Journal of Agricultural Science, Cambridge (1991), 116, 145-157.

Printed in Great Britain 145

Prediction of intake and digestion in ruminants by a

model of rumen kinetics integrating animal size and
plant characteristics
A. W. ILLIUS 1 AND I. J. GORDON 2
1
Division of Biological Sciences, University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, UK
2
Macaulay Land Use Research Institute, Pentlandfield, Roslin, Midlothian EH25 9RF, UK
(Revised MS received 2 My 1990)

SUMMARY
Simulation modelling was used to investigate interactions between forage degradation characteristics,
rumen processes and body weight, and to predict the voluntary food intake and digestion of a range
of forages. Predicted voluntary intake and digestion agreed well with empirical data, explaining 61
and 70%, respectively, of variance in observed values. Since the data covered a wide range of animal
weights and forage qualities, these results suggest that the model is a useful means of integrating the
effects of animal and forage variables. Interactions were examined between animal weight and diet
quality, as defined by the proportion of potentially digestible cell contents and cell walls and their
rates of digestion. Retention time of food in the digestive tract was shown by regression to scale with
W 027 . The time taken to comminute large fibre particles also scaled with W 027 . Longer retention of
digesta by large ruminants increases digestive efficiency compared with small animals and would
permit them to survive on lower-quality foods. The model showed that maximum intake of
metabolizable energy scales with c. W 0 8 ', greater than the scaling of maintenance with W 073 .

Fermentation characteristics of forages determined in

INTRODUCTION
The principal functions of simulation modelling are
to predict the behaviour and dynamics of complex
systems, and to reveal where knowledge is lacking.
Applied to comparative studies of nutritional physi-
ology, a modelling approach has an additional benefit
of allowing the results of empirical studies of a few
species to be generalized to other species in the form
of quantitative predictions. The strength of the
comparative approach is that the underlying mechan-
isms can be used to establish general relationships
across species, and this may throw into relief the
deviations (assumed to be specific adaptations) of
individual species from interspecific trends. In order
to investigate these interspecific trends in the nu-
tritional physiology of ruminants, a model of the
relationships between ruminant body size, food
characteristics, voluntary food intake and digestion
was developed.
Thefirstaim was to develop a dynamic mechanistic
model to simulate rumen processes and the fer-
mentation characteristics of forages in order to predict
voluntary intake and the extent of forage digestion.
situ have been shown to be useful predictors of
voluntary intake in ruminants. For example, 0rskov
et al. (1988) obtained regression equations relating
the potential digestibility and cell wall digestion rate
of forages to independently estimated intake of cereal
straws by cattle, explaining 79% of variation in
intake. Such empirical relationships apply strictly
only to these straws, and, given the dynamic nature of
rumen processes, may not apply to other forages. To
be able to generalize across all forages and animal
types, a model of digestion kinetics and digestaflowis
required, combining an adequate description of forage
and animal characteristics such as body weight and
digesta load.
The second aim of this paper was to use the
simulation model to test the hypothesis that body
weight influences the ability of an animal to obtain its
energy requirements from forages of low digestibility.
Support for this hypothesis was advanced by Van
Soest (1982) and Demment & Van Soest (1985), who
argued that, since metabolic needs scale with weight
(W) as W075 and gut contents scale with W1, large
animals should have a superior capacity to process
6-2
146 A. W. ILLIUS AND I. J. GORDON

Process Component

Intake:
Lag phase LIQCCo CELCQ) LDCW, SDCW, LINDFo S1NDF,,

Rumen: L1QCC, CELCC, LDCW, SDCW | L1NDF S1NDF,

Breakdown k4
Escape
Passage
Digestion

Digested
DM

MICR,

Small
intestine:
Digestion
Digested
DM

Large CELCO LDCW, SDCW^ SINDF-,

intestine:
Passage
Digestion

Digested
DM

MICR^
Excretion Undigested residues

Fig. 1. Flow diagram of forage digestion in ruminants (see Table 1 for definition of symbols). Compartments in the lag phase,
rumen and large intestine are distinguished by suffix 0, 1 and 2, respectively.

foods relative to their requirements, and so should be
able to tolerate poorer-quality forages than small-
bodied animals.
MODEL DESCRIPTION
The objectives were (a) to develop a mechanistic
model of food intake, whereby the quantity of food
eaten balances the loss from the rumen of food and
microbial dry matter (DM) through digestion and
passage; (b) to determine, from digestion in the
rumen and large intestine, the energy supplied by the
food; and (c) to extend the model to animals of
different body weight using allometry. Components
of the digesta, such as the cell contents, particles of
cell wall and microbial matter, are depicted as a
number of compartments, with unidirectional flow
between them, in the rumen and large intestines (Fig.
1). The modelling of fibre kinetics is similar to the
approach adopted by Mertens (e.g. Mertens 1977;
Allen & Mertens 1988). The rumen is taken to be the
site of control of physical intake. When, because of
digestion and passage, the total rumen digesta load
falls below a threshold, further ingestion takes place
to refill the rumen and maintain a specified average
fill. The model determines stable solutions to the
combination of meal size, number of meals/day and
the semisteady-state digesta composition by iteration
over 20 days. Copies of the program may be obtained
from A. W. Illius or I. J. Gordon.
Quantifying rumen clearance of digesta
Forages consist of cell contents (CC) which are
wholly or largely digestible (Van Soest 1982), di-
gestible cell wall (DCW) and an indigestible residue
(INDF), most of which is cellulose and hemicellulose
encrusted with lignin. The digestible and indigestible
 Prediction of intake and digestion in ruminants 147

