickening and the cell walls increase at the 
expense of the nitrogen-rich protoplasm and other components of the 
neutral detergent solubles. The thickened walls contain more cellulose 
and because of increased lignin content, are less digestible (Klock,
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6Schank and Moore 1975» Sullivan 19^ 9» 1973)*
Vicente-Chandler £t aJ. (1959a,b) in Puerto Rico, attributed 
much of the lowering of nutrient value as herbage matures, to the fall 
in leafiness and corresponding increase in the less nutritious stems. 
They reported that elephant grass (Pennisetum purpureum) had 55» 42 
and 30$ leaf at 40, 60 and 9° days respectively while guinea grass 
(Panicum maximum Jacq.) had 53j 53 and 36$. Similar trends have been 
reported by Arias and Butterworth (1965) in Brazil for elephant grass 
and for buffel grass by Juko and Bredon (1961) in the Caribbean,
Taerum (l970a,b,c) in East Africa and Burton (1976) in Georgia. On the 
other hand, '/icente-Chandler et al. (1959a,b) reported that pangola 
grass cut at 30, 45 and 60 days contained 38, 39 and 40$ foliage res­
pectively and varied little in leafiness during growth over the 30 to 
60 day period. The differences between pangola and the former group 
can be attributed to differences in growth habit. Pangola is stoloni- 
ferous and develops new foliage continually if growth conditions are 
favourable. Initially, it has a high proportion of stem and this pro­
portion does not change with age because leaves arise every few centi­
metres along stolons. On the other hand, the stool forming bunch 
grasses of the former group are initially leafy and develop stems with 
maturity. The decline in leafiness of these stool forming grasses is 
most rapid just before inflorescence emergence.
Nitrogen levels in tropical forages have received much 
attention: French (1939) in East Africa studied giant star grass,
Paterson (1933, 1935, 1938) in The West Indies examined guinea grass 
and elephant grass, among others; while in Ghana, Lansbury et al,
(1965) and Sen and Mabey (1965) investigated several natural grassland
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7species including buffel and guinea grass. However few of these 
studies involved separate leaf and stem fractions. It is apparent from 
the data of Vicente-Chandler et al^  (1959a,b), Norman (19^ 3) and Arias 
and Butterworth (1965) "that despite the growth differences among 
grasses, and even though nitrogen is higher in the leaf than in the 
stem, the rate of decline is the same in both. In pangola grass, 
guinea grass and elephant grass, the leaf nitrogen fell from 2.45% at 
30 days to 1.5% at 90 days while the stem decreased from 1.68% to 0.8% 
at 30 and 90 days respectively (Vicente-Chandler et al. 1959a,b). 
Similarly Arias and Butterworth (1965) reported that elephant grass 
had leaf and stem nitrogen content averaging 3*37% and 2.76% respec­
tively at 20 days but these dropped to 1 .14% and 0.45% respectively 
by the 90th day; the decrements being 2.23 percentage points in the
leaf and 2.31 in the stem.
Changes in cell wall components by detergent fibre analysis 
in tropical grasses have been reported by Johnson, Guerrero and Pezo 
(1973) in Peru, by Olubajo and Van Soest (1974) in Nigeria, Klock et al. 
(l975)i Ventura et al. (1975) in Florida, and by Reid, Post and Olsen 
(1975) in Uganda. Although differences in the proportions of the 
various fibre components occurred among species, cellulose and lignin 
increased sharply with plant maturity whereas silica and hemicellulose 
were erratic. Kamstra, Stanley and Ishizaki (1966) reported that per­
cent cellulose rose from 33% "to 36% and hemicellulose from 30 to 37%
in the tops of Kikuyu grass (Pennisetum clandestinum) during a 10 week 
growth period. Van Biljohn and LeRoux (1969) in South Africa, obtained 
an increase of 25 to 33% in foliage cellulose of Themeda triandra during 
maturation in the rainy season. Gomide et al. (1969) in Brazil reported
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8two groupings of species in the rate of change of fibre. Meiinis 
minutiflora Beauv. (molasses grass), elephant grass and guinea grass 
increased sharply from 25 to 40$ in cellulose in 36 weeks. Whereas 
only slight increases occurred in the procumbent species pangola,
Kikuyu, and C.ynodon dact.ylon (Bermuda grass): 36 to 40$, 20 to 30$
and 31 to 32$ respectively. Percent lignin in the leaves underwent 
very little change with maturity whereas stem lignin soared from about 
7*5$ at 30 days to 12.5$ at 90 days (Vicente-Chandler et al. 1959a,b).
Digestibility of forages declines as the herbage matures 
(Duckworth 1946, Minson and McLeod 1970, Danley and Vetter 1973, Mba,
Oke and Oyenuga 1973, Engdahl and Ellis 1974, Olubajo and Van Soest 
1974, and Chenost 1975)* The main cause of this decline is thought to 
be the increase in ligno-cellulose as the plant matures (Crampton and 
Maynard 1938, Drapala, Raymond and Crampton 1947, Kamstra et al. 1958, 
Sullivan 1959, 1964, Butterworth 1967, Moore and Mott 1973, Cross,
Smith and DeBarth 1974, an<i Johnson and Pezo 1975)* However this con­
tention has been challenged by Kayongo-Male and Thomas (1972), OLubajo 
et al. (1973) and Olubajo and Van Soest (1974), who have found low 
correlations between fibre components and digestibility, and Carza,
Hertel and Jolliff (1976) who found inconsistent relations between cell 
wall and digestibility in C.ynodon sp.
Silica has been implicated in the decline of digestibility 
with age in tropical herbage (Minson 1971, Van Soest, Arroyo-Aguilu 
and Tessema 1974, Johnson and Pezo 1975)* However, recent work by 
Olubajo and Van Soest (1974), Johnson and Pezo (1975) and Ventura et al. 
(1975) suggest that silica cannot be the sole factor responsible for 
decreases in digestibility because silica levels are erratic with herbage 
maturity.
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9Similarly, the reduction in leafiness cannot fully explain 
decreases in digestibility as the leaf is not always more digestible 
than the stem; Raymond (1969) found immature stems more digestible than 
leaves in Lolium perenne.
A further complication in attempting to explain the causes 
of the decline in digestibility with growth arises from the fluctua­
tions during the maturation process. Johnson and Pezo (1975) working 
with Brachiaria reported that digestibility decreased from 82% at 2 
to 4 weeks to 64% at 10 weeks but rose to 71% at 12 weeks, down to 64% 
again at 13ir weeks and up to 67% at 17g- weeks. In the same study, 
the digestibility of Hyparrhenia at 2 weeks was 79%; this fell to 75% 
at 8 weeks but rose to 79% at 12 weeks to fall again to 74-79% at 
13lr to 17-4- weeks (Reid et al. 1973)-
Weather changes could complicate the effect of growth on the 
changes in quality. A severe dry spell during the growing season may 
temporarily arrest growth and cause premature senescence of lower 
leaves, thus reducing leafiness. There may also be a drop in cell 
solubles and an accelerated secondary thickening, both of which would 
tend to reduce digestibility (Haggar and Ahmed 1970, 1971)• On 
resumption of the rains, growth would commence, leafiness would increase, 
soluble cell contents would rise, cell expansion would occur and 
digestibility could increase.
In contrast with the reduction of digestibility with age,
Grieve and Osbourne (1965) working on pangola grass, reported that 
digestibility increased with maturity. They found a rise in diges­
tibility from 58.8% at 3 weeks to 62.4% at 4 weeks and 64.5% at 5 weeks. 
Butterworth and Butterworth (1965) found the same trends from preflowering
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10
to flowering in pangola grass. This may be due to changes in the 
polymers of hemicellulose in the cell walls, since the different poly­
mers of hemicellulose have different digestibilities (Bailey 1973)* 
Another possible factor may be a change in the physical configuration 
of the cell wall constituents in the rapidly expanding stem cells, 
such that they became more susceptible to effective enzyme attack.
Such wall "loosening" is known to occur (Brady 1973) but has not been 
linked with digestibility. During maturation, as the cells become 
thickened, they would decline in digestibility.
With the exception of silica, all the forage quality para­
meters being investigated herein are usually affected by growth. 
Hemicellulose may change little in quantity but undergoes changes in 
monoglyceride composition (Bailey 1973)- Cellulose and lignin increase 
with maturity, but silica rises and falls erratically while leafiness 
declines as the plant matures. Digestibility generally declines as 
the plant matures but there are differences due to species and environ­
mental changes, particularly moisture supply to the plant.
Both introduced and natural grassland species behave similarly 
as regards the effects of maturity and plant part on herbage quality 
but the introduced grasses, because of better nurture under cultivation 
and possibly because they have been selected for these characteristics, 
tend to outyield and are higher in nitrogen, leafiness and digestibility. 
Thus, these introduced species may be used to replace the natural 
grassland species which have poor dry matter yield and quality.
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GENERAL MATERIALS AND METHODS
To assess the productivity and quality of a natural grass­
land and the three introduced grasses - pangola, giant star and buffel 
- in Ghana, two experiments were undertaken. One, on a natural grass­
land site located at the University of Ghana, Legon and the other on 
the introduced grasses cultivated at the University of Ghana Research 
Farm, Legon. Each experiment was conducted in 1974 and 1975* The 
results are reported in four separate papers: 1. Dry matter accumula­
tion in natural grassland in the Accra Plains; 2. The effect of 
maturity on the quality of Sporobolus pyramidalis and Heteropogon 
contortus in the Accra Plains; 3- Dry matter accumulation in pangola, 
giant star and buffel grasses in the Accra Plains; 4* The effect of 
maturity on the quality of pangola, giant star and buffel grasses in the 
Accra Plains.
Natural Grassland
For the natural grassland trial, a split plot design was used. 
Three pretreatments (slashing, grazing and burning) formed the main 
plots and the harvest dates constituted the subplots. There were four 
replications. The pretreatment slashing was conducted with machettes, 
the herbage being cut to a stubble height of about 5 cms. For grazing, 
80 wethers from the University of Ghana Agricultural Research Station, 
Legon, grazed the appropriate plots for 12 hours. Burning was applied 
by an oxyacetylene torch. These pretreatments were imposed at the start 
of the main rainy season (April 1, 1974, April 9, 1975 (Table 0.1).
At each of the ten weekly harvesting dates, the appropriate plots were
11
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12
Table OJ. Schedule of activities in the study of natural grassland.
Item
Dates
1974
Herbage 
maturity 
in weeks
Dates
1975
Herbage 
maturity 
in weeks
Pretreatment April 1 April 9
Fertilization April 11 April 12
Harvest 1 April 29 4 May 21 6
2 May 6 5 May 28 7
3 May 13 6 June 4 8
4 May 20 7 June 11 9
5 May 27 8 June 18 10
6 June 3 9 June 25 11
7 June 17 11 July 3 12
8 July 1 13 July 16 14
9 July 15 15 July 28 16
10- July 29 17 August 13 18
Regrowth to end 
of rainy season July 29 August 13
Regrowth to mid 
dry season August 12 August 27
Regrowth to late 
dry season August 26 September 10
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13
mowed b.y means of a Jari mower with a 92 cm sickle bar attachment. 
Harvesting commenced when the herbage was about 10 cm high. To esti­
mate recovery yields for dry season use, the subplots were subdivided 
randomly into three sections and cut at three dates to span the ensuing 
dry period: the end of the major rainy season (July 29 and August 13
in 1974 and 1975 respectively), the middle of the dry period (August 
12 and 27, 1974 and 1975, respectively) and end of the dry period 
(August 26 and September 10, 1974 and 1975» respectively).
Composite soil samples of each main plot were sent to the 
Department of Land Resource Science, University of Guelph, for analysis. 
Fertilizer was broadcast at 50 kg N, 40 kg P and 50 kg K per hectare 
after the pretreatments.
In the repeat experiment in 19751 "the pretreatments and har­
vesting order were applied to the same plots to which they had been 
assigned in 1974*
Introduced Grasses
Two trials were established in March-June, 1974 (Table 0.2) 
one for harvesting during the minor rainy season 1974 (September 17 - 
December 31) and the ensuing dry period (December 31 - May 5), and the 
second for the major rainy season (April 29 - August 12) 1975 and the 
ensuing dry period (August 12 - September 9» 1975)* Pangola and giant 
star grasses were planted from sprigs, while buffel grass (variety 
Biloela*) was seeded. Fertilizer was broadcast at planting, at the
* Buffel grass (variety Biloela) from Australia by courtesy of the 
Director of Veterinary Services, Accra.
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14
Table 0.2. The schedule of activities in the study of introduced 
grasses.
Item 1974 1975
Slashing September 16 April 29
Soil sampling and 
fertilizing September 17 April 30
3 week harvest October 8 May 20
4 week harvest October 15 May 27
5 week harvest October 22 June 3
6 week harvest October 29 June 10
7 week harvest November 5 June 17
8 week harvest November 12 June 24
9 week harvest November 19 July 1
11 week harvest December 3 July 15
13 week harvest December 17 July 29
15 week harvest December 31 August 12
Regrowth to start 
of dry season December 31 August 12
Regrowth to middle 
of dry season March 4 August 26
Regrowth to end of 
dry season May 5 September 9
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15
rate of 50 kg N, 40 kg P and 80 kg K per hectare; when full ground 
cover had been achieved, a further 5° N/ha was applied.
The plots were subdivided into ten subplots randomly assigned 
to ten weekly harvest dates. To estimate regrowth yields from these 
harvests, three recovery cuts were taken as in the natural grassland 
trial. In the first trial these were cut on December 311 March 4 and 
May 5 , while in the major rainy season trial, they were cut on August 12, 
August 26 and September 9) 1975*
Sampling
Two samples of herbage were taken along the length of each 
harvested plot, and weighed together with the remaining harvest for the 
fresh yield, in all experiments. The first sample was weighed fresh 
and dried at 80°C for 48 hours in a forced draught oven, for dry matter 
determination. The second sample was stored at -4°C and later 
fractionated into leaf and stem portions of each species. The separated 
samples were dried as before and all dried samples were ground in a 
Wiley mill using a 1 mm mesh screen.
Data gathered included rainfall from a rain gauge on site, 
phenotypic notes at harvesting, dry matter yield of each plot at each 
harvest, species composition of the natural grassland, the yield of 
leaf dry matter of each species, percent total nitrogen (N) content 
and percent In Vitro dry matter digestibility (IVD) of the two dominant 
species in the natural grassland (Sporobolus and Heteropogon) and of the 
introduced species over the ten initial cuts. IVD was determined using 
the Tilley-Terry method modified by Mowat et al. (1965). N was deter­
mined using a Technicon Autoanalyzer.
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PAPER 1: DRY MATTER ACCUMULATION IN NATURAL GRASSLAND
IN THE ACCRA. PLAINS
Abstract
In order to study the accumulation of dry matter of a 
natural grassland community in the Accra Plains and the effect of time 
of cutting on the availability and growth of sward during the ensuing 
dry period, a four replicate split plot trial with three pretreatments 
and ten harvest dates was conducted during the major rainy seasons of 
1974 and 1975* The regrowth from each harvest date was harvested at 
three times, end of rainy season, mid-dry season and end of dry season.
Slashing, grazing and burning at the beginning of the rainy 
season, did not affect botanical composition or dry matter accumulation 
during development. Dry matter yield increased from 24 kg/ha at week 4 
to 5003 kg/ha at week 18.
Regrowth following the rainy season harvests was also not 
affected by pretreatment. Recovery yields increased with increasing 
regrowth periods up to 10-12 weeks to the end of the rainy season, 13-14  
weeks to mid-dry season and 12-16 weeks to the end of the dry season.
The major components of the sward were Sporobolus pyramidalis 
Beauv., and Heteropogon contortus (L.) Beauv., while small erratic pro­
portions of other grasses included guinea grass (Panicum maximum Jacq.), 
buffel (Cenchrus ciliaris L.), gamba (Andropogon ga.yanus Kuntb), 
Bothriochloa sp. Kuntze, Setaria sphacelata (Schumach) Stapf and 
Hubbard, '/etiveria fulvibarbis (Trin.) Stapf. Sporobolus started 
growth and flowered at 3-6 weeks and dominated the community until week
16
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17
7 when Heteropogon became dominant. Heteropogon grew slowly and 
flowered from week 6. Buffel, Bothriochloa sp. and Setaria sp. 
flowered within 4-8 weeks but gamba grass did not flower during the 
study. Dry matter accumulation continued after flowering until the 
end of the rainy season.
Introduction and Literature Review
The Accra Plains of Ghana is composed of approximately 
242,000 hectares of rolling savanna grassland and is located in the 
south-eastern region of Ghana. This natural grassland area provides 
the only source of feed for about 60,000 cattle.
Although many trees and forbs, including several legumes, 
are present throughout this area, grasses are the main source of feed 
(Pianu 1966, B.Sc. Dissertation, University of Ghana, Legon). The 
dominant grasses include Vetiveria fulvibarbis (Trin.) Stapf., 
Andropogon spp. L., Schizachyrium spp. Nees., Hyparrhenia spp. Anderss 
ex Pourn., Sporobolus spp., R. Br., Cenchrus spp. L., and Heteropogon 
contortus (L.) Beauv. (Loxton 1955? Rose Innes 1962, Brammer 1967).
The inadequate distribution of production from these species 
throughout the year is one of the major problems confronting the live­
stock industry in Ghana and indeed in most tropical areas. A cyclic 
pattern of live weight gain during the wet season followed by a loss 
during the dry season results in prolonged animal maturity (Lansbury 
I960, Rose Innes 1961).
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18
Two peaks of herbage production occur: in May—June during
the major rainy season and in September and October, during the minor 
rainy season. Lansbury, Rose Innes and Mabey (1965) found the growth 
of the natural grassland from the beginning of the major rainy season 
(May) through to the end of the minor rainy season (November) to be 
from 2400 to 3000 kg/ha.
There is little information on dry matter accumulation in the 
species that compose the sward, and the effect of harvest management 
during the rainy season on sward regrowth for use during the dry period. 
However Lansbury (i960) and Blair (1963) noted that production during 
the dry season is grossly deficient in supply and quality. As a result, 
herds are most frequently driven to distant areas in search of ungrazed 
herbage. Such unused herbage is coarse, dry and of poor quality (Blair 
1963, Rose Innes 1963). Frequently, burning of the ungrazed and also 
the grazed areas, is practised during the dry season to stimulate new 
growth (Hopkins 1963) Rose Innes 1971).
Although the use of stored feed during the dry season has 
been suggested as a means of overcoming the problem of feed supply and 
quality (Lansbury i960) the use of aftermath pasture should be con­
sidered, as suggested by Whyte, Moir and Cooper (1959). The provision 
of aftermath pasture is a function of cutting or grazing practises that 
are employed during the rainy season and the species of grasses in the 
sward.
The present study was undertaken to study the accumulation of 
dry matter of a natural grassland community in the Accra Plains and the 
effect of time of cutting on the availability and growth of the sward 
during the ensuing dry period.
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19
The site chosen for the experiment was a relatively uniform 
area of natural grassland on the campus of the University of Ghana.
The soil was coarse in texture and the pH varied from 5*5 6.1 with
a low phosphorus content (4-6 ppm) and a high potassium level (125-236 
ppm). Fertilizer at the rates of 50 kg N/ha, 40 kg P/ha and 50 kg K/ha 
was applied after pretreatments had been imposed.
The experiment was conducted throughout the major rainy 
seasons (April - August) of 1974 and 1975 an<i 'the ensuing dry seasons 
(August - September). It consisted of: three pretreatments (slashing,
grazing and burning) applied at the beginning of each rainy season 
(April 1, 1974; April 9» 1975) to ascertain their effects on subse­
quent species composition and production; ten cutting dates during the 
major rainy season (4- 17 weeks in 1974> 6-18 weeks in 1975) "to deter­
mine the accumulation of dry matter and changes in species composition; 
and three recovery harvests, one at the end of the major rainy season 
(July 29, 1974; August 13, 1975)? "the second at the middle (August 12, 
1974; August 26, 1975) an<i 'the third at the end of the ensuing dry 
season (August 26, 1974; September 10, 1975) to assess the effect of 
time of cutting during the rainy season on the herbage regrowth in the 
subsequent dry season.
A four replicate, split-split plot design was employed where 
pretreatments formed the main plots (20 m x 8 m) separated by paths 2 m 
wide. The ten cutting dates were randomly assigned to 2 m x 8 m sub­
plots. Each subplot was further divided into three sections 2.67 m 
x 8 m each, corresponding to the three recovery harvests.
Materials and Methods
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20
For the pretreatments of slashing and burning, machettes and 
an acetylene torch were used respectively. West African Forest wethers 
which had been fasted for 24 hours were used for the grazing pretreat­
ment. Twenty wethers were permitted to graze the respective main plot 
for a period of 12 hours.
At each harvest date, the paths were cleared and the appropriate
plots were harvested with a Jari mower. Using a 92 cm sickle bar
attachment to the mower, a swath was cut throughout the length of each 
plot. Prior to raking and weighing the cut material, two samples of 
herbage were taken along the length of the cut area. These samples 
were weighed with the remaining harvest. One sample was stored
in a freezer at - 4°C and later separated into the various grass species
components. The second sample was dried at 50,0'C for 48 hours in a forced
draft oven and used for dry matter determinations.
All data are presented in kilograms per hectare of dry matter. 
The total yield of the natural grassland, the contribution of species 
and regrowth yields were analyzed across harvest dates as well as 
within harvest dates. Differences among means were tested with Duncan's 
Multiple Range Test (Steele and Torrie i960).
Results and Discussion
In 1974, 730 mm of rain fell during the major rainy season
(April 15 to July 29) whereas in 1975, 400 mm fell from May 21 to August
13 (Appendix LIX and LX). In the dry season of 1974 (July 29 to August 
26) a total of 12 mm rain fell, all of which occurred in the week of
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21
August 5 to 12. In 1975i the total rainfall during the dry period,
August 13 to September 10, was 6 mm. In both years, rainfall increased 
to a peak in June and declined sharply in July.
The data obtained from this study (Table 1.1, Appendices IV 
and V) tend not to support the contention that burning of natural grass­
land increases yield (Doyne 1937j Semple 1970). Only during the 11th 
and 14th weeks of 1975 was the result of burning superior to slashing 
or grazing. Burning, likewise, did not affect the species composition 
(Appendices I, II, III, IV and V). Birch (i960) and Daubenmire (1968) 
attribute the superiority of burning to enhanced nitrification by 
briefly exposing the soil to drying. All pretreatments in this study 
possibly received this exposure which may explain the lack of differences.
In both years, growth commenced following the onset of rains 
at the beginning of the major rainy seasons (Fig. 1.1,, Appendices IV 
and V). Initially growth was relatively slow in both years but a marked 
increase occurred from week 4 (24 kg/ha in 1974 and 268 kg/ha in 1975) 
through to week 13 in 1974 (3839 kg/ha) and week 14 in 1975 (3870 kg/ha). 
These progressive increases contrast with reports of Lansbury et_ al.
(1965) in the Accra Plains, Norman (1963) in North Australia and 
Cassady (1973) and Taerum (1970a,b,c) in East Africa. They reported 
that herbage growth was erratic as dry matter accumulation was inter­
spersed with dry matter losses as the rainy season progressed. They 
attributed this to soil moisture depletion during intervening brief dry 
spells (Cassady 1973i Taerum 1970a,b,c), to senescent leaves being 
pounded off by heavy rainfall (Cassady 1973) and to plot variability 
(Lansbury et al. 1965).
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22
Table 1.1. Analyses of variance of yield of dry matter from natural
grassland during the major rainy seasons of 1974 and 1975*
1974 1975
Degrees Sums of Degrees Sums of
of squares of squares
Source freedom x 10^ freedom x 104
Reps 3 301.4 3 239.6
Pretreatment 2 287.7 2 284.O
Error (a ) 6 113.2 6 416.6
Dates 7 18,349-0** 9 28,302.0**
Pretreat. x 
dates 14 287.6 18 933.1
Error (b) 63 1,174.0 81 3,723.0
** Significant (P<Co.Ol).
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23
Fig. 1*1* Dry matter accumulation during the major rainy seasons of 
1974 and 1975 in natural grassland in the Accra Plains.
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24
The natural grassland in this study consisted largely of 
Sporobolus pyramidalis and Heteropogon contortus (Figs. 1.2 and 1-3* 
Appendices IV and V). There were several minor components that were 
classified as 'others'. The contribution from each of: Panicum maximum
Jacq. (guinea grass), Andropogon gayanus Kunth., (gamba grass),
Vetiveria fulvibarbis (Trin.) Stapf., Bothriochloa spp. Kuntze.,
Setaria sphacelata (Schumach) Stapf. et Hubbard., Pennisetum pedicelatum 
Trin., Galactia tenuifolia (Willd) Wight et Arn and Uraria picta Jacq. 
was variable throughout the major rainy seasons of both years.
Sporobolus dominated the community at the early growth stages 
(up to seven weeks), in both 1974 and 1975 (Figs. 1.2 and 1.3, Appendices 
I, II, IV and V). This species matured early and had flowered and set 
seed within 3 to 6 weeks. However by the 7th and 8th weeks Heteropogon 
had become dominant. It started blooming from the 6th week. Both 
species continued to accumulate dry matter after flowering as 
Brockington (1961) observed in various tropical grasses in Rhodesia.
The yield of Sporobolus increased from 13 kg/ha at 4 weeks to 1212 kg/ha 
at 13 weeks in 1974 and 1975 from 158 kg/ha at 6 weeks to 1009 kg/ha 
at 18 weeks. In contrast, Heteropogon contributed 5*6 kg/ha at week 4 
and increased to 2156 kg/ha at week 13 in 1974 and 46 kg/ha at week 6 
to 2558 kg/ha at week 18 in 1975* Among the other species, Cenchrus. 
Bothriochloa and Setaria flowered within 4 "to 8 weeks but Andropogon 
did not flower during the observation period and Vetiveria was not found 
in mature samples. The main forbs found were the legumes Uraria,
Galactia and Rhynchosia which did not occur in mature samples. Loxton 
(1955) also found these species in a survey of the natural grassland 
near the Agricultural Research Station, Nungua.
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25
M A T U R I  T Y  I N  W E E K S
Fig. 1.2. Accumulation of dry matter of species components in
natural grassland during the major rainy season of 1974 
in the Accra Plains.
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26
m a t u r i t y  i n  w e e k s
Pig. 1.3* Accumulation of dry matter of species components in 
natural grassland during the major rainy season of 
1975 in "the Accra Plains.
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27
The regrowth yields following the major rainy season harvests 
are shown in Tables 1.2 and 1.3 and Appendices VI, \TII, VIII, IX and X. 
The greatest amount of regrowth of herbage at the end of the 1974 rainy 
season (July 29) was obtained from rainy season harvests of April 29 
(5367 kg/ha) and May 6 (3927 kg/ha) after 12 and 13 weeks of recovery 
respectively (Table 1.2). The highest mid-dry season regrowths in 1974 
also came from harvests of April 29 and May 6 and amounted to 5796 and 
5163 kg/ha respectively, the recovery period being 14-15 weeks.
Recovery to the end of the dry season, on the other hand, was greatest 
for harvests of April 29 to June 3 (with the exception of May 13) and 
the dry matter regrowth ranged from 4047 kg/ha to 5221 kg/ha (except 
May 13) for the regrowth periods of 12-17 weeks. In contrast with 
these trends, the 1975 recovery yields to the end of the rainy season 
(August 13) was greatest for harvests of May 21 to June 4: 3098 to
3489 kg/ha for recovery periods of 10-12 weeks (Table 1.3). Mid-dry 
season recovery yields from May 21 and 28 were the greatest: 508 3 kg/ha
and 4747 kg/ha for 14 and 13 weeks recovery respectively while end of 
dry season recover:/ from May 21 was the greatest recovery yield:
6107 kg/ha for 16 weeks recovery. Over the two years, it would appear 
that for the highest dry matter accumulation up to the beginning of the 
dry season, a minimum of 10-12 weeks should be allowed, while for 
regrowth up to the middle of the dry season at least 13 to 14 weeks 
recovery period is required and maximum recovery to the end of the dry 
season requires 12 to 16 weeks.
Growth occurred in both years during the dry period, this 
being more pronounced in 1975 (Tables 1.2 and 1.3). In the first half 
of the dry period of 1974 (July 29 - August 12) growth was erratic but
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Table 1.2. Regrowth yield from natural grassland during the dry period (July - August) 1974*
(Kg/ha dry matter)
Harvest date 
in
End of rainy season
(July 29)
Mid dry 
(Aug.
season
12)
End of dry season 
(Aug. 26)
rainy season Recovery"1- Yield Recovery4- Yield Recovery-*- Yield
April 29 13 5367S 15 5796|° 17 4499y°
May 6 12 3927y 14 5163^ 16 52215
May 13 11 2362^ 13 2153* 15 1399°
May 20 10 295l£ 12 3357xC 14 43505
May 27 9 2093J 11 4322° 13 41455
June 3 8 2123^ 10 2894$ 12 40475
June 17 6 923£ 8 1724$ 10 3875x
July 1 4 548° 6 1723^ 8 2955*
July 15 — 4 655d 6 1678c
July 29 — — 4 1623c
+ Weeks from date of harvest during the major rainy season.
o Data followed by the same letter are not different (P>0.05): columns - a,b,c,d; rows - x,y.
