Livestock Science 107 (2007) 253 – 264

www.elsevier.com/locate/livsci

Nutritive value of forage species grown in the warm temperate

climate of Australia for dairy cows: Grasses and legumes
W.J. Fulkerson a,⁎, J.S. Neal a , C.F. Clark a , A. Horadagoda a ,
K.S. Nandra b , I. Barchia b
a
University of Sydney, Camden, New South Wales 2570, Australia
b
Elizabeth Macarthur Agricultural Institute, Menangle, New South Wales 2570, Australia
Received 28 March 2006; received in revised form 18 September 2006; accepted 18 September 2006

Abstract

The objective of this study was to quantify the differences in the nutritive value over 4 seasons, of 7 C3 temperate grasses, 2 C4
tropical grasses and 11 clover species used as forages for dairy cows. The nutritive value was assessed in terms of nutrient content
and the availability of effective rumen degradable protein, rumen by pass protein, metabolisable protein (MP) and fermentable
metabolisable energy.
All species were grown in plots as monocultures under conditions of non limiting nutrients and moisture and harvested by
mechanical means. All species had a high crude protein content and this resulted in a high effective rumen degradable protein:
fermentable metabolisable energy ratio varying from 15, for cowpeas (Vigna unguiculata) to 29 for birdsfoot clover (Lotus
corniculatus), and all were above the ratio of 11 required for optimal microbial protein synthesis in the rumen of dairy cows. The
calculated availability of MP varied from 105 g/kg dry matter (DM) for cowpeas to 173 g/kg DM for berseem clover (Trifolium
alexandrinum) indicating that all forages would be able to meet the requirements of dairy cows producing up to 30 L/milk/day,
provided they were able to consume over 19 kg DM of forage/cow/day.
Grasses had much higher hemicellulose (neutral detergent fibre minus acid detergent fibre) content than legumes. Kikuyu
(Pennisetum clandestinum), a C4 grass, had a higher proportion of hemicellulose content than the C3 temperate grasses. Perennial
ryegrass (Lolium perenne) and kikuyu had a similar metabolisable energy (ME) density (9.9 MJ/kg DM) in summer. The mean ME
density of perennial ryegrass, prairie grass (Bromus wildinowii) and short rotation ryegrass (Lolium multiflorum) winter was similar
at 10.6 MJ/kg DM and slightly higher than cocksfoot (Dactylus glomeratus), phalaris (Phalaris tuberosa) and fescue (Fescue
arundunicea) which had a mean ME density of 10 MJ/kg DM.
All forages grown were able to satisfy MP and ME requirements of dairy cows producing up to 30 L milk/cow/day, provided
they were able to consume sufficient forage to achieve this level of production.
© 2006 Elsevier B.V. All rights reserved.

Keywords: Rumen degradability; Metabolisable protein; Nutritive value; Forages; Dairy cows

1. Introduction

⁎ Corresponding author. Tel.: +61 2 93511635; fax: +61 2 46552374. Grazed pasture is the predominant feed source for
E-mail address: billf@camden.usyd.edu.au (W.J. Fulkerson). dairy cows in Australia. However, most cows are fed
1871-1413/$ - see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.livsci.2006.09.029
254 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264