cell wall fractions are each treated as pools containing Table 1. Definition of symbols used in the model of
large and small particles (the latter defined as those rumen kinetics
capable of passing through a 1 mm screen). This is
regarded as a reasonable assumption for modelling State variables: expressed in dry matter (DM), proportion
purposes (Ulyatt et al. 1986) and was adopted in the of standard* rumen DM load
model of Poppi et al. (1981 c). Food ingested as large
particles is selectively retained and broken down to LIQCC Cell contents released into the liquid phase
small particles at fractional rate kr This is a CELCC Cell contents trapped in cell wall
prerequisite of passage for all but a very small LDCW Large particles of digestible cell wall
percentage of the cell wall fraction, which escapes SDCW Small particles of digestible cell wall
LINDF Large particles of indigestible cell wall
from the rumen at rate kb. Mastication of the food SINDF Small particles of indigestible cell wall
during eating releases some of the cell contents and MICR Rumen and large intestine microbial DM
breaks down some of the fibre to small particles,
leaving a proportion, p, of the cell wall fraction to Rate variables: fractional rate/h
enter the rumen as large particles. It is assumed that k0 Liquid passage rate
the release of cell contents into the liquid phase ky Digestion rate of cell contents in rumen
(LIQCC) occurs only from small particles during ks Digestion rate of cell contents in large intestine
eating, or subsequently during breakdown at rate kv k2 Digestion rate of cell wall in rumen
and that the remaining fraction of cell contents k7 Digestion rate of cell wall in large intestine
(CELCC) trapped within the cell wall of large fibre k3 Passage rate of small particles in rumen
particles pending particle breakdown must therefore ks Passage rate of small particles in large intestine
be subject to the escape rate, /c5, of large particles. ki Breakdown rate of large particles to small
k& Escape rate from the rumen of large particles
Expressing food constituents ingested as proportions
of food DM, and using symbols defined in Table 1: Additional variables