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Table 1.3. Regrowth yield from natural grassland during the dry period (August - September) 1975*
(Kg/ha dry matter)
Harvest date 
in
End of rainy season 
(Aug. 13)
Mid dry 
(Aug.
season
27)
End of dry season 
(Sept. 10)
rainy season Recovery+ Yield Recovery"*" Yield Recovery"*" yield
May 21 12 3489z° 14
a°
5083y 16 6l07x
May 28 11 3456* 13 47475 15 5071^
June 4 10 30982 12 3947^ 14 5090^
June 11 9 2226y 11 3 ° !6 c 13 5110h
June 18 8 12660 10 250l£ 12 2819$
June 25 7 895 ° 9 I837y 11 2384$
July 3 6 575* 8 978y 10 2233CCI
July 16 4 321* 6 81 ly 8 2042®d
July 28 128* 4 690e 6 1342deX
August 13 - — 4 690e
+ Weeks from date of harvest during the major rainy season.
o Data followed by the same letter are not different (P>-0.05): columns - a,b,c,d,e; rows - x,y,z.
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30
in the second half (August 12 - 26), there was growth in herbage 
harvested after June 3 (Table 1.2). Thus herbage that was 9 weeks or 
less in maturity, continued to grow during the second half of the dry 
season. In 1975, growth occurred throughout the dry period (Table 1.3) 
although there was less rainfall during the 1975 dry period (6 mm) than 
in the 1974 dry period (12 mm). This difference may be related to the 
lower rainfall during the major rainy season of 1975 (400 mm) than in 
1974 (730 mm). In 1975 the herbages may have been conditioned to grow 
with low rainfall whereas in 1974 they may have been conditioned to 
grow with high moisture supply.
Dry matter losses occurred in some instances during the dry 
season. In 1974> following the harvest of April 29, a loss of 1297 
kg/ha occurred between the middle and the end of the dry season. Also 
after the May 13 harvest, 754 kg/ha of dry matter was lost. These 
losses may be related to the induction of senescence by dry spells 
during the rainy season and the stimulation of growth during the dry 
season by light rainfall. The rapid decline of precipitation following 
the higher rainfall regime during the major rainy season 1974 may have 
induced senescence in the mature herbage during July - August. As rain­
fall in 1975 was lower during the major rainy season than it was in 
1974, the plants may have been conditioned to moisture stess in 1975* 
Thus, the light showers of the dry season of 1975 may have been better 
utilized by the herbage for growth than in 1974* These fluctuations in 
dry matter yield in response to drought spells alternating with rainfall 
are in agreement with reports of Cassady (1973) and Taerum (1970a,b) in 
East Africa and Lansbury et al. (1965) in the Accra Plains.
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Table 14* Seasonal total yield from natural grassland during 1974 (upper) and 1975 (lower).
(Kft/ha dry matter)
1974
__ tf.-.- . _.... .
Maturity
4 +
Apr. 29*
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 17
13
July 1
Main
end
season + 
of rainy season 5391ac
) 3979bc 2528e 3438cd 2886de 3598bcd 4084bc 4387b
Main season + 
mid dry season 5820a 5215ab 2329d 3844c 5115ab 4396bc 4685ab 5562a
Main
end
season + 
dry season 4523b 5273b 1575c 4837b 4938b 5522b 7036a 6794a
1975
Maturity
6+ 
May 21*
7
May
8
28 June 4
9
June 11
10 
June 18
11 
June 25
12 
July 3
14 16 
July 16 July 28
18 
Aug. 13
Main
end
season + 
of rainy season 3757'bc0 3954bc 3800bc 3695bc 3352c 3633bc 36O3bc 4191b 3974bc 5003a
Main season + 
mid dry season 5351a 5245ab 4649abc 4485^0 4587abc 4575abc 4006c 468labc 4536abc -5003ab
Main
late
season +
: dry season 6375ab 5569bcd 5792abcd 6579a 4905d 5122cd 526lcd 58l9ab<3 5l88cd 5688bcd
+ Weeks from pretreatment: April 1, 1974? April 9» 1975-
+ Harvest date during the major rainy season.
o Data followed by the same letter are not different (P^ - 0.05): rows - a,b,c,d.
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33
Summary
Slashing, grazing and burning pretreatments did not affect 
yield of species composition of the sward during the major rainy 
season, nor did they affect herbage recovery after harvesting in the 
major rainy season.
Growth was slow initially until week 9» at which stage 1400 
kg/ha of dry matter had been accumulated. This period of grand growth 
continued until week 13 in 1974 when dry matter accumulation amounted 
to 3838 kg/ha. In 1975» the grand growth period continued until week 
18 and 5003 kg/ha of dry matter accumulated.
Sporobolus dominated the community at the early stages in 
both years; from week 7 Heteropogon became dominant. Sporobolus 
flowered in week 6. Dry matter accumulation continued after flowering. 
Several minor species occurred in erratic minute quantities in the 
sward. Among these were Cenchrus ciliaris, Bothriochloa bladhii, and 
Setaria sphacelata, which flowered in 4-8 weeks; and Andropogon gayanus, 
which did not flower during the observation period.
Regrowths following the rainy season harvests amounted to 
about 3000 - 5300 kg/ha at the end of the rainy season, 4700 - 58OO 
kg/ha by the middle of the dry period and 4000 - 6000 kg/ha at the end 
of the dry period. During the dry period (August) 1974 herbage growth 
ceased after 8-9 weeks of regrowth whereas in 1975 growth continued 
during the dry period.
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PAPER 2: EFFECT OF MATURITY ON QUALITY OF SPOROBOLUS PYRAMIDALIS
AND HETEROPOGON CONTORTUS IN THE ACCRA PLAINS
Abstract
The effects of slashing, grazing and burning pretreatments 
and ten sampling dates on leaf production, In Vitro dry matter diges­
tibility (IVD) and nitrogen (N) of leaves, stems and whole plants of 
Sporobolus pyramidalis Beauv. and Heteropogon contortus (L.) Beauv. 
during the major rainy seasons of April 1 - July 29, 1974 and April 9 
- August 13, 1975 were studied in Legon, Ghana.
The pretreatments did not affect leaf production, IVD or N. 
Leaf production increased with maturity, reaching a plateau between 
11 and 13 weeks in 1974} in 1975, Sporobolus increased in leaf production 
until 16 weeks, while Heteropogon reached a plateau at week 10. During 
the early stages of growth Sporobolus produced more leaf dry matter than 
Heteropogon but between the 8th and 9'th weeks of maturity, this was 
reversed. Sporobolus tended to have a higher proportion of leaves than 
Heteropogon, particularly at the mature stages. Heteropogon was higher 
in whole plant IVD (57*0 - 62.0$ in weeks 7—9> falling to 27.0 - 38.0$ 
in weeks 17-18) than Sporobolus (51-6 - 59*0$ declining to 29-8 - 34*3$). 
Leaves were more digestible than stems in both species.
Nitrogen declined with maturity in leaves, stems and whole 
plants. Leaves contained more N (1.80 - 2.00$ at 7-9 weeks to 0.88 -
1 .00$ at 17-18 weeks) than stems (0 .82 - 1 .74$ a-t 7-9 weeks to O .34 - 
0.4&$ at 17-18 weeks). Species differences in whole plant and leaf N 
content were not consistent but Sporobolus stems were higher in N than
34
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35
Heteropogon stems. The correlation between IVD and N was 0.61. This 
was significant.
Introduction and Literature Review
The rapid deterioration in quality of tropical natural grass­
land species as the herbage matures has been reported by Paterson (1933) 
and Anon. (1941) in the West Indies, French (1957) in East Africa, and 
Rose Innes (1961) and Blair (1963) in Ghana, and Miller (i960) in 
Nigeria. Such decreases in quality pose one of the problems confronting 
the livestock industry in Ghana (Lansbury, Rose Innes and Mabey 1965).
In Ghana, Sen and Mabey (1965) reported that the nitrogen (n) 
content of Sporobolus pyramidalis Beauv. decreased from 2.29% at 4 weeks 
to O.69% at 36 weeks maturity. The decrease in N content however may 
not be constant, for with Heteropogon contortus (L.) Beauv., they 
reported that N decreased from 1.15% at 4 weeks to 1.01% at 8 weeks 
and increased to 2.50% at 36 weeks. They attributed this to high rain­
fall.
In temperate forages, the importance of leafiness of species 
to quality of forages has been documented (Mowat al. 1965) because 
the leaves contribute a large fraction of digestible nutrients. Few 
reports of the importance of the role of leafiness on tropical natural 
grassland species are available. Norman (1963) in Northern Australia, 
reported a decline in leafiness from 58% and 74% at 6 months for 
Themeda triandra Forsk., and Chrysopogon sp. Trin., respectively, to 
38% and 42% at 8 months. The importance of leaf proportion to quality
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36
was not outlined. Likewise information on the digestibility of leaves 
and its importance to total quality of tropical species is limited.
Palvey (1977) in Northern Australia, compared in vitro dry 
matter digestibility (IVD) of natural grassland species with that of 
introduced ones and found that the native species were lower in diges­
tibility than the introduced species. Also, Gohl (1975) reporting work 
on Sporobolus pyramidalis in Rhodesia, gave the dry matter digestibility 
as 63.4fo, but the stage of maturity was unspecified. No research has 
been conducted on the changes with maturity in the digestibility of the 
dominant grasses in the natural grasslands of Ghana, nor has the impor­
tance of leaf to quality been investigated in Ghana.
This study examined the changes in quality of Sporobolus 
pyramidalis Beauv., and Heteropoffon contortus (L.) Beauv., two dominant 
species in a natural grassland community in the Accra Plains, during 
growth and development, in the major rainy seasons of 1974 and 1975*
The parameters investigated included leaf production, IVD, and N content.
Materials and Methods
The study was conducted at the University of Ghana in the major 
rainy seasons of 1974 and 1975* A four replicate, split plot design 
was used with three pretreatments: slashing, grazing and burning forming
the main plots and ten weekly harvesting dates the subplots. Pretreat­
ments were applied on April 1, 1974 and April 9> 1975* Cutting began 
on the 4'th week following application of pretreatments and continued 
through to the 17th week in 1974 and in 1975 began on the 6th week and
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37
terminated, on the 18th week.
At each of these cutting dates, the appropriate subplots were 
cut to about 5 cms stubble height, using a Jari mower with a 92 cm 
sickle bar attachment. Samples of the cut material were taken at random 
along the length of the subplots. Each sample was separated into three 
groups - Sporobolus, Heteropogon and other species. Sporobolus and 
Heteropogon were further separated into leaf and stem fractions. These 
fractions were dried in a forced draught oven at 80°C for 48 hours, 
weighed to determine leaf-stem proportions and ground using a Wile/ mill 
with a 1 mm screen. The leaves included sheaths while the inflorescence 
and seeds were included in the stem portions.
In Vitro dry matter digestibility (IVH) and total nitrogen (N) 
were determined on the leaves and stems of Sporobolus and Heteropogon 
at the Crop Science Department, University of Guelph. A modified Tilley 
and Terry method (Pritchard, Polkins and Pigden 1963> Mowat et al. 19^ 5) 
and a Technicon autoanalyzer were used for the IVD and N respectively. 
Owing to small sample sizes, N determinations were not conducted on the 
first three cutting dates. The data were analyzed as a split plot 
design (Steele and Torrie i960). The analyses of variance were conducted 
on the dates, plant part or species separately to bring out differences 
across dates, between species and between leaf and stem fractions. 
Regressions of ^  IVD and $ N on maturity dates were calculated and the 
lowest order equations of best fit at the 5% level adopted.
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38
The dry matter yield of the leaves of Sporobolus and 
Heteropogon at each harvest date did not appear to' be affected by the 
pretreatments of slashing, grazing and burning (Appendices XIII and XIV). 
In 1974i the yield of leaf progressively increased reaching 878 kg/ha 
and 1315 - 1360 kg/ha for Sporobolus and Heteropogon respectively, 
between 11 and 13 weeks (Table 2.1). In 1975, however, the highest 
leaf yield was attained for Sporobolus and Heteropogon in the 11th and 
9th weeks, respectively, reaching a plateau of yield that extended to 
the 18th week (Table 2.2). Sporobolus appeared to have a higher leaf 
percentage in its dry matter yield than Heteropogon. especially at the 
mature stages. In 1974, leaf dry weight, as a percentage of dry matter 
yield fell from 85 at week 4 to 72 at week 13 in Sporobolus and from 83 
to 63 in Heteropogon. In 1975, leafiness declined from 92$ at 6 weeks 
to 74$ at week 18 in Sporobolus while in Heteropogon it decreased from 
100$ to 47$.
IVD was not affected by pretreatment in a consistent manner 
(Appendices XV, XVI, XVII, XVIII, XIX, XX, XXI, XXII, XXIII and XXIV).
In 1974, burning resulted in higher IVD values than slashing and grazing 
in Sporobolus whole plants in weeks 8, 9 and 11 whereas in Heteropogon, 
grazing gave superior IVD’s in weeks 8 and 15* There were no pretreat­
ment effects in the other weeks. In 1975, there were no significant 
IVD differences among the three pretreatments. Therefore the IVD 
averages of each species for all pretreatments were used to determine 
the effect of maturity on IVD of leaves, stems or whole plants separately.
Results and Discussion
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39
Table 2.1. Yield of leaves of Sporobolus and Heteropogon during the 
major rainy season of 1974*
(Kg/ha dry matter)
Maturity"1" Sporobolus Heteropogon
4 11 d° ■ 5 c
5 27 d 8 c
6 75 d 61 c
7 172 cd 203 C
8 247 cd 337 be
9 394 be 677 b
11 554 b 1315 a
13 878 a 1360 a
+ Weeks from pretreatment: April 1, 1974
o Data followed by the same letter are not different (P_> 0.05): 
columns - a,b,c,d.
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40
Table 2.2. field, of leaves of Sporobolus and Heteropogon during the 
major rainy season of 1975*
(Kg/ha dry matter)
Maturity"1” Sporobolus Heteropogon
6 137 e.° 46 c
7 188 e 130 c
8 260 de 290 be
9 302 cde 65O ab
10 484 cde 978 a
11 704 abed 1095 a
12 725 abc 1187 a
14 917 ab 1195 a
16 1006 a IO36 a
18 821 abc 1221 a
+ Weeks from pretreatment: April 9, 1975
o Data followed by the same letter are not different (PJ>0.05): 
columns - a,b,c,d,e.
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41
The leaves of both Sporobolus and Heteropogon were generally 
more digestible than their stems (Pigs. 2.1 and 2.2). This agrees with 
reports by Tilley and Terry (1963) and Mowat et al. (1965) on mature 
temperate grasses and by Laredo and Minson (1973) on Panicum coloratum 
and P. maximum, in Australia. The contention that stem digestibility 
is higher than leaf digestibility at very early growth stages as repor­
ted by Mowat et al. (1965) in orchard and brome grasses could not be 
fully confirmed in this study because of inadequate stem tissue at the 
early stages. However the higher values for stem IVD than leaf IVD in 
Heteropogon at weeks 7, 9 and 11 in 1974 and at weeks 10 and 11 in 1975 
(Pigs. 2.1 and 2.2, Appendices XXII and XXIII) tend to confirm this 
contention at least for some species.
The digestibility of the leaves of the two species did not 
differ consistently until weeks 7 and 9 in 1974 and 1975 respectively.
Prom then on, the Heteropogon leaves tended to be more digestible than 
those of Sporobolus. The stems of Heteropogon, however, tended to be 
more digestible than those of Sporobolus at all harvest dates in both 
years. Heteropogon whole plant IVD tended to be higher than whole 
Sporobolus throughout the growing season (Table 2.3, Appendix XXIV), 
this apparently being a reflection of the higher proportion of stem 
tissue in Heteropogon than in Sporobolus and the higher IVD of Heteropogon 
stems.
With advancement in maturity, IVD of leaf and stem and whole 
plants declined (Pigs. 2.1 and 2.2, Table 2.3). This observation agrees 
with those of Reid et al. (1975)» Johnson and Pezo (1975)1 Danley and 
Vetter (1973) and Moore and Mott (1973). However, there was an increase 
in leaf IVD at the early stages of growth in 1974 (Fig* 2.1) before the
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% 
IN 
VI
TR
O 
DI
GE
ST
IB
IL
IT
Y
42
MATURITY IN WEEKS
Fig. 2.1. Percent In Vitro dry matter digestibility of leaves and
stems of Sporobolus and Heteropogon during the major rainy
season of 1974.
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IN 
VI
TR
O 
DI
GE
ST
IB
IL
IT
Y
43
MATURITY IN WEEKS
Fig. 2.2. Percent In Vitro dry matter digestibility of leaves and
stems of Sporobolus and Heteropogon during the major rainy
season of 1975-
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Table 2.3. Percent In Vitro digestible dry matter of whole plants of Sporobolus and Heteropogon
during the major rainy seasons of 1974 (upper) and 1975 (lower).
1974 Maturity
7+ 8 9 11 13 15 17
May 20* May 27 June 3 June 17 July 1 July 15 July 29
Sporobolus 58.9a0
X
59.7a 53. 49-2° 46.7$ 40.9® 34-3^
Heteropogon 6l.9| 59.8a 56.9* 53.2° 43*4d 44-7* 37*7®
1975 Maturity
9+ * 10 11 12 14 16 18June 11* June 18 June 25 July 3 July 16 July 28 Aug. 13
Sporobolus 51.6a° 51.8a 46.8 f 43-7x° 4 1.2£d 37-8^ 29-81
Heteropogon 56-5|b 57-9* 53.2$ 46.5x 45-3£ 38.7| 27.21
+ Weeks from pretreatment: April 1, 1974» April 9» 1975*
+ Harvest date during the major rainy season.
o Data followed by the same letter are not different (PJXO.O5): rows - a,b,c,d,e; between species -
x,y.
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45
decline began. Such initial rise has been ascribed to rapid growth by 
Drapala, Raymond and Crampton (1947) and Haggar and Ahmed (1970). In 
the present study, however, while the initial increase in IVD fell within 
the period of "grand growth", the decline started before the rapid 
growth phase was over. The decline in IVD with herbage maturity has 
been attributed to increasing lignification by Moore and Mott (1973) 
and Crampton and Maynard (1938), to an increase in silica by Van Soest 
and Jones (1968), and to a rise in hemicellulose content by Danley and 
Vetter (1973)• Owing to inadequacy of samples, these cell wall com­
ponents were not studied.
In 1974» the relations between percent IVD and maturity was 
quadratic in the leaves of Sporobolus (Table 2.4) but linear in stems 
and whole plants. However in the leaves, stems and whole plants of 
Heteropogon, IVD was linearly related to maturity. In 1975t the regres­
sions were linear in both plant parts as well as whole plants of both 
species (Table 2.4). With Heteropogon in both years stem IVD tended to 
decline faster than leaf IVD whereas in Sporobolus such differences 
were not evident (Table 2.4 , Figs. 2.1 and 2.2). Heteropogon stems 
appeared to fall in IVD more rapidly than those of Sporobolus in both 
years. With leaves, however, the converse was true. On the other hand, 
differences in whole plant IVD of the two species did not show any con­
sistent trend in the two years (Table 2.3). In 1974i the IVD of 
Sporobolus whole plant appeared to decline at a similar rate to that of 
Heteropogon whole plant, whereas in 1975 the rates of decline tended 
to be lower in Sporobolus than Heteropogon.
The pretreatment of slashing, grazing and burning did not 
appear to have any consistent effect on N content (Appendices XXV, XXVI,
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Table 2.4. Regressions of percent In Vitro digestibility on maturity in Sporobolus and Heteropogon during
the major rainy seasons of 1974 and 1975*
1974
2r 1975 2r
Sporobolus
Whole plant Y = 76.91 - 2.44x O.85** Y = 73.63 - 2.32x O.57**
Leaf IVD Y = 49.83 + 2.57x - 0.20x2 0.78** Y = 73-09 - 2.13x O.59**
Stem IVD Y = 70.13 - 2.14x 0 .60** Y = 61.66 - 2.29x 0.39**
Heteropogon
Whole plant Y = 78.63 - 2.34 0.83** Y = 87.81 - 3.22x 0 .82**
Leaf IVD Y = 65.98 - 1.23x 0 .51** Y = 73.22 - 2.00x O.65**
Stem IVD Y = 83.42 - 2.83x O.87** Y = 90.68 - 3.63x O.52**
** Significant (P < 0.01). 
x Weeks.
Y In Vitro dry matter digestibility.
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47
XXVII, XXVIII, XXIX, XXX, XXXI and XXXII). Thus, this study did not 
confirm the view that burning herbage increases nitrogen content 
(Plowes 1957, Mes 1958, Smith i960). Similarly, species differences in 
N content were not consistent.
Percent N generally fell as the plants matured (Pigs. 2.3 
and 2.4, Tables 2.5 and 2.6, Appendices XXVIII and XXXIX) in the leaf, 
stem and whole plant of both species, thus agreeing with reports by 
Brockington (1961), Lansbury et al. (1965), and Sen and Mabey (1965). 
Linear relations were found between percent N content and maturity in 
the leaves and stems of both species in 1974* On the other hand, in 
1975» quadratic and cubic regressions were given by stem N of 
Sporobolus and Heteropogon respectively, while their leaf N contents 
had linear relations with maturity (Table 2.6). In both species leaf 
N, like the IVD, tended to fall more rapidly than stem N in both 
years. Furthermore, N levels in the leaves of both Sporobolus and 
Heteropogon were higher than in the stem (Pigs. 2.3 and 2.4, Appendices 
XXVIII and XXIX) as was the case in IVD's. Thus, percent IVD and per­
cent N of the two species seemed to be associated. The correlation 
coefficient of IVD on N in this study was 0.61 and was highly sig­
nificant (P < 0.01). This association between IVD and N agrees with 
findings by Sullivan (1969), Maynard and Loosli (1969), Leng (1973) 
and Ventura et_ al. (1975) who explained the association between IVD 
and N by nitrogen being a necessity for proper rumen microbial fer­
mentation. However, this contrasts with the results of Sullivan (1964) 
and Oh, Baumgardt and Scholl (1966) who had low correlation coefficients 
of 0.24 and 0.37 respectively. The inconsistency in the relations 
between IVD and N probably arises from the different behaviour of the
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NI
TR
O
GE
N
49
MATURITY IN WEEKS
Fig- 2.4* Percent nitrogen content of leaves and stems of Sporobolus
and Heteropocon during the major rainy season of 1975*
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Table 2.5. Percent nitrogen in whole plants of Sporobolus and Heteropogon during the major rainy 
seasons of 1974 (upper) and 1975 (lower).
1974 Maturity
7+ * May 20
8
May 27
9
June 3
11 
June 17
13
July 1
15
July 15
17 
July 29
Sporobolus 1.87| 1.65$ 1.26° I.i9£d 1.10$ 0.82| O.85I
Heteropogon 1.845 1.35J 1.42$ 1 .1 35 1.00$ 0.781 0.72|
1975# Maturity
*June 11
10 
June 18
11 
June 25
12 
July 3
14
July 16
16 
July 28
18 
Aug. 13
Sporobolus 1-785 1-25$ 1.20* 1.01$ 0.93§d 0.92yd O CO 0
Heteropogon 1-731 1.271 1.20* o.92£
Oco0 0.71de 0 .64®
+ Weeks from pretreatment: April 1, 1974! April 9i 1975*
+ Harvest date during the major rainy season.
# Average of two pretreatments: slashed and grazed.
o Data followed by the same letter are not different (P_>-0.05): rows - a,b,c,d,e; columns within
rows - x,y.
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Table 2.6. Regressions of percent nitrogen on maturity in Sporobolus and Heteropogon during the major
rainy seasons of 1974 and 1975*
1974
2r 1975
2r
Sporobolus
Whole plant N Y = 2.36 - 0.097* 0.67** Y = 2.20 - 0.08x O.52**
Leaf N Y = 2.49 - 0.097* 0.71** Y = 2.56 - 0.102x O.54**
Stem N Y = 1.78 - 0.082x 0.71** Y = 5.26 - 0.620x + 0.020x2 O.74**
Heteropogon
Whole plant N Y = 2.42 - 0.107x 0.80** Y = 2.41 - O.llx 0.71**
Leaf N Y = 2.27 - 0.076x 0.70** Y = 2.56 - 0.097x O.67**
Stem N Y = 0.99 - 0.035x 0.51** Y = 9-34 - 1.837x + 1 .246x2 O.84**
- 2.798x3
** Significant (P < 0.01). 
x Weeks.
Y Percent nitrogen.
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two parameters in the plant. In this study for instance, N declined 
more rapidly in the leaf than in the stem whereas IVD fell faster in 
the stem than in the leaf. Furthermore, while IVD levels tended to 
be higher in Heteropogon than Sporobolus, the latter had higher nitrogen 
content.
As regards the nitrogen and digestible dry matter production 
of the natural grassland sward and herbage intake by ruminants in the 
grassland, more information is required on the quality of the other 
species in the community (e.g. gamba and guinea grasses), during the 
major rainy season, and on the quality of the component species during 
the regrowth period, to help determine when these natural grassland 
swards should be grazed. Certain general inferences may however be 
drawn from the data on the two dominant species in these communities.
From the viewpoint of dietary nitrogen for ruminants, both 
Sporobolus and Heteropogon were too low in nitrogen content to meet the 
requirements for high levels of animal production, such as the 2.5$ 
suggested by Sullivan (1969). These grasses contained enough nitrogen 
to support moderate levels of livestock production at the early stages 
of growth but the yield of the sward was low at these stages. On the 
other hand as the yield rose, nitrogen fell, and if the 0.8 nitrogen 
suggested by Fianu, Attakrah and Koram (1972) as the minimum for main­
tenance requirements in yearling wethers be accepted, then whole plant 
Sporobolus would barely continue to meet this minimum requirement until 
17 weeks of maturity and Heteropogon would only do so until week 13.
To meet the needs of the rapidly developing livestock industry 
in Ghana therefore, introduced species may be examined. Where natural 
grassland is the only forage, supplementation with nitrogenous concen­
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trates such as urea may be necessitated. Overseeding with legumes 
would also furnish an alternative method of meeting the dietary 
nitrogen requirements of ruminants in the Accra Plains.
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Summary
The pretreatments: slashing, grazing and burning did not
differ consistently in their effects on leaf production, nitrogen 
content or In Vitro dry matter digestibility. However, burning, being 
relatively inexpensive, would seem preferable to slashing as a tool 
for removing mature growth from small holdings. Long term effects on 
the sward and safety precautions would, however, have to be considered 
before vegetation is burned.
Sporobolus produced more leaf dry matter than Heteropogon 
until weeks 5 to 6 when this trend was reversed. Leaf production rose 
with maturity up to 554 - 878 kg/ha in Sporobolus and 1315 - 1360 kg/ha 
in Heteropogon at 9-11 weeks of maturity. Sporobolus appeared to have 
a higher proportion of leaves than Heteropogon especially at later 
stages of growth.
The leaves of both species were more digestible and contained 
higher levels of nitrogen than the stems. Although nitrogen content 
of whole plants showed no consistent differences between the two species, 
Heteropogon was more digestible than Sporobolus.
Quality declined with maturity in both species. The rate of 
decline tended to be higher in Heteropogon than in Sporobolus and the 
latter would probably continue to meet the maintenance nitrogen require­
ments of the grazing ruminant until week 17 whereas Heteropogon would 
only do so until week 1 3.
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PAPER 3: DRY MATTER ACCUMULATION IN GIANT STAR, BUFFEL
AND PANGOLA GRASSES IN THE ACCRA PLAINS
Abstract
In two separate trials, giant star (C.ynodon plectostachyus 
(K. Schum.) Pilger, buffel (Cenchrus ciliaris L. cv. Biloela and 
pangola (Digitaria decumbens Stent.) grasses were harvested at ten 
maturity dates during (a) the minor rainy season of 1974i and (b) the 
major rainy season of 1975* To determine the amount of aftermath 
herbage for use in the ensuing dry period, one-third of each plot was 
harvested at end of rainy season (December 31, 1974 or August 12, 1975)» 
another third at mid-dry season (March 4» 1975 or August 26, 1975)* 
and the last third at end-dry season (May 5? 1975 °r September 9j 1975)*
During early growth, pangola was sensitive to moisture stress 
and failed to grow during the minor rainy season trial. Buffel grass 
on the other hand, being drought tolerant, grew under the light showers.