some supplement to address quantity and quality deficits whilst the CP content of kikuyu (Pennisetum clandes-
in pasture availability. The predominant pasture species tinum), another pasture species of major importance to
used to graze dairy cattle is perennial ryegrass (Lolium the dairy industry, can be over 20% (Fulkerson et al.,
perenne) for reasons of high nutrient quality and, under 1998). Thus, these pastures can provide protein levels in
an appropriate environment, a long growing season. excess to cow requirements (NRC, 2001). While ME
However, the climate in most dairy regions of Australia requirements of ruminants can be calculated reasonably
is marginal for the growth of perennial ryegrass with accurately (ARC, 1980), determination of protein re-
many farmers reestablishing pastures every 2–4 years or quirements has proven more difficult and an under-
oversowing on an annual basis. Previous studies by standing of ruminant requirements lag behind those of
Fulkerson et al. (2003), confirmed the lack of mono-gastric species.
persistence of perennial ryegrass in diverse dairy regions Protein digestion in ruminants is a two-stage process
of Australia. Furthermore, these studies indicated that from (1) microbial protein synthesized in the rumen
management could only go so far to improve persis- from feed degradation (and non-protein-N (NPN)), and
tence. In this context, the dairy industry was interested (2) feed protein which reaches the small intestine intact
in alternative forage species to perennial ryegrass with i.e. by passing degradation in the rumen (UDP).
selection based on more industry-relevant criterion such Recognition of these processes led to development of
as water use efficiency, nutrient content and palatability, the rumen degradable protein (RDP)/UDP system
than yield which has been the predominant selection (ARC, 1980, 1984). Recent feeding standards (e.g.
criteria in the past. AFRC, 1993) suggest that protein requirements of
Nutrient (protein, carbohydrates, minerals and vita- ruminants should be based on metabolisable protein
mins) content and metabolisable energy (ME) density (MP). The major advance in using MP over the RDP/
changes in relation to season (Fulkerson et al., 1998; UDP system is that microbial protein availability is
Smith et al., 1998; Stockdale, 1999), stage of growth
(Reeves et al., 1996; Mulholland et al., 1996; Nandra
et al., 1998; Ayres et al., 1998; Fulkerson et al., 1999),
Table 1
time of day (Minson, 1990; Fulkerson et al., 1995; Reeves Details of forage species evaluated for nutrient content
et al., 1996; Lindgren and Lindberg, 1998), soil fertility
Forage species Cultivar
or fertilizer application rate (particularly nitrogen (N))
(Reeves et al., 1996) and probably soil moisture status. Common name Scientific name
Thus, an awareness of the factors influencing nutrient Temperate grasses
content of forage is required to allow more efficient Perennial ryegrass Lolium perenne Bronsyn
Cocksfoot Dactylus glomerata Kara
supplementation of animals.
Fescue Fescue arundinacea Quantum
In a pasture-based farming system, energy is usually Phalaris Phalaris tuberosa Holdfast
the first limiting nutrient for milk production of dairy Prairie grass Bromus wildenowii Matua
cows (van Vuuren, 1993). The nonstructural carbohy- Short rotation Ll. multiflorum Concord and Tetila
drates (NSC) (such as low molecular-weight sugars, ryegrass
starch and fructosans), and structural polysaccharides or
Perennial legumes
dietary fibre (consisting of cellulose, hemicellulose and Lucerne Medicago sativa Spectre
lignin) which comprise the principal components of cell Red clover Trifolium pratense Astred
walls (Theander et al., 1993) are considered to be the White clover Trifolium repens Kopu
primary energy source for ruminant animals. However, Birdsfoot clover Lotus corniculatus Dawn
the fibre content of ryegrass can vary from being deficient
Annual legumes
when the plant is in a vegetative state (between May and Sulla Hedysarum coronarium Aokan
August) (Fulkerson et al., 1998), to excess (Delagarde Lablab Lablab purpureus Highwarth
et al., 2000), after seed set or where moisture is deficient in Cow pea Vigna unguiculata Red Caloon
summer. Non-structural carbohydrates are important in Persian clover Trifolium resupinatum Shaftal
Berseem clover Trifolium alexandrenum Elite-11
that they are a readily fermentable source of energy in the
Balansa clover Trifolium michelianum Paradena
rumen and may also be important in synchronizing with Melilotus Melilotus indicus Vicia
rumen ammonia from the high protein intakes common in
cows grazing pasture (Trevaskis et al., 2004). Tropical grasses
The crude protein (CP) content of perennial ryegrass Kikuyu Pennisetum clandestinum Whittet
Paspalum Paspalum dilatatum Common
is commonly over 25% and can be as high as 35%,
 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264
related to fermentable metabolisable energy (FME) or
the portion of ME which can supply energy-yielding
ATP to rumen microbes. However, this improved sys-
tem is still not widely used because reliable information
on protein and organic matter (OM) degradation char-
acteristics of forages is lacking.
The objective of this study was to quantify the dif-
ferences in the nutritive value of forages based on chem-
ical composition and digestibility, and also the availability
of RDP, UDP and MP, calculated from the in sacco
degradation characteristics of CP and OM in the rumen of
various species of grasses and legumes commonly fed to
dairy cows.
2. Materials and methods
In this study, 19 grass and legume species (see
Table 1) were compared during the 12 month period in
2004, when grown under identical climatic and edaphic
conditions and under best practice management (see
Fig. 1. A – Long-term (143 years) average, B – actual for 2004, monthly rainfall (mm) and monthly minimum and maximum temperature (°C) for the
site.
255
later) with soil nutrient and moisture availability
predicted to be non-limiting to growth.
2.1. Climatic conditions
Fig. 1 shows the actual and long term monthly
rainfall and maximum and minimum temperature for the
experimental site during the year when the herbage
samples were taken for nutrient analysis.
2.2. Experimental design
The experimental layout was a randomized block
design with 3 replicates. The plots were 5 ⁎ 5 m with
herbage samples taken from the 2 × 2 m in the middle of
the plots. Sprinklers at each corner of the 2 × 2 m area
ensured maximum uniformity of water coverage.
Irrigation supplemented rainfall such that the soil was
refilled when 30 mm of water had been used in the top
30 cm soil profile. The moisture in the soil profile was
256 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264
estimated from weather data (rainfall, evaporation, crop
factor) and then monitored by a neutron probe with
measurements taken from access tubes placed to a depth
of 2 m in the centre of each plot to confirm the estimates
obtained from weather data.
2.3. Application of fertilizers
At the commencement of the study, a capital dressing
of 600 kg/ha of mixed fertilizer (22:4: 23:10 N;
phosphorus (P); potassium (K); sulphur (S), respective-
ly) was applied to all plots. The N (80% there of and not
on plots with legume species), P, K and S removed at
each harvest, was then returned as mixed fertilizer with
the amount applied derived from the actual DM yield at
each harvest multiplied by value indicating adequate
availability viz. N = 4%: P = 0.35%, K = 3.5% nutrient to
the plant. Plant tissue and soil were monitored at regular
intervals to determine adequacy of fertilizer application.
These practices attempted to ensure nutrients and
moisture did not limit growth as far as possible whilst
ensuring nitrate (NO3) and K did not accumulate to the
extent where they would be toxic to animals grazing
such forages.
2.4. Establishment and defoliation management
A challenging issue for this study was to determine the
most appropriate establishment and management needs
for such a diverse range of forage species. For most