CELCC = p.CC, LIQCC = (1 -p).CC, p Proportion of DM entering the rumen as large

LDCW = p.DCW, SDCW = (1 -p).DCW,
LINDF = />.INDF, SINDF = (1 -/O.INDF.
(1)
After ingestion there is a time lag, L, before
digestion commences, during which wetting and
microbial attachment take place. It was assumed that
food components are not subject to passage during
the lag phase. The quantity of DM in each of the
compartments representing the lag phase (denoted as
LIQCC0 etc.) is determined by the composition of the
food and the size of each meal. In common with the
size of the model compartments, meal size, y, is
expressed as a proportion of the mean daytime rumen
DM load, which is defined as the standard DM load.
Then at time, t, after ingestion:
Lag phase DM = .y.LIQCC0+.y.CELCC0
+.y.SINDF0, (2)
where y = meal size when 0 < t < L and y = 0 when
t> L.
After L has elapsed (i.e. at t = L+\), the food
components leave the lag phase and are added to the
active compartments (LIQCC, etc). Therefore, at any
time:
Total rumen DM =
Lag phase DM + LIQCC,
+CELCC, + LDCW, + SDCW,
+ LINDF, + SINDF,+ MICR,. (3)
The clearance of food components from the rumen
particles
y Meal size, expressed as proportion of standard*
rumen DM load
* See text and Eqn 18 for details.
is a function of the rates of digestion (A:,,/c2), passage
(ko,k3), breakdown (ks) and escape (A;5). The flow of
material from and between compartments is con-
tinuous and simultaneous, and can be represented by
a series of linear differential equations:
d(CELCC,)/d<
= -A:,.CELCC,-/fc 4 .CELCC,-/c 5 .CELCC,
d(LIQCC,)/d/
= -kv LIQCC, -fe o .LIQCC, + A:4.CELCC,
d(LDCW,)/dr
= -fc 2 .LDCW 1 -/c 4 .LDCW 1 -)t 6 .LDCW 1
d(SDCW,)/d/
d(LINDF,)/dr = -A^.LINDF^^.LINDF,
d(SINDF,)/df = -A:3.SINDF, + A:4. LINDF,. (4)
The rate of DM digestion in the rumen,
d(RDDM)/d/, is:
d(RDDM)/d? = ^.LIQCQ + A^CELCC,
C tSDCW (5)
Note that the fraction of LIQCC leaving the rumen
at the liquid passage rate, k0, is assumed to be totally
digested in the small intestine (Fig. 1) since only trace
amounts of cell solubles reach the large intestine
(Ulyatt et al. 1975).
148 A. W. ILLIUS AND I. J. GORDON
The DM load of the rumen is composed of food
constituents and microbial biomass (MICRj). Quanti-
fying the feed intake that replaces rumen dry matter
digested or passing from the rumen therefore depends
on quantification of the total rumen DM space
occupied by microbial DM undergoing growth and
passage. In each time step, the microbial growth rate
was calculated as 0-26 kg microbial OM/kg fermented
OM (Mathers & Miller 1981), microbial DM con-
taining 8 5 % OM (Czerkawski 1976). Ulyatt et al.
(1984) found that microbial outflow to the duodenum
occurred at the same fractional outflow rate as that of
fibre. John (1984) estimated that this only accounted
for c. 0 6 of the decrease in size of the rumen microbial
pool, and assumed that the remaining decrease was
due to microbial death and lysis. Therefore, the
modelled outflow from the microbial pool assumes
that 0-6 of microbial biomass flows with fibre particles
at rate k3, and that the remainder is lysed and flows
with the liquid phase at k0. Rumen microbial DM
turnover is thus:
d(MICR,)/dr = 0-3 OMC.d(RDDM)/dr
,) (6)
where OMC is the organic matter content of the
digested DM.
Numerical methods and determination of meal size
and frequency
The equations representing compartmental flow (Eqns
4 to 6) were solved using Euler's method of numerical
integration. The time step length, dt, was chosen to be
01 h, after examination of steps in the range 001-1 h,
as an interval which minimized errors at acceptable
computing speed.
Due to the differential clearance of digesta constitu-
ents, arising from differences in their rates of digestion
plus passage, the ratio of these constituents in the
rumen changes over time. This leads to a build up of
the slow-digesting and indigestible fraction in the
rumen. With continued intermittent ingestion of the
food, a semisteady state in the contents of the rumen
is reached where clearance of food constituents from
the rumen is balanced by each meal. The model
determines this state by hourly and daily iterations for
5 days with 3 meals/day. Over the remainder of the
20-day iteration, numbers of meals/day increase by 1
meal/day from 3 to a maximum of 8 if the rumen DM
load at the start of the day is < 073 of the average
daytime load, GC, defined below (Eqn 18). Meal
intervals are thus determined from numbers of
meals/day, and meal size is adjusted during iteration
to that required to refill the rumen to the extent that
maintains the mean daytime digesta load at the
content specified. This method of determining the
combination of number of meals and size gave stable
solutions, and an appropriate balance between these
variables by avoiding, for example, very large and
infrequent meals.
Since meal size is defined in terms of the fraction of
the mean daytime.^or standard DM load, the sum of
the day's meals is the daily fractional turnover time,
RT, and daily DM intake (DMI, kg/day) is given by:
DMI = GC/RT, (7)
where GC is the standard rumen contents of DM.
Contribution of digestion in the large intestine to
total true and apparent digestion
It is known that further microbial digestion occurs
outwith the rumen, chiefly in the caecum and colon
(Ulyatt et al. 1975). This was accounted for in the
model by additional hindgut pools of the food
components, with inflow determined by passage from
the rumen. Compartment turnover by digestion (ke
and &,) and passage (&8) are described as in Eqns (4)
and (5), except that particle breakdown is not
considered because it is virtually completed before the
large intestine (Ulyatt et al. 1986). Large and small
particles are assumed to be retained to the same
extent, and thus their flow is described by a single
passage rate. Total dry matter digestibility is thus the
sum of the digestion in the rumen, the small intestine
and the large intestine as a fraction of DMI. To relate
true digestion predicted by the model to apparent
digestion in vivo requires an estimate of metabolic
faecal output. This was determined by Uden & Van
Soest (1982) to be 10% of DM intake over a wide
range of intakes, animal weights and species.
Food requirements for maintenance
The model of digesta turnover incorporates a de-
scription of energy needs to equate with the energy
yield from digestion. The daily net energy require-
ments (NE) for maintenance are taken to scale with
W 073 (Brody 1945), allowing for the costs of activity:
NE R E Q = 0-4 W ° " (MJ/day). (8)
The NE concentration of a food, NEC, for
maintenance is determined by the proportion ap-
parently digested, D t , the organic matter content,
OMC, and the efficiency of utilization of metaboliz-
able energy (ME) for maintenance, km, (Agricultural
Research Council 1980). The ME content (MJ/kg
DM) is:
M E = 15-6D t .OMC (9)
NEC = ME.fcm, (10)
where: km = 0-503 + 0019 ME. (11)
To this is added the contribution from enzymic
digestion of cell contents by assuming that 66 % of
cell contents provides an energy substrate which is
 Prediction of intake and digestion in ruminants
converted to glucose with an energy content of
15-6 MJ/kg, used for maintenance with an efficiency
close to 100% (Blaxter 1971).
Estimation of parameter values and extension of the
model to animals of different body mass
The description of how parameter values were
estimated follows roughly the order in which they
were introduced. Some of the parameters (k3,kt,k0,
GC) are related to body weight (W), and it is by these
that animal size effects are modelled. Accordingly, W
is a driving variable, as are the food characteristics
CC, DCW, INDF and k2.
The proportion of large particles entering the
rumen, p, was estimated to be 0-66 (c.v. 17 %, n = 20)
from the data presented by Van Soest (1982); Lee &
Pearce (1984); Pond et al. (1984); Ulyatt et al. (1986)
and Chai et al. (1988). Poppi et al. (1981c) found
virtually the same: 0-65-(c.v. 14-2, n = 16). Exam-
ination of the data upheld the conclusion of Lee &
Pearce (1984) that the variables controlling com-
minution during eating have not been established.
The data cover a wide range of dry and fresh forages,
and Ulyatt et al. (1986) regard some of the variation
between reports as due to methodological differences.
Given that, the degree of variation is not great, and
this perhaps suggests some uniformity in the process
of bolus formation.
The factors affecting the duration of the lag phase
(L) for fibrous feeds have not been well quantified,
especially for food which has been masticated.
Mastication reduced lag times from 15-5 to 3-1 h
(Poppi et al. 1981 b). The lag times of three masticated
forages were estimated using the method of Dhanoa
(1988) from the data of Playne et al. (1978). From the
relationship of cell wall L (h) to CC content, the
following approximation was used in the model for all
but LIQCC, for which zero lag was assumed:
L=\ CC 5= 0-5, L = 2 0-5 > CC =s 0-3,
L = 3 0-3 > C C S s 0-25, L= 4 0-25 > CC ^ 0-2,
L = 5 0-2 > 0-15, L = 6 015 > CC.
(12)
Published estimates of the digestion rate of cell
contents (/:,) are scarce, and do not distinguish
between digestion rates applying to LIQCC and
CELCC. Therefore, kt was assumed to be 0-3
throughout (Van Soest 1982).
Digestion rate of cell wall (fc2) may possibly differ
between particle sizes. However, while in principle
there may be a range of digestion rates associated
with the size spectrum of particles (see Mahlooji et al.
1984), in practice no consistent differences have been
associated in situ with fineness of grind, provided
disruption of the epidermis is adequate for microbial
access (Ford et al. 1987; Kyle & Hovell 1987).
149
Therefore, the conservative assumption was adopted
that digestion rates of large and small particles are
equal. Regarding digestion rates in the large intestine,
Ulyatt et al. (1975) noted that the cellulolytic activity
of caecal digesta was as high as that of rumen digesta.
Therefore the caecal cell wall digestion rate, &,, was
taken to be the same as the rumen digestion rate, kv
Passage rate, k3, is the inverse of retention time,
which was assumed to scale with W 027 , in common
with the scaling of the duration of other time-related
physiological variables (Taylor 1980; Peters 1983).
Empirical support for this was obtained by allometric
regression of mean retention times on body weight
using the data of Warner (1981) and Foose (1982).
This gave total mean retention time for ruminants
(MRT, h ) a s :
log e (MRT) = 2-725(± 0-147)
+ 0-251( ±0-028) loge(W) (13)
2
(r = 0-76, n = 26, R.S.D. 0197).
The range in weights was from 3-7 kg (suni,
Nesotragus moschatus) to 1000 kg (water buffalo,
Bubalis bubalis). Since the exponent is not significantly
different from 0-27, the relationship was recalculated
as 14-1 W 0 2 ', and hence passage rate from the whole
gut is 0071 W-° 27 . Warner (1981; Table 4) and Uden
et al. (1982) presented data for a wide range of species
relating rumen MRT to total MRT in the whole gut.
In hay-fed animals, the particle MRT in the rumen,
expressed as a proportion of the total retention time,
was 0-74 (c.v. 15-5%). This was used as a first
approximation to estimate rumen passage rate, k3, as
the inverse of 7 5 % of the total MRT, which was
obtained from the allometric relationship described
above. Similarly, the large intestine passage rate, ks,
was estimated as the inverse of 20% of total gut MRT
(Warner 1981).
The rate of breakdown of large particles to small
particles, k4, was estimated from two sources, each in
two animal species and a range of forages. Data
presented by Poppi et al. (1981c) on large-particle
retention time in cattle and sheep was adjusted to
account for losses from the large-particle pool by
digestion. The data of Spalinger et al. (1986) on
breakdown of large- to medium-sized particles in
mule deer (Odocoileus hemionus) and elk (Cervus
elaphus) were adjusted to equate with particles of
> 1 mm (D. E. Spalinger, unpublished). Then re-
gression of large-particle retention time (RL, h) on the
proportion of INDF and W after logarithmic trans-
formation gave:
logc(RL) = 2-12(±0-34) + 0-46(±0065)log e (INDF)
+ 0-235(± 0-064) loge(W) (14)
(r2 = 0-616, n = 40, R.S.D. 0-377).
Since the allometric coefficient of weight is not
150 A. W. ILLIUS AND I. J. GORDON