It flowered from week 3 in both seasons while giant star flowered only 
during the minor rainy season trial, at week 6, and pangola flowered 
in week 6 during the major rainy season. In all three grasses growth 
continued after flowering. In buffel, senescent leaves tended to 
remain on the plant whereas in the stoloniferous grasses, they were 
stripped off by rainfall.
During the minor rainy season, September 16 - December 31, 1974, 
giant star and buffel produced similar dry matter yields (maxima of 
4715 and 5707 kg/ha respectively at week 13)* In the major rainy season 
(April 29 - August 12), however, buffel was superior to the prostrate
55
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56
grasses which did not differ consistently. Yields rose to 5^96, 6212 
and 4566 kg/ha in giant star, buffel and pangola grasses respectively 
at week 9 and levelled off.
Herbage regrowth from the minor rainy season harvests to the 
end of the rainy season, was highest after the harvests of October 8 
and 15: 2384 - 2812 kg/ha in giant star and 4560 kg/ha in buffel. For
mid-dry season, the highest regrowth herbage was obtained after harvests 
of October 22-29 in giant star (3037 - 3789 kg/ha) and October 8-29 in 
buffel (3400 - 455° kg/ha). At the end of the dry season, the regrowth 
herbage yields from all rainy season harvest dates were the same in 
both species: 1976 - 3756 kg/ha in giant star and 2890 - 5207 kg/ha
in buffel.
In the major rainy season trial the highest recovery yields 
at the end of the rainy season (August 12) were 4110 kg/ha for giant 
star after May 20 harvest, 5951 - 6508 kg/ha for buffel and 4998 - 5456 
kg/ha after May 20-27 harvests. At mid-dry season (August 26) giant 
star had accumulated regrowth of 46OO - 5106 kg/ha after May 20 - 
June 3 harvests; buffel had 5791 kg/ha after the May 20 harvest and 
pangola had 4554 - 5602 kg/ha after the harvests of May 20-27. At end- 
dry season (September 9) the maximum recovery yields were 6379 kg after 
May 20 for giant star, while for buffel and pangola it occurred between 
May 20 and June 10: 4572 - 5646 kg/ha for buffel and 4588 - 5626 for
pangola.
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Introduction
Herbage on the natural grasslands in the Accra Plains has 
been reported to deteriorate rapidly in both quality and dry matter 
production in the dry season (Lansbury i960). This deterioration is 
responsible for retardation in the growth of grazing livestock (Rose 
Innes 1961)* Several grass introductions have been made since the 
1920’s in attempts to solve this problem. Among these are: pangola
(Digitaria decumbens Stent.); giant star grass (c.ynodon plectostach.yus 
(K. Schumach) Pilger) and buffel (Cenohrus ciliaris L. syn. Pennisetum 
ciliare L.); these were judged to be promising on the basis of their 
ease of propagation, and vigorous growth (Evans 19611 Thompson 1961, 
Rose Innes 1961, Asare 1972).
Several studies have been conducted on these grasses in East 
Africa (French 1957, Cassady 1973» Taerum 1970a,b,c), in Rhodesia 
(Brockington 1961), in the Caribbean (Nestel and Creek 1962, Vicente- 
Chandler et_ al. 1964j Sallette 1965), in Australia (Humphreys 1969), 
and in the forest zone of Ghana (Asare 1970). However little infor­
mation is available to describe their dry matter accumulation during 
growth and development. Furthermore, little is known about their pro­
ductivity in the Accra Plains.
This study was conducted to appraise these three species for 
dry matter accumulation during growth in the major and minor rainy 
seasons and to assess the regrowth yields of herbage for use in the 
ensuing dry season after cutting in the rainy season.
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Pangola grass is a sod. forming stoloniferous perennial
adapted to the humid tropics and subtropics over a wide range of soils
(Hosaka and Goodell 1954 i Nestel and Creek 19 6 2 , Sallette 1 9 6 5>
Vicente-Chandler 19 75 )*  Because of its creeping habit, pangola is more
suited to grazing than cutting (Nestel and Creek 1962 ) and Vicente- 
Chandler et_ al. (1964) advise that it should be grazed with rest periods 
of 20 to 30 days, while Brown et al. ( 1966b ) recommend 30 to 40 days 
for proper regrowth in the rainy season, and 60 days in the dry season. 
Pangola responds to nitrogen fertilization, and in Puerto Rico, 
Vicente-Chaiidler &t al. ( 1964) obtained dry matter increases from 
1 3 ,4 4 0  kg/ha at zero N to 3 0 ,2 4 0  kg/ha at 448 kg/ha N and Sallette 
( 1965)  and Plucknett (1 9 7 0 ) obtained increased response to higher N 
level than those of Vicente-Chandler et al. ( 1964) .
Giant star grass is a perennial occurring naturally in associa­
tion with buffel grass, guinea grass (Panicum maximum Jacq.) and 
Brachiaria brizantha (Hochst) Stapf. in East Africa (Edwards 1956)
Rattray i 960) .  Ahlgren et al. ( 1 9 5 9 )» Ruthenberg (1 9 7 4 ) and Chedda
(1 9 7 1 ) recommend it as suited to West Africa. At Avetonou in Togo, 
giant star grass under fertilization yielded 50 tonnes/ha fresh herbage 
per year (Ruthenberg 1 9 7 4 )*  Caro-Costas et al. (1 9 7 3 ) and Vicente- 
Chandler (1 9 7 5 ) considered giant star superior to pangola as it is 
leafier, higher in N and produces more dry matter per hectare.
Buffel grass (Cenchrus ciliaris L. synonym Pennisetum ciliare 
L.) is a tufted, and often weakly rhizomatous, perennial that occurs in 
a r id  areas under as little as 300 mm rainfall per year. Though
Literature Review
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59
susceptible to fungal attack in humid environments, Asare (1972) found 
that this grass produces well in the West African forest zone if the 
soil is well drained. He obtained 26.1 tonnes/ha and 11.9 tonnes/ha 
dry matter in the first and second years of establishment, with 67 kg 
N, 90 kg P and 45 kg K per hectare while at Bouake, Ivory Coast, 
Ruthenberg (1974) reported 14*2 tonnes/ha.
Materials and Methods
Two trials each containing the three species were established 
at the University of Ghana Research Farm, Legon, in March to June 1974- 
A split plot design with four replicates was used, the species forming 
the main plots 5 x 20 m in size, and ten harvesting dates were randomly 
assigned to the subplots. These subplots were further randomly divided 
into three sub-subplots for harvesting at the end of the rainy season, 
mid-dry season and at the end of the dry season. The first tiral 
commenced at the start of the rainy season (October 8 - December 31) 
1974, and continued into the ensuing dry period through May 5* The 
second trial commenced during the major rainy season of 1975 (May 20 - 
August 12) and continued into the following dry period - through 
September 9*
Pangola grass and giant star grass were established from 
cuttings planted at 15 x 15 cm. The cuttings were obtained from the 
Agricultural Research Station (ARS), Legon. Buffel grass cv. Biloela 
was established using seed obtained from Australia*. The seed was
* By courtesy of the Director of the Veterinary Services, Accra.
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planted in 15 cm rows, 5 cm apart. Only giant star and buffel grass 
were harvested from the first trial due to poor growth of pangola 
during the minor rainy season. At establishment 50 kg N, 40 kg P and 
80 kg K per hectare were broadcast over the trial. Following full 
ground cover an additional 50 kg N/ha was broadcast over each trial.
On September 17> 1974 the growth in the first trial was removed by 
means of machettes to a stubble height of about 5 cms. Harvesting 
began at week three and continued at- weekly intervals for 10 weeks.
The three regrowth cuts were harvested on December 31 (end of minor 
rainy season), March 4, 1975 (mid-dry season) and May 5» 1975 (end of 
the dry season). All plots were harvested by means of a Jari mower 
with a 92 cm sickle bar attachment. After weighing, two samples of 
herbage were taken from the harvested material. The first was dried 
at 80°C for 48 hours and used for dry matter determination and the 
second stored in a freezer at -4°C and later separated into leaf and stem.
On April 29, 1975* the treatments on the second trial commen­
ced. The growth was slashed with machettes and harvesting commenced on 
May 20 (3 weeks following slashing) and continued through to August 12, 
1975- Three regrowth cuts were taken on August 12, August 26 and 
September 9i 1975*
The minor rainy season and major rainy season data were both 
analyzed as a split plot design with species as main plots and maturity 
dates as subplots. Dry season harvest data were also analyzed as a 
split plot experiment with species as the main plots and regrowth dates 
as subplots but data from each rainy season maturity date were treated 
separately.
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During the minor rainy season, rainfall was light, scarcely 
more than 20 mm falling in any one week. In the subsequent dry period, 
however, although there was no rain in the first five weeks, it rained 
more intensively from February 25 to May 6 than during the minor rainy 
season. Rainfall during the major rainy season was more intensive than 
in the minor rainy season while the short dry period of August 13 -
September 9 was quite dry (Appendices LXI and LXIl).
Dry matter accumulation of the grasses during the minor rainy 
season and the major rainy season trials are shown in Fig. 3«1, and 
Appendices XXXIII, XXXIV, and XXV. During the minor rainy season of
1974t both giant star and buffel grasses grew rapidly from 1686 and
1633 kg/ha respectively at week 3, to 4079 and 4665 kg/ha respectively 
at week 6. Thereafter, growth slowed down and reached a peak of 4715 
kg in week 13 for giant star grass and 5707 kg/ha in week 13 for buffel. 
Giant star then declined to 4467 kg/ha and 5360 kg/ha in week 5* This 
difference between the two species appeared to be due to the loss of 
senescent leaves in giant star under the pounding action of the showers 
of November 18 (week 8); on the other hand, in buffel grass the senes­
cent leaves tended to stay on the plant. The development of new shoots 
tended to compensate for the loss of senescent leaves in giant star.
The level of dry matter production observed in this study 
was below that reported by Asare (1972), Caro Costas et_ al. (1965, 1973) 
and Vicente-Chandler (1975) for these species. However, the levels are 
of the same magnitude as those reported by Ruthenberg (1974) in the 
Ivory Coast and Togo. The superior yields reported by the former
Results and Discussion
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M A T U R I T Y  I N  W E E K S
1 9 75
M A T U R I T Y  I N W E E K S
Fie-, 3.1. Accumulation of' dry matter in giant star, buffel and pangola grasses
during the minor rain;/ season 1974 and the ronjor rain/ season -.975'
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63
authors were obtained under higher rainfall conditions than that which 
occurred during this trial or those reviewed by Ruthenberg (1974)*
Buffel grass was headed by week 3 although the flowers had not opened, 
while giant star began flowering during week 4 . In both species dry 
matter accumulation continued after heading which agrees with the 
findings of Brockington (1961) in Rhodesia. Seed setting and shatter­
ing continued throughout the season and this together with leaf senes­
cence and the development of new shoots during the light showers of 
November to early December (Fig. 3*1) may have contributed to the small 
fluctuations in yields.
Although there appeared to be a trend for buffel to produce 
more dry matter than giant star grass during the light rainfall period 
of November and December, the differences were not significant (P>0.05). 
The tendency of buffel to utilize marginal rainfall more efficiently 
than other grasses tends to confirm the observations of Rattray (i960), 
Fitzgerald (1955)» Humphrey (1969)1 Pereira and Beckley (1953) and 
Edwards (1956). During the last fortnight, the yield of both species 
was reduced by 300 kg each, from week 13 to 15 weeks of maturity 
(December 17 to 31). This was primarily due to leaf senescence in 
giant star, while in buffel grass seed shattering was increased by the 
dry weather and this may have been a factor in reducing yield. Such 
observations were made on buffel by Brockington (1961) and Cassady 
(1973). A further factor that may have contributed to this reduction 
in yield in both species may have been translocation of organic matter 
to roots (Norman 1963, Taerum 1970a,b,c).
During the major rainy season 1975 trial the accumulation of 
dry matter of the three species tended to follow a pattern similar to
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64
the 1974 minor rainy season. However, while buffel grass flowered 
from week 3, pangola from week 5» giant star grass did not flower during 
the major rainy season. Dr.y matter accumulation continued in pangola 
grass and buffel grass after heading and because of the ability of 
buffel grass to retain the senescent foliage, it continued to accumu­
late dry matter, rapidly at first, then more slowly from week 9 "to week 
13* Pangola and giant star grasses also grew rapidly from week 3 to 
weeks 8 and 9 respectively. Thereafter the yield of giant star fluc­
tuated; pangola also seemed to fluctuate (Fig. 3«l) but the changes were 
not significant (Appendic XXXIII). This pattern was apparently deter­
mined by the balance between the shedding of senescent leaves and 
development of new ones. Prom week 11 (July 15), the shattering of 
seed tended to reduce buffel yield but this was compensated for by the 
new flush of leaves induced by the rainfall of July 8 to 22. As leaf 
development slowed down and seed shattering continued, buffel yields 
declined after week 13 (July 29). In the stoloniferous grasses on the 
other hand, the net formation of new foliage from week 11 tended to 
raise dry matter yields. In pangola grass, the proliferation of a new 
crop of inflorescence from week 11 enhanced the increase in dry matter 
accumulation. In all three species, however, the changes in dry matter 
yield after week 11 were not significant (P>0.05).
The major rainy season of 1975 yields of the stoloniferous 
grasses were below those reported in giant star by Arias (1965) over 
a 45 day period (900 kg/ha at 10 days to 3000 kg/ha at 40 days).
However Virquez (1965) in Brazil obtained similar dry matter yields to 
those obtained in pangola in this study. In our study, buffel tended to 
outyield the other two grasses particularly at the later stages of
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development during the major rainy season. This finding thus agrees 
with Asare (1974) who reported that buffel was superior to giant star 
and other grasses in the humid zone of Ghana.
Herbage regrowth from the minor rainy season 1974 and major 
rainy season 1975 harvests until the end of the rainy season, middle of 
the ensuing dry season and the end of the dry season, are shown in 
Tables 3.1 and 3*2 and Appendices XXXVI and XXXVII.
In 1974> although buffel grass generally appeared to produce 
higher yields than giant star grass at all harvest dates, differences 
were not consistent (Table 3-1 and Appendices XXXVI and XXXVII), buffel 
being significantly superior (P K 0.05) only in regrowths after the 
harvests of October 8 , 15 and 22 and November 19 at each of the harvests 
at the end of the rainy season, mid-dry season and the end of the dry 
season. In both years all grasses tended to grow during the dry period. 
Within regrowth harvest dates, generally the amount of herbage accumu­
lated during the dry period increased with the lengthening of the 
recovery period (Tables 3»1 and 3-2).
At the end of the minor rainy season of 1974 (December 31) 
recovery yield of giant star was highest for the rainy season harvest 
of October 8 and 15 (2812 and 2384 kg/ha respectively). The low rainfall 
immediately before and after the October 22 harvest (Fig. 3-1) would 
appear to be a factor for the decreased recovery from the October 22 
harvest. In buffel on the other hand, recovery from the minor rainy 
season to the end of the rainy season was greatest for the first rainy 
season harvest of October 8 (45^0 kg/ha). Buffel was not affected by 
the dry period before and after October 22.
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______________________________________ (kg/ha dry matter)_________________________________
Table 3.1. Regrowth yields from giant star and buffel grasses during the dry period (December 1974 to
April 1975).
Harvest 
date in
End rainy season 
(December 31. 1974)
Mid dry 
(March 4
season
, 1975)
End dry
(May 6,
season
1975)
rainy season Giant star Buffel Giant star Buffel Giant star Buffel
October 8 3+ 2812a0 45601° 2733xpd° 36105'° 2970abcO 52071°
15 4 2384f:b 39275 289l£c 34005p 3075ab° 46l4®b
22 5 1922hc 35675C 37891 4550^ 3756f 3787'5°d
29 6 2262|bc 297l£d 30375 35885 3320^b 36275°*
November 5 7 1702C 248lde 2731|ed 2691d 3122abe 3858bcd
12 8 1677S 2506de 2242yde 34805 340 2|b 35795°'d
19 9 909y 2233® 2052def
y
3230bcd 2843|bc 2890^
December 3 11 593d 1926® I66i|fe 2722od 2002°y
3942b°d
17 13. — 826f I4l5yg 2031® 1976° 40965°
31 15 — — 1136f 1918® 23875^ 3202|4
+ Weeks of maturity from beginning of trial to the harvest date during the rainy season, 
o Data followed by the same letter are not different (P ^  0.05): columns - a,b,c,d,e,f,g; rows within
regrowth harvests - x,y.
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_______     (Kg/ha dry matter)_________________________________
Table 3.2. Regrowth yields from giant star, buffel and pangola grasses during the dry period (August -
September) 1975-
Harvest 
date during
End rainy season 
August 12 , 1975
Mid dry season 
August 26 , 1975
End dry season 
September 9 . 1975
rainy season Giant star Buffel Pangola Giant star Buffel Pangola Giant star Buffel Pangola
May 20 4+ 4110| ° 6508f 54561 51061 57911 5602^ 6379£ 56465 4907^
27 5 3511$ 59515 4998-$ 4 9 1 0 | 4867^ 4554* 5449^ 5342$ 5626a
June 3 6 2543$ 3344* 2348$ 4600| 4184° 31415 4565$ 5 0 9 1 f* 4586*
10 7 19971° 3 1 5 5 ^ 2197^ 1924^ 2897-x 27835* 3492^ 4572% 4 8 7 9 !
17 8 18915 2220|e 1765* 14 71*° 2377*® 2249^ 2166* 23495 22675
24 9 H 68y 2668cd 1040- l424bcd 2 8 l l d 1595® 17475 20425 16104
J u ly  1 10 99!d 1947® 794y 1178y de 2786d lOOlf 1915® 21745 938®
15 n 627y 993^ 570cd 864$ef 1904® 759y 9 5 8 f 1099^ 6991
29 13 - — 45d 694£f 662| 6 0 1 ^ 693x 755de 6755
August 12 15 - — — 573x 3 3 8 f 75y 5 6 lx 350^y 75f
+ Weeks of maturity from beginning of trial to the harvest date during the rainy season, 
o Data followed by the same letter are not different (P >0.05): columns - a,b,c,d,e,f,g; rows within
regrowth harvests - x,y,z.
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68
At the mid-dry season harvest (March 4), giant star grass 
had reached maximum recovery yields after the harvest of October 22 
(3789 kg/ha). This contrasts with recovery to the end of the rainy 
season and may have been due to the rainfall of February 4 - March 4 
stimulating new growth (Fig. 3*1 and Table 3*1). Buffel grass also 
made its maximum recovery after the October 22 harvest (4550 kg/ha) 
but the loss in dry matter following October 8 and 15 harvests may have 
been due more to seed shedding than leaf losses.
By the end of the dry period giant star recovery from harvests 
of October 22 was the highest (3756 kg/ha) while in buffel recovery fro m 
October 8 was the highest (5207 kg/ha).
In 1975» buffel grass again tended to produce more regrowth 
herbage at each of the recovery harvest dates than giant star and 
pangola grasses (Table 3*2 and Appendices XXXVI and XXXVIl). Across 
the three recovery harvest dates (end of rainy season, mid-dry season 
and end-dry season) there generally was an increase in herbage recovery 
in all grasses, although buffel grass fluctuated - probably because of 
its shedding of seed. Within the regrowth period until August 12 (end 
of the rainy season) recovery from harvests of May 20 and 27 were the 
highest yields in all three species. At the mid-dry period harvest of 
August 26, on the other hand, while giant star made its maximum yields 
from recovery after rainy season harvests of May 20 to June 3, buffel 
and pangola made their greatest recovery after May 20. At the end of 
the dry period (September 9) giant star yielded its highest herbage 
recovery from the May 20 harvest (6379 kg/ha) while in buffel grass the 
highest recovery yields were from May 20 to June 3.(5646 kg/ha - 5091 
kg/ha) and in pangola grass, the highest recovery yield was 5626 kg/ha
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69
To obtain the greatest amount of herbage recovery for use 
in the (August - September) dry period therefore, the major rainy 
season harvest may be taken in late May in all species. Harvesting in 
early mid-October would give the maximum recovery herbage growth in 
both giant star and buffel for use at the end of December, while by 
harvesting at mid- to late-October, both species would produce their 
highest recovery yields for use in early March. Giant star may be 
harvested from October to mid-November to produce the maximum regrowth 
for use in early May, but buffel would yield its highest recovery for 
early May when it is cut in early to mid-October. However, harvesting 
in early October during the minor rainy season or in May to early June 
during the major rainy season would give poor rainy season yields.
The total seasonal yields would therefore- need to be considered.
Seasonal total herbage yields obtained from the rainy season 
and dry season cuts are given in Tables 3-3 and 3«4» In 1974> giant 
star maximum totals came from the harvests of October 29 (shortly before 
the rains) and their aftermaths, while buffel being less susceptible to 
drought, gave its maximum herbage totals for each season from the 
harvests between October 15 and December 17* On the other hand, timing 
of major rainy season harvests (1975) f°r maximum total seasonal yields 
was the same in all three species and fell from May 27 to June 3 and 
from June 4 to July 15 of 1975*
The overall superiority of buffel to pangola and giant star 
seems to be partly in its ability to grow under low rainfall as reported 
by Taerum (l970a,b,c) in East Africa, Humphreys (1969) in Australia, 
Rattray (i960) in the Sahel, Brown, Layman and Rotar (1966a) in Hawaii,
from May 27.
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Table 3 .3. Seasonal total yields from giant star and buffel grasses during 1974*
_______________________________(Kg/ha dry matter)________
_____________________________________ Maturity___________________________________
3 + 4  5 6 7 8 9 11 13 15
Oct. 8* Oct. 15 Oct. 22 Oct. 29 Nov. 5 Nov. 12 Nov. 19 Dec. 3 Dec. 17 Dec. 31
Minor season + end 
of rainy season:
Giant star 4498c° 4854bc 50 20abc 6341a 6l76ab 6l69ab 5176abc 4903bc 4715c 4467c
Buffel 6l93ab 6800a 7080a 7636a 7218a 7246a 7390a 7297a 6533ab 5360b
Minor season + mid 
dry season:
Giant star 4419e 5361de 6887ab 7116ab 7205a 6534abc 6319abcd 5971bcd 6l31abcd 5603de
Buffel 5243c 6273bc 8063a 8253a 7428ab 8220a 8387a 8093a 7738a 7278ab
Minor season + end 
of dry season:
Giant star 4656c 5545bc 6854ab 7399a 7596a 7894a 7110ab 6312abc 6691ab 6854ab
Buffel 6840c 7487bc 7300c 8292abc 8595abc 8319abc 8047abc 9313ab 9803a 8562abc
+ Weeks of maturity from major rainy season harvest.
+ Harvest date during the major rainy season.
o Data followed by the same letter are not different (pP^0.05): rows - a,b,c,d,e.
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Table 3r4- Seasonal total yields from giant star, buffel and pangola grasses during 1975*
(Kg/ha dry matter)
Maturity
3+ 4 5 6 7 8 9 11 13 15
May 20 May 27 June 3 June 10 June 17 June 24 July 1 July 15 July 29 Aug. 12
Major season +
end rainy season:
Giant star 4513cd° 4558cd 4266d 4513cd 5985ab 5982ab 6287a 5239hc 4516cd 4759cd
Buffel 7106ab 7972a 5562c 5993bc 6486abc 7283ab 7456ab 7737a 6833abc 6040bc
Pangola 5647ab 5849a 3728e 4419de 4507cde 5l82abcd 5360abc 4836bcd 4712cd 5241abcd
Major season + 
mid dry season:
Giant star 5509abc 5957abc 6323ab 4440d 5565abc 6238ab 6474a 5476abc 5210cd 5332bcd
Buffel 6389bc 6888bcd 6402bcd 5735cd 6643bcd 7426ab 7867ab 8648 a 7505ab 6378bc
Pangola 5793a 5405ab 4521b 5005ab 4991ab 5737a 5567a 5025ab 5269ab 5418ab
Major season + 
end dry season:
Giant star 678 2ab 6496abc 6288abcd 6008bcde 6260abcde 656labc 7211a 5570cde 5209e 5320de
Buffel 6244d 7363abcd 7309abcd 7410abcd 66l5bcd 6657bcd 8386a 7843ab 7598abc 6390cd
Pangola 5098c 6477ab 5968abc 7001a 5009c 5752bc 5504bc 4965c 5343bc 54l8bc
+ Weeks of maturity at minor season harvest.
+ Harvest date during minor or major rainy season.
o Data followed by the same letter are not different (P ,>0.05): rows - a,b,c,d,e.
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72
Pereira and Beckley (1953) in Southern Africa, Edwards (1956) in Kenya 
and Mufti and Kaul (1972),in Iraq. Giant star also seemed to be drought 
tolerant, although not as good as buffel grass. Pangola was very sensi­
tive to drought especially at the very early stages. In all the three 
grasses, fluctuations occurred in dry matter accumulation during 
maturation both during the rainy season and in the dry season recovery, 
probably reflecting the balance between the formation and shedding of 
leaves and seeds. Buffel showed more fluctuations particularly during 
the recovery periods apparently because of its ability to utilize light 
rainfall for growth and reproduction.
It would therefore appear that pangola grass on account of its 
sensitivity to drought, especially during its early developmental stages, 
should not be grown in the drier parts of the Accra Plains. However, 
it may be suitable for the wetter areas to the north of the plains.
Giant star grass and buffel grass, on the other hand, would probably 
be suitable for the drier as well as the more humid parts of the Accra 
Plains. The stage to harvest these grasses would appear to depend on 
the season of the first growth harvest as well as the date the aftermath 
is required in the ensuing dry season. During the minor rainy season, 
giant star grass would be best harvested at 6 weeks for maximum com­
bination of the minor rainy season yield and regrowth to the beginning 
of the dry season. If the aftermath is to be harvested at the middle 
of the long dry season, then the minor rainy season harvest should be 
taken at 7 weeks. Similarly the harvest date would be 8 weeks where 
the aftermath is required at the end of the dry season. During the 
major rainy season, however, the best stage to harvest giant star would 
appear to be 9 weeks, regardless of the date at which the aftermath is
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73
required during the ensuing short dry period. Buffel grass would seem 
to be best harvested at 6, 9 or 13 weeks during the minor rainy season 
if the aftermath is required for the beginning of the subsequent dry 
season, the middle of the dry season or at the end of the dry season 
respectively. The harvest dates during the major rainy season could be 
11 weeks whether the aftermath is harvested at the beginning or middle 
of the ensuing short dry period; if the regrowth is required for the 
end of the dry period, then the major rainy season harvest would be at 
9 weeks. For pangola grass, the corresponding stages of growth for 
major rainy season harvest are 4, 3* and 6 weeks.
In addition to the total herbage yields considered above, the 
quality of herbage during growth and development in both the rainy season 
and the aftermath should be taken into account to ensure that optimal 
nutrient levels are harvested.
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Summary
Giant star, buffel and pangola grew rapidly for the first 6 to 
8 weeks' and then slowed down their rates of growth. Buffel commenced 
flowering by the third week in both years and had set seed by the sixth 
week. Giant star flowered at week 6 in 1974 but did not flower in 
1975* Pangola began flowering from week 6 during the major rainy 
season 1975 hut made little growth after slashing at the beginning of 
the minor rainy season 1974* In all three grasses, growth continued 
after flowering.
Buffel grass was outstanding in its ability to accumulate 
dry matter even with light showers. Much of this dry matter appeared 
to be in the seed and the yield was reduced when the seeds were shed. 
Unlike leaves of the prostrate giant star and pangola, senescent leaves 
of buffel were resistant to decay and were not easily stripped off by 
rainfall.
Aftermath yields did not follow a consistent trend in buffel 
grass but in pangola and giant star, there was an increase in dry matter 
yield during the dry period. Fluctuations were attributable to leaf 
losses, seed shedding and growth and because buffel produced seed 
copiously, it fluctuated markedly.
The most drought sensitive of the three species - pangola - 
made its highest total seasonal yield of herbage when it was cut 
shortly before the heavy rains of the major rainy season. The drought- 
hardy buffel and giant star grasses, on the other hand, made their 
highest totals after harvesting late in the major rainy season.
During the minor rainy season, however, maximum total production of
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herbage in giant star grass was derived from harvests at the middle 
of the minor rainy season and their aftermaths, while buffel gave its 
highest seasonal total herbage dry matter yields from harvests of weeks 
4 to 1 3.
It would appear that pangola should not be grown in the drier
parts of the Accra Plains but buffel and giant star are suitable for
these areas.
To obtain maximum seasonal totals of herbage yield, giant star
grass may be harvested in the minor rainy season at 6, 7 or 8 weeks
maturity if the aftermath is required at the beginning, the middle or 
the end of the subsequent dry season. The corresponding growth stages
for harvesting buffel, during the minor rainy season, are 6, 9 and. 13
weeks.
