Animal Science 2002, 74: 503-515 1357-7298/02/11020503$20·00

© 2002 British Society of Animal Science

Influence of dietary fibre on digestive utilization and rate of passage

in growing pigs, finishing pigs and adult sows
G. Le Goff, J. van Milgen and J. Noblet†

INRA, Unité Mixte de Recherches sur le Veau et le Porc, 35590 St-Gilles