significantly different from 0-27, the equation was their supply is adequate, or that inadequacies are
recalculated to give: reflected in lower measured rates of digestion, and are
144 27 therefore accounted for. Nor does the model describe
kt = 0144 INDF-° .W-° . (15)
how metabolic regulation of nutrient intake may limit
It is reasonable to suppose that the escape rate of intake of high-quality forages according to appetite
large particles, k5, is proportional to the small-particle or physiological state (see Forbes 1980), nor how
passage rate, k3, and thus kb was estimated to be c. digesta load may respond to those factors.
12% of k3. This was based on the mean (11-85, c.v. The need to apply the model to a wide range of
27-4%, n = 4) of values obtained from data presented foods restricts the degree of detail with which food
by Van Soest (1982); Egan & Doyle (1984); Ulyatt characteristics are specified to the variables which are
et al. (1986) and Chai et al. (1988). fairly readily accessible. The model has been de-
Liquid passage rate (k0) was estimated by regression veloped and tested on gramineous forages, but there
in a manner similar to that of Faichney et al. (1981), is no reason to suppose that the same principles
who argued that the fractional outflow rate of water governing digesta flow do not apply to other classes of
should be a function of dietary factors which tend to forage, such as legumes or woody browse plants. It is
increase rumen osmolality. Data on liquid passage assumed that compartments are physically distinct
rates in animals fed a range of forages were collated and that kinetic processes are first-order. A wide
from Weston & Hogan (1968), Hogan & Weston range of ruminant species and body weights was used
(1969), Egan et al. (1975), Uden et al. (1982) and to develop the expressions for k3, k4, k0 and GC,
Renecker & Hudson (1990), covering five ruminant permitting generalizations across that range of body
species over the weight range: 24-716 kg body mass. sizes. Predictions for a single species will be valid to
Terms for CC, DCW and W accounted for 45-2 % of the extent that they conform to the allometric scaling
variation in k0: employed. Notwithstanding the view that specific
adaptations unrelated to weight (e.g. stomach anat-
A:o = -0-0487(± 0-022) omy) distinguish the digestive physiology of the
+ 0-176( ± 0029) CC + 0-145( ± 0033) DCW various ruminant species (reviewed by Kay et al.
+ 00000231 (±0000011)W (16) 1980), we found no significant differences between
grazers, intermediate feeders and browsers in passage
(r> = 0-45, n = 44, R.S.D. 00158).
rate (k3) or in rumen DM load after weight had been
For comparison, inclusion of terms for crude accounted for. This is contrary to the differences
protein increased the explanation of variance to 54 %, assumed to exist in rumen load and passage rate
as in the model of Faichney et al. (1980), who based on liquid volume (or wet weight) and observed
estimated that individual differences between animals feeding preferences.
account for a substantial part of the residual variance.
The scaling of the weight of dry matter in the
rumen (GC, kg) with body weight was determined MODEL TESTING AND VALIDATION
from data on eighteen species ranging from 3-7 to
The model was tested by examining the effect of
720 kg W (I. J. Gordon & A. W. Illius, unpublished):
forage characteristics on predicted intake and di-
loge(GC) = - 4-37( ± 0-186) + 1 • 10( + 0-039) loge(W) gestion in modelled ruminant animals of different
(17) weight. Comparisons of predictions with published
2 observations were made by regression. Details of 23
(r = 0-98, n = 18, R.S.D. 0105).
gramineous forages were collated to test the model,
This was not significantly different from an allo-
from data presented by Smith et al. (1972), Poppi
metric coefficient of 1, and the regression was
et al. (1981a), Hovell et al. (1986) and 0rskov et al.
recalculated as:
(1988). The forages covered a wide range of chemical
GC = 0021 W. (18) composition and of cell wall digestion rates. These
include the temperate species cocksfoot (Dactylis
glomerata), perennial ryegrass (Lolium perenne), reed
Summary of assumptions and limitations to use of
canarygrass (Phalaris arundinaced), smooth brome
the model
(Bromus racemosus), barley (Hordeum vulgare) and
Emphasis is placed on modelling the limiting con- wheat (Triticum aestivum) and the tropical species
dition of maximum intake to determine how forage pangola (Digitaria decumbens) and Rhodes grass
characteristics restrict nutrient intake from abundant (Chloris gayana). Values for ash, cell contents,
foods. The case of survival on depleted resources has digestible and indigestible cell wall, and cell wall
been considered elsewhere (Illius & Gordon 1987). digestion rate (k2) are presented in Table 2.
Nitrogen and mineral contents of foods may also Mature sheep and cattle of 45 and 485 kg average
impose limits on digestion and survival, but have not weight, respectively, fed ad libitum at hourly intervals
been included in the model. It is assumed that either were used in the study of Poppi et al. (1981 a). Hovell
 Prediction of intake and digestion in ruminants 151
Table 2. Composition of foods used in the model of rumen kinetics