During the major rainy season giant star may be harvested at 
9 weeks regardless of the aftermath harvest date. Buffel, on the other 
hand, may be cut at 11 weeks of growth if the aftermath is required at 
the beginning or the middle of the short dry period; if the aftermath 
is required at the end of the dry period, then the major rainy season
harvest should be at 9 weeks of growth.
These indications would need to be modified as information 
becomes available on the quality of herbage during growth in the rainy 
season as well as the quality of regrowth herbabe during the subsequent 
period.
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PAPER 4; EFFECT OF MATURITY ON QUALITY OF GIANT STAR, BUFFEL 
AND PANGOLA IN THE ACCRA PLAINS
Abstract
Giant star (c.ynodon plectostachyus (K. Schum) Pilger and 
buffel (Cenchrus ciliaris L.), cv. Biloela, were harvested at ten dates 
during the minor rainy season (September 16 - December 31, 1974)*
During the major rainy season (April 29 - August 12, 1975)» pangola 
grass (Digitaria decumbens Stent) was added to the experiment.
Leaf dry matter production increased to a plateau at weeks 
8-9: 2112 and 2104 kg/ha in giant star and buffel respectively in 1974
and 3153» 3212 and 2374 kg/ha in giant star, buffel and pangola grasses 
respectively in 1975* Giant star and buffel were not different but 
both were superior to pangola in leaf dry matter production. Leaf pro­
portions were higher in buffel at the early stages but at mature stages, 
buffel was stemmier. Leaf proportions were higher in the major rainy 
season than in the minor rainy season.
Whole plant IVD and N were similar in giant star and buffel 
during the minor rainy season (60.6 - 61.0$ at week 3 and 40.8 - 41*7$ 
at week 15)• In the major rainy season, however, whole plant IVD was 
higher in buffel (70 .3 - 38.6$) than pangola (64*7 “ 46.2$) and giant 
star (57*7 - 36.2$).
IVD of leaves was higher than stem: the disparity being wider
in buffel than in the creeping grasses; buffel leaves were the most 
digestible (70*4 - 7 1.2$ at 3 weeks to 53-7 - 60.7$ at 15 weeks), while 
the stems had the lowest IVD values (48.6 - 61.3$ at 3 weeks to 26.3 -
76
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77
30.3$ at 15 weeks).
Whole plant N differed among species only in weeks 3-5 during
the major rainy season: pangola and buffel were similar (2.62 - 2.12^)
but were superior to giant star (1*54 - 1*43$)• During both seasons, 
leaf N was higher than stem N in all grasses.
The N content of whole plants would be below maintenance
requirements after week 7 in the minor rainy season 1974 and week 11 in 
the major rainy season 1975* However, maximum digestible dry matter 
and nitrogen yields were obtained in weeks 6-7 in the minor rainy season
and weeks 7 to 15 in the major rainy season.
IVD was highly correlated with N in leaves, stems and whole
plants of all species (r = 0 .714 - O.989) except in giant star stems
(r = 0 .024).
Introduction
Giant star (c.ynodon plectostachyus (K. Schum.) Pilger), buffel 
(Cenchrus ciliaris L.) and pangola (Pigitaria decumbens Stent.) are 
three of the grasses reported by Rose Innes (1966) and Asare (1972) as 
suited to Ghanaian conditions to help solve the poor supply and low 
quality of natural grassland fodder. Asare (1974) studied the nitrogen 
content and In Vitro digestibility (IVD) of buffel grass grown in mix­
tures with other grasses and legumes and found that, harvested at six- 
week intervals throughout the major and minor rainy seasons, the IVP of 
buffel averaged 49-0 and 43*9/& in the first and second years respectively, 
while corresponding nitrogen content was 1.91 and 1.68% respectively.
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The changes in quality during growth and development of these or other 
grasses have not been studied in Ghana.
In this study, the changes in leafiness, IVD and nitrogen 
content of these three grasses during growth in the minor rainy season 
of 1974 and the major rainy season of 1975 were investigated.
Literature Review
Nutrients are mostly located in the leaves of forages (Sullivan 
1969, Bailey 1973)* Funes and Yepes (1974) and Chenost (1975) attributed 
much of the reduction in nutrient value of pangola and other grasses as 
they matured, to the decline in percent leafiness and corresponding 
increase in stem tissue. Thus several studies employ percent leafiness 
as an index of quality. In buffel grass Taerum (l970a,b,c) reported a 
decline in leafiness from 55*6 to 44*8$ in the first two and a half 
weeks in East Africa. Vicente-Chandler et al. (1959a,b) reported little 
change in percent leaf in pangola grass during maturity in contrast with 
the bunch type grasses - Pennisetum purpureum Schumach. (elephant grass) 
and Panicum maximum Jacq. (guinea grass). However, Minson, Raymond and 
Harris (i960) and Mowat et al. (1965) showed that stem tissue, at early 
stages of development, may be more digestible than leaf tissue. 
Furthermore Minson, Raymond and Harris (i960) reported that percent 
leafiness was higher in cocksfoot than in ryegrass yet digestibility 
was lower in the former. Thus the importance of percent leafiness per se 
as an index of quality was weakened. Nonetheless, Minson and Laredo
(1972) showed that voluntary intake was highly correlated with percent
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79
leafiness among panicums of similar growth form.
That herbage digestibility declines as the plant matures is 
widely documented (Minson and McLeod 1970, Mba, Oke and Oyenuga 1973* 
Olubajo and Van Soest 1974, Falvey 1977)- In pangola grass, Johnson 
and Pezo (1975) in Peru, reported 81$ IVD at 2 weeks declining to 66$ 
at 16 weeks, while in Florida, Ventura et al. (1975) reported 68.4$
In Vitro organic matter digestibility at 2 weeks, falling to 54*5$ at 
12 weeks. In a second trial Ventura et al. (ibid) reported 90.2$ at 
4 weeks and 39*8$ at 10 weeks. In giant star grass, on the other hand, 
Olubajo and Van Soest (1974) reported 47» 47 and 37$ IVD at 6, 8 and 
10 weeks respectively in Nigeria. Coward-Lord, Arroyo-Aguilu and 
Garcia Molinari (1974) reported that in Puerto Rico, buffel grass IVD 
fell from 69.1$ at 30 days to 40.3$ at 180 days.
Nitrogen content has been reported by Minson (1973) and 
Barton et al. (1976) to be a more important index of quality in tropical 
grasses than in temperate ones. Thus total nitrogen (N) is a widely 
used parameter in studies on quality of tropical forages, and is gener­
ally reported as crude protein (N x  6.25) according to the Weende 
system of proximate analysis (Church 1972). However, Ferguson (1969) 
has reported that the conversion factor of 6.25 is too high for forages.
In giant star grass, French (1939) obtained 2.51$ N at one 
month, 1.6 - 1.8$ for the next three months and 1 .15$ at 6 months.
Similar data have been furnished by Virquez (1965) on pangola and star 
grass in Brazil. It is apparent from the data of Vicente-Chandler 
et al. (1959a,b) in Puerto Rico, Norman (19^ 3) in Northern Australia, 
and Arias and Butterworth (1965) in Brazil, that although leaf N content 
is higher than stem N, their rates of decline with plant maturity are the
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80
same. Thus, Vicente-Chandler et al. (1959a,b) reported 0.11$ decline 
per week in the levels of guinea, pangola and elephant grasses while in 
the stem, they had 0.103$ per week. Arias and Butterworth (1965) 
reported the faster rates of decline of N in elephant grass: 0.223 and
0.231$ per week. Brockington (1961) in Rhodesia, reported nitrogen 
content of buffel grass decreased from 1 .76$ to 0 .64$ in the first 4 
weeks and to O .48 - 0 .64$ by the end of 12 weeks. Similar studies have 
been reported by Taerum (1970a,b,c) in buffel in Kenya. French (1939) 
in Tanzania reported that giant star grass had 2.51$ N at one month and 
1.15$ at 6 months; while in Brazil, Virquez (1965) reported 3-33$ at 
10 days, declining to 1 .05$ at 40 days and in pangola grass he had 
2.47$ and 1.28$ at 10 and 4° days respectively. Similar reports have 
been made by Vicente-Chandler et al. (1964) in Puerto Rico, Ventura 
et al. (1975) and Johnson and Pezo (1975) on pangola grass.
Materials and Methods
In March-June 1974, giant star, buffel and pangola grasses 
were established in each of two separate trials at the University of 
Ghana Research Farm, Legon. The first trial commenced during the minor 
rainy season of 1974 and the second during the major rainy season of 
1975* Each trial was established in a four replicate, split plot design 
with the three species as the main plots and ten harvesting dates as 
subplots. The subplots were harvested at 3 , 4> 5» 6, 7, 8, 9, H, 13 
and 15 weeks. Pangola and giant star were established by sprigging at 
15 x 15 cms while buffel grass was seeded at 5 cms in rows 15 cms apart.
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Fertilizer was applied at the rate of 50 kg N, 50 kg P and 80 kg N/ha, 
broadcast at 6 weeks.
The 1974 trial was slashed on September 17* As pangola did 
not make much growth thereafter, only giant star and buffel were harves­
ted, starting at three weeks of growth. The second trial was slashed 
on April 29, 1975 and harvesting began on May 20.
At each harvest, a sample of herbage was taken by randomly 
picking out material along the length of the plot after it was cut with 
a Jari mower. The sample was separated into leaves and stems, weighed 
fresh and dried in a forced draught oven at 80°C for 48 hours to deter­
mine leaf/stem proportions of each species. The dried leaf and stem 
samples were ground in a Wiley mill (l mm screen). These were, analyzed 
in the Crop Science Department, University of Guelph for In Vitro dry 
matter digestibility (IVD) using a modified Tilley and Terry method 
(Mowat et al. 1965) and for total N using a Technicon autoanalyzer.
The data were analyzed as a split plot design with species as 
the main plots and plant part in the subplots for each harvest date; 
for the effect of maturity on plant part, species were in the main 
plots and dates in the subplots for each plant part separately. Differ­
ences were tested by Duncan’s Multiple Range Test (Steele and Torrie 
i960). Regressions of IVD and N on maturity stages were calculated 
and the lowest order equations of best fit were taken at the 5$ level.
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Leaf production at the ten harvest dates in the minor rainy 
season of 1974 and the major rainy season of 1975 is shown in Tables 
4-1 and 4.2 and the analyses of variance in Appendix XXXIX. Daring the 
minor rainy season of 1974» giant star grass tended to produce more 
leaf dry matter than buffel, whereas in the major rainy season of 1975» 
buffel generally produced more than giant star. Pangola gave the lowest 
leaf yield of the three species, except for the final harvest date.
Leaf production during the minor rainy season of 1974 increased 
rapidly in both giant star and buffel up to the 4th week (1462 and 1488 
kg/ha in giant star and buffel respectively). Giant star started 
flowering during week 4 and continued to flower and set seed throughout 
the growth period. Seed production was poor, however. Similarly, 
buffel grass began flowering by week 3, although the flowers had not 
opened. Seed development had commenced by week 5 and by week 8, the 
large quantities of seed coupled with the dry weather appeared to depress 
leaf yield (1564 kg/ha). The showers of November 19 (week 9) nay have 
induced a flush of foliage (2104  kg/ha). Thereafter dry weather appeared 
to have retarded leaf development. Thus senescence tended to cause a 
steady but slight and nonsignificant drop until week 15 (1849 kg/ha) 
in buffel.
The 1975 data (Table 4.2) show similar features. Giant star 
did not flower during this trial. Leaf production rose gently until 
week 8 (3153 kg/ha) but from week 1 1, leaf yield dropped slightly to 
2375 kg/ha by week 1 5, apparently because of leaf senescence. Buffel 
trends were similar to 1974s flowering by week 3 and seed filling to
Results and Discussion
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83
Table 4.1. Yield of leaves of giant star and buffel grasses during 
the minor rainy season of 1974•
________________________(Kg/ha dry matter)_____________________
Maturity 
in weeks Giant star Buffel
3+ 920 1° 849 }
4 1462 1488 ab
5 1518 ab 1844 $
6 1958 5b 1731 y
7 1986 $b n&i a
8 1756 xb 1564 fb
9 2112 a 2104 |
11 2077 $ 2073 a
13 2744 $ 2009 a
15 2162 a 1849 p
+ Weeks from beginning of trial, September 16, 1974* 
o Data followed by the same letter are not different (P>-0.05): 
columns - a,b; rows - x,y.
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Table 4.2. Yield of leaves of giant star, buffel and 
pangola grasses during the major rainy season
(Kg/ha dry matter)
of 1975-
Maturity 
in weeks Giant star Buffel Pangola
3+ 307 $ 542 § 143 g
4 799 £d 1651 bc 681 bc
5 !285 0 I442 bc 1129 J
6 1653 bc 1388 Jo 154° b
7 2555 |b 2201 1593 \
8 3153 | 2252 |b 2374 f
9 3098 a 3212 a 2333 f
11 2596 2826 a 2109 |
13 2348 2979 | 2194 |
15 2375 f 2174 2537 |
+ Weeks from beginning of trial, April 29, 1975* 
o Data followed by the same letter are not different (P-> 0.05): 
columns - a,b,c,d; rows - x,y,z.
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week 6 at the expense of the leaves. The continuing rainfall sustained 
buffel leaf development so its leaf yield rose to 3212 kg/ha by week 9. 
Seed filling and the shedding of senescent leaves under declining rain­
fall apparently resulted in a drop in leaf yield by week 15 (2174 kg/ha). 
Pangola grass on the other hand started flowering in week 5 in 1975• 
However, as the rain continued, more stolons were proliferated and new 
foliage developed reaching a plateau by week 8 (2374 kg/ha). There­
after there was a lull in rainfall, slowing down leaf development until 
August 5 - 1 2  when about 10 mm of rain fell and leaf yield tended to 
increase. Thus shifts between leaf development and flower formation 
and the influence of rainfall on them, seem to explain the pattern of 
leaf dry matter accumulation among the three species.
Since the diet of the grazing ruminant consists largely of 
leaves (Zemmelink, Haggar and Davies 1972), leaf production may be 
considered a basis for determining stage of maturity at which herbage 
may be grazed. It would appear then that during the minor rainy season, 
giant star could be grazed from week 9 and buffel from week 5» as these 
are the dates of highest leaf production. In the major rainy season 
giant star, buffel and pangola may be grazed at 8-9, 9-13 and 8-15 
weeks respectively. The yield of digestible nutrients as well as 
regrowth potential would, however, need to be considered for the precise 
date to be determined.
During the major rainy season, the grasses had higher leaf 
proportions in their dry matter yield (36 .0 to 90.6$) than they had in 
the minor rainy season (33.0 to 59.2$). Buffel grass and giant star 
grass were similar in leaf proportions at the early stages of growth 
during the minor rainy season but buffel tended to become stemmier than
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giant star as it matured. During the major rainy season, buffel had 
higher leaf proportions at the early stages (90.6$) than giant star 
(76.^5) and pangola (75.1$); but buffel declined more rapidly in leaf 
proportions than the stoloniferous grasses. These differences in leaf 
proportions may be related to the growth habits of the two types of 
grasses; being erect, buffel developed higher proportions of stem 
tissue to maintain its upright habit in contrast with the prostrate 
grasses. These findings agree with those of Vicente-Chandler et al. 
(1959a,b), Brockington (1961) and Taerum (1970a,b,c).
Generally IVD of leaves and whole plants decreased as the 
grasses matured (Figs. 4.1 and 4«2, Tables. 4*3 and 4.4 , Appendices XL, 
XLI, XLII, XLIII, XLIV and XLV). These data conform to those reported 
for temperate and tropical grasses by Minson and McLeod (1970) and 
for giant star grass by Mba, Oke and Oyenuga (1973)* They however are 
in contrast with those reported by Butterworth and Butterworth (1965) 
and Grieve and Osbourn (1965) who found an increased digestibility as 
pangola matured. The stems of buffel increased in IVD only in week 4 
of the major rainy season of 1975 (Appendix XLV). This increase, 
however, agrees with the findings of Haggar and Ahmed (1970, 1971) 
who reported that IVD increased in growing stems in Andropogon gayanus 
Kunth. The decrease in IVD as the grasses matured was more pronounced 
than the differences among species at the same stage of maturity as 
reported by Blaser, Skrdla and Taylor (1952) and Oyenuga (1957)•
The regressions of IVD on maturity are given in Table 4.4. 
During the mingr rainy season of 1974» giant star gave linear relations 
in both its leaves and its stems but in buffel grass, the relationship 
was quadratic in the leaves and linear in the stems. During the major
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IN 
VI
TR
O 
D
IG
ES
TI
BI
LI
TY
67
m a t u r i t y  i n  w e e k s
Fig. 4*1* In Vitro dry matter digestibility of leaves and stems
of buffel and giant star grasses during the minor rainy 
season of 1974.
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IN 
VI
TR
O 
DI
GE
ST
IB
IL
IT
Y
88
MATUR IT Y  IN WEEKS
Fig. 4*2. In 7itrc dry matter digestibility of leaves and stems
o* giant star, 'buffel and pangola grasses during the
major rainy jeason of 1975.
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Table 4.3. Percent In Vitro dry matter digestibility of whole plants of giant star and buffel during the minor
rainy season of 1974 (upper) and of giant star, buffel and pangola grasses during the major rainy
season of 1975 (lower).
1974 - Minor rainy season
Maturity
3+ .
Oct. 8+
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Giant star 61.O f 59.3$' 57.l£ 51.6£ 48-3$ 47.7| 43.3d 40.5d 40.3d 40.8*
Buffel 60.6£b 62.0| 57-Ob 53-l£ 50.00 46.9gd 47.8§de 43•8de 44-6de 41-71
1975 - Major rainy season
Maturity
3+ . 4 5 6 7 8 9 11 13 15
May 20+ May 27 June 3 June 10 June 17 June 24 July 1 July 15 July 29 Aug. 12
Giant star 57.7| 53.,8f 5 1.6abc 42.3def 46 * 5xde 47•2bcd 4 1.8def 43*7de 40.0ef 36.
Buffel 70.35 69-.9? 65-51 59-2b 53-9bc 5l.3°d 5 1. If 46.2d.e 42.6« 38.6$
Pangola 64.7^b 67-•?x 60.5bc 55•6£d 54-5xde 52.9def 50-3def 48.7®f 49.8def^  X 46.2f^ X
+ Weeks from beginning of trial: September 16, 1974; April 29, 1975*
+ Harvest date during the minor rainy season of 1974 °r major rainy season of 1975*
o Data followed by the same letter are not different (P^0.05): rows - a,b,c,d,e; columns within years - x,y,z.
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Table 4.4. Regressions of percent In Vitro dry matter digestibility on maturity of giant star and buffel
grasses during the minor rainy season of 1974 and of giant star, buffel and pangola grasses
during the major rainy season of 1975*
1974 Minor rainy season 2r 1975 Major rainy season 2r
Giant star
Whole plant Y = 75-9 - 5 .09x2 + 0 .l8x 0 .32** Y = 59-1 - 1 .58x O.65**
Leaves Y = 73-2 - 1.34* 0.81** Y = 57-5 - l-33x O.4I**
Stems Y = 55-7 - 2.04x 0 .69** Y = 58.7 — 1 .69x 0 .87**
Buffel
Whole plant Y = 64.3 - I.64X 0.68** Y = 77.2 - 2.75* 0 .90**
Leaves Y = 90.0 - 7.35x + 0.27x2 O.94** Y = 72.1 - 0 .03x 0 .60**
Stems Y = 55-8 - 1 .21x 0.74** Y = 74-4 - 3-53x O.84**
Pangola
Whole plant Y = 68.4 - I.64X 0.67**
Leaves Y = 72.4 - 2.06x 0.82**
Stems Y = 63.4 - l.lOx O.42**
** Significant (P < 0.01). 
x Weeks.
Y Percent In Vitro dry matter digestibility.
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rainy season of 1975, however, IVD was linearly related to maturity in 
both leaves and stems of giant star, buffel and pangola grasses (Table 
4*4). The regressions in the two years were therefore not consistent.
The rates of decline in IVD in giant star tended to be higher 
in the leaves than in the stems during the minor rainy season (Pig. 4*1 
and Table 4.4). In buffel grass, the stems tended to deteriorate faster 
in IVD than the leaves in both seasons. In pangola grass, leaf IVD 
declined faster than stem IVD during the major rainy season. The rates 
of decline were about 0 .21 - 0 .31 percentage points per day which are 
lower than 0 .5 points per day reported by Reid (1959) in temperate grasses.
Whole plants of giant star and buffel did not differ in IVD 
during the minor rainy season (Table 4.3> Appendix.XVIV). This appeared 
to be due to the fact that the leaves of buffel were more digestible 
than giant star leaves from weeks 6 to 1 5, whereas the stems of giant 
star were superior to buffel stems during this period (Pig. 4*1? Appendix 
XLIV). During the major rainy season of 1975» pangola whole plants were
similar to buffel whole plants in IVD and both were superior to giant
star whole plants (Table 4»3> Appendix XLV). The leaves of buffel were 
however superior to pangola, which was also higher in IVD than giant star 
(Fig. 4*2, Appendix XL’/). In the stems on the other hand, buffel tended 
to be more digestible than the other two grasses at weeks 4 and 5*
However, buffel stem IVD fell rapidly (Fig. 4*2). From week 6, it was 
below pangola and after week 9 below giant star.
Leaf IVD was higher than stem IVD in all species in both years
(Figs. 4*1 and 4-2, Appendices XLIII, XLIV and XLV). This agrees with
reports by Mowat et_ al. (1965), Sullivan (1969), Raymond (1969), Minson 
and McLeod (1970), Bailey (1973) and Moore and Mott (1973)- This is
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in contrast with Minson (1973) who reported higher stem digestibility 
than leaves in guinea grass (Panicum maximum Jacq.). The differences in 
leaf IVD and stem IVD were most striking in buffel grass both in the 
minor and the major rainy seasons, in contrast with the prostrate pangola 
and giant star, especially at the mature stages. This difference may be 
related to the amount of stem lignification that is required to support 
the erect buffel grass. This aspect of the differences among the three 
grasses merits investigation.
There was progressive decline in the nitrogen levels in whole 
plants, leaves and stems of all three grasses as they matured (Figs. 4*3 
and 4*4» Tables 4*5 and 4*6, Appendices XLVI, XLVIII, L and Li). This 
agrees with reports by Paterson (1933)» French (1942), Oyenuga (1957)» 
Arias and Butterworth (1965) and Virquez (1965). The rates of decline 
of N are given in the regressions in Table 4 -6. The nitrogen levels in 
the leaves generally fell faster than in the stems of all the three 
grasses in both years. This disagrees with Vicente-Chandler et al. 
(1959a,b) who reported similar rates of decline in both leaves and stems 
of pangola, signal, elephant and guinea grasses.
Nitrogen had a quadratic relationship with maturity in the 
leaves of giant star grass and both leaves and stems of buffel grass in
1974. Giant star stems, on the other hand, gave a linear regression.
The relations were linear in the leaves and the stems of all the three 
grasses in 1975 (Table 4.6).
The whole plants of giant star and buffel did not differ in 
nitrogen levels during the minor rainy season of 1974 except in week 3 
when buffel was higher (Table 4 .5 , Appendices XLVII, XLVIII, L and Li).
In weeks 3 to 5 of the major rainy season of 1975» however, buffel and
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ea f
MATURITY IN WEEKS
Fig. 4'3* Percent nitrogen content of leaves and stems of giant
star and buffel grasses during the minor rainy season
of 1974,
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MATUR I TY  IN  WEEKS
1975.
Fig. 4»4* Percent nitrogen content of leaves and stems of giant star,
buffel and pangola grasses during the major rainy season of
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Table 4.5. Percent nitrogen of whole plants of giant star and buffel during the minor rainy season of
1974 (upper) and of giant star, buffel and pangola during the major rainy season of 1975 (lower).
1974 - Minor rainy season
Maturity
3+ .
Oct. 8+
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Giant star l-47f 1.30gb l.l8^bc 0 .90? 0 .96? 0.81£* 0.63|i 0.83£d 0.73$ 0.66$
Buffel 1.61* 1.44f$ 1 .3o£c 1 .00? 0 .98° O.78? 0 .80° 0.91? 0 .92g 0.77?
1975 - Major rainy season
Maturity
May 20^
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11
July 15
13 
July 29
15
Aug. 12
Giant star 1.54* 1-571 1-43| 1 .29f:* 1.13r 1 .05*? 0 .90*cd 0 .80*cd 0 .69?d 0.67d
Buffel 2.62^ 2.46% 2.12* 1.62? 1.30$ I.l6de 1.07de 0 .85®f 0.67£ 0.67f
Pangola 2.25* 2.49^ 2.26* 1.87r 1.51? 1.3l| l*15de 0.89®f 0 .64^ 0.76^
+ Weeks from beginning of trial, September 16, 1974 and April 29, 1975*
+ Harvest date during the minor rainy season of 1974 or major rainy season of 1975*
o Data followed by the same letter are not different (pJ^O.05): rows - a,b, c, d, e,f; columns within years -
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Table 4.6. Regression of percent nitrogen on maturity of giant star and buffel grasses during the minor
rainy season of 1974 and of giant star, buffel and pangola grasses during the major rainy
season of 1975*
1974 Minor rainy season 2r 1975 Major rainy season 2r
Giant star
Whole plant Y = 2 .33 - 0 .287x + 0.012x2 0.76** Y = 1.78 - 0.083x 0 .86**
Leaves Y = 3.02 - 0.320x + 0.013*2 O.64** Y = 2.02 - 0.074* O.79**
Stems Y = 0.90 - 0.035x O.47** r = 0.84 - 0 .041x O.78**
Buffel
Whole plant Y = 2.67 - 0.337x + 0.015x2 O.74** Y = 4-06 - 0 .508x + 0.019x2 O.95**
Leaves Y = 2.46 - 0.390x + O.Ol6x2 O.84** Y = 3-12 - 0 .152x O.85**
Stems Y = 1.42 - 0.170x + 0.008x2 0.47** Y = 1.66 - 0 .098x 0.73**
Pangola
Whole plant Y = 2.82 - 0.l62x 0.77**
Leaves Y = 3-06 - 0.157x 0.80**
Stems Y = 1.92 - O.lllx 0.60**
** Significant (P <,0.0l). 
x Weeks.
Y Percent nitrogen.
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pangola were similar in their whole plant nitrogen content but were 
superior to whole plant giant star grass. The former declined very 
rapidly and after week 6, the three grasses were not different in the 
nitrogen content of their whole plants.
Leaf and stem nitrogen levels were similar in giant star and 
buffel in 1974 but in 1975» buffel and pangola were superior to giant 
star grass in both leaf and stem nitrogen from weeks 3 to 7* There­
after, the three grasses were not different in the nitrogen content of 
leaves and stems (Figs. 4*3 and 4.4, Appendices L and Li).
Leaves had higher levels of nitrogen than stems in all the 
three grasses at all stages of maturity (Figs. 4*3 and 4*4j Appendix 
XLIX) in agreement with reports by Vicente-Chandler et al. (1964) on 
pangola, guinea and elephant grasses.
Because nitrogen influences the extent to which rumen microbes 
can ferment forages (Sullivan 19^9» Church 1972), high correlations were 
obtained between IVD and N of leaf and stem fractions of all grasses in 
both years (Table 4*7) except in giant star stems in 1975* Perhaps 
this anomaly in giant star stems could be linked with its low nitrogen 
content, 0.76$ at week 3 and 0.28$ at week 15 compared with 1.35$ to
O .43 - 0.49$ in the other two grasses. The high correlation agrees with 
the report by Barton «st al. (1976). Thomas and McLaren (1971) on the 
other-hand, reported low correlations between IVD and N in pangola.
Minson (1967) reported that N limits digestibility only when 
it is deficient in the diet. He suggested 1 .4 4 $  as the minimum limit 
as this was the lowest level required for positive N balance in the 
ruminant. From work on nitrogen supplementation to tropical ruminants 
however, it would appear that tropical ruminants may adapt to declining
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Table 4.7 . Correlation coefficients between In Vitro dry matter
digestibility and nitrogen in leaves and stems of giant 
star and buffel grasses during the minor rainy season of 
1974 and of giant star, buffel and pangola grasses 
during the major rainy season of 1975 -
1974 1975
Giant star Leaves 0 .92** C .02
Stems 0 .86** 0.99**
Buffel Leaves 0 .71* O.96**
Stems O.79** O.98**
Pangola Leaves 0.94**
St ems O.85**
* Significant (P < 0.05).