species, recommendations on establishment were gener-
ally available. Likewise, the optimal time and height of
defoliation were known for species. In these species, the
correct stage for defoliation could be related to in field
indicators. For example, the optimal time to defoliate
perennial ryegrass and fescue is when 3 new leaves have
grown (Fulkerson and Donaghy, 2001); for prairie grass it
is 4 leaves (Fulkerson et al., 2000); for cocksfoot
(Rawnsley et al., 2001) and kikuyu (Reeves et al., 1996)
it is 4–5 leaves; for lucerne it is when flowering com-
mences and for others it is when the oldest leaf begins to
senesce at the bottom of canopy. For both phalaris and
prairie grass, defoliation is deferred during seed set in late
spring/early summer; for phalaris this builds up plant
reserves to cope with stress of hot dry summer conditions;
and for prairie grass this allow seedling recruitment in
autumn to replace plants that have died over summer.
Defoliation height varied but was between 5 and 8 cm,
with height increasing in late spring to reduce evaporative
loss.
For other species we had to be guided by principles
based on allowing sufficient time for the plant to re-
plenish its reserves (soluble carbohydrates), but before
the oldest foliage began to senesce, leading to a fall in
herbage quality and palatability. The application of these
principles to development of optimal grazing manage-
ment strategies as they apply to ryegrass is outlined in
Fulkerson and Donaghy (2001). This ensured regrowth
would be maximized while optimizing nutrient contents
and palatability.
2.5. Sample collection and preparation
When the central 2 × 2 m plot was mown, representa-
tive herbage samples were taken and dried at 70 °C for
48 h in a forced-draught oven to determine DM yield at
each harvest. These samples were pooled within each of 4
seasons (Spring: September 1 to November 30, Summer:
December 1 to February 28, Autumn: March 1 to May 30
and Winter: June 1 to August 30) and analysed for nutrient
content and rumen degradability characteristics. Thus, the
values for nutrient content represent the seasonal herbage
analysed in duplicate samples.
Herbage samples, for determination of rumen
degradability, were ground through a 2.5 mm screen
and fine particles removed by sieving through a 45 μm
screen. A portion of herbage sample was also ground
through a 1 mm screen for chemical analysis and in-
vitro dry matter digestibility (DMD).
2.6. Rumen degradability
The rumen degradability of CP and OM for forages
were determined, using six rumen cannulated castrate
Merino sheep (4–5 years old, mean live weight 55 kg).
These sheep were housed in individual crates and fed a
basal diet of lucerne hay and crushed barley (ratio of 4:1)
to predicted maintenance ME requirements in two equal
amounts at 08.30 and 17.00 h. The sheep were randomly
divided into 2 groups of 3 and two forage species were
tested on any one occasion, thus sheep formed replicates.
Samples of the test feed (4 g) were weighed into each
of eight nylon bags, seven of which were then placed
into the rumen of sheep before the morning feed and one
bag retrieved after 2, 4, 8, 16, 24, 48 and 72 h of
incubation in the rumen. After retrieval, the bags were
washed through the cold rinse cycle of a washing
machine. The value at time 0 h was obtained by washing
one bag containing test feed before placing bags in the
rumen (time = 0 h). The bags were then dried at 70 °C for
72 h in a forced-draught oven. The disappearance of DM
was measured as the loss of weight of each bag. The
residual material in each bag was then ground through a
1 mm screen and analysed for N and ash content.
 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264
Data on CP and OM loss from the bags at different
times in the rumen were fitted to Eq. (1) (McDonald,
1981):
P ¼ A for tbt0 and P ¼ a þ bð1−e−ct Þ for tNt0 ð1Þ
The term A represents washing loss and t0 lag time.
Potential degradability of CP (PDGCP) and OM
(PDGOM) was calculated as (a + b) and insoluble but
degradable fractions (BCP and BOM) as (a + b − A). The
coefficient c represents degradation rate of the insoluble
but potentially degradable fraction B. In the case where
t0 ≤ 0, the effective degradability's of CP (EDGCP) or
OM (EDGOM) were calculated as EDG = a + bc / (c + k)
(Orskov and McDonald, 1979) and in the case where
t0 N 0 the EDG was calculated as EDG = a + bce−(c+k)to /
(c + k) (McDonald, 1981). A particle outflow rate (k) of
0.08/h was used as this is typical of outflow rates found
in lactating cows.
2.7. Calculation of various protein components and
fermentable metabolisable energy
The following protein fractions were calculated
according to AFRC, 1993: effective rumen degradable
protein (ERDP) – the portion of RDP captured and
utilized by rumen microbes for growth and microbial
protein synthesis; digestible undegraded protein (DUP) –
the portion of UDP digested and absorbed in the lower
intestine; and MP – the total digestible true protein
available to the animal for metabolism after digestion and
absorption.
The pepsin–cellulase digestibility assay of Clark
et al. (1982) was used to determine in-vitro DMD of
forages. The ME was then calculated from DMD using
Eq. (2) (SCA, 1990):
MEðMJ=kg DMÞ ¼ ð%DMD⁎0:17Þ−2:0 ð2Þ
Fermentable metabolisable energy was calculated
using Eq. (3).
FMEðMJ=kg DMÞ ¼ Total ME−MEfat −MEDUP ð3Þ
To calculate FME, account had been taken of the
contribution of ME from dietary fat to total ME and the
ME attributable to a portion of UDP which was digested
(DUP) after by passing from the rumen. Digestible
undegradable protein was calculated from UDP and acid
detergent insoluble N (ADIN) using Eq. (4) (AFRC,
1993).
DUPðg=kg DMÞ ¼ 0:9  ðUDP−6:25⁎ADINÞ ð4Þ
257
2.8. Chemical analysis
Total N content of forages was determined by
combustion (990.03; AOAC, 1990) using a Leco® FP-
428 Nitrogen Determinator (Leco® Corporation St.
Joseph, Michigan, U.S.A.).Crude protein was then
calculated by multiplying the N content by 6.25.
True protein-N was determined according to Marias
and Evanwell (1983). The method was based on the
precipitation of protein with trichloroacetic acid and the
separation of the insoluble protein from the soluble non-
protein fraction by filtration and true protein N content
in precipitate was determined using Leco. Non-protein
N was calculated by the difference between total and
true protein-N.
The determination of NO3–N was based on the
nitration of salicylic acid under highly acidic conditions
and colorimetric determination of the resulting colored
complex, which absorbs maximally at 410 nm in basic
solution (Catalado et al., 1975).
Forages were also subjected to the fibre analyses:
acid detergent fibre (ADF) (973.18; AOAC, 1990) and
neutral detergent fibre (NDF) (adapted from van Soest
et al., 1991). The ADIN content of forages was deter-
mined on the ADF residues. Neutral detergent fibre was
analysed without the use of alpha amylase but with
sodium sulphite and the results were expressed with
residual ash. The hemicellulose content was calculated
as the difference between ADF and NDF.
The water soluble carbohydrate (WSC) content of
forages was determined using a modification of the
method described by Smith (1969). The dried samples
were ground through a 1 mm sieve and then extracted in
a reciprocal shaker for an hour using 0.2% benzoic
acid–water solution, following which the carbohydrates
soluble in cold water was hydrolysed by hydrochloric
acid. The resultant solution was heated with alkaline
potassium fericyanide at 90 °C, and then read at 420 nm
using an auto analyser. Crude fat in forages was
determined using Soxhlet apparatus (920.39; AOAC,
1990).
2.9. Statistical analysis
The rumen degradability parameters and protein com-
ponents were compared between pasture species using the
following model:
Response ¼ sheep þ pasture species þ error
The sheep and error are assumed to have random
effects. Because the sheep and pasture species effects
258 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264