Cell Digestible Indigestible Cell wall

Food Plant species Growth contents cell wall cell wall digestion Ash
number or component stage (g/kg) (g/kg) (g/kg) rate (k2) (g/kg) Reference

1 Dactylis glomerata Vegetative 640 317 43 0128 100*'

2 Preflowering 380 459 161 0-077 100*
3 Mature 350 383 267 0046 100*
Smith et at. (1972)
4 Phalaris Vegetative 620 342 38 0183 100*
5 anmdinacea Mature 330 370 300 0053 100*
6 Bromus racemosus Mature 320 340 340 0073 100*;
Digitara
decumbens
7 Leaf 6 weeks 343 468 189 00219 112 \
8 Stem 6 weeks 306 440 254 00195 103
9 Leaf 12 weeks 378 340 282 00219 101
10 Stem 12 weeks 319 376 305 0022 90
Chloris gayana Poppi elal. (1981a)
11 Leaf 6 weeks 300 520 180 00236 115
12 Stem 6 weeks 279 478 243 0023 86
13 Leaf 12 weeks 270 449 281 00208 107
14 Stem 12 weeks 232 417 351 00203 71 ,
Barley
15 Straw Mature 125 264 611 0-0337 37
16 Straw- Mature 136 297 567 00391 37
17 Straw Mature 160 361 479 0-0483 43 0rskov et al. (1988)
18 Straw Mature 150 405 445 00303 48
Wheat
19 Straw Mature 193 293 514 0-0345 61
,
Lolium perenne
20 Hay Mature 338 424 238 0-0824 66 1
21 Hay Mature 264 400 336 0-1172 58 1 - Hovell et al. (1986)
22 Hay Mature 245 290 465 0-0747 69
23 Hay Mature 251 206 543 00685 96

* Assumed.

et al. (1986) used lambs of 40 kg. 0rskov et al. (1988)
used growing cattle of 360 kg to measure intake, and
sheep of 60 kg for digestibility.
A sensitivity analysis was performed by examining
the effect on predicted intake and digestion of varying
the value of parameters selected in turn. The results
were calculated as the percentage change expressed
for a unit percentage change in the parameter, termed
a sensitivity elasticity (Gilchrist 1984).
RESULTS
= 25, R.S.D. 8-49). The regression of best fit was:
observed intake = 0-98 (±0028) predicted intake,
with intercept not significantly different from zero
and thus omitted (Fig. 2). Digestibility appeared to be
underpredicted for low-quality forages (Fig. 3), but
was highly correlated with observed values. The
regression equation was: observed digestibility = 0-26
(±0-03)+ 0-57 ( ± 0 0 7 ) predicted digestibility (r2 =
0-698, n — 25, R.S.D. 0039). These discrepancies are
thought to arise from a number of sources outwith the
model (see Discussion).