** Significant (P < 0.01).
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nitrogen levels to as low as 0.8$ if the rate of decline in nitrogen 
is gradual enough for the rumen microbes to adapt in kinds and numbers 
(Topps and Slliott' 1965 in Rhodesia, Alhassan 1970 at Kumasi, Ghana, 
Attakrah 1971, B.Sc. Dissertation, Faculty of Agriculture, Legon, Ghana, 
Tuah and Tetteh 1972, Kumasi, Ghana). On this basis, nitrogen levels 
from giant star, buffel and pangola grasses in this study would be 
inadequate for maintenance after week 7 in the minor rainy season of 
1974 and week 11 in the major rainy season of 1975*
The yields of digestible dry matter in the three grasses are 
shown in Table 4.8 and Appendices LII, LIII, LIV and LV. Digestible 
dry matter accumulated in the whole plants of all species with advancing 
maturity to a plateau at weeks 6 and 7 in the minor rainy season of 
1974 and weeks 7 to 9 in the major rainy season of 1975* The plateau 
was reached 1 to 2 weeks earlier in the prostrate grasses than in buffel. 
grass in 1975 (Table 4*8, Appendices LII, LIV and LV).
No species differences were discerned in whole plant diges­
tible dry matter production in 1974 although buffel tended to yield more 
digestible foliage during the season than giant star grass (Table 4*8, 
Appendix LIV). During the major rainy season of 19751 on the other 
hand, buffel produced more digestible dry matter in the whole plants, 
leaves and stems than the prostrate grasses which did not differ from 
each other (Table 4-8, Appendix LV). Although there were occasional 
exceptions to this trend, this tendency agrees with the report by 
Falvey (1977) that buffel produced more digestible dry matter than 
pangola grass in Australia.
The leaves tended to contribute more than the stems to whole 
plant digestible dry matter during the minor rainy season of 1974 but
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Table 4.8. Yield of In Vitro digestible dry matter of whole plants of giant star and buffel grasses during
the minor rainy season of 1974 (upper) and of giant star, buffel and pangola grasses during the
major rainy season of 1975 (lower).
(Kff/ha dry matter)
1974 - Minor rainy season
p/ —_
Maturity
3+
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Giant star 1032g° l490bc 1763£b 21251 2162^ 2046|b 1868;* 1756£b 1928|b, 18255b
Buffel 978g 1741$ 2024gb. 2366a 2274|b 2l40|b 2287ab 2274|b 2149!b 2051|b
1975 _ Major rainy season
Maturity
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11
July 15
13 
July 29
15
Aug. 12
Giant star 233c ° xy 564J° 899j 1064b 1890^ 22701 2212| 2016* 18071 1720a
Buffel 420c 14115 14535 1681cX 2304b 2369^ 31835 30455 29195 2337b
Pangola !34« 577de 834^ 1300bcy 15241 2190S 2299^ 2064a 2327* 24205
+ Weeks from beginning of trial, September 16, 1974 and April 29, 1975.
+ Harvest date during the minor rainy season 1974 or the major rainy season of 1975.
o Data followed by the same letter are not different (P ✓*‘0.05): rows - a,b,c,d,e; columns within years —
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in the major rainy season of 1975> this superiority of leaf contribution 
to the digestible dry matter harvest was more striking (Appendices LIII, 
LIV and LV). The digestible dry matter yields obtained by Vicente- 
Chandler et al. (1964) under humid and high fertilizer conditions in 
Puerto Rico, were higher than those found in this study.
Assuming leaf digestible dry matter production as the basis 
for grazing the sward and whole plant digestible dry matter yield for 
harvesting stored feed, it would seem that during the minor rainy 
season giant star should be grazed in weeks 6-7 and buffel in week 6 
for maximum digestible dry matter yield; during the major rainy season 
giant star could be harvested at 7-15 weeks, buffel at 9-15 weeks and 
pangola at 8-I5 weeks.
The yields of nitrogen from the three grasses are shown in 
Table 4*9 and Appendices LII, LVI, LVII and LVIII. The yield of total 
nitrogen of the whole plants, leaves and stems of the three grasses 
appeared to increase to a peak and then fall in both years. In the minor 
rainy season, giant star reached a peak in whole plant nitrogen (45*3 
kg/ha) at week 7i while in buffel it occurred at 5 weeks (51*1 kg/ha).
In the leaves, giant star accumulated its maximum levels in weeks 4j 7 
and 1 3, while buffel leaves reached their peak at week 5i in the stem the 
peaks were obtained in weeks 7-8 (l4»8 - 1 5*7$ kg/ha) in giant star and 
week 11 in buffel grass. In the major rainy season, the whole plants 
reached their maximum nitrogen yield in weeks 7~9 in giant star, week 
9 in buffel and .weeks 8-9 for pangola grass, while the leaves reached their 
maximum nitrogen yields in weeks 8-9 (Table 4«9» Appendices LVII and 
LVIII).
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Table 4-9. Yield of nitrogen of whole plants of giant star and buffel grasses during the minor rainy season 
of 1974 (upper) and of giant star, buffel and pangola grasses during the major rainy season of 
1975 (lower).
__________________________________ (Kpjha. dry matter) _____________________________
1974 - Minor rainy season Maturity
3+ .
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5 Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Giant star 
Buffel
28.4f
29.3q^
3 7 .3 d
42.4°d
35.53
51-lf
39 -9 §
41.43
4 5 -3 |  
4 3 •5qc
3 8 .3 ° d 
3 4 .8 f
35 -0 |  
3 7 -9 |
39-1^
4 4 -7 *
42.9* 
4 0 .7 ^
32.8^ 
35 -8 |
1975 - Major rainy season
Maturity
3+ +
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Giant star 6- 2® 16*5g 3 4 -4 * c 41 .3^ 44-6^ 50 .3^ 47-8^b 3 7 .2 b c 3 1 .1 ° 3 1 .9 °
Buffel 15 -7 d 4 9 .8 J 47.0*0 4 4 .8 ° 5 5 -3 * 53 *4q r 66 . 2a q 5 5 . lbq 4 9 .4 b 4 0 .5 °
Pangola 4 -3 ? 21-3® 3 1 .2 - 43-1^ 4 1 -9 * 54.25 52 . 6* 3 7 -4 * c 2 9 .7 * 39* 6^
+ Weeks from beginning of trial, September 1 6 , 1974 and April 2 9 , 1975*
+ Harvest date during the minor rainy season 1974 o r  the major rainy season 1975*
o Data followed by the same letter are not different (P > 0.05): rows - a,b,c,d,e,f,g; columns within
years - q,r,s.
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Although no species differences occurred in nitrogen yields 
of leaves, stems or whole plants at any harvest date during the minor 
rainy season of 1974, buffel grass generally yielded more nitrogen in 
its whole plants, leaves and stems than the.stoloniferous grasses during 
the major rainy season of 1975 (Table 4*9, Appendices LVII and LVIIl). 
The latter trend agrees with Falvey (1977) who reported that buffel 
outyielded pangola in nitrogen production in North Australia. Vicente- 
Chandler et al. (1964) on the other hand, found that bunch type grasses 
yielded similar nitrogen levels to pangola while in other tests 
(Vicente-Chandler et al. ibid) they reported that pangola was more 
efficient in recovering fertilizer nitrogen than bunch type grasses.
Vicente-Chandler (1975) reported that giant star grass was more
efficient than pangola in the yield of nitrogen. This is in contrast 
with the findings of this study.
For maximum nitrogen yield, giant star would be grazed at week
7 and buffel at week 5» the minor rainy season, but for mowed stored
feed the stages indicated are 5 an<i 7 weeks for giant star and buffel 
respectively. During the major rainy season, the maximum nitrogen 
yields in leaves for grazing were obtained in weeks 8, 9 and 8 for giant 
star, buffel and pangola respectively; while for mowing the dates for 
highest yields were 7-8 weeks for giant star, 9-13 weeks for buffel and 
8-I5 weeks for pangola grass. The harvest dates inferred from leaf 
production, IVD yields and N yields would need to be combined with their 
effects on regrowth potential and on longevity of the sward among other 
factors (e.g., animal management system and economics) to determine the 
harvesting schedule.
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The leaves contributed, more nitrogen than the stems to whole 
plant nitrogen yield in all the grasses during both seasons, thus 
confirming the importance of the leaves in the supply of quality feed 
in grasses as stated by Minson and Laredo (1972) and Bailey (1973)*
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Summary
Giant star and buffel gave similar leaf dry matter accumu­
lation during the minor rainy season of 1974 while pangola grass was 
drought sensitive and failed to grow. During the main rainy season, 
all the three grasses gave similar leaf dry matter accumulation. Leaf 
yield was depressed by flowering and seed setting in all species.
Senescent leaves of buffel were held on the plant while pangola and 
giant star dropped their old leaves.
Buffel tended to be leafier than the prostrate giant star 
and pangola at the early stages but at mature stages it was more stemmy. 
Leaf proportions appeared to be higher during the major rainy season 
than in the minor rainy season in all grasses.
Percent IVD and N decreased with maturity in all species.
Leaves were higher in N and were more digestible than the stems. Rapid 
growth seemed to raise stem IVD in buffel grass during the early stages.
Both total nitrogen and digestible dry matter production rose 
to a peak in 7-8 weeks in giant star and pangola, and 9 weeks in buffel 
and then tended to fall slightly. Buffel accumulated more digestible 
dry matter and N than pangola and giant star grasses.
The stages of maturity to harvest the grasses would appear to 
be 7 and 5 weeks for giant star and buffel during the minor rainy season.
In the major rainy season, the harvesting stages would be 8 weeks for giant 
star and pangola and 9 weeks for buffel.
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GENERAL DISCUSSION
In view of the studies conducted on natural grassland and the 
three introduced grasses certain observations ma.y be made regarding the 
comparative usefulness of the two groups of forages in the Accra Plains.
In the natural grassland, Sporobolus began growth and reached 
its peak production early and then declined. On the other hand, 
Heteropogon began its period of grand growth later than Sporobolus.
The total yield in the natural grassland therefore continued to rise 
reaching a peak by week 18. The introduced grasses on the other hand, 
grew more rapidly than the two natural grasses and reached their peak 
of production by weeks 9 to 13 in the minor rainy season and week 9 in 
the major rainy season. The introduced grasses appeared to be higher 
in yield than the natural grassland at the same stages of growth during 
the rainy season. Recovery rates were of similar magnitude although 
natural grassland appeared to recover slightly faster than the culti­
vated grasses. In both types of grassland, a minimum of 9 to 12 weeks 
recovery period was required to reach maximum recovery yield of dry 
matter for use in the dry season. Growth occurred in both types of 
sward during the dry period.
Sporobolus was similar to buffel in earliness to flower: less
than four weeks, while Heteropogon was similar to pangola grass and giant 
star grass in maturing later. However, while Sporobolus declined 
quickly, buffel sustained its dry matter accumulation over a long period. 
At the same stages of growth, Sporobolus and Heteropogon had higher leaf 
proportions in their dry matter yields than the introduced species.
The quality of natural grassland herbage was similar to the 
introduced grasses at the early stages of growth: IVD was about 60$ in
106'
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107
the whole plant. The erect species, both natural and cultivated, tended 
to deteriorate in IVD faster than the prostrate cultivated grasses.
The nitrogen content of the natural grassland species appeared 
to be slightly above the cultivated grasses at the same stages of growth. 
However by the 15th week the N levels had fallen to about the same in 
both planted and natural species. It would therefore seem that both 
natural grassland and cultivated grasses had similar quality character­
istics. However, the cultivated species grew faster and produced more 
fodder at the start of the rainy season. These findings are the reverse 
of Falvey's (1977) report that natural grassland produced higher yields 
than pangola, buffel and other introduced grasses in the rainy season 
although the quality of herbage was inferior to the introduced species. 
The poorer quality of natural grassland compared with cultivated grass­
land has also been noted by Reid et al. (1973)• On the other hand, 
recovery yields after the rainy season harvest in the present study were 
similar in both groups of forages. Palvey (1977) also found that dry 
matter yields during the dry season were comparable in both native and 
introduced species. Thus the cultivated grasses could be provided for 
use early in the season before switching to natural grassland, thereby 
permitting the cultivated grasses to recover. The natural grassland 
and cultivated grasses would therefore be used alternately.
The nitrogen content of both the natural grassland and the 
cultivated grasses fell too low at the mature stages, even for main­
tenance. To maintain high production at the levels of performance 
demanded by modern livestock systems, the nitrogen requirements should 
be about 2.5$ (Sullivan 1969). Such levels of nitrogen could only be 
furnished in the minor rainy season by the leaves of buffel at 3 weeks,
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and in the major rainy season by leaves of buffel and pangola at 3 to 
5 weeks or by whole plants of buffel at 3 to 4 weeks, but at these 
stages, dry matter production was low.
There is little information on the nutrient requirements of 
tropical ruminants, and although their minimum requirements may be lower 
than temperate animals as suggested by Topps and Elliott (1965) and 
Fianu et al. (1972), the levels required for the full expression of 
their genetic potential production may be of similar magnitude to those 
of temperate animals with due allowance for ambient temperatures, etc. 
Thus the nitrogen levels obtained in the forages in this study would 
probably be inadequate for high animal production if mowed at the 
suggested 7 to 9 weeks of maturity. To raise the dietary nitrogen 
levels in the use of these and such mature coarse herbage, various sug­
gestions have been made: irrigation and fertilization of pastures for
supplementing natural grazing (Adjei 1972, B.Sc. Dissertation, Faculty 
of Agriculture, Legon, Ghana), ensilage (Lansbury 1959« Tuah 1971)*
Hay (Adjei 1972 ibid), supplemental feeding with concentrates (Wharton, 
Shepard and Buamah 1967, Burton and Asiedu 1972), the use of urea in 
supplemental feeds (Fianu et al. 1972, Tuah and Adenku-Tetteh 1972), 
cultivation of shrubby legumes in natural grasslands (Rose Innes 1965), 
cultivation of grasses mixed with herbaceous legumes (Asare 1974)*
Other possible ways of solving this nitrogen deficiency problem in 
feeding low quality herbage include (a) the use of manure treated with 
sterilants (Smith et al. 1977) or ensiled with herbage (Berger et al. 
1977), (b) sodseeding natural grassland with herbaceous legumes.
With highly mature grasses, avitaminosis-A and low energy in the herbage 
could compound the nutrient deficiency. These could be overcome by
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109
vitamin A supply in the mineral lick and by delignifying the coarse 
forage by alkali treatment (Oji and Mowat 1977). Delignification could 
also render more useful the straw of rice, sorghum, maize, etc. as 
energy supplement to coarse pasture grasses, possibly with urea or 
manure provided a more readily fermentable energy source such as 
molasses or cassava chips is added. The diet of cattle grazing in the 
rangelands of the Accra Plains includes a variable proportion of high 
quality browse plants which raise the animal's nutrient intake (Rose 
Innes and Mabey 19^4* Fianu 1966, B.Sc. Dissertation, University of 
Ghana, Legon). Such browse plants as well as other supplements are fed 
to backyard sheep and goats to upgrade their diet (Fianu et al. 1972).
From the point of view of the yields of digestible nutrients 
and of nitrogen in herbage dry matter, it would appear that buffel and 
giant star could provide fodder for use during the minor rainy season. 
They were similar in digestible dry matter yield and nitrogen yield.
For the major rainy season, buffel grass appeared to be the superior 
species. However buffel being erect in growth habit would lend itself 
more readily to harvesting by mowing. The stoloniferous grasses, par­
ticularly pangola, would be more suited to grazing than mowing by virtue 
of their prostrate growth habit. This use was suggested by Nestel and 
Creek (1962) for pangola.
During selective grazing, the ruminants diet is largely foliar 
(Laredo and Minson 1975? Zemmelink, Haggar and Davies 1972), while 
herbage harvested by mowing consists indiscriminately of leaves and stems. 
Thus the stage of herbage maturity for grazing may be based on leaf pro­
duction, leaf IVD and leaf N, while the stage to mow for stored feed may 
be determined from total herbage dry matter yield as well as the yields
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110
of IVD and. N of whole plants.
It would appear that during the minor rainy season of 1974 
giant star could be grazed at 6 weeks and buffel at 5 weeks, because at 
these stages the grasses combined high yields of digestible foliage 
(1071 kg/ha and 1210 kg/ha for giant star and buffel respectively) with 
high yields of leaf nitrogen (29*6 kg/ha in giant star and 37•5 kg/ha 
in buffel grass). In both species, some leaf dry matter yield would be 
sacrificed to obtain the highest levels of nutrient production.
During the major rainy season, the optimum stage for grazing 
giant star appeared to be week 8, which satisfied the three criteria - 
maximum leaf dry matter yield (3153 kg/ha), maximum leaf - IVD yield 
(I56I kg/ha) and maximum leaf - N yield (43.2 kg/ha). In buffel grass, 
the stage of maturity indicated for grazing during the major rainy 
season of 1975 was 9 weeks. At that stage, the yields of leaf dry 
matter, leaf - IVD and leaf - N were 3212, 2046 and 48.8 kg/ha respec­
tively. The grazing date for pangola grass during the major rainy 
season would also appear to be week 8, for this was the date of the 
second highest leaf dry matter production (2374 kg/ha), the highest 
leaf - IVD yield (1318 kg/ha) and the greatest leaf - N yield (38.5 
kg/ha) during the season.
At the stages of maturity suggested for each of the three 
species, percent IVD and N in the leaves were fairly high. During the 
minor rainy season, nitrogen concentrations in the leaves of giant star 
and buffel were 1 .38$ and 2.01$ respectively at the recommended grazing 
stage; the corresponding IVD levels were 54*7 and 65.6$. During the 
major rainy season, giant star, buffel and pangola grasses had 1 .3 7 * 
I.52 and 1.62$ N respectively in their leaves, while the respective IVD
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Ill
levels were 49-5, 63.7 and 5 5 - 5 The l°w IVD (49*5$ - 55*5$) of
giant star and pangola during the rainy season may be remedied by sup­
plementation with concentrates.
For mowing during the minor rainy season as stored feed, 
giant star grass would supply the highest amounts of whole plant diges­
tible nutrients (2162 kg/ha) and nitrogen (43*5 kg/ha) at 7 weeks.
These are slightly lower than the levels obtained at 6 weeks, the graz­
ing stage. However, the difference in digestible dry matter was not 
significant. On the other hand, in buffel grass, the mowing date indi­
cated was the same as the grazing stage of 5 weeks, during the minor 
rainy season. In the major rainy season, however, both the grazing date 
and mowing date coincide in all the grasses, i.e., 8 weeks in giant star 
and pangola and 9 weeks in buffel. For a rule of thumb, therefore, 
mowing and grazing dates may be the same and the earlier of the two 
dates would seem preferable, for in utilizing giant star at 7 weeks 
(mowing date) instead of 6 weeks (grazing date) during the minor rainy 
season, leaf IVD fell from 54»7 to 49*6$ which was significantly dif­
ferent (P < 0.05), although percent leaf nitrogen levels were not dif­
ferent. The percent IVD and N in whole plants were also not different 
at the two dates.
From the totals of seasonal herbage dry matter production 
harvesting giant star at 6 weeks during the minor rainy season would 
provide regrowth to result in seasonal total herbage yields of 6341 
kg/ha (highest to the end of the rainy season), 7116 kg/ha at mid-dry 
season (not different from the highest seasonal total of 7205 kg/ha for 
7 weeks) and 7399 kg/ha at the end of the dry season (not different from 
the highest total of 7894 kg/ha at 8 weeks). In buffel grass, too, the
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112
seasonal sums of the yields at the recommended 5 week harvest stage 
during the minor rainy season (3513 kg/ha) and the corresponding re- 
growth until the end of the rainy season (3567 kg/ha), to mid-dry 
season (4550 kg/ha) and to end dry season (3787 kg/ha) were not dif­
ferent from the highest seasonal total herbage production for these dry 
season harvest dates (7636 at 6 weeks, 8387 kg/ha at 9 weeks and 9803 
kg/ha at 13 weeks respectively). Thus for the optimum combination of 
regrowth potential and quality, giant star grass may be mowed or grazed 
at 6 weeks and buffel grass at 5 weeks during the minor rainy season. 
Similarly during the major rainy season, the regrowth potential and the 
quality of giant star, buffel and pangola grasses were generally 
optimal at the suggested dates for mowing or grazing.
In giant star, the total seasonal herbage yields when mowed 
at 8 weeks during the major season were 5982, 6238 and 656I kg/ha for 
end rainy season, mid-dry season and end dry season respectively; 
buffel gave 7456, 7867 and 8386 kg/ha respectively, while pangola pro­
vided 5182, 5737 and 5752 kg/ha respectively. In pangola grass regrowth 
to end of the dry season at the recommended 8 weeks was 2267 kg/ha which 
gave a lower seasonal total herbage yield (5752 kg/ha) than harvesting 
earlier (7001 kg/ha at 6 weeks). Thus about 1250 kg/ha dry matter bulk 
would be sacrificed for quality. The quality of regrowth herbage would 
however be needed for a more precise determination of these grazing or 
mowing dates.
Successive mowing or grazing at these stages could affect their 
regrowth potential by reducing their root reserves (Fulkerson 1970,
Sheard 1973)* A rest period would therefore be required periodically.
The timing and duration of this rest period requires investigating.
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113
Pangola grass and giant star grass under grazing would develop 
long stolons which would be defoliated by the grazing animals. These 
bare stolons could accumulate and detract from the quality of the pas­
ture by impeding the movement of the grazing animals. To eliminate them, 
fire may be employed as it has the same effect on yield as slashing, or 
mowing. A mower would not remove the trailing stolons adequately 
although it may be safer than fire. However, it is not known how these 
introduced grasses would withstand burning. Thus, the effect of fire 
on their regrowth should be explored. As in grazing or mowing for 
fodder, frequent defoliation by burning may quickly lead to pasture 
degradation. Thus burning, whether in natural grassland or cultivated 
pastures, may have to be employed once in one to two years. Some 
authors have recommended once in three years where herbage is sparse 
(Edwards 1942). The appropriate burning frequency would have to be 
determined for the particular pasture.
The timing of the fire is a crucial determinant of its effects 
on the plant community (Rose Innes 1971). Burning at the beginning of 
the rainy season after the first showers would minimize the period of 
exposure of bare soil to erosion and sun scorch as new sprouts could 
then cover the ground within two to three weeks. This brief exposure 
of the soil could enhance nitrification (Doyne 1937) Diamond 1937, 
Daubenmire 1968).
The use of fire to remove rank herbage in large fields would 
require special precautions to prevent wild fires consuming valuable 
forage and destroying property. A high capital outlay could be involved 
in using fire on such large estates, e.g., in tractor drawn water tanks 
and sprayers or even the use of aircraft to control fires. Fire should 
therefore only be used after due consideration of such problems.
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GENERAL CONCLUSIONS
1. Pretreatment slashing, grazing and burning did not affect the dry 
matter yield and quality of natural grassland. Sporobolus 
dominated the sward for the first 7-9 weeks and Heteropogon took 
over thereafter.
2. Flowering was early in both the natural grassland sward and the 
introduced grasses but dry matter accumulation continued after 
flowering. The introduced grasses grew faster than the natural 
grassland.
3. Fluctuations in dry matter yield in the prostrate grasses giant 
star and pangola seemed to be due mainly to loss of senescent 
leaves but in buffel grass senescent leaves were retained, seed 
production and shedding being the apparent major cause of fluc­
tuations in dry matter yield in buffel grass.
4- Leaves contained more nitrogen and were more digestible than stems, 
this difference being most striking in buffel grass. Species 
differences in nitrogen were not consistent but the natural grass­
land whole plants appeared to be as digestible as the introduced 
species at comparable stages of growth. Among the introduced 
grasses, giant star and buffel grasses had similar digestibilities 
during the minor rainy season but in the major rainy season, buffel 
was the most digestible followed by pangola and then giant star 
grass.
114
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115
5- Optimum harvest dates for giant star grass appeared to be 7 weeks 
during the minor rainy season and 8 weeks in the major rainy 
season. In buffel gras§, it was 5 weeks during the minor rainy 
season and 9 weeks in the major season, while in pangola grass, 
the most desirable harvest stage seemed to be 8 weeks during the 
major rainy season. Pangola would seem to be inappropriate for 
the drier parts of the Accra Plains as it was drought sensitive 
particularly at the early stages of growth and grew poorly in the 
minor rainy season. Buffel was the most hardy and giant star 
intermediate between the two.
6. Both natural and introduced grasses would need to be supplemented 
with concentrates or leguminous forage for high levels of animal 
production.
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B.L. and M.J. Creek. 1962. Pangola grass. Herb. Abst. 32: 
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Norman, J.T. 1963. Pattern of dry matter and nutrient content changes 
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Oji, U.I. and D.N. Mowat. 1977- Nutritive value of storke treated corn 
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64I.
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intake of Andropogon gayanus hay by cattle, as affected by 
level of feeding. Anim. Prod. 15: 85-88.
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APPENDICES
128
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Appendix I. Analyses of variance of weekly yields from natural grassland during the major rainy
season of 1974*
(Sums of squares x 10^ )
Degrees Maturity
Source
of
freedom 4+ * April 29+
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 17
13
July 1
Replications 3 0.0190 0.0390 0.9294 17.940 16.96 10.50 141.10 50.84
Pretreatment 2 0.0250 0.1139 O.56OO 4-998 29.07 18.79 91.90 46.25
Error (a ) 6 O.O53O 0.1806 2.4070 9 .110 19.67 30.30 102.60 I25.40
Species 2 0.0475* 0 .4328** 2.3260* 12.560** 38.26 231.40** 1358.00** 1911.00**
Species x 
pretreatment 4 O.O63O 0.4420 1.6850 10.320 24.98 138.60 396.00 157.60
Error (B) 18 0.1019 O .4187 5-9760 15.830 102.00 241.80 I856.OO 1862.00
+ Weeks from pretreatment: April 1, 1974*
+ Harvest date during the major rainy season. 
* Significant (P'1'- 0.05).
** Significant (P-C. 0.01;.
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Appendix II.. Analyses of variance of weekly yields from natural grassland during the major rainy season
of 1975-
(Sums of squares x 10^ )
Degrees Maturity
Source
of
freedom 6 + * May 21*
7
May 28
8
June 4
9
June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Repli­
cations 3 1.023 1.834 3-168 19.09 48.88 114.30 69.81 155.20 279.70 1095.00
Pretreat­
ment 2 3.187 3.626 15.820 25.68 4.50 2-55 12.65 9.01 108.00 62.10
Error (A) 6 2.296 17.450 23.180 55.80 151.90 150.80 306.80 389.20 463.OO 676.80
Species 2 5-591* 3.965* 12.320* 232.00** 415.50** 785.30** 1203.00 579.90* 829.50* 2002.00*
SPP x 
pretreat. 4 10.990 23.700 44.580 122.50 I44.OO 279.90 323.20 557.50 729.00 84O.6O
Error (b) 18 15.700 51.410 82.530 IO9.4O 495.00 735-60 1005.00 1199.00 1457.00 3896.00
+ Weeks from pretreatment: April 9* 1975*
+ Harvest date during the major rainy season. 
* Significant (P<:-0.05).
** Significant (P<^0.0l).
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131
Appendix III. Analyses of variance of dry matter of species components
of natural grassland during 1974 (upper) and 1975 (lower).
1974
Degrees
of Sums of scruares x 10^
Source freedom Sporobolus Heteropogon Others
Replications 3 12 ;11 . 217.5 120.00
Pretreatment 2 12.68 233.5 85.07
Error (A) 6 65,13 572.8 349.10
Dates 7 1404.00** 6603.0** 300.90
Date x pretreat. 14 96.53 263.2 230.30
Error (B) 63 437.70 1903.0 934.80
1975
Source
Degrees
of
freedom
Sums of squares x 10^
Sporobolus Heteropogon Others
Replications 3 216.0 1447.00 113.30
Pretreatment 2 923.8* 17.59 214.90
Error (A) 6 325.5 1993.00 117.40
Dates 9 1342.0** 7438.00** 173.80
Date x pretreat. 18 290.4 IO84.OO 32.07
Error (b) 81 IO48.O 4922.00 25.63
* Significant 0.05).
** Significant (P^O.Ol).
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Appendix IV. Weekly yield of species components of natural grassland during the major rainy seaGon of 1974.
Maturity
Pre­
treatment Species
4+
Apr. 29+
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 17
13
July 1
Slashed Sporobolus
Heteropogon
Others^
Total
11
10
10
31
59
13
4
75
113
90
24
227
290
210
48
548
336
370
155
861
386
1054
135
1575
586
2272
203
3062
1265
2643
162
4070
Grazed Sporobolus
Heteropogon
Others^
Total
9
6
7
21
26
1
19
46
111
29
0
141
225
71
34
330
285
128
20
433
696
293
186
1175
1126
1380
119
2625
1117
1706
531
3359
Burned Sporobolus
Heteropogon
Others?