Table 2a
Neutral detergent fibre (NDF), acid detergent fibre (ADF), crude protein (CP), non-protein nitrogen (NPN), nitrate–nitrogen (NO3–N), acid detergent
insoluble nitrogen (ADIN), water soluble carbohydrate (WSC) and fat expressed in g/kg DM and metabolisable energy (ME) in MJ/kg DM of grasses
during summer and autumn
Species NDF ADF CP NPN NO3–N ADIN WSC Fat ME
Summer
Perennial ryegrass 515 313 221 8.6 3.3 1.6 57 27 9.9
Cocksfoot 594 323 228 17.6 3.1 2.8 49 34 9.7
Fescue 554 305 216 17.0 3.8 2.2 77 24 9.9
Phalaris 483 290 241 7.8 – 1.8 41 28 9.6
Prairie grass 548 299 246 8.0 2.8 2.1 38 26 10.7
Kikuyu 503 259 253 11.4 5.2 2.3 37 23 9.9

Autumn
Perennial ryegrass 497 266 240 8.8 3.5 1.8 53 – 10.0
Cocksfoot 556 299 227 8.0 3.2 21 – 9.3
Fescue 508 263 269 10.5 3.5 1.8 10 – 9.5
⁎S.R. ryegrass (Concord) 428 234 265 11.2 4.9 2.9 34 23 11.0
⁎S.R. ryegrass (Tetila) 411 233 285 10.0 4.7 4.2 47 25 11.0
Phalaris 417 206 291 10.3 2.9 1.8 25 – 10.4
Prairie grass 519 270 285 9.7 3.2 – 26 – 9.7
Kikuyu 535 231 293 5.6 4.9 – 20 – 9.2
Paspalum 604 304 198 2.2 1.7 3.2 33 23 9.0
⁎S.R. Short rotation.

were not independent, the residual maximum likelihood comparisons. All the analyses were run on Genstat for
estimation was used to estimate the parameters and their Windows 7th edition (VSN International Ltd, 2003).
variance, which allowed the computation of the least Data on the nutritive value of different groups of forage
significant difference at the 5% level for feed type species were analysed using the conventional analysis of

Table 2b
Neutral detergent fibre (NDF), acid detergent fibre (ADF), crude protein (CP), non-protein nitrogen (NPN), nitrate–nitrogen (NO3–N), acid detergent
insoluble nitrogen (ADIN), water soluble carbohydrate (WSC) and fat expressed in g/kg DM and metabolisable energy (ME) in MJ/kg DM of grasses
during winter and spring
Species NDF ADF Protein NPN NO3–N ADIN WSC Fat ME
Winter
Perennial ryegrass 489 232 243 8.9 1.1 1.1 78 27 11.4
Cocksfoot 485 257 319 21.5 1.1 1.1 21 34 10.5
Fescue 488 233 253 18.9 1.7 1.2 53 24 10.6
Phalaris 553 254 285 6.6 3.2 1.3 22 28 10.2
Prairie grass 437 240 310 19.3 2.5 1.2 141 26 11.2
⁎S.R. ryegrass (Concord) 422 236 264 23.3 4.0 2.9 23 23 11.0
⁎S.R. ryegrass (Tetila) 459 241 271 24.5 4.2 4.0 35 25 10.5

Spring
Perennial ryegrass 552 259 263 12.9 1.3 1.8 63 – 11.1
Cocksfoot 586 279 267 9.5 1.4 2.2 44 – 10.2
Fescue 542 284 267 16.5 1.4 2.4 56 – 9.7
Paspalum 672 327 229 9.8 1.4 3.7 57 22 8.9
Phalaris 524 267 271 14.9 1.5 1.7 32 – 10.5
Prairie grass 555 270 264 13.8 0.9 1.9 76 – 10.4
⁎S.R. ryegrass (Concord) 531 277 247 17.8 1.3 2.3 63 – 9.7
⁎S.R. ryegrass (Tetila) 495 262 256 17.6 2.1 2.0 75 – 10.4
Kikuyu 639 266 228 12.8 1.7 2.7 31 – 9.5
⁎S.R. Short rotation.
 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264 259

Table 3a
Neutral detergent fibre (NDF), acid detergent fibre (ADF), crude protein (CP), non-protein nitrogen (NPN), nitrate–nitrogen (NO3–N), acid detergent
insoluble nitrogen (ADIN), water soluble carbohydrate (WSC) and fat expressed in g/kg DM and metabolisable energy (ME) in MJ/kg DM of
legumes during summer and autumn
Species NDF ADF Protein NPN NO3–N ADIN WSC Fat ME
Summer
Lucerne 358 283 206 8.2 1.3 1.6 81 13 9
Red clover 395 262 242 22.2 0.8 3.5 59 18 9.2
Sulla 378 296 218 21.1 – 2.7 49 25 9.3
White clover 276 210 242 16.1 0.8 2.0 41 18 10.0
Lab Lab 453 404 178 – 1.3 – 88 – 8.4
Cow pea 353 311 175 3.4 1.6 3.1 51 13 8.5