Assessment of model predictions

The intakes and digestibilities predicted by the model
showed good agreement with observed intakes and
digestibilities of foods fed to cattle and sheep by
Poppi et al. (1981a), Hovell et al. (1986) and
0rskov et al. (1988) (Table 3). For intake, which was
expressed as g D M / k g W 0 7 3 , predicted values were
highly correlated with observed values (r2 = 0-606, n
Sensitivity analysis
The only parameters to which the model showed
appreciable sensitivity were the digesta DM load
(one-to-one effect on intake, none on digestibility),
the content of DCW, and passage rate, k3 (Table 4).
The R.S.D. of the equation used to predict passage rate
was 18-25%, and so a change of 1 S.D. in k3 would
 152 A. W. ILLIUS AND I. J. GORDON

Table 3. Mean observed and predicted intake and digestibility in cattle and sheep

Intake
(g DM/kg W 073 ) Digestibility

Observed Predicted Observed Predicted Reference

Cattle 70-5 72-2 0-54 0-51 Poppi et al. (1981a)

Sheep 49-2 57-0 0-51 0-44 Poppi et al. (1981a)
Cattle 52-3 480 0rskov et al. (1988)
Sheep 0-42 0-28 0rskov et al. (1988)
Sheep 61-9 55-8 0-53 0-46 Hovell et al. (1986)
All 58-7 59-9 0-50 0-44

80- Table 4. Sensitivity elasticity* of predicted intake and

digestion, or percentage change, of model parameter
values
pT 70-
o
Sensitivity elasticity
00

§ 60" Parameter Intake Digestibility

Lag time -0095 0

° 50-| + 0-025 + 0023
K + 0086 + 0-055
ki 0 + 0-037
6 40- k3 + 0-632 -0055
k> + 0010 0
*s + 0014 0
30-1 + 0089 0
30 40 50 60 70 80 DCWt + 0-489 + 0-606
0 73 Proportion of long particles + 0-037
Predicted DMI (g/kg W ' ) -0129
Microbial growth rate -0129 0
Fig. 2. Observed and predicted dry matter intake (DMI) in Proportion of microbes -001 0
cattle ( # ) and sheep ( • ) , with regression (—) and line passing with solid phase
representing y = x ( ). Gut DM contents +1 0
Percentage change
Allow passage during lag + 2-4% -5-1%
0-7- Reduce k2 of long particles to - 2 - 8 7 % -1-3%
half that of short particles
Change proportion of mean + 10% + 0-9%
0-6- retention time in rumen and
hindgut to 0-65 and 0-3,
respectively (cf. 0-75 and 0-2,
0-5- respectively)
-a
* Percentage change in output from 1 % increase in
a 0-4- parameter value; food 6 (Bromus racemosus), for an animal
of body weight 240 kg.
t Increase in the amount of the cell wall constituents
6 0-3- which are digestible; corresponding decrease in indigestible
cell wall.

0-2-1
r
0-2 0-3 0-4 0-5 0-6 0-7
Predicted digestibility
Fig. 3. Observed and predicted apparent DM digestion in
cattle ( # ) and sheep (H), with regression (—) and line
representing y = x ( ).
 Prediction of intake and digestion in ruminants 153