Total
19
1
0
20
13
10
12
35
50
85
25
160
122
359
101
582
338
572
178
1087
422
1047
210
1679
760
2134
894
3788
1255
2118
712
4085
Average Sporobolus
Heteropogon
Others^
Total
131°
%
24 f
32$
8y
120
52f
92$
68®
17°
176ef
212de
2135
<
487de
320£d
357x°
yabc 
1 z
794a
505£y
798*
177^b
147bc
826*y
1928$
405^
3158b
1212*
2156*
468^
3838 a
Weeks from pretreatment: April 1, 1974*
Harvest dates during the major rainy season.
Other species consisting of variable proportions of Panicum maximum, Andro 00 fro u ga.yanus, Cenchrus ciliaris, 
Vetiveria fulvibarbis, Setaria sphacelata, Pennisetum sp., Uraria picta, Astragalus sp., Desmodium 
triflorum, Galactia tenuifolia and Rhynchosia minima.
Data followed by the same letter are not different (pj?>0.05): rows - a,b,c,d,e,f; columns - x,y,z.
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Appendix*V. Weekly yields of species components of natural grassland during the major rainy season of 1975- 
 _______ __________________________________ (Kg/ha dry matter)_______________________
Maturity
Pre­
treatment Species 6+ 4= May 21+
7
May 28
8 9 
June 4 June 11
10 
June 18
11 
June 25
12 14 
July 3 July 16
16 18 
July 28 Aug. 13
Slashed Sporobolus
Heteropogon
Others*
Total
262
40
61
363
254
222
58
535
473
328
89
890
655
776
143
1573
716
846
335
1897
974
1213
391
2578
1143
1258
363
2764
1463
1841
244
3548
1543
2183
153
3879
1051
178?
1382
4220
Grazed Sporobolus
Heteropogon
Others*
Total
170
39
73
282
243
83
173
498
279
143
143
565
996
448
52
1495
738
744
390
1873
1213
1307
104
2624
1074
1930
222
3226
1174
1913
542
3629
1368
1166
570
3104
985
3009
1427
5421
Burned Sporobolus
Heteropogon
Others^
Total
14
59
96
168
89
85
235
408
84
405
206
695
220
832
288
1340
437
1934
67
2438
449
1947
629
3025
473
2413
440
3326
777
2311
1374
4462
736
2472
1341
4549
992
2969
1408
5369
Average Sporobolus
Heteropogon
148|°
46®
196^
130«
2791
2925
339de
685^
63055
H75Sa
879bcxy
1489bc
897l£c
I867b
1138$
2022^b
1216^
1940lb
l009|b
2588!
Others^ 77y 155y !46| I6ly 264J 37 5y 341° 720y 688b I406y
Total 271 f 48lf 7l8f 1469e 2069d 2742c 3105c 3876b 3844b 5003a
+ Weeks from pretreatment: April 9* 1975*
* Harvest date during the major rainy season.
Other species consisting of variable proportions of Panicum maximum, Andropogon gayanus, Cenchrus ciliaris, 
tfetiveria fulvibarbis, Setaria sphacelata, Pennisetum pedicellatum, Uraria picta, Astragalus sp., Desmodium 
triflorum, Galactia tenuifolia and Rhynchosia minima, 
o Data followed by the same letter are not different (P>0.05): rows - a,b,c,d,e,f; columns - x,y,z.
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Appendix VI.. Analyses of variance of weekly regrowth yields from natural grassland during 1974.
(Sums of squares x 10^ )
Degrees Maturity
Source
of
freedom 4+ * Apr. 29
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 17
13
July 1
Replications 3 225.6 8OI.5 297.2 235-0 235-4 335-0 27-4 160.7
Pretreatment 2 I5.8 157-9 145-9 162.2 97-7 316.5* 5-0 102.4
Error (A) 6 1034-0 1200.0 281.4 618 .3 166.2 180.2 295-1 770.7
Regrowth dates 2 IO4 7.O* 1283.0** 614.9** 1243.0 3683.O** 2249.O** 5595.O** 3477-0**
Dates x pretreat. 4 308.8 151.5 139.4 442.4 123.8 373-4 150.9 326.2
Error (b) 18 1761.0 I884.O 494.3 3620.0 1843.0 1288.0 1211.0 1989.0
+ Weeks from pretreatment: April 1, 1974*
$ Harvest date during the major rainy season. 
* Significant (P'c<0.05).
** Significant (P-C.0.01).
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Appendix VII. Analyses of variance of weekly regrowth yields from natural grassland during 1975* 
 ____    (Sums of squares x 10^ )_________________________
Degrees __________________________________ Maturity
Source
of
freedom
6+ 
May 21*
7
May 28
8
June 4
9
June 11
10
June 18
11
June 25
12 
July 3
14 
July 16
Replications 3 242.30 40.98 325.80 268.50 54.67 12.67 44.90 82.16
Pretreatment 2 2.256 0.8297 83.15 119.90 66.48 25.15 71.73 46.83
Error (A) 6 553.70 519.70 686.10 108.50 225.80 209.60 91.15 15.43
Regrowth dates 2 4176.00** 1752.00** 2399.OO** 5331.00** 1615.00** 1361.00** 1795.00** 1887.00**
Dates x pretreat. 4 35-39 315.60 55-26 73.43 120.10 38.87 49-86 30.48
Error (B) 18 832.50 1081.00 719.70 77-16 750.90 275.70 177.20 191.20
+ Weeks from pretreatment: April 9> 1975*
+ Harvest date during the major rainy season.
* Significant (P-=^ . 0.05).
** Significant (PC.O.Ol).
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Appendix VIII. Analyses of variance of regrowth yields during the dry periods of 1974 (upper) and 1975
(lower).
1974 End of rainy season Mid dry season End of dry season
Source
Degrees
of
freedom
Sums of 
squares 
x 104
Degrees
of
freedom
Sums of 
squares 
x 104
Degrees
of
freedom
Sums of 
squares 
x 104
Replications 3 143.00 3 81.09 3 1988.00
Pretreatment 2 71.58 2 29.74 2 542.60
Error (A) 6 328.10 6 638.70 6 2185.00
Dates 7 20488.00** 8 28552.00** 9 20335-00**
Dates x pretreat. 14 447.70 16 1004.00 18 1186.00
Error (b) 63 3500.00 72 5506.00 81 7163.00
1975 End of rainy season Mid dry season End of dry season
Source
Degrees
of
. freedom
Sums of 
squares 
x 104
Degrees
of
freedom
Sums of 
squares 
x 104
Degrees
of
freedom
Sums of 
squares 
x 104
Replications 3 IO8.5O 3 35-82 3 21.65
Pretreatment 2 180.20 2 41.36 2 134-50
Error (a ) 6 92.61 6 268.00 6 257.70
Dates 8 17983.00** 8 27397.00** 9 383I7 .OO**
Dates x pretreat. 16 131.80 16 449.70 18 288.70
Error (b) 72 826.00 72 2842.00 81 4225.OO
** Significant (P^_O.Ol).
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Appendix IX. Yield of regrowth from natural grassland during the dry period of 1974*
Harvest date during 
major rainy season 
Maturity at major
rainy season harvest
4+
Apr. 29+
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 17
13
July 1
15 
July 15
17 
July 29
End of major rainy season (July 29) 
Maturity 13^ 12 11 10 9 8 6 4 2 0
Slashed 5037 4243 2100 2475 2163 1803 969 566 0 0
Grazed 5546 3964 2272 3082 2078 2233 1060 289 0 0
Burned 5518 3574 2714 3295 2040 2333 738 789 0 0
Mid dry season (Aug. 12) 
Maturity 15* 14 13 12 11 10 8 6 4 2
Slashed 6088 5312 1952 3770 4285 3015 1497 1091 745 0
Grazed 6047 5196 2020 3112 4795 2584 1595 2440 601 0
Burned 5296 4981 2487 3188 3887 3038 2080 1638 618 0
End of dry season (Aug. 
Maturity
26)
17* 16 15 14 13 12 10 8 6 4
Slashed 4302 4969 1036 3635 3869 3084 4084 2859 1664 1374
Grazed 4299 5793 I85O 4458 4377 4412 4007 2927 1430 2204
Burned 4897 4903 1312 4956 4189 4645 3570 3079 1941 1290
Average end rainy season 5367$/ 3927f 2352$ 295lfe 2093$
4322g
2123]?
2894y
4047$
923y 548f O' 0
Average mid dry season 5796$ 5163I 2153£ 33575° 1724$ 17239y 654d 0
Average end dry season 4499y 5221| 1399£ 4350$ 41451 3875$ 2955^ 1678c 1723c
+ Weeks from pretreatment: April 1, 1974-
^ Harvest date during the major rainy season.
 ^Weeks from major rainy season harvest to recovery harvest.
o Data followed by the same letter are not different (P ^>0.05): columns - a,b,c,d; rows - x,y,z.
University of Ghana          http://ugspace.ug.edu.gh
Appendix X. Yield of regrowth from natural grassland during the dry period of 1975*
Harvest date during 
rainy season 
Maturity at rainy 
season harvest
6+ 
May 21^
7
May 28
8
June 4
9
June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
End of major rainy season
(Aug. 13)
Maturity 12$ 11 10 9 8 7 6 4 2 0
Slashed 3465 3206 2870 1907 1221 740 567 290 120 0
Grazed 3449 3297 3122 2128 1166 796 550 301 98 0
Burned 3554 3869 3301 2643 1411 1151 608 373 166 0
Mid dry season (Aug. 27)
Maturity 14* 13 12 11 10 9 8 6 4 2
Slashed 4949 5057 4018 2789 2161 1890 938 704 696 0
Grazed 5045 5016 3808 2910 2326 1757 880 851 632 0
Burned 5256 4168 4OI4 3348 3015 1865 1115 879 737 0
End of dry season
(Sept. 10) 
Maturity 16* 15 14 13 12 11 10 8 6 4
Slashed 6I64 4973 5010 5026 2942 2207 2213 1782 1279 527
Grazed 6207 5025 4804 5238 2780 2516 1874 1986 1181 548
Burned 5949 5216 5457 5O6 6 2736 2430 2611 2359 1567 995
Average end rainy season 3489§° 3456f 
47471 
5071“
30981 
3947 y 
5090'g
2226 y 1266|
25015
895 § 575 f 321 £ 128* 0
Average mid dry season 5083 f 3016 S 
5110|
1837*
2384$
978| 8H y
20425*
690 9 0
Average end dry season 6107^ 2819° 2233x 1342 5e 690 e
+ Weeks from pretreatment: April 9> 1975*
+ Harvest date during the major rainy season.
$ Weeks from major rainy season harvest to recovery harvest.
o Data followed by the same letter are not different (P >0.05): columns - a,blc,d,e; rows - x,y,z.
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139
Appendix XI. Analyses of variance of seasonal total yield from 
natural grassland during 1974.
Degrees  Sums of squares (x 104)
Source
of
freedom
End of 
rains
Mid dry 
season
End dry 
season
Replications 3 179-5 680.1 3330.0
Pretreatment 2 425-6 121.5 505.2
Error (a) 6 525.8 1124.0 3287.1
Dates 7 6675.4** 1067.3** 22890.1**
Date x pretreat. 14 712 .1 1234-9 1121.9
Error (B) 63 4974-2 7002.2 8326.9
** Significant (P^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
140
Appendix XII. Analyses of variance of seasonal total yield from 
natural grassland during 1975*
Degrees  Sums of squares (x 104)
Source
of
freedom
End of 
rains
Mid dry 
season
End dry 
season
Replications 3 525.2 400.9 121.7
Pretreatment 2 8OO.7 470.2 737.1
Error (A) 6 764.2 492.3 1169.6
Dates 9 213.9* 1565.3 3052.2**
Date x pretreat. 18 578.8 1354.1 1245.7
Error (b ) 81 394.7 6445.7 7520.6
* Significant (P^0.05).
** Significant (P^.0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XIII. Analyses of variance of weekly yields of leaves of Sporofrolus and Heteropogon during the
major rainy seasons of 1974 (uPP®r) and 1975 (lower).
(Sums of squares x 10 )^
1?74
Source
Degrees
of
freedom
Maturity
4+ 
Apr. 29*
5
May 6
6
May 13
7
May 20
8
May 27
9
June 3
11 
June 14
13
July 1
Replications 3 O.OO46 0.0527 0.5746 9.4890 11.340 13-74 58.910 19.320
Pretreatment 2 0.0097 0 .3415* 0.3731 6.OO4O 24.840 23.86 2.843 35.530
Error (a ) 6 0.0119 0 .1310 1.8006 8.0125 18.120 32.18 137.000 133.500
Species 1 0 .0312* 0.2992* 0.1033 O.56II 9.277 51.90 302.700 159.000
SPP x Pretreat. 2 O.OI87 0.1377 1.3470 6.1450** 19.080 81.17 109.700 1.564
Error (b) 9 0.0354 O.IO84 3.9012 2.0870 51.790 97-44 312.700 538.600
i m
Degrees __________________________________ Maturity
Source
of
freedom
6+ 
May 21*
7
May 28
8
June 4
9
June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 0.351 3.168 6.75 23.43 75-01 10.93 9-43 25.24 84.67 150.10
Pretreatment 2 4.863 10.890* 30.97 32.17 8.08 31.30 72.73 122.40 I48.8O 133.70
Error (a) 6 5.132 3.135 12.12 88.69 123.80 110.90 152.90 161.60 102.90 392.00
Species 1 3.240* 1.862 .82 2.57 76.00 71.20 87.72 3.77 2.88 158.40
SPP x Pretreat. 2 3.724* 3.639 6.87 68.86** 53.64 94.41 80.36 25.86 79.09 72.16
Error (B) 9 2.847 9.462 10.33 24.92 145.50 215.90 284.60 251.90 394.50 284.00
+ Weeks from pretreatment: April 1, 1974* April 9» 1975*
 ^ Harvest date during the major rainy season.
* Significant fP<0.05).
** Significant (P<. 0.01).
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142
Appendix XIV. Analyses of variance of yield of leaves of Sporobolus 
and Heteropogon during the major rainy seasons of 
1974 (upper) and 1975 (lower).
1974
Source
Degrees
of
freedom
Sums of 
Sporobolus
squares x 10^
Heteropogon
Replications 3 5.458 88.27
Pretreatment 2 4.315 I H .40
Error (A) 6 50.160 273.10
Dates 7 725.600** 2594.OO**
Date x pretreat. 14 69.940 124.30
Error (B) 63 282.600 751.20
1975
Source
Degrees
of
freedom
Sums of 
Sporobolus
squares x 10
Heteropogon
Replications 3 170.70 394.0000
Pretreatment 2 61.72 0.2959
Error (A) 6 241.40 593.3000
Dates 9 858.7O** 1821.0000**
Date x pretreat. 18 193.80 273.3000
Error (b) 81 729.40 1136.0000
* Significant (P<=i.0.05).
** Significant (P^C.0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XV. Analyses of variance of weekly percent In Vitro dry matter digestibility of leaves (upper) and
stems (lower) of Sporobolus and Heteropogon during the major rainy season of 1974*
(Sums of squares)
Leaves
Degrees Maturity
of 4+ . 5 6 7 8 9 11 13 15 17
Source freedom Apr. 29t May 6 May 13 May 20 May 27 June 3 June 14 July 1 July 15 July 29
Replications 3 46.74 34.92 98-73 87.84 40.10 64.O5 128.70 78.28 96.97 136.80
Pretreatment 2 22.11 66.57 I.44 28.73 61.92* 47.30 17.89 54.25 42.02 4-28
Error (a) 6 98.87 13.36 32.36 67.49 26.14 48.81 52.88 111.30 51.20 66.53
Species 1 31.05* 33.61 12.61 33.84 9-88 23.60 53.70** 21.09 24I.30** I8O.4O**
SPP x pretreat. 2 103.00** 29.05 33.44 9.27 44.21* 58.90 0 .23 28.87 23.12 11.50
Error (b) 9 44-57 11.90 64.12 76.39 24.35 75-96 39.52 167.20 62.79 151.70
Stems
Degrees _______________________________Maturity
Source
of
freedom 7+ * May 20
8
May 27
9
June 3
11 
June 14
13 
July 1
15 
July 15
17 
July 29
Replications 3 97.26 111.70 121.60 46.82 111.30 120.20 317.70
Pretreatment 2 I64.6O** 190.60** 285.30* 136.80* 40.79 2.44 79.91
Error (A) 6 43-57 34-81 102.20 36.48 28.45 38.65 219.90
Species 1 552.00** 32.43 897.90** 236.90** 43-47* 35-28* 26.25
SPP x pretreat. 2 94.65 72.78 57.78 44.08 29.21 46.59 47.32
Error (b ) 9 130.90 75.93 23.62 32.83 64.07 53-53 129.10
+ Weeks from pretreatment date: April 1, 1974*
4= Harvest date during the major rainy season.
* Significant (P"C 0.05).
** Significant (P<. 0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XVI. Analyses of variance of weekly percent In Vitro dry matter digestibility of leaves (upper) and
stems (lower) of Sporobolus and Heteropogon during the major rainy season of 1975*
(Sums of squares)
Leaves
Source
Degrees
of
freedom
Maturity
6+ ± 7 May 21* May 28
8
June 4
9
June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 75-87 36-31 20.58 125.00 19.66 198.60 300.70 11.91 83.50 221.20
Pretreatment 2 250.50** 64.8I 168.90* 179.20* 65.66 7.76 83.99 28.66 121.70 177.40
Error (a ) 6 51.27 50.24 82.92 68.31 53.46 236.20 254.70 120.30 158.60 358-30
Species 1 17.17 35.04* 19.82 49-88 67.33 135.80** II.48 117.00 10.53 19.08
SPP x pretreat. 2 161.50 44*74* 1 75.60** 13.06 43-88 31.51 287.70 20.73 32.30 162.60
Error (b) 9 170.70 32.11 71.86 99.03 158.90 213.90 897.90 298.40 395-00 621.00
Stems
Degrees  Maturity
Source
of
freedom 9+ * June 11^
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 91.40 54.97 15.14 149.60 98.73 8.63 35.66
Pretreatment 2 1371.00** 21.39 46.81 192.80 133.00 140.53 288.70
Error (A) 6 77.83 421.60 177.40 I87.50 193.90 186.40 567.00
Species 1 1549.00** 1332.00** 1350.00** 84O.ll** 735.90** 1114.00**' 2.60
SPP x pretreat. 2 622.20 472.50 124.50 28.33 336.40* 130.00 162.50
Error (B) 9 643.OO 737-00 164.30 328.50 272.90 868.70 208.40
+ Weeks after pretreatment: April 9i 1975-
Harvest date during the major rainy season. 
* Significant (P<-0.05).
** Significant (P<_0.0l).
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145
Appendix XVII. Analyses of variance of percent In Vitro dry matter 
digestibility of leaves (upper) and stems (lower) 
of Sporobolus and Heteropogon during the major rainy 
season of 1974*
Leaves
Source
Degrees
of
freedom
Sums
Sporobolus
of squares
Heteropogon
Replications 3 I84.6O** I65.9O*
Pretreatments 2 81.59* 34.22
Error (A) 6 33.78 51.72
Dates 9 7729.00** 4697.00**
Date x pretreat. 18 315.10* 257.20
Error (B) 81 710.40 954-00
Stems
Degrees
of  Sums of squares
Source freedom Sporobolus Heteropogon
Replications 3 315.00 328.2
Pretreatments 2 244.70* 569.8**
Error (A) 6 91.20* 122.2
Dates 6 4743.00** 8621.0**
Date x pretreat. 12 111.70 358.5**
Error (B) 54 666.80 414.6
* Significant (P^0.05).
** Significant (P^O.Ol).
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146
Appendix XVIII. Analyses of variance of percent In Vitro dry matter 
digestibility of leaves (upper) and stems (lower) 
of Sporobolus and Heteropogon during the major rainy 
season of 1975*
Leaves
Source
Degrees
of
freedom
Sums
Sporobolus
of squares
Heteropogon
Replications 3 183.00 69.04
Pretreatments 2 52.24 194.00
Error (a ) 6 335.70 120.90
Dates 9 8759.00** 7663.OO**
Date x pretreat. 18 I424.OO* 452.10
Error (b ) 81 3044.00 1733.00
Stems
Degrees
of  Sums of squares
Source freedom Sporobolus Heteropogon
Replications 3 25.41 230.0
Pretreatment 2 28.13 190.0
Error (A) 6 100.00 518.0
Dates 6 5225.00** 1 1,112.0*
Date x pretreat. 12 1298.00** 1840.0**
Error (b) 54 2408.00 3351.0
* Significant (P^-0.05).
** Significant (P-^-0.01).
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147
Appendix XIX- Analyses of variance of percent In Vitro dry matter 
digestibility of whole plants of Sporobolus and 
Heteropogon during the major rainy seasons of 1974 
(upper) and 1975 (lower).
1974
Source
Degrees
of
freedom
Sums
Sporobolus
of squares
Heteropogon
Replications 3 183.64 226.76
Pretreatment 2 187.98* 156.03
Error (A) 6 81.52 77.08
Dates 6 6165.30** 5566.36**
Date x pretreat. 12 75.48 75-74
Error (b) 54 337.06 539.78
1321
Degrees
of  Sums of scruares
Source freedom Sporobolus Heteropogon
Replications 3 47.17 36.28
Pretreatment 2 55.82 124.15
Error (A) 6 325.79 105.73
Dates 6 4379-60** S496.5O**
Date x pretreat. 12 452.89 182.80
Error (b ) 54 2962.75 881.01
* Significant fP“=^0.05).
** Significant (P=^I0.0l).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XX. Percent In Vitro digestible dry matter of leaves, stems and whole plants of Sporobolus and
Heteropogon during the major rainy season of 1974*
Maturity
4+ 5 
Apr. 29* May 6
6
May 13
7
May 20
8 9 
May 27 June 3
11 13 
June 17 July 1
15
July 15
17 
July 29
^Sporobolus
Leaves 57-7t° 52.7? 59.9ab 59-7|b 61.51 54-5? 49-6de 48.4S 43-3^ 34- 6g
Stem t+ ~ — — 55.3£ 53.5f 48.5b 48.7 J 42.3$ 38.0d 33-31
Whole +++ - — — 58.91 59-71 5 3.2b 4?-2£ 46.7g 40.9® 34-3^
Heteropogon 0
Leaves 55.45 55*1? 61.3| 62.0| 60.2fb 56.5^c 52.6£d 50.3^ 49- 7t 40.1|
Stem ++__+ — — 64.9$ 55-8b 60.7° 55.00 45-0^ 40.5§ 35-4?
Whole r - --- --- 61.9| 59-85 56.9b 53.2C 43.4d 44*7x 37-7®
+ Weeks from pretreatment: April 1, 1974*
+ Harvest date during the major rainy season.
Not determined.
o Data followed by the same letter are not different (P^>0.05): within rows - a,b,c,d,e,fg;
between species: within leaves - t,u; within stems - v,w; within whole plants - x,y.
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Appendix XXI. Percent In Vitro digestible dry matter of leaven, stems and whole plants of Sporobolus
and Heteropopon during the major rainy season of 1975*
Maturity
6+ 4 7 8 9 10 11 12 14 16 18May 21* May 28 June 4 June 11 June 18 June 25 July 3 July 16 July 28 Aug. 13
M
Sporobolus
Leaves 58.6f*° 61.9| 53•9$c 53-9^ 54-l|bS 47.6Sd 45.7^ 44- if 41.0^ 33.5?
Stems t±- — — 34-7*c 43.6^ 40.8a* 33-7bc 29.1S 20.8d 22.5$
Whole 1
+1++ - — 51.6* 51.8* 46.8ab 43-7bc 41 * 2yd 37.8| 29.81
^Heteropogon
Leaves 60.3|° 59- An 52.l£c 56.7fb 57-5tb 52*4bc 47-o| 48.5? 42.4? 33-5?
Stems — — 50.8*° 58.5^ 55-8^ 45-5vd 40.2de 34-5^ 22.
Whole *±- --- --- 56.5*b 57-9$ 53-2* 46.5° 45-3° 38.7x 27.2 J
+ Weeks from pretreatment: April 9i 1975*
+ Harvest date during the major rainy season.
4^  Wot determined.
o Data followed by the same letter are not different(P^> 0.05): within rows - a,b,c,d,e,f; between 
species: within leaves - t,u; within stems - v,w; within whole plants - x,y.
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXII. Analyses of variance of percent In Vitro dry matter digestibility (leaves, vs. stems) in
Sporobolus (upper) and Heteropogon (lower) during the major rainy season of 1974.
(Sums of squares)
Sporobolus
Source
Degrees
of
freedom
Maturity
May 20*
8
May 27
9
June 3
11 
June 14
13
July 1
15 
July 15
17 
July 29
Replications 3 3-81 77.75 82.38 70.55 87-76 184.30 273.00
Pretreatment 2 38.52 101.40* 158.20** 54.72** 17.16 59.63 87.36
Error (a) 6 41.61 27.78 48.99 13.20 68.88 40.73 149.00
Leaves vs. stems 1 114.00** 380.80** 214.80** 4.86 225.70** 167.50** 10.93
L-S x  pretreat. 2 3-66 2.67 .5908 7.57 16.69 5.30 40.09
Error (b) Qy 95.36 51.86 86.43 53.00 65.67 28.81 219.20
Heteropogon
Degrees ____________________________ Maturity
Source
of
freedom ?+ * May 20
8
May 27
9
June 3
11 
June 14
13
July 1
15 
July 15
17 
July 29
Replications 3 117.70 65.87 114.70 89.01 77-39 55.00 236.70
Pretreatment 2 122.90 225.30** 187-60* 89.85 115 .10 40.24 II.56
Error (A) 6 146.80 55-46 72.52 80.35 200.30 65.81 50.00
Leaves vs. stems 1 49.02 114.00** 109.20** 34.32** 169.60** 507.80** 134.90**
L-S x pretreat. 2 132.10* 4O.I4* 102.90** 46.88** 4.21 9.01 4.00
Error (b ) 9 98.24 34.36 31.22 31.06 60.70 48.59 94.00
+ Weeks from pretreatment: April 1, 1974*
+ Harvest date during the major rainy season. 
* Significant (P<0.05).
** Significant (P<L0.0l).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXIII. Analyses of variance on percent In Vitro dry matter digestibility (leaves vs. stems) in
Sporobolus (upper) and Heteropogon (lower) during the major rainy season of 1975.
(Sums of squares)
Sporobolus
Source
Degrees
of
freedom
Maturity
9+ ±
June 11*
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 19-18 76.63 199.10 427.30 54.07 95-22 57-06
Pretreatment 2 163.80 I64.6O 113.60 231.60 178 .10 175.60 5.89
Error (a) 6 193.60 I25.6O 255.80 450.80 280.60 270.50 563.70
Leaf vs. stem 1 2214.00** 669.90** 277.40* 856.8O* 1350.00** 2444.00** 439.40**
L-S x pretreat. 2 36.90 266.40 33.95 284.20 279.90 85.86 5.06
Error (b) 9 449.40 290.90 277.70 883.10 374.40 373.40 193.30
Heteropogon
Degrees ____ Maturity
Source
of
freedom 9+ * June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 98.22 67.92 27.51 137.60 37.64 127.80 218.30
Pretreatment 2 1287.00** 57-39 .46.03 18.61 58.86 67.51 121.60
Error (A) 6 177.40 311.00 119.00 172.60 172.60 254.30 578.10
Leaf vs. stem 1 217.80** 5.80 71.07 13.50 417.50** 372.90* 726.00**
L-S x pretreat. 2 698.20** 115 .10 17.06 58.44* 1.97** 95.56 658.30*
Error (b) 9 166.70 573.50 126.50 47.55 76.94 579.50 401.10
+ Weeks from pretreatment: April 9> 1975*
+ Harvest date 
* Significant (P<C0.05).
** Significant (P <1,0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXIV. Analyses of variance for weekly percent In Vitro dry matter digestibility in whole plants
of Sporobolus and Heteropogon during the major rainy seasons of 1974 (upper) and
1975 (lower).
(Sums of squares)
i m
Source
Degrees
of
freedom
Maturity
7+ *May 20*
8
May 27
9
June 3
11 
June 14
13
July 1
15
July 15
17 
July 29
Replications 3 68.80 51-44 75.04 98.17 83.79 93.69 130.60
Pretreatment 2 42.19 8 3.24** 85.79 45.00 53.96 19.17 7.16
Error (a) 6 68.90 16.30 49.48 35.12 68.06 41.43 48.42
Species 1 55.28 0.17 82.68** 96.12** 16.81 85.07** 66.97*
SPP x pretreat. 2 11.43 48.31 56.93 2.11 15.79 17.48 9-93
Error (b) 9 95-34 41.67 51.40 25.54 128.30 58.96 64.06
i m
Degrees _____________________________ Maturity
Source
of
freedom 9+ * June 11
10 
June 18
11 
June 25
12 
July 3
14
July 16
16 
July 28
18 
Aug. 13
Replications 3 115.70 23.34 126.60 297.50 5.03 51.31 130.90
Pretreatment 2 138.60* 43-29 17.20 74.52 1 .70 63.67 26.65
Error (A) 6 79.20 34.13 228.70 174.70 110 .70 122.80 317.50
Species 1 139.80* 224.00** 312.20** 48.28 101.80* 5.12 42.35
SPP x pretreat. 2 9.45 11.66 70.72 169-10 35.31 67.27 36.31
Error (b) 9 147.60 104.50 129.40 578.60 186.00 262.40 238.60
+ Weeks from pretreatment: April 1, 1974» April 9i 1975*
+ Harvest date during the major rainy season.