Autumn
Meli Lotus 254 213 277 14 2.6 – 54 – 9.5
Sulla 331 211 273 9.5 1.1 – 84 – 9.5
Persian clover 291 214 234 12.5 2.0 2.8 59 – 9.8
Balansa clover 307 231 253 15.5 1.0 2.0 44 32 9.4

variance and the difference between groups were tested
using least significant difference at 0.05 level.
3. Results
3.1. Seasonal changes
The nutrient content and ME values of temperate
and tropical grasses in summer and autumn are given in
Table 2a.
In summer, the NDF content of the grasses varied
from a high of 594 g/kg DM for cocksfoot to a low of
483 g/kg DM for phalaris with kikuyu also having a
relatively low value of 503 g/kg DM. Surprisingly, the
ADF comprised about 520 g/ kg NDF for both kikuyu
and perennial ryegrass and the ME at 9.9 MJ/kg DM
was also similar. The highest ME value was found in
prairie grass (10.7 MJ/kg DM). The lowest WSC values
were found in kikuyu in summer (and also in autumn)
with highest values for fescue and perennial ryegrass.
In autumn, the growth of newly established short
rotation (SR) ryegrass had a high ME value with low
NDF and ADF contents (Table 2a). As expected, by
autumn the ME of the two C4 grasses – kikuyu and

Table 3b
Neutral detergent fibre (NDF), acid detergent fibre (ADF), crude protein (CP), non-protein nitrogen (NPN), nitrate–nitrogen (NO3–N), acid detergent
insoluble nitrogen (ADIN), water soluble carbohydrate (WSC) and fat expressed in g/kg DM and metabolisable energy (ME) in MJ/kg DM of
legumes during winter and spring
Species NDF ADF Protein NPN NO3–N ADIN WSC Fat ME
Winter
Lucerne 472 246 301 19.4 0.2 – 55 – 9.3
Red clover 353 254 303 15.7 0.4 – 54 – 10.0
Sulla 416 381 288 15.0 0.2 2.7 47 25 8.6
White clover 291 193 298 15.5 0.7 1.0 90 18 10.5
Persian clover 414 219 310 13.1 0.3 4.2 54 22 10.1
Balansa clover 234 180 284 12.6 0.1 2.1 144 32 10.1
Berseem clover 327 175 274 24.6 0.1 3.1 63 25 9.9

Spring
Balansa Clover 382 286 257 16.1 0.4 2.8 41 – 9.0
Berseem clover 461 242 289 15.0 0.7 3.4 39 – 9.1
Birdsfoot clover 347 240 286 12.1 1.3 2.7 62 32 9.8
Lucerne 322 263 290 16.1 1.0 1.0 55 – 9.7
Meli lotus 341 227 261 12.9 1.9 0.9 70 – 9.4
Persian clover 392 257 266 13.5 0.9 1.4 48 – 9.4
Red clover 412 375 289 8.4 0.6 2.7 44 – 9.5
White clover 339 233 281 14.5 0.8 2.5 75 – 9.3
Sulla 489 248 283 7.9 0.5 3.4 27 – 8.1
260 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264