maintenance, and of animals of 360 kg from 21 to 1-2

2-5-
times maintenance.
The shape of the response curve relating intake to
weight is influenced by the content in a food of
indigestible cell wall and the cell wall digestion rate.
The superiority of large animals over small is greater
2-0-
on a food with low indigestible residues and slow cell
wall digestion (e.g. food 7) than on a food with large
amounts of indigestible components and high rates of
cell wall digestion (e.g. food 5). This is because it is
1-5-
only advantageous to retain particles for longer, as
large animals do, if they are digestible. Small animals,
on the other hand, can utilize the rapidly fermenting
cell wall without long retention of the indigestible
component.
The effects of these different foods on intake and
i 1-0- digestion can be summarized by examining how these
animal functions scale with weight for the various
foods, bearing in mind that only the modelled physical
limits are considered. The scaling of DM intake
0-5- ranges from W 093 to W 081 on foods differing in
content of indigestible cell wall from 38 to 611 g/kg
(data for each food plotted in Fig. 5 a). The theoretical
15 relationship between indigestible content and the
scaling of DMI with W is also shown on this graph.
This was derived by running the model with no lag
rr—i 1— 1 —1—
960 phase. Without this constraint, highly digestible foods
30 60 120 240 480
Body size (kg)
could be eaten in nearly direct proportion to weight,
their clearance from the rumen being primarily a
Fig. 4. Modelled energy intake, expressed as multiples of function of the digestion rate (see Discussion). Fixed
maintenance requirements, in ruminants ranging from 30 to lag times impose differential constraints according to
960 kg body weight. The numbers refer to forages listed in animal size, resulting in the deviation of the plotted
Table 2.
data points from the theoretical line obtained under
simplified conditions. Figure 5 b shows how digestion
change intake by 11-5% and digestibility by - 1 % . scales with weight in relation to cell wall digestion
Similarly, an increase of 1 S.D. in rumen DM load rate. Large animals have a greater advantage when
would increase intake by 24-3 % without changing cell wall digestion rates are low (D cc W 006 ) because
digestibility. of their longer retention time and hence more extensive
digestion. Conversely, the shorter retention times of
Effect of animal size on intake and digestion small animals allow a lower extent of digestion of
Selected results from the foods in Table 2 are shown slowly fermenting forages.
in Fig. 4, which relates animal weight to energy intake The proportion of cell wall digested in the large
in terms of the multiples of maintenance provided by intestine was 7-15 % for good and poor foods with W
a food at maximum intake. Foods 1 and 4 are not = 30, and 3-12% with W = 960. Ulyatt et al. (1976,
shown, because they are predicted to provide > 3 fig. 7) reported a similar effect of food quality. In
times maintenance at maximum intake, and thus small animals, the more rapid passage of partially
consumption would be unlikely to be subject to digested residues from the rumen to the hindgut is
physical limits. There is a marked effect of body predicted to support more extensive digestion in the
weight on energy intake: increases in body weight distal site.
from 30 to 960 kg enable energy intake relative to
maintenance to increase by up to 1-7 times. Similar DISCUSSION
increases in energy intake occur on high-quality and Adequacy of the model and predictive ability
poor-quality foods (e.g. food 2 cf. 14).
The effect of increasing forage maturity on food The first objective of modelling rumen kinetics was to
intake and digestion is demonstrated by comparison evaluate the prediction of intake and digestion of
of D. glomerata at three stages of growth (foods 1, 2, forages from fermentation characteristics determined
3), increasing maturity (foods 2 and 3) depressing the in situ. Intake can be predicted by the model with an
intakes of animals of 30 kg from 1-35 to 0-8 times R.S.D. of 14-5% of the mean intake observed, which is
 154 A. W. ILLIUS AND I. J. GORDON
(a)
0-6- 1 •
0-5- \ •
:cel
X) 04- \ I 0-16-1
•3
\ \.\»
v:
of in
0-3-
co
rti
8.
o
0-2-
01-
0 0
0-75 0-8 0-85 0-9 0-95
b
Fig. 5. Relationship between (a) the indigestible content of forages and the allometric coefficient, b, relating forage intake
to weight in the expression dry matter intake = a W (data from forages in Table 2), and theoretical relationship (
text); and (b) cell wall digestion rate and the allometric coefficient, d, relating digestion to weight in the expression dry matter
digestion = c W .
surprisingly close given the generalized specification
of animal characteristics from allometric equations.
The equivalent R.S.D. for digestibility is 7 7 % . Both
are a considerable improvement over models where
animal characteristics have not been defined or have
been held constant (e.g. Mertens & Ely 1982).
Digestibility was underpredicted in foods with
observed apparently digestible DM content < c. 0-45,
and the model performed worst at predicting di-
gestibility on the low-quality foods used by 0rskov
et al. (1988), who used sheep to estimate digestibility;
low intake combined with the ability of sheep to select
the more digestible components (leaves) of the straw
may explain the high digestibilities observed (E. R.
0rskov, unpublished), as illustrated by the observed
apparent digestibility being close to, and in one case
higher than, the stated potential true digestibility. The
sensitivity analysis indicates that the cell wall digestion
rate and potentially digestible cell wall content are the
only parameters which, if varied, would effect the
required increase in predicted digestibility without
reducing intake. Therefore, one interpretation of the
model's bias is that in situ degradation characteristics
underestimate in vivo degradation. This is supported
by the finding that microbial attachment to fibrous
samples in situ can significantly increase the weight of
the sample (Mathers & Aitchison 1981). The resulting
underestimation of degradation rate constants and
extent of potential degradability would be more
(ft)
g 0-2 -\
i 014-
= 0-12-
^ 0-1-
| 0-08-
I 006
"
.V
1 0-04-
S 0-02-
10 0 0-01 002 003 0-04 0-05 006
; see
marked on low-quality forages. The data of Varvikko
& Lindberg (1985) and Negi et al. (1988) suggest that
7-15% of the residual DM (but not NDF) of fibrous
materials may be of microbial origin after 24-48 h of
in situ incubation. This suggests a source of error in
model predictions based on degradation character-
istics which do not distinguish N D F from DM, as in
the case of the data of 0rskov et al. (1988). A
potentially greater source of error in the data of
0rskov et al. (1988) arises from their not accounting
for and estimating the lag time and degradation
constant simultaneously. This can reduce estimated
degradation rates markedly (Dhanoa 1988), especially
if the first bags are removed from the rumen during
the lag phase. When 0rskov's published data were
analysed using Dhanoa's (1988) method, estimates of
degradation rates nearly doubled, and predicted mean
digestibility for the data set increased to 032.
The use of the data of Poppi et al. (1981 c) as part
of the data set for both deriving the breakdown rate,
k4, and testing the model is open to criticism. However,
predicted intake is in good agreement with the
independent data sets of Hovell et al. (1986) and
0rskov et al. (1988), and the model is not very
sensitive to varying kt ( 1 1 % higher intake for a
doubling of k4; Table 4), so bias is unlikely to have
occurred.
 Prediction of intake and digestion in ruminants 155