* Significant (Pc^0.05).
** Significant (P-cO.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXV. Analyses of variance of weekly percent nitrogen of leaves (upper) and stems (lower) of
Sporobolus and Heteropogon during the major rainy season of 1974*
(Sums of squares)
Leaves
Source
Degrees
of
freedom
Maturity
7+
May 20*
8
May 27
9
June 3
11 
June 14
13
July 1
15
July 15
17 
July 29
Replications 3 0.0163 0.2376 O.3OO4 0.0790 0.1477 O.O4I3 O.OI46
Pretreatment 2 0.2288 0.2666 0.2032 0.0061 0.1290 0.0335 O.II63
Error (A) 6 0.3289 0.2728 0.1599 0.2960 0.0875 0.0979 0.0767
Species 1 0.0150 O.528I** 0.1536 0.0247 0 .0171 0.1751** 0 .0610**
SPP x pretreat. 2 0.0802 0 .1256* 0 .1047 0.0076 O.OO44 0.0129 0.0119
Error (b) Q 0.0768 0.1506 0.3034 O.II65 O.O325 O.O665 0.0218
Stems
Source
Degrees
of
freedom
Maturity
7+ * May 20*
8
May 27
9
June 3
11 
June 14
13
July 1
15 
July 15
17 
July 29
Replications 3 0.0501 0.0997 0.1698 0.0120 0.0837 0 .0177 0.0276
Pretreatment 2 0.2228** 0.1444 0.0503 0.0223 O.O488 0.0253 0.0097
Error (A) 6 O.O459 0.1324 0.2340 0.1670 0.0593 0.0240 0.0270
Species 1 I.659O** 1.4702** 0.0794** 0.8932** 0.2542** 0.0280** 0.0012
SPP x pretreat. 2 O.O484* 0.1043** 0.0663* 0.0097 0.0052 O.OI44** 0.0176*
Error (b) 9 0.0717 0.0690 0.1728 0.0260 O.OI46 0.0150 0.0426
+ Weeks from pretreatments April 1, 1974*
+ Harvest date during the major rainy season. 
* Significant (P^C0.05).
** Significant (P<d0.01;>
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXVI. Analyses of variance of weekly percent nitrogen of leaves (upper) and stems (lower) of
Sporobolus and Heteropogon during the major rainy season of 1975*
(Sums of squares)
Leaves
Source
Degrees
of
freedom
Maturity
9+ * June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 0.3024 0.1277 0.2540 0.1829 0.0382 O.O466 0.0049
Pretreatment 1 0.0827 0.0256 0 .0600** O.O38O O.OI56 0.0077 0.0100
Error (A) 3 0.1381 O.OI98 0.0095 0.0310 0.0118 O.O87O 0.0244
Species 1 0.0011 0.0600 O.OI44 0.0042 0 .0756* 0.0039 0 .0342*
SPP x pretreat. 1 0.0248 0 .1225* 0.0002 O.O484* 0.0306 0.0077 0 .0182*
Error (B) 6 0.4447 0.0822 0.1308 0.0339 0.0521 0.0715 0.0176
Stems
Degrees  Maturity
Source
of
freedom 9+ * June 11+
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 0.2902 0.0245 0.0616 0.0105 0.0221 0.0152 0.0099
Pretreatment 1 0.0729 0.0218 O.O248 0.0176 0.0031 0.0014 0.0072
Error (a) 3 0.1729 0.0398 0.0098 0.0061 0.0125 0.0118 0.0390
Species 1 1 .5500** O.3752** 0 .2328** 0 .1871** 0 .1702** 0.0333** 0.0600
SPP x pretreat. 1 0.0040 0.0495** 0.0086 0.0060 0.0011 0.0028 0.0169
Error (b) 6 O .5246 0.0159 0.0151 0.0077 0.0266 0.0069 0.1329
+ Weeks from pretreatment: April 9t 1975*
+ Harvest date during the major rainy season. 
* Significant (P<^0.05).
** Significant (P<C,0.0l).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXVII. Analyses of variance of percent nitrogen in leaves,:stems:and whole plant of Sporobolus
and Heteropogon during the major rainy seasons of 1974 (upper) and 1975 (lower).
1974
Source
Degrees
of
freedom
Sums of squares
Sporobolus Heteropogon
Leaf Stem Whole Leaf Stem Whole
Replications 3 0.1376 0.0347 O.I54I 0.2350 0.0909 O.4O8O
Pretreatments 2 0.0887 0.1287 0.2251 O.O965 0.0040 0.1247*
Error (A) 6 0.4035 O.IO83 0.4237 O.I476 O.O443 0.0837
Dates 6 IO.84OO** 7 .94OO** II.O45O** 6.648O** I.533O** 12.5930**
Date x pretreat. 12 0.9491 0.3419 1.1592 0.2909 0.2943* 0.3013
Error (b) 54 1.3150 0.7677 1.2832 0.7482 0.4423 0.7330
1975
Degrees  Sums of scruares
of  Sporobolus__________  Heteropogon
Source freedom Leaf Stem Whole Leaf Stem Whole
Replications 3 0.6812 0.0945 0.5959 0.1891 0.0915 0.1824
Pretreatment 1 0.2379 0.0106 0.1029 0.0412 O.O463 0.0818
Error (a ) 3 0.0920 0.0660 0.0897 0.0831 O.I84O 0.0778
Dates 6 5.0282** 5.8I5O** 5 .1112** 4.1190** 6.9570** 7.1293
Date x pretreat. 6 O.O858 0.2017 0.1075 0.2455* 0,2007 0.3205
Error (b) 36 0.8537 1.1910 1.1377 0.7H5 0.2930 0.4272
* Significant (P<10.05).
** Significant (P<c0.0l).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXVIII. Percent nitrogen in leaves,, stems and whole plant of Sporobolus and Heteropogon during
the major rainy season; of.1974*
Maturity
7+ * 8 9 11 13 15 17May 20 May 27 June 3 June 17 July 1 July 15 July 29
Sporobolus
Leaves
Stems
Whole plant
2.00f°
1.34^
1.87|
l*78t
1.18$
1 .65*
1.39?
0.80$
!-26y
1.32|d
0.93y
1.19£d
1.25f
0 .69^
l*10x
0.99^
0.49$
0 .82®
0-95S
0-47^
O.85®
Heteropogon
Leaves
Stems
Whole plant
l.95|
0.82jJ
1.84|
l-49uc
0.68*
1.35*
l*55t
0.68b
1.42b
1.40?
0-55S
1 .13°
1.30f
0 .48«d
1.00d
1.16?
0.42$
0.78|
1-oof
0.48®d
0.72®
+ Weeks from pretreatment: April 9i 1975*
+ Harvest date during the major rainy season.
o Data followed by the same letter are not different (P ^ >0 .05): rows - a,b,c,d,e,f; between species:
within leaves - t,u; within stems - v,w; within whole plants -. x,y.
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Appendix XXIX. Percent nitrogen in leaves, stems and whole plants of Sporobolus and Heteropogon during
the major rainy season of 1975*^
Maturity
9+ 10 11 12 14 16 18
June 11* June 18 June 25 July 3 July 16 July 28 Aug. 13
Sporobolus
Leaves
Stems
Whole plant
1.84£°
l*43v
1.78®
1*37%
0.92*
1.25$
1.31$
0.77*°
1.20*
1-12?
0.6l$d
1.01$
1.02gd
0.57$
0.93®d
l.Ol^d
0-45$
0.92cd
0.88d
0.44$
0.80d
Heteropogon
Leaves
Stems
Whole plant
1 .80*
0.74ft
1.735
1*39^
0*57b
1.27*
1.40$
0-57^
1.20*
1.20$
°-4l5
0.92^
l.l3f
0.37®
0.80d
i.o o fe
0-355
0.71de
°-95u
0-34$
0.64®
+ Weeks from pretreatment: April 9» 1975*
+ Harvest date during the major rainy season.
# Average of 2 pretreatments: slashed and grazed.
o Data followed by the same letter are not different (P>0.05): rows - a,b,c,d,e; between species: 
within leaves - t,u; within stems - v,w; within whole plants - x,y.
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Appendix XXX. Analyses of variance of percent nitrogen of leaves vs. stems in Sporobolus (upper) and
Heteropogon (lower) during the major rainy season of 1974*
(Sums of squares)
Sporobolus
Degrees ____ Maturity
of 7 . 8 9 11 13 15 17
Source freedom May 20 May 27 June 3 June 14 July 1 July 15 July 29
Replications 3 O.O645 0.2857 0.1541 0.0729 0.1226 0.0121 O.OI56
Pretreatment 2 0.2784 0.5489 0.3328* 0.0089 0.0991 0.0268 0.0277
Error (A) 6 0.2738 0.4129 0.1603 0.3234 0.0769 0.0984 O.O5O3
Leaves vs. stems 1 2.5091** 2.2510** 2.0827** O.96OO** 0.8816** 1 .5150** I.0584*'
L-S x pretreat. 2 0.0059 0.0275 0.0369 0.0095 0.0104 0.0016 0 .0120*
Error (B) 9 0.1230 0.0924 0.2422 O.O585 0.0226 0.0395 0.0210
Heteropoffon
Degrees _____________________________Maturity
Source
of
freedom 7+ * May 20T
8
May 27
9
June 3
11
June 14
13
July 1
15
July 15
17
July 29
Replications 3 0.0104 O.O76I 0.3236 0.0278 0.1057 0.0570 0.0241
Pretreatment 2 0 .1296* 0.0240 0.0388 0.0174 0.0731 0.0530 0.0899
Error (a ) 6 O.O654 0.0573 0.2943 0.1719 O.O545 0.0223 0.0328
Leaves vs. stems 1 7 .5600** 3.9450** 4.4807** 4.335O** 4.0262** 3.3000** 1.7174**
L-S x pretreat. 2 0.1662** 0 .0405** 0.0162 0.0099 O.OO48 O.OO46 0.0257
Error (B) 9 0.0526 0.0376 O.I659 0.0398 0.0429 0.0329 O.O665
+ Weeks from pretreatment: April 1, 1974*
4= Harvest date during the major rainy season. 
* Significant (P-<0.05).
** Significant (P<_0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXI. Analyses of variance of percent nitrogen of leaves vs. stems in Sporobolus (upper) and
Heteropogon (lower) during the major rainy season of 1975*
(Sums of squares)
Sporobolus
Source
Degrees
of
freedom
Maturity
9+ * June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 1.0837 0.1318 0.2279 0.0839 0.0721 O.O488 O.O483
Pretreatment 1 O.OO64 0.1936* 0.0218 0.0977* 0.0306 0.0120 0.0010
Error (A) 3 0.6298 0.0256 0.0220 0.0109 0.0375 0.0491 0.0399
Leaves vs. stems 1 0.6889** 0 .7056** 1 .2155** 1.0353** 0.7921** 1.2660** 0.7744**
L-S x pretreat. 1 0.0441 0.0049 0.0068 0.0105 O.OI56* 0.0020 0.0506
Error (b) 6 0.0968 0.0597 O.O848 O.O55I 0.0140 O.O448 0.0806
Heteropogon
Degrees _________________________■____ Maturity
Source
of
freedom 9+ * June 11
10 
June 18
11 
June 25
12 
July 3
14 
July 16
16 
July 28
18 
Aug. 13
Replications 3 0.0201 0.0359 0.0967 0.0512 0.0084 0.0166 0.0202
Pretreatment 1 0.0095** 0.0176 0.0650 0.0002 0.0011 O.OO64 0.0020
Error (A) 3 0.0010 0.0397 O.O34O 0.0280 O.OI45 0.0828 0.0092
Leaves vs. stems 1 4.4420** 2.8820** 2.9070** 2.2952** 2.4885** 1 .8769** 1 .7161**
L-S x pretreat. 1 0.1207** 0.0033 0.0020 0.0016 0.0003 0.0090 0.0016
Error (b) 6 O.O4I6 0.0172 O.OI48 0.0437 0.0168 0.0269 0.0300
+ Weeks from pretreatments April 9i 1975*
$ Harvest date during the major rainy season. 
* Significant (P<.0.05)*
** Significant (P-C,O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXII. Analyses of variance of weekly percent nitrogen of whole plants of Sporobolus and
Heteropogon during the major rainy seasons of 1974 (upper) and 1975 (lower).
(Sums of squares)
1974
Source
Degrees
of
freedom
Maturity
7+ *May 20
6
May 27
9
June 3
11 
June I4
13
July 1
15
July 15
17 
July 29
Replications 3 0.0103 0.2687 0.4011 0.0375 0.2192 0.0213 0.0605
Pretreatment 2 O.4OIO 0.0361 0.3609 0.0189 0.0886 O.OI65 0.0923
Error (A) 6 0.2853 0.3185 0.2250 0.2884 O.O585 0.0731 0.0376
Species 1 O.OO54 0.2541* O.I454* 0.0252 0.0572 0.0109 0.0973
SPP x pretreat, 2 0.0017* 0.1100 0.1243 0.0014 O.O45I 0.0086 0.0175
Error (b) 9 0.0017 0.2311 O.I413 0.0789 0.0617 O.O54I 0.0104
m i
Degrees ____ Maturity
Source
of
freedom 9+ * June 11*
10 
June 18
11 
June 25
12 
July 3
14
July 16
16 
July 28
18 
Aug. 13
Replications 3 0.6189 0.0815 0.3540 0.0781 0.0371 0.0743 0.0098
Pretreatment 1 0.1892 0 .1278* 0.0105 0.0163 0.0136 0.0072 0.0049
Error (a ) 3 0.4172 0.0176 0.0379 0.0100 0.0067 0.1253 0.0193
Species 1 0.0072 0.0011 0.0005 0.0371 0.0495 O.I849* 0 .1122**
SPP x pretreat. 1 0.1406 0.0028 0.0068 O.O495* 0.0126 0.0009 0 .0272*
Error (b) 6 0.2346 0.0643 O.O646 O.O48I 0.0777 0.1119 0.0217
+ Weeks after pretreatment: April 9* 1975-
* Harvest date during the major rainy season.
* Significant (P<1 0.05).
** Significant (P<,0.0l).
University of Ghana          http://ugspace.ug.edu.gh
 _______________________________ (Sums of squares x 10^ )__________ _________________
1974-Minor rainy season
Appendix XXXIII. Analyses of variance of weekly yields from giant star, buffel and pangola grasses during
the minor rainy season of 1974 (upper) and the major rainy season of 1975 (lower).
Degrees ___________________________________ Maturity
Source
of
freedom
3+ .
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Replications 3 10.83 11 .72 53-96 33-26 43-34 76.92 346.10 125.60 56.81 39.62
Species 1 O .56 32.52 34.36 68.68 13.89 12.35 158.20 225.10 196.90 159.60
Error 3 42.61 11.49 10.79 419.20 258.00 90.09 114.80 149.90 603.20 458.00
1975-Major rainy season
Degrees Maturity
of
Source freedom
3+ +
May 20*
4
May 27
5
June 3
6
June 10
7 8 
June 17 June 24
9
July 1
11
July 15
13 
July 29
15
Aug. 12
Replications 3 0.35 14-89 20.92 87-96 123.60 73 .83 87.26 201.60 98.49 386.80
Species 2 33.18** 314.20* 142.20 75-81 557.50* 95-20 544.20* 1471.00** 1343.00 334.90
Error 6 3.86 42.26 87.39 103.20 221.70 248.20 254.30 112.90 238.30 321.00
+ Weeks from beginning of trial.
+ Harvest date during minor rainy season of 1974 and major rainy season of 1975* 
* Significant (P^ O.C^).
** Significant (P^O.Ol).
161
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162
Appendix XXXIV. Analyses of variance of yields from giant star, buffel 
and pangola grasses during the minor rainy season of 
1974 and the major rainy season of 1975*
Source
Minor rainy season 
of 1974
Major rainy season 
of 1975
Degrees
of
freedom
Sum of 
squares 
x 104
Degrees
of
freedom
Sum of 
squares 
x 104
Replications 3 380.30 3 301.90
Species 1 649.6O 2 3126.00**
Error (A) 3 443.30 6 374-80
Dates 9 9637.00** 9 39062.00**
Date x species 9 252.60 18 1785.00
Error (b) 54 2132.00 81 2055.00
* Significant (P^-0.05).
** Significant (P^^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXV. Accumulation of dr.y matter of giant star, buffel and pangola grasses during the minor
rainy season of 1974 (upper) and the major rainy season of 1975 (lower).
1974 Maturity
3+ .
Oct. 8T
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Giant star 1686A0 2470dc 3098^ 4079bc 44745 44925 42675 4310'x 47151 4467b
Buffel 1633d 2873£d 3513$P 4665b 47375 47405 51571 5371S 57071 5360|
1.975. Maturity
3+ .
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11
July 15
13 
July 29
15
Aug. 12
Giant star 403® 1047yd 1723£d 2516° 4094b 48145 5296* 4612* 45165 4759b
Buffel 598d 2021° 2218C 2838$ 4266* 46155 6212® 6744J 68331 6040!
Pangola 191® 851°d 1380|c 22225° 2742b 4142| 4566* 4266ay 4668| 5241«
+ Weeks from beginning of trial: September 16, 1974; April 29, 1975*
+ Harvest date during minor rainy season of 1974 or major rainy season of 1975.
o Date followed by the same letter are not different (P >-0.05): rows - a,b,c,d,e; columns within
years - x,y.
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXVI. Analyses of variance of regrowth yields from giant star, buffel and pangola grasses during
the dry periods of December 1974 _ May 1975 (u PPe*0 and August - September 1975 (lower).
1974
Source
End rainy season Mid dry season End dry season
Degrees of 
freedom
Sums of 
squares x 104
Degrees of 
freedom
Sums of 
squares x 104
Degrees of 
freedom
Sums of 
squares x 104
Replications 2 54.16 3 176.40 3 396.70
Species 1 1921.00** 1 1196.00** 1 1981.00**
Error (A) 2 202.30 3 276.90 3 444.70
Harvest dates 8 4853.OO** 9 4430.00** 9 1436.00
Date x species 8 201.80 9 300.70 9 1414.00
Error (b) 32 637.20 54 2219-00 54 5601.00
1975
Source
End rainy season Mid dry season End dry season
Degrees of 
freedom
Sums of 
squares x 104
Degrees of 
freedom
Sums of 
squares x 104
Degress of 
freedom
Sums of 
squares x 10
Replications 3 1101.00 3 51.17 3 263.50
Species 2 2708.00** 2 983.OO** 2 199.20
Error (A) 6 1076.00 6 109.60 6 168.80
Harvest dates 7 23127.00** 9 32972.00** 9 45478.00**
Date x species 14 1234.00 18 1621.00 18 1213.00
Error (b) 63 2278.00 81 2466.OO 8 1- 2553.00
* Significant (p-''0.05) •
** Significant (P ■< .0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXVII. Analyses of variance of weekly regrowth yields from giant star, buffel and pangola
grasses across regrowth dates during 1974 (upper) and 1975 (lower).
(Sums of scruares x 10^ )
1974 - Minor rainy season 
Degrees Maturity
of 3+ 4 5 6 •7 8 9 11 13
Source freedom Oct. 8+ Oct. 15 Oct. 22 Oct. 29 Nov. 12 Nov. 19 Dec. 3 Dec. 17 Dec. 31
Replications 2 29-31 61.28 48.92 88 .13 74.55 67.74 192.40 I64.7O 99.25
Species 1 1570.00* 831.40* 633.60 457.50* 99-83 293.50* 195.60 984.30 478.30
Error (A) 2 25.15 76.64 253.20 28.80 85.87 9.25 242.10 194.10 I67.8O
Regrowth dates 2 326.50 520.00 646.40* 168-70 801.60* 568.90* 532.80 880.60* 2078.00*
Date x species 2 191-90 84.89 55.66 64-93 131.00 O .58 270.90 426.90 101.80
Error (B) 8 464-50 48O.5O 368.70 430.30 369.90 498.20 860.60 556.50 130.70
1975 - Major rainy season
Degrees ___ Maturity
Source
of
freedom
3 +
May 20
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11
July 15
Replications 3 111.40 32.79 176.90 276.20 131.70 170 .70 56.36 19.47
Species 2 425.90 350.80 442.30 749.90* 134.40 927.20 1209.00 286.20*
Error (a ) 6 363.50 284.90 302.90 336.40 266.70 229.50 236.90 79.23
Regrowth dates 2 49.05 363.60 2450.00* 2661.00* 59.52 60.79 142.30 119.90*
Date x species 4 1263.00 906.30 303.20 207.00 109.00 224.80 208.30 109.10
Error (b) 18 I857.OO 2227.00 626.70 432.90 344.50 637.90 609.70 105.70
+ Weeks from beginning of trial.
* Harvest date during the minor rainy season of 1974 and the major rainy season of 1975*
* Significant (P-^ 0.05).
** Significant (P ‘:--0.0l) .
University of Ghana          http://ugspace.ug.edu.gh
Appendix XXXVIII. Analyses of variance of seasonal total yields from giant star, buffel and pangola grasses
during 1974 (upper) and 1975 (lower).
(Sums of squares x 10^ )
1974
Source
Degrees
of
freedom
End rainy season 
December 311 1974
Degrees
of
freedom
Mid dry season 
February 4. 1975
Degrees
of
freedom
End dry season 
April 6, 1975
Giant star Buffel Giant star Buffel Giant star Buffel
Replications 2 103.1 1075.6 3 624.2 708.0 3 415.72 I864.3
Dates 9 1023.4** 1797.0** 9 2631.5** 3701.7** 9 3437.50** 2914.8**
Error 27 899-7 IO54.I 27 1360.0 2166.1 27 3285.90 3746.0
1975 End rainy season Mid dry season End dry season
Degrees ______ August 12, 1975______   - August 26, 1975______   September 9» 1975
of Giant Giant Giant
Source freedom star Buffel Pangola star Buffel Pangola star Buffel Pangola
Repli­
cations 3 825.70 3216.50 162.90 292.60 659.30 21.30 62.00 520.00 334.50
Dates 9 2044.30** 2860.50** 1570.00** 1355.80** 3980.40** 649.OO** 1520.90** 1704.30** I784.6O**
Error 27 745.10 2350.00 741.60 1021.90 2164.50 1241.70 1158.00 1701.50 1817.50
** Significant (P~=^  0.01) .
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167
Appendix XXXIX. Analyses of variance of yield of leaves of giant star 
and buffel grasses during the minor rainy season of 
1974 and of giant star, buffel and pangola during 
the major rainy season of 1975*
Source
1974 Minor rainy season 1975 Major rainy season
Degrees
of
freedom
Sums of 
squares 
x 104
Degrees
of
freedom
Sums of 
squares
Replications 3 2.634 3 2.503
Species 1 66.558* 2 14.868**
Error (A) 3 8.847 6 2.554
Dates 9 7.230** 9 15.492**
Date x species 9 5 .592** 18 8.688**
Error (b) 54 5-187 81 1.277
* Significant (P^0.05).
** Significant (P^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
1974 and the major rainy season of 1975*
Appendix XL. Analyses of variance of percent In Vitro dry matter digestibility of leaves, stems and
whole plants of giant star, buffel and pangola grasses during the minor rainy season of
1974 Minor rainy season 1975 Major rainy season
Source
Degrees of 
freedom Leaves Stems
Whole
plants
Degrees of 
freedom Leaves
Whole 
Stems plants
Replications 3 27.90 6O .85 3.72 3 96.70 1.59 41.81
Species 1 2060.01** 958.40** 59-28* 2 6337.00** 2161.00'* 2106.00**
Error (a) 3 14.38 79-57 7.53 6 94-33 155.80 122.40
Dates 9 4352.00** 3466.00** 452.OO** 9 4012.00** 8346.00**7126.00**
Date x species 9 636.10** 338.30** 7-34* 18 1008.00** 2272.00** 761.50**
Error (b) 54 341.50 534.40 10.38 81 857.90 785.IO 744.40
* Significant (P^O.05).
** Significant (P<_0.0l).
University of Ghana          http://ugspace.ug.edu.gh
   __________________________ (Sums of squares)__________________ _______
Appendix XLI. Analyses of variance of weekly percent In Vitro dry matter digestibility of leaves, stems
and whole plants of giant star, buffel and pangola grasses during the minor rainy season
of 1974.
Degrees  Maturity
Source
of
freedom
3
Oct. 8+
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves
Replications 3 10.97 31.42 17-44 19 .18 50.30 4.01 57.64 35-33 5.83 25.48
Species 1 O .85 4-96 22.78 199.00** 444.02** 421.95** 433.65** 38.92** 457.50** 322.58**
Error 3 3.38 6.89 6.91 15.77 4-67 5.76 11-39 16.78 21.51 33.12
Stems
Replications 3 19.80 2.80 21.63 14.07 10.75 2.92 4.90 187.40 5.37 6.91
Species 1 10.55 I .44 i7 .ll IOI.53** 172.05** 252.00** 139.44** 67.28 306.28** 228.98**
Error 3 96-92 42.12 9-96 13.96 4*16 17.25 0.49 192.83 9-27 11.25
Whole plants
Replications 3 12 .11 17.58 4-44 1-35 16.89 5.83 11.71 88.90 15.20 23.90
Species 1 O .36 14.39 0.02 4-99 4.29 1.47 40.41 20.64 37.30 1.45
Error 3 9.13 19.00 2.12 7.16 12.97 1-57 63.83 111.68 34.40 28.98
+ Weeks from beginning of trial, April 1, 1974*
+ Harvest date during the minor rainy season of 1974>
* Significant (p-^0.05).
** Significant (p-^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
__________________________________ (Sums of squares)___________________________________
Degrees ___  _______________________________ Maturity__________________________________
of Jf 4 5 6 7 8 9 H  13 W
Appendix XLII. Analyses of variance of weekly percent In Vitro dry matter digestibility of leaves, stems and
whole plants of giant star, buffel and pangola grasses during the major rainy season of 1975.
Source freedom May 20^ May 27 June 3 June 10 June 17 June 24 July 1 July 15 July 29 Aug. 12
Leaves
Replications 3 32.14 19.96 34-12 3.63 125.36 42.07 15.93 8.28 57-89 11.37
Species 2 330.62** 64I.5I** 48I.57** 1713.40** 462.18 346.65** 973.01** 426.30** 571.92** 1378.28**
Error 6 33-36 12.77 16.52 38.67 361.06 54-77 34.48 30.49 27.66 86.45
Stems
Replications 3 51.17 13.45 14.81 6.96 6.61 13.81 38.96 97.24 8.66 22.03
Species 2 112.36 554.77** 265.85** 257.61** 211.83* 166.03* 224.70* 469.08** 903.84** 1266.33**
Error 6 84.62 19.27 29.04 31.67 103-44 80.37 81.87 106.89 100.87 30.90
Whole plants
Replications 3 15.70 13.76 27.94 22.94 59.07 26.58 23.36 25.17 24.65 4.56
Species 2 317.84** 607.45** 392.39** 638.52* 159.58 69.80 212.82** 49.35 205.53* 213.90
Error 6 13.74 12.47 12.52 177.99 237.72 58.88 35-98 35-09 73.16 7.38
+ Weeks from beginning of trial, April 29, 1975*
+ Harvest date during the major rainy season of 1975* 
* Significant (P^IO.05).
** Significant (P'-- 0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLIII. Analyses of variance of weekly percent In Vitro dry matter digestibility (leaves vs. stems) of
giant star, buffel and pangola grasses during the minor rainy season of 1974 (upper) and the
major rainy season of 1975 (lower).
(Sums of squares)
1974
Source
Degrees
of
freedom
Maturity
3+ .