paspalum – was low; although an ME value of 9.2 MJ/ ADF) content in legumes (see Tables 2a and 3b).
kg DM for kikuyu is still reasonable. Similarly, the mean NDF content of the legume species
In winter (Table 2b), all grasses had a relatively low was 29% lower than perennial ryegrass but legumes had
NDF and ADF content, but phalaris was still the highest a higher proportion of NDF as ADF (66% in the
and this was reflected in ME values (11.4 MJ/kg DM for legumes and 52% for perennial ryegrass) giving a lower
perennial ryegrass and 10.2 MJ/kg DM for phalaris). hemicellulose content. For example, the estimated
Cocksfoot and prairie grass accumulated very high hemicellulose content of red clover in winter–spring
levels of protein, although the NPN and NO3 content of of 69 g/kgDM was 25% of the content in perennial
prairie grass was not abnormally high (see Table 2b). ryegrass. The ME values for legumes were significantly
In spring, temperate grasses had a high NDF content lower (P b 0.05) than for temperate grasses in winter but
(N500 g/kgDM) and a low CP, NPN and NO3 compared did not differ (P N 0.05) in other seasons (Table 3a).
to winter. In spite of its higher NDF content, kikuyu still All pasture legumes were higher in CP (mean of
had a reasonably high ME value of 9.5 MJ/kg DM, 294 g/kg DM) and ME (N10 MJ ME/kg DM except
possibly associated with the presence of a higher hemi- sulla and lucerne) in winter than in other seasons (see
cellulose (60% of NDF) content than other grasses. Table 3b).
As expected, the NDF content of the legumes was Interestingly, the legumes did not differ significantly
significantly lower (P b 0.05) than for grasses, but they (P b 0.05) in CP and NPN contents which were just as
had a much higher proportion of NDF as ADF (see high as in the C3 grasses in summer, autumn and winter,
Table 4) reflecting very low hemicellulose (NDF minus although the NO3–N content was significantly lower in
legumes (see Table 4).
Table 4
Neutral detergent fibre (NDF), acid detergent fibre (ADF), crude protein 3.2. Rumen degradability and protein components
(CP), non-protein nitrogen (NPN), nitrate–nitrogen (NO3–N), and water
soluble carbohydrate (WSC) expressed in g/kg DM and metabolisable The rumen degradability parameters of OM and pro-
energy (ME) in MJ/kg DM for various groups tein and the availability of FME, ERDP, UDP and MP
Groups Nutrient content (g/kgDM) contents of grasses and legumes are shown in Tables 5a
NDF ADF Protein NPN NO3–N WSC ME and 5b, respectively.
Summer
C3 grasses 539a 306a 230a 11.8a 3.25b 52a 9.96a 3.3. Temperate and tropical grasses
C4 grasses 503a 259a 253a 11.4a 5.20a 37a 9.90a
Legumes 369b 294a 210a 14.2a 1.16c 61a 9.07a Paspalum had a significantly lower (P b 0.05) content
lsd1 126 119 58 16.9 0.9 43 1.28
of readily soluble fraction of OM (AOM) and a higher
lsd2 70 67 32 9.3 0.5 24 0.72
fraction of protein (ACP) than kikuyu in summer. The
Autumn UDP and MP content of kikuyu was significantly higher
C3 grasses 477b 253a 266a 9.8a 3.70a 30.9b 10.13a (P b 0.05) than paspalum.
C4 grasses 570a 268a 246a 3.9b 3.30a,b 26.5b 9.10b In summer, prairie grass also had the highest UDP
Legumes 296c 217a 259a 12.9a 1.68b 60.3a 9.55a,b
and MP content, but ERDP content and the ratio of
lsd1 90 52 54 3.2 1.83 27.1 0.98
lsd2 70 41 43 2.5 1.43 21.2 0.76 ERDP to FME was lower in prairie grass than in other
C3 grass species. The ERDP content of C3 grasses in
Winter winter was higher than summer, resulting in a higher
C3 grasses 476a 242a 278a 17.6a 3.83a 53.3a 10.77a ERDP to FME ratio. The ratio of ERDP:FME was very
Legumes 358b 235a 294a 16.6a 0.29b 72.4a 9.79b
high (31:1) in cocksfoot in winter and lowest in
lsd2 76 59 26 6.7 2.47 46 0.63
perennial ryegrass (17:1).
Spring The species with the lowest UDP content in summer
C3 grasses 557a 278a 258b 14.1a 1.41a 58.3a 10.11a was fescue and in autumn was paspalum. The two C4
C4 grasses 639a 266a 228b 12.8a 1.70a,b 31.0a 9.50a,b summer grasses were very different in nutrient content
Legumes 387b 263a 278a 12.9a 0.90b 51.2a 9.26b
and this is exemplified by the fact that kikuyu had
lsd1 125 82 30 7.0 0.92 34.3 1.31
lsd2 58 38 14 3.24 0.42 15.8 0.61 nearly double the UDP content and 45% more MP than
paspalum in summer. The mean MP content of prairie
Note: lsd1 is least significant difference value at 5% level for com-
paring C3 grasses and C4 grasses and lsd2 is for comparing C3 grasses
grass (26% more) and SR ryegrass (Tetila) (16% more)
and legume. Means with different letters are significantly different at was significantly greater (P b 0.05) than perennial rye-
P b 0.05. grass in winter.
 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264 261

Table 5a
Rumen degradability parameters; water soluble component (A), water insoluble component (B), and rate of degradation of B component (c); effective
rumen degradable protein (ERDP), undegradable protein (UDP) and metabolisable protein (MP) in g/kgDM; ratio of ERDP: fermentable
metabolisable energy (FME) in g/MJ FME at out flow rate of 0.08 of grasses during summer and winter
Species AOM BOM cOM ACP BCP cCP ERDP ERDP:FME UDP MP
(g/g) (g/g) (/h) (g/g) (g/g) (/h) (g/kgDM) (g/MJ) (g/kgDM) (g/kgDM)
Summer
Perennial ryegrass 0.26d 0.62a 0.05a,b 0.49c 0.46a 0.05a,b 132b,c 19b 89c,d 120b,c
Cocksfoot 0.20b 0.69a,b 0.04a,b 0.29a 0.65d 0.05a 134c 20c,d 94c,d 121c
Fescue 0.24c,d 0.63a 0.07b 0.35b 0.56b,c 0.09b 140c 19b 76a,b 111b
Phalaris 0.30e 0.59a 0.05a,b 0.51c,d 0.46a,b 0.05a,b 144c,d 22e 97d 124c
Prairie grass 0.22b,c 0.76b,c 0.03a,b 0.34b 0.66d 0.03a 115a 16a 131e 157e
Kikuyu 0.34f 0.59a 0.05a,b 0.32a,b 0.65c,d 0.05a,b 132b,c 20c 121e 145d
Paspalum⁎ 0.16a 0.79c 0.02a 0.54d 0.46a 0.02a 128a,b 19b 69a 101a
s.e.d. 0.01 0.05 0.02 0.02 0.05 0.02 6.67 0.54 6.67 4.98

Winter
Perennial ryegrass 0.36c 0.59a,b 0.03a 0.49e 0.48a 0.02a 142a,b 17a 101a 139a,b
Cocksfoot 0.31b 0.62a,b 0.07a 0.36a,b,c 0.64b,c 0.01a 224d 31e 96a 127a
Fescue 0.32b 0.62a,b 0.06a 0.42d 0.52a,b 0.06a 152b,c 21c 101a 132a
Phalaris 0.36c 0.55a 0.06a 0.41c,d 0.52a,b 0.08c 166c 23d 119b 147b,c
Prairie grass 0.35c 0.53a 0.04a 0.38b,c,d 0.51a,b 0.04a 162c 22d 148c 173d
S.R. ryegrass Tetila 0.21a 0.78c 0.02a 0.34a,b 0.61b,c 0.03a 130a 19b 141c 164d
S.R. ryegrass Concord 0.23a 0.70b,c 0.03a 0.33a 0.65c 0.03a 166c 19b 123b 153c
s.e.d. 0.01 0.05 0.02 0.02 0.05 0.02 6.67 0.54 6.67 4.98
Note: Means with different superscripts are significantly different at P b 0.05. ⁎Sampled in autumn; s.e.d. denotes standard error of difference.