temperature. Hence, intake tends to a function of

Body size effects on intake and digestion rumen digesta pool size which scales as W1. Con-
Provided cautious use is made of data from low- versely, as food quality declines, passage tends to
quality forages, the performance of the model when predominate over digestion in clearance, and the
073
compared with empirical data suggests that it is a scaling of intake tends to W , or the ratio of gut
027
sufficient description of particle kinetics and digestion contents to passage rate (W'/W ). At low digestion
to satisfy the second objective of examining the effects rates the retention of fibre particles becomes more
of body size. This is clearly supported by the slope of significant, and thus larger animals have higher
1 relating predicted intake to that observed in animals digestive efficiency, with digestibility scaling at c.
006 073
of 40, 45, 360 and 485 kg live weight over a range of W , offsetting the scaling of intake tending to W .
foods. Moreover, any bias in the prediction of At high digestion rates, digestibility scaling tends to
digestibility is not related to weight. W° as weight effects disappear, and as intake scaling
1
The model clearly supports the argument that large tends to W . The scaling of energy intake with W is
animals can obtain a greater proportion of their the product of the scaling of DMI and digestion with
energy requirements from abundant poor-quality W. The tendency for an inverse correlation between
foods than small animals. The longer retention time these produced a scaling of potential ME intake
087
in large animals allows more extensive digestion and which showed little variation, with a mean of W .
it is this which permits higher DMI and higher energy Therefore the scaling of ME intake in relation to
087 073
intake relative to W° ™. The essential property of the maintenance is W " = W° 14 , equivalent to a
model is the description of retention time as scaling 13-4% difference in ME intake over maintenance for
with W 0 2 ', in the manner of other temporal variables, a doubling in weight. This results in selection pressure
such as the time between successive hearts beats or for the evolution of large body size under conditions
intestinal contractions (Clark 1927). The duration of of abundant food.
physiological events is simply longer in larger animals. Demment & Van Soest (1985) argue along similar
The time taken to comminute large particles obeys the lines, though without explicitly linking the processes
same rule, as demonstrated by the relationship of in a single model relating animal and food properties.
large-particle retention time and W (Eqn 14). It is Criticism of Demment (1983) and Demment & Van
assumed that reticular contraction rate is an important Soest (1985), as expressed by Taylor & Murray
determinant of the rate of passage from the rumen, (1987), arises from the method of derivation of
and our empirical analysis strongly supports the retention time, the definition of digestibility and their
expected scaling rule. Other factors may influence causal relationship. Demment (1983) and Demment
retention time, notably the rate of intake (Van Soest & Van Soest (1985) derived the theoretical scaling of
1982), but this is irrelevant to the conditions of retention time as varying with the actual digestibility
maximum intake addressed by the model. of a food in an animal of a given weight and with the
0 76
The scaling of DMI and digestion with weight ratio of gut contents to metabolic rate, W'/W or
025
shows that the complex interaction of food character- W . At first sight this appears to be circular and to
istics with animal mechanisms cannot be described ignore the different rates of digestion and passage of
adequately by an analytical approach, as was pro- each food component and the higher digestive
posed by Demment & Van Soest (1985). Figures 5a efficiency with increasing W. However, actual di-
and b show that the main effects are functions of the gestibility can be defined to account for weight effects
proportion of indigestible cell wall and digestion rate (as kj(ica + kp) where ka and kp are digestion and
of digestible cell wall in relation to rumen volume and passage rates, respectively) provided passage rate is a
passage rate. The shape of the theoretical relationship known function of W, and ideally is not derived from
between digestible content and the allometric co- the difference between exponents of animal require-
efficient by which DMI scales with W results from ments and gut contents. Our model employs passage
clearance from the rumen following two routes: the rate as an external driving variable which scales with
weight-related route of passage, and the weight- W by a general rule (Taylor 1980) and is specified
independent route of digestion. The relative im- empirically. Given this, predicted energy yields of
portance of these alternatives is a food variable. foods to animals differing in weight agree with the
Given a food of low indigestible content the scaling of prediction which can be obtained from the for-
potential DMI (assuming no metabolic regulation) mulation of Demment & Van Soest (1985). However,
tends to W1 (Fig. 5a). This is because clearance of the aggregated retention time of all components of a
00G
food from the rumen is dominated by the digestion single food scales from W to W 022 , depending on
rate, and the size-scaling effects on passage rate are the properties of the food, and therefore the premise
diminished. Digestion rate, determined by the rate of of Demment & Van Soest (1985), that potential intake
025
microbial fermentation, would not be expected to scales with W'/W = W 075 , does not hold.
differ much with animal size, given that microbes' The advantage of a simulation modelling approach
homeothermic hosts show little variation in body is that it can integrate the causal relationships and
156 A. W. ILLIUS AND I. J. GORDON

provide a description of the response to food variables
over a range of weights, taken to be the major variable
underlying animal differences (Taylor & Murray
1987). The disadvantage of a modelling approach are
that simplifying assumptions have to be made where
data are lacking. In the present model, these included
assumptions about the growth and passage rate of
microbial biomass, the factors affecting lag times, and
the assumption that the daily mean rumen digesta
load is the same in proportion to body weight on all
foods. These relationships need a more dynamic
description. Potential errors were identified with DM
degradation rates of cell wall contaminated with
microbes, and the differences in digestion rates due to
particle size and site of digestion are unclear. An
additional problem is that' validation' of a generalized
model by comparison with experimental data is
dependent on the data being representative of general
trends and not being unduly specific to the ex-
perimental circumstances. For example, a sheep may
possess physiological adaptations which limit its
usefulness as a representative of the average ruminant
from the same part of the weight range. Insight into
these adaptations could be gained from modelling,
but distinguishing adaptations from model inaccur-
acies must be the subject of further work.
We are very grateful to the following for their
helpful comments during development of the model:
A. Chesson, M. S. Dhanoa, D. A. Elston, F. D. DeB.
Hovell, G. Hughes, A. B. McAllan, E. L. Miller, J.
A. Milne, C. A. Morgan, J. D. Oldham, E. R.
0rskov, D. E. Spalinger.

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