Oct. 8T
4
Oct. 15
5
Oct. 22
6
Oct. 29
7 8 
Nov. 5 Nov. 12
9
Nov. 19
11 13 
Dec. 3 Dec. 17
15
Dec. 31
Reps 3 3-92 13-80 1.24 4 .10 49.90* 4.33 27.86 103.70 7-94 22.99
Species 1 2.72 5.88 0.20 8 .12 3 1.64* 10.89 40.64* 66.42 7.56 4.00
Error (A) 3 58.14 36.14 8.21 15.24 3-61 7-55 7-98 149.30 14.02 40.48
Leaves vs. 
stems 1 I652.OO* 706.20** 753.50** 737.10** 822.30** 951.70**1121.00** 622.50**1014.00** 524.40**
L-S x species 1 8.70 0.53 39-69 292.40** 584.40** 663.IO** 532.50** 390.10** 756.20** 547.60**
Error (B) 6 69.00 33.30 46.51 43.65 16.39 18 .07 38.58 179.40 20.04 13.29
1975 Degrees Maturity
Source
of
freedom May 20*
4
May 22
5
June 3
6
June 10
7
June 17
8
June 24 -
9
July 1
11 13 
July 15 July 29
15
Aug. 12
Reps 
Species 
Error (A)
3
2
6
9.14
359-80**
17.23
13.32
1132.00**
24.54
40.46 5.O3 
729.80** 1229.00** 
24.79 64.86
54.80
396.10
328.70
51.42
187.90*
94.17
48.89
4O4.IO**
52.46
49.66 15.67 
142.90* 392.40** 
72.37 58.98
7-46
134-10**
47.87
Leaves vs. 
stems 1 130.70** 90.87** 104.60** 0 .15 240.00* 624.80** 563.60** 885.70** 777.50** 68O.4O**
L-S x species 2 
Error (B) 9
83.17
174-90
64.39*
27.58
17.58
29.25
742.30**
11 .03
297.90*
212.90
324.80**
45.42
793.70**
69.90
752.40**1083.00** 
120.90 120.40
67.03
91.41
+ Weeks from beginning of trial, September 16, 1974» April 29, 1975*
+ Harvest date during minor rainy season 1974 °r major rainy season 1975* 
* Significant (P~i0.05).
** Significant (P'<-O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLIV. Percent In Vitro dry matter digestibility of leaves, stems and whole plants of giant star
and buffel grasses during the minor rainy season of 1974*
Maturity
3+
Oct. 8+
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves
Giant star 
Buffel
69.8|°
70.45
65.6®*
67-2|b
62.3^
65.6§*c
54-7£
64.7^c
49.6J
64-5*°
49. 5y 
64.!*°
46*4ye
6l.l5d
42ayf
56.1*
41*8yf
56.9de
40.7f y
53.45
Stems
Giant star 
Buffel
50.9a
48.65*.
52.7*
53.5|
51.7|
48.85*
49.75
42.5*
47-35b
38.1cdy
47.05*
35.8cde
41.2*c 
32.8|ef
39.55
33.7def
39-65
27.2ef y
41-0*c
» - k
Whole plants
Giant star 
Buffel
61.0* 
60.65*
59.3$
$2.0*
57. l£ 
57-0*
5I.6®
53.lS
48.35
50.05
47.75
46.95*
43.3x 
47*85d-
40.5d 
43.85^
40.3x
44•6de
40.8^ 
41.75
+ Weeks from beginning of trial, September 16, 1974- 
+ Harvest date during the minor rainy season 1974*
o Data followed by the same letter are not different (P ^ 0.05): rows - a,b,c,d,e; columns within plant
part; leaves - x,y; stems - q,r.
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLV. Percent In Vitro dry matter digestibility of leaves, stems and whole plantB of giant star,
buffel and pangola grasses during the major rainy season of 1975*
Maturity
3+ +
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11
July 15 ,
13 
July 29
15
Aug. 12
Leaves 
Giant star 
Buffel 
Pangola
58.4*° 
71.2* 
65.2 ^
54.3 f  
70. lg 
69.51
52.1|b
67-5*
6l.4*bc
38.3§d 
66. 9 |
57 -6f>c
46.5|bcd
61.9*
56.4bc
49-5zbc
62.7*
5 5 - 5 j°
4 1.6bcd
63.7*
52.2°d
47-7*bcd
60.9*
49-0°d
42.8bcd
59.6a
49*0°d
36.0* 
60.7£ 
40.7d
Stems
Giant star
Buffel
Pangola
56.1* 
61 *3q
63- 4|
52.3|b 
68.9* 
61.Of*
50. 3fb
61-8q
56-3*bc
49-9*b
51.9?
60.6*b
45*6bc
45-6?d
54-5qCd
42.7°d 
40.5de 
49.35
42.0°d 
37-9|f 
48-4d
38.6d 
33.8fe 
48.7d
37-Od
29.7fh
50.6cd
q
36.3d 
26.3b 
51*3qd
Whole plants 
Giant star 
Buffel 
Pangola
57-7* 
70.3^ 
64-7fb
53-8*b
69-9|
67.7|
51.6ahc 
65.5f 
60.5bc
42.3def 
59.2b 
55-6£d
46.5^de
53*9bc
54-5?de
47.2bcd
5l.3?d
52.9tef
41.8def
5 1 * l? d
50.3^ef
43-7§e
46.2^
48.7ff
40.0ef
42-6®f
49-8fef
36.2^ 
38.6^  
46.2{
+ Weeks from beginning of trial, April 29, 1975*
+ Harvest date during the major rainy season 1975*
o Data followed by the same letter are not different (P_>0.05); rows - a,b,c,d,e,f,g,h; columns within 
plant part: leaves - x,y,z; stems - q,r,s; whole plant - t,u,v.
University of Ghana          http://ugspace.ug.edu.gh
_____   (Sums of squares)__________________________________
Appendix XLVI. Analyses of variance of percent nitrogen in leaves, stems and whole plants of giant star,
buffel and pangola grasses during the minor rainy season of 1974 and the major rainy
season of 1975*
_______ 1974 Minor rainy season_______________  1975 Major rainy season___
Degrees of Whole Degrees of Whole
Source freedom Leaves Stems plants freedom Leaves Stems plants
Replications 3 0.9309 0.0861 0.2868 3 0.1495 O .4674 0.2386
Species 1 0.1361 0.0001 0.2152 2 4.545O** 5.3950** 3.8210*
Error (a) 3 0.2340 0.1100 O.O843 6 1.1890 1.2770 1.1770
Dates 9 13.5170** 2.2840** 5.924O** 9 30.1850** 13.3130** 35.7280**
Date x species 9 0.3380 0.1147 O.O73I 18 2.8370* 2.8040** 3.6490**
Error (b) 54 2.3690 1.0680 2.2100 81 2.7660 2.2240 2.1830
* Significant (P<^,0.05).
** Significant (P<^ 0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLVII. Analyses of variance of weekly percent nitrogen content of leaves, stems and whole plants
of giant star, buffel and pangola grasses during the minor rainy season of 1974-
(Sums of squares)
Source
Degrees
of
freedom
Maturity
3+
Oct. 8*
A
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves
Replications 3 0.4829 0.2047 0.0793 O.O465 0.0276 0.1755 0.0945 0.2195 O.4O49 O.8O85
Species 1 0.0861 0.0112 0.3120 O.O465 O.OO84 0.0098 0.0005 0.0003 0.0004 0.0026
Error 3 0.0356 0.5570 0.0724 0.1374 O.O566 0.1307 0.0007 0.0006 0.0013 0.0041
Stems
Replications 3 O.O483 0.1072 0.0914 0.0334 0.0244 0.0878 0.1185 0.0121 0.0285 0.1749
Species 1 0.0162 0.0018 0.0420 0.0231 0.0231 O.OO84 0.0001 0.0001 0.0011 0.0039
Error 3 0.0675 0.2396 0.1256 0.0318 0.0201 0.0520 0.0004 0.0034 0.0071
Whole plants
Replications 3 O.O658 0.0978 0.0995 0.0305 0.0180 0.2052 O.O434 O.O669 O.I856 0.4475
Species 1 0 .0778* O.O496 0.1143 0.0200 0.0011 0.0010 0.0080 0.0122 0.0669 0.0222
Error 3 0.0206 0.3560 0.1052 0.0812 0.0176 0.0922 0.0215 0.0248 0.0335 0.0243
+ Weeks from beginning of trial, April 1, 1974*
+ Harvest date during the minor rainy season of 1974-
* Significant (P<L_0.05).
** Significant (Pc^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLVIII. Analyses of variance of weekly percent nitrogen content of leaves, stems and whole plants
of giant star, buffel and pangola during the major rainy season of 1975*
(Sums of squares)
Degrees ____________________________________ Maturity
Source
of
freedom
3+
May 20*
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15 
Aug. 12
Leaves
Replications 3 0.1630 O.O715 0.1123 O.O554 0.1477 0.0908 0.0169 0.0417 0.0319 0.0121
Species 2 2.0713** 1.7877** 1.5921** I.O863* 0.3225 0.2267 0.1705 O.O546 0.0617 0.0087
Error 6 0.1038 0.2843 0.1509 0.5713 0.2515 0.5091 0.5039 O .5887 O.I465 0.2521
Stems
Replications 3 0.0459 0.1668 O.O446 0.4194 0 .1017 O.O74O 0.1279 0.1161 O.OO59 0.0723
Species 2 0.9048** 2.9520** 1 .5221** I.325I** O.5482** O.4OO8 O.33IO 0.1232 0.0026 O.O894
Error 6 0.1625 0.0827 0.2093 O.565I 0.1004 0.3383 0.4828 0.4726 0.0115 0.3693
Whole plants
Replications 3 0.0738 0.0730 O.O983 0.1271 0.0915 0.1082 0.0359 O.O65O 0.0091 O.O384
Species 2 2.4181** 2.2046** I.5877** 0.6703 0.2867 0,1331 0.1307 0.0132 O.OO58 0.0197
Error 6 0.1143 0.1174 0.1595 0.5067 0.2065 0.5273 O .4825 0.4754 0.0347 0.2558
+ Weeks from beginning of trial, 
$
* Significant (P^-0.05).
** Significant (P" 0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix XLIX. Analyses of variance of weekly percent nitrogen (leaves vs. stems) of giant star, buffel and
pangola grasses during the minor rainy season of 1974 (upper) and major rainy season of 1975
(lower).
(Sums of squares)
1974 - Minor rainy season
Degrees Maturity
Source
of
freedom
3+ .
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Reps 
Species 
Error (A)
3
1
3
0.4139
O.O885
0.1001
0.2737
0.0110
0.7479
0.1555
0.2916
0.1898
O.O653
0.0020
0.1472
0.0495
0.0018
O.O685
0.2552
0.0182
0.1709
0.0681
0.0370
0.2750
0.1757
0.0808
0.2911
0.3051
0.0051
0.1687
0.0962
O.OO47
0.0897
Leaves vs. 
stems 1 8.7470** 5.4760** 4.3470** 3.2760** 3.5630** 2.544O** 4.2950** 2.6090** 2.3790** 3.3420**
L-S x species 
Error (b)
1
6
0.0138
0.1204
0.0020
0.0869
0.0625**
0.0236
O.O676*
O.O367
O.O298**
0.0109
0.0002
0,0200
0.0157
O.O836
0.0032
0.0290
0.0006
0.1286
0.0112
0.1256
1975 ~ Major rainy season
Maturity
Source
of
freedom
3+ 
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Reps 
Species 
Error (A)
3
2
6
0.1679
2.8320
0.2126
0.1935
4.5750**
0.1802
O.I487
3.1080**
0.2748
0.3068
I.904.0*
0.0323
0.1819
O.7866*
0.3171
0 .1§36
0.5271
0.8330
0.0999
0.4028
0.9379
0.1207
0.0902
1.0250
O.O324
0.0241
O.IO4O
0.0391
0.0368
0.7862
Leaves vs. 
stems 1 8 .9060** 6.8800** 6.7950** 5 .9OOO** 4.996O** 4.9699** 3.7520** 3.1030** 2.8630** 1.7550**
L-S x species 
Error (B)
2
9
O.I444*
O.O948
0.1639
0.2315
0.0059
0.0938
O.5O67*
0.3721
0.0842
0.1023
0 .1005** 0 .0988* 
0.0155 0.0938
0.0877*
0.0728
0.0401
O.O594
0.0167
0,0329
+ Weeks from beginning of trial, September 16, 1974! April 29, 1975*
+ Harvest date during the minor rainy season of 1974 or the major rainy season of 1975*
* Significant (p~~0.05).
** Significant (P-~~ 0.01).
University of Ghana          http://ugspace.ug.edu.gh
Appendix L. Percent nitrogen of leaves, stems and whole plants of giant star and huffel grasses during
the minor rainy season of 1974*
Maturity
3+ 4 5 6 7 8
Oct. 8+ Oct. 15 Oct. 22 Oct. 29 Mov. 5 Nov. 12
9 11 
Nov. 19 Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves 
Giant star 
Buffel
2.33|°
2.53|
1.98|b
2.05b
1.615?
2*01b
1.38^
l-54£
l*49xd
1.565
1.34£de
1.27$
I.l8de 
1 -12x
1.30gde
1.30C
1.23xde
1 .20g
1.091
1.08^
St ems
Giant star 
Buffel
0-91q
1.00|
0.83^
0.86^
0.70|bc
0.84|
0.6iabe
0.50^
0.63qbc
0-53^
0•54qC 
0-47q
°*47q
0*47b
0.545c
0-54^
0-46®
0.45'§
0*45q
0*45q
Whole plants 
Giant star 
Buffel
1-475
1.61$
1.30|k 
1.44fb
1.18f!?g 
1•30bc
0.90i
1.00?
0.96?
0.98?
0.8l£d
0.78?
0.63t
0 .80?
0.83td
0.91?
0.73?
0.92?
0.66$
0.77?
+ Weeks from beginning of trial, September 6, 1974*
+ Harvest date during the minor rainy season of 1974*
o Data followed by the same letter are not different (P_>0.05): rows - a,b,c,d,e; columns within
plant part: leaves - x,y; stems - q,r; whole plants - t,u.
University of Ghana          http://ugspace.ug.edu.gh
Appendix LI. Percent nitrogen of leaves, stems and whole plants of giant star, buffel and pangola
grasses during the major rainy season of 1975*
Maturity
3+ 
May 20*
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Leaves 
Giant star 
Buffel 
Pangola
aO
1.79y
2.76®
2.56|
1.86*
2.68*
2.67|
1.70*
2.48ab
2.46a
1.61*
2.32*60
2.!4$
1.415
1 .81abc
1 .85a)3
1*37£
1 .70.ab,c 
1.62|b
1 • 24x 
1.52*^c 
1 .46.*b
1.155
1 .30b(?
I.l6b
1.02 a 
1 .22$ 
0.95$
1.06 a 
1 .10g
1.04b
Stems
Giant star
Buffel
Pangola
0.76®
1-35*
1 * 35qC
0.66*bc
l* 5 1q r
1 .84*
0 .65abc
1.44*
1.35q°
0.66§b
0.95^
l*48q
0 .56|bc
0.75r°d
1 .08°d
0.43|bP
0 .64bcd
q
0 .88d
0.43*bc
°*58qd
0.83^
0.36bc
0'47qd
0 -6lq
0 .33q°
0.34d
0 -36|
0 .28°
0-43qd
0.49§
Whole plants 
Giant star 
Buffel 
Pangola
1-54*
2.62a
2.25|
l*57v
2.46*
2.49|
1*435
2>12t
2.26|
1.29|b
1-62?
1.87$
1*135
1-30|
1-51?
1 .05$c
l . l 6 f e
1-31?
0.90^cd
1.07te
l . l 5f
0.80^°d
0.85|f
0.89|f
0.69^d
0.67f
0 .64^
0 .67^
0.67f
0.76f
+ Weeks after beginning of trial, April 29, 1975.
\ Harvest date during the major rainy season of 1975*
o Data followed by the same letters are not different (P>-0.05): rows - a,b,c,d,e,f; columns within
plant part: leaves - x,y,z; stems - q,r,s; whole plants - t,u,v.
University of Ghana          http://ugspace.ug.edu.gh
Appendix LII. Analyses of variance of yield of In Vitro digestible dry matter and nitrogen of whole
plants of giant star, buffel and pangola grasses during the minor rainy season of 1974
and the major rainy season of 1975*
(Sums of squares x 104)
Minor rainy season 1974 Ma.ior rainy season 1975
Source
Degrees of 
freedom
Digestible 
dry matter Nitrogen
Degrees of 
freedom
Digestible 
dry matter Nitrogen
Replications 3 97-90 0.0015 3 53.07 0.0063
Species 1 238.60 0.0002 2 995.90** 0.0973
Error (A) 3 119.10 0.0015 6 99.72 0.0493
Dates 9 1118.80** 0.0040 9 6668.00** 0 .2318**
Date x species 9 106.70 0.0013 18 424.IO** 0.0346
Error (B) 54 655.IO 0.0082 81 539.10 0.1088
* Significant (P <10.05).
** Significant (P^O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix LIII. Analyses of variance of weekly yields of In Vitro digestible dry matter (leaves vs. stems) of
giant star, buffel and pangola grasses during the minor rainy season of 1974 (upper) and the
major rainy season of 1975 (lower).
(Sums of squares x 104)
1974 Degrees Maturity
of 3+ 4 5 6 7 8 9 11 13 15
Source freedom Oct. Oct. 15 Oct. 22 Oct. 29 Nov. 5 Nov. 12 Nov. 19 Dec. 3 Dec. 17 Dec. 31
Reps 3 1.09 3.29 9-35 4.01 6.18 6.19 93.54 12.17 18-35 10.87
Species 1 0.19 10.02 5.28 13-93 3.17 O .56 44-41 34.78 43.46 16.83
Error (a) 3 8.85 4.31 1-55 49.58 26.94 12.83 35-66 24.43 74-22 32.72
Leaves vs. 
stems 1 24-39* 56.48* 28.15* 0.03 1.31 24.97* 53.24 8 .I4 62.10 0.20
L-S x species 1 0.30 11.69 0.01 72.64 63.57* 61.25** 137.70 95.61* 330.70** 138.20**
Error (B) 6 15.56 41.43 23.82 46.74 62.69 I6.65 276.40 48.41 89.93 56-49
1975 Degrees Maturity
Source
of
freedom
3+ +
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Reps 
Species 
Error (a)
3
2
6
0-12
9.10**
O .84
3.02
94-44*
9-12
1.98
46.57*
14.34
15.52
40.52*
18.07
18.11
67.62*
22.84
5.74
3-08
56.32
2.75
113.40*
39.42
33-62
159.50**
19.98
8.55
II6.4O**
23.66
27.60
56.19**
23.03
Leaves vs. 
stems 1 19.74** I65.7O** 147.80** 59.31** 88.18** 212.50** 132.20** 8O.65** 26.72* 0-35
L-S x species 2 
Error (B) 9
9.21**
1.08
43.20**
5-90
1.09
4-29
15.27*
6.30
4.00
19.89
20.26
25.88
65.23**
10 .70
43.70**
13.50
63.50*
51.72
42.53*
35.92
+ Weeks from beginning of trial, September 16, 1974* April 29, 1975*
+ Harvest date during the minor rainy season 1974 °r the major rainy season 1975* 
* Significant (P-<-0.05).
** Significant (P 0.01).
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________________________________   (gg/ha)_________________________
Appendix LIV. Yield of In Vitro digestible dry matter of leaves, stems and whole plants of giant star and
• buffel grasses during the minor rainy season of 1974*
Maturity
3+ .
Oct. 8+
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves
Giant star 642f 9592 946® 1071| 985y 869* 980® 874y H47x 880*
Buffel 598* 1000®* 1210®* 1120®* 1148®* 1003®* 1286® 1163^* 11435* 987|*
Stems
Giant star 390* 531^ 817^ 1054®*° 1177a* 1286® 888° 882° 781® 945*°
Buffel 380* 741° 814*c 1246® 1126® 1137® 1001q*c 1111® 1006®*°q IO64**
Whole plants
Giant star 1032^ 1490*° 1763|-* 2125| 2l62f 2046|* 1868®* 1756|* 1928®* 1825|*
Buffel 978£ 1741? 2024^* 2366® 2274®* 2140|* 2287®* 2274®* 2149®* 2051I*
+ Weeks from beginning of trial, September 16, 1974*
+ Harvest date during the minor rainy season of 1974*
o Data followed by the same letter are not different (P.^0.05): rows - a,b,c,d; columns within plant
part: leaves - x,y; stems - q,r; whole plants - t,u.
University of Ghana          http://ugspace.ug.edu.gh
____________   (Kg/ha)__________________________________________________
Appendix LV. Yield of In Vitro digestible dry matter of leaves, stems and whole plants of giant star, buffel
and pangola grasses during the major rainy season of 1975*
Maturity
3+ t
May 20+
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Leaves 
Giant star 
Buffel 
Pangola
m f
386*
93*
434yf 
1157xc 
47 3y
669*e
973$
693*°
633*e
928°
887*
1188*°
1362*
898*
1561a
1412b
1318a
1289a* 
2046 a 
1218a
1238ab
172ia
1033|
1005*°
1775|
1075a
855|*
1320*°
1033a
Stems
Giant star 
Buf f el 
Pangola
5<
34q
30|
130°*
254*e
104*e
220°*
480°d 
q
141 *e
431*°
753*°
413*
70-2®*
942®*
626°
70 gab
q
957qb
872*°
923? 
1137^ 
1081ab
778|
1324|
103ia*
802*
H44|
1252|
865t
1017|
13875
Whole plants 
Giant star 
Buffel 
Pangola
233?u
420?
123«
564*°
1411?
577*e
889* 
1453? 
834°*
IO64*
1681$
1300*°
l890g
2304$
1524*
2270|
2369$
2190a
22125
3183|
2299g
20165
3045|
2064g
1807*
2919|
23275
1720g
2337$ 
2420 a
+ Weeks from beginning of trial, April 29, 1975*
+ Harvest date during the major rainy season.
o Data followed by the same letter are not different (P/^0.05): rows - a,b,c,d,e; columns within plant
part: leaves - x,y,z; stems - q,r,s; whole plants - t,u,v.
University of Ghana          http://ugspace.ug.edu.gh
_______ _______________________________________ (Sums of squares)_____________________  _________
Maturity
Appendix LVI. Analyses of variance of weekly yields of nitrogen (leaves vs. stems) of giant star, buffel and
pangola grasses during the minor rainy season of 1974 (upper) and the major rainy season of
1975 (lower).
Source
of
freedom
3 ±
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Reps 
Speed es 
Error (a)
3
2
6
54.11
6.83
79-76
54.93
97-47
176.90
198.80
177.90*
43.82
47-77
103.30
25I.4O
35-43 
10.60 
141.90
240.20
0.24
250.90
61.09
114.10
99.16
119.60
I6O.4O
95-10
415.OO
333-80
425.10
686.20
158.10
264.OO
Leaves vs. 
stems 1 804.70** 1609.00** 1295.00** 776.60** 668.20** 235.3O** 667.80 712.10** I486.OO 488.30
L-S x species 2 
Error (B) 9
7.44
132.50
94.92 
251.70
9-70
169.10
279.90
167.86
106.80
261.90
20.66
69-07
104.10
674.90
I46.9O
307.60
680.70
752.70
235.90
466.70
1975 Degrees Maturity
Source
of
freedom
3+ +
May 20*
4
May 27
5
June 3
6
June 10
7
June 17
8
June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Reps 3 O .96 29-35 21.09 100.90 131.10 103-00 134.50 137.50 31.23 24.28
Species 2 146.IO** 1313.00** 491.10* 180.90 230.60 18.52 381.20 601.60 325.90* 81.05
Error (A) 
Leaves vs.
6 10.21 120.30 144.30 318.70 298.60 935-60 931.40 517.40 114.20 352.70
stems 1 324.20** 2972.00** 3094.00** 2791.00** 3250.00** 4690.00** 3784.00** 2137.00** 1531.00** 957.90**
L-S x species 2 I4I.8O** 825.30** 34.59 26.95 I65.7O* 203.30 362.40* 182.00* 56.20 128.40
Error (b) 9 8.92 97.41 125.60 253.60 103.70 338.60 277.60 142.20 87.84 146.30
+ VJeeks from beginning of trial, September 16, 1974* April 29• 1975-
+ Harvest date during the minor rainy season 1974> °r the major rainy season 1975*
* Significant (p- O.O5).
** Significant (P O.Ol).
University of Ghana          http://ugspace.ug.edu.gh
Appendix LVII. Yield of nitrogen of leaves, stems and whole plants of giant star and buffel grasses during
the minor rainy season of 1974*
(Kg/ha)
Maturity
3+ .
Oct. 8*
4
Oct. 15
5
Oct. 22
6
Oct. 29
7
Nov. 5
8
Nov. 12
9
Nov. 19
11 
Dec. 3
13 
Dec. 17
15
Dec. 31
Leaves
Giant star 21.4$ 28.9* 24«4y° 27.05* 29.65 23.55* 24.9*0 27.05* 33.8| 23.6od
Buffel 21.5f 30.5* 37.15 26.75* 27.80 19.9x 23.65 26.95 24.1*e 20. o£
Stems
Giant star 7-0® 8>4qe n.i*° 12.9$ 15.75 14 .85 10.15* 12.1* 9-1* 9.2*
Buffel 7-8* i4.o§ 14.7*c 15.7JO 14.9*0 14.35C 17.85 16.6* 15.8JO
Whole plants
Giant star 28.4f 37-3? 35.5| 39.9? 45- 3| 38,3?d 35.0? 39*1^* 42.9* 32.8(
Buffel 29.3f 42.4?d 51.if 41.4? 43*5*° 34.8{ 37.91 44.7? 40.7* 35-8f
+ Weeks from beginning of trial, September 16, 1974*
+ Harvest date during the minor rainy season of 1974*
o Data followed by the same letter are not different (P.^0.05): rows - a,b,c,d,e,f,g; columns within 
plant part: leaves - x,y; stems - q,r; whole plants - t,u.
University of Ghana          http://ugspace.ug.edu.gh
 _______________________________________(Kg/ha)_____________________________________
Appendix LVIII. fields of nitrogen of leaves, stems and whole plants of giant star, buffel and pangola
grasses during the major rainy season of 1975*
Maturity
3+ 4 5 6 7 8 
May 20+ May 27 June 3 June 10 June 17 June 24
9
July 1
11 
July 15
13 
July 29
15
Aug. 12
Leaves 
Giant star 
Buffel 
Pangola
p°
5-5$ 
14-9| 
3-71
14-9|
44-2|b
18-2y
31'6y°
35-8cd
27-8bc
35-4xb
31-Od 
33-0|b
36.oab
39-8^
' 29-s5
43-2|
38.3bc
38.5|
38.4fb
48.8-
34.iab
29-9y
36-7bc
24-5°
23-9°
7
36-3bc 
20.8?
25.2° 
23.9* 
26.4bc
Stems
Giant star
Buffel
Pangola
°*7b
0.8-
0 .6c
l.6®b
5.6£c
3-lqr
2.8®b
1 1.2ab
q
3-4°
5-9jib
13-8-
10.1%
8.6® 
15*5q 
12.4b°
7-I f
15.lq 
I5.7|b
9-4*
17'4qr
I8.5®
7*3®b 
18.4| 
12-9qb
' * r
13*lqb
8.9b
6.7pb 
16.6® 
13.2qb
Whole plants 
Giant star 
Buffel 
Pangola
6.2®
15-7?
4-4
16.5v 
49-8$ 
21.3g
34-45°
47.o£c
31-2°
41-35 
44-8?
43*lbu
44.6g
55-3$ 
41.9b
50 .35
53.4L
54.2a
47-8£b
66.23-
52.4
37-2^°
55. l£
37.4bc
31. lg 
49-4$ 
29-7*
31.9g
40.5?
39.6^
+ Weeks from beginning of trial, April 29, 1975*
+ Harvest date during the major rainy season 1975*
o Data followed by the same letter are not different (P^>0.05): rows - a,b,c,d,e,f; columns within
plant part: leaves - x,y,z; stems - q,r,s; whole plants - t,u,v.
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Major rainy season 3 974
I
Weekly distribution of rainfall during 1974-
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Dry season 1974
i i t
L._
1 8  1 5  2 2  2 9  5  1 2  1 9  2 6  
JULY AUGUST
University of Ghana          http://ugspace.ug.edu.gh
M
M
120 •
80 c
Major rainy season 1975
I
Dry season 1975i i i
9  16 2 3  3 0
APRIL
4  II
JUNE
LEGEND J  COMMENCEMENT  OF HARVESTS REGROWTH HARVEST DATES.
■ipendix L/.. '•teekly distribution of rainfall during 1975-
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Rpc-ndix I,XI. Heekly distribution of rainfall during the minor rainy season of 1974 and the ensuing
dry period.
189
University of Ghana          http://ugspace.ug.edu.gh
Aivendix  I )
R
A
N 120.
80 •
40*
MAJOR RAINY SEASON 1975 DRY SEASON 1 9 75
i i ±
2  9  16
SEPTEMBER
LEGEND: ^ COMMENCEMENT  OF HARVESTS.  ^  REGROWTH HARVEST DATES.
vOO
CTI. Weekly distribution of rainfall during the major rainy season of 1975 arid the ensuing 
dry period.
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