3.3.1. Pasture legumes highest content of UDP and MP was found in sulla. This
Birdsfoot clover had the highest ERDP value and this was probably due to the presence of a high value of Bpro
resulted in a high ERDP to FME ratio of 29 well above and a slower rate of degradation of protein (cCP) in the
the optimum of 11. Among the summer legumes, the rumen. White clover and perennial ryegrass did not

Table 5b
Degradability parameters; water soluble component (A), water insoluble component (B), and rate of degradation of B component (c); effective rumen
degradable protein (ERDP), undegradable protein (UDP) and metabolisable protein (MP) in g/kgDM; ratio of ERDP: fermentable metabolisable
energy (FME) in g/MJ FME at out flow rate of 0.08 of legumes and perennial ryegrass during summer and winter
Species AOM BOM cOM ACP BCP cCP ERDP ERDP:FME UDP MP
(g/g) (g/g) (/h) (g/g) (g/g) (/h) (g/kgDM) (g/MJ) (g/kgDM) (g/kgDM)
Summer
Perennial ryegrass 0.26b 0.62c 0.05a 0.49d 0.46b,c 0.05a,b 132b 19b 89b,c 120b
Birdsfoot clover⁎ 0.35e 0.52a,b 0.08a 0.61e 0.34a 0.07b,c 209d 29e 77a,b 109a
Cow pea 0.18a 0.61b,c 0.09a 0.25a 0.60d 0.10c 100a 15a 75a 105a
Lucerne 0.29c 0.46a 0.09a 0.53d 0.40a,b 0.07b,c 131b 19b 76a 107a
Red clover 0.29c 0.62c 0.08a 0.26a 0.69e 0.08b,c 134b 21c 108d 132c
Sulla 0.25b 0.56a,b,c 0.07a 0.31b 0.59d 0.03a 95a 16a 123e 143d
White clover 0.32d 0.60b,c 0.09a 0.42c 0.55c,d 0.07b,c 166c 23d 93c 126b,c
s.e.d. 0.01 0.05 0.02 0.02 0.05 0.02 6.67 0.54 6.67 4.98

Winter
Perennial ryegrass 0.36e 0.59a,b 0.03a 0.49c 0.48b 0.02a 142b 17a 101a 139a
Bansa clover 0.31c 0.60a,b 0.09a 0.36b 0.55b,c 0.08a 171c,d 26c 113a,b 138a
Berseem clover 0.27b 0.62b 0.04a 0.28a 0.65c 0.03a 115a 20b 159c 173b
Persian clover 0.22a 0.62b 0.08a,b 0.25a 0.59b,c 0.06a 160c 26c 150c 168b
Sulla⁎⁎ 0.35d,e 0.48a 0.06a,b 0.40b 0.28a 0.07a 131b 30d 157c 163b
White clover⁎⁎ 0.43f 0.54a,b 0.07a,b 0.39b 0.59c 0.06a 181d 26c 117b 144a
s.e.d. 0.01 0.05 0.02 0.02 0.05 0.02 6.67 0.54 6.67 4.98
Note: Means with different superscripts are significantly different at P b 0.05. ⁎Sampled in autumn, ⁎⁎sampled in spring; s.e.d. denotes standard error
of difference.
262 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264
differ significantly (P N 0.05) in their MP content
(Table 5b).
The ratio of ERDP: FME in all legumes in winter was
significantly higher (P b 0.05) than for perennial rye-
grass. In winter, the content of MP in berseem clover,
persian clover and sulla were significantly higher
(P b 0.05) than the other legumes and perennial ryegrass.
4. Discussion
The aim of the present study was to compare the
innate nutritive value of different species of grasses and
legumes relative to perennial ryegrass, the species
most commonly used to graze dairy cows in Australia.
Broadly speaking, nutritive value was high during
the cool-season months of autumn and winter with a
progressive decline in value through late spring to
summer, associated with reproductive development in
temperate species and the growth period for tropical
species of relatively lower nutrient value. The presence
of a lower content of NDF in phalaris and kikuyu in
summer probably reflects the plants are actively
growing at this time. However, the low fibre content
in SR ryegrass in the early vegetative state in autumn is
regarded as a problem in terms of dairy cow nutrition
associated with acidotic conditions in the rumen and
‘low milk fat’ syndrome in dairy cows. In winter,
cocksfoot and prairie grass accumulated very high levels
of protein, however, NPN and NO3 content was normal
and therefore poses no problems to animal health. In
spring, temperate grasses would have contained plants
setting seed, and this is reflected in high NDF (N 500 g/
kgDM). Reproductive development would also be a
reason for the rapid decline in CP, NPN and NO3 in
spring compared to winter.
The CP content of all grasses was high and reflects
the non-limiting N conditions under which the forages
were grown. Application of nitrogenous fertilizer in-
creases the CP content of grasses and increases CP yield
due to the simultaneous stimulation of growth. The
results are in line with those of Minson (1990) who
showed that the CP content was lowest in mid-summer
(170 g/kg DM) and highest in autumn (230 g/kg DM),
due to the increased proportion of leaf in the forage. The
CP content also decreases with herbage maturity (Ayres
et al., 1998; Fulkerson et al., 1998). It is interesting
though, that the CP content of legumes in this study was
just as high as grasses in most cases, despite the fact that
no N fertilizer was applied, but the NO3 content was
substantially lower in the legumes.
Oldham (1984) reviewed the results of many studies
on dairy cattle nutrition, and found that, over the range
of 100 to 180 g CP/kg DM in the diet, significant
positive responses in forage intake, total diet digestibil-
ity, and hence milk production could be observed. The
effects varied in size but increases of 0.2 to 0.4 kg DM
intake per 1% (10 g/kg) increase in dietary CP were
common, ranging from 0.1 to 0.8, depending on whether
the basal forage was grass or maize silage. Alderman
(1987) suggested that 160 to 170 g CP/kg DM is
required to meet rumen microbial N needs. All pasture
species investigated in the present study, including the
legumes, were well above these requirements and
provided excess amounts of rumen degradable protein.
For every MJ of FME it is possible to produce 9 to
11 g of microbial crude protein (MCP) (AFRC, 1993).
Effective rumen degradable protein would be expected
to be converted completely to microbial protein, pro-
vided FME is non-limiting; therefore 9 to 11 g of ERDP/
MJ of FME would be required to adequately feed the
microbes in the rumen. In the present study, MCP
supply is calculated from the limiting factor, either
ERDP or FME. Thus, when the ERDP: FME ratio was
higher than 11, MCP was based on FME. Conversely,
when this ratio was lower than 11, the MCP was based
on ERDP. For all grasses and legumes, the concentration
of ERDP was greater (relative to available FME), and
the ratio of ERDP: FME varied from 14 to 31, well
above the ideal ratio of 11 required for MPS (AFRC,
1993) for dairy cows. This indicates that there was more
protein coming from forages available in rumen than the
microbes could convert to MCP. Any excess protein
must be converted to urea and excreted in the urine. The
effect of this, on feed efficiency, is a matter of some
conjecture, although, in theory, there is an energy cost of
converting toxic ammonia to urea and its excretion in
urine. However, there is anecdotal evidence that cows
on a largely grazed pasture diet can adapt to very high
(above requirements) levels of protein intake.
The requirements for maintenance of a typical 600 kg
cow producing 30 kg milk with 4.04% milk fat and
3.28% milk protein are 1697 g MP and 220 MJ/cow/day
(AFRC, 1993). To meet this requirement, the cow would
have to eat 19.1 kg DM/cow/day of a feed containing
89 g MP/kg DM and 11.5 MJME/kgDM. The present
study has shown that all forage species would provide
sufficient MP (95–173 g MP/kgDM). However, energy
is the primary limitation for milk production in a
predominately pasture-based system, even if top quality
pastures can be provided, and van Vuuren (1993)
suggests a maximum milk yield of 27 L/cow/day based
on energy as the limitation. In the present study, the ME
value of all grasses and clovers were below 11.5 MJ/kg
DM, therefore ME density of pastures would be the
 W.J. Fulkerson et al. / Livestock Science 107 (2007) 253–264
primary limitation to milk production of dairy cows on
pasture. This is supported by the findings of Miejs
(1981) who found that even top quality ryegrass-white
clover pasture was deficient in energy for high yielding
dairy cows (N30 L milk/cow/day). The high content of
MP and UDP in various forage species may rule out
the need to supplement with expensive bypass protein
sources in the ration of relatively high yielding dairy
cows. The species with the lowest UDP content was
fescue in summer and paspalum in autumn. This was
probably because both species were actively growing at
that time of year, whilst other species were not.
Amongst the C3 grasses, prairie grass, perennial
ryegrass and SR ryegrass were found to be high in
digestibility with a mean ME density of 10.6 MJ/kg DM
over the whole year. Kikuyu especially in summer, and
all legumes except Sulla in winter, were found to be
highly digestible with a ME density of N 10 MJ/kg DM.
Maximum herbage DMI of around 4.2% of live
weight has been reported for cows grazing pasture
(Cohen et al., 2000). Similar findings were reported by
Fulkerson et al. (2006) from their studies when dairy
cows were grazing either good quality kikuyu (27% CP,
73% DMD) or ryegrass (22.3% CP, 78% DMD)
pastures. In this respect, a 600 kg dairy cow should be
able to eat up to 25 kgDM/day and would meet the ME
requirement for more than 30 L milk/cows/day when
offered good quality pasture (SCA, 1990).
The digestion of the fibre (cell wall) component of
forages can be suppressed due to adverse effect of low
rumen pH on cellulolytic bacterial populations in the
rumen, when high amounts of grain are fed (Huhtanen,
1991). Wedekind et al. (1986) found that rumen micro-
organisms that favor hemicelluloses as a substrate may
be more sensitive to the negative affect of low pH on
digestion than those that prefer the cellulose component
of the diet. This demonstrates that the feeding of grain-
based supplements to dairy cows is likely to have a
greater deleterious effect on digestibility of the forage
component of the ration when the content of hemicel-
lulose in pasture species grazed is high. In this regard,
the grasses investigated in our study had much higher
hemicellulose content than the legumes and therefore
grain supplementation could be expected to have a more
negative effect on the digestibility of grasses.
5. Conclusions
The results of this study highlight that under optimal
conditions of growth, the availability of MP and ME
from the grasses and clovers examined were able to
meet requirements for even relatively high producing
263
dairy cows (N 30 L milk/cow/day), provided animals
could consume sufficient DM intake to achieve this
level of production.
The ERDP to FME ratios in the rumen of the grasses
and legumes studied, were well above those required for
microbial growth and would have resulted in wastage of
protein. Prairie grass had the highest MP value of any
grass species in summer and winter. The ME density of
perennial ryegrass, prairie grass and SR ryegrasses
were similar (10.6 MJ/kg DM) and slightly higher than
cocksfoot, phalaris and fescue with a mean value of
10 MJ/kg DM, over the whole year.
A logical extension of this work would be to examine
the voluntary intake of dairy cows grazing these forage
species (varying in digestibility), when grazed at the
optimum stage of growth, in order to determine the
potential ME intake of the modern dairy cows under
these conditions.
Acknowledgement
We are grateful to Mr. Robin Dobos for the technical
advice on the interpretation of degradability data. We
would also like to thank Dairy Australia for financial
support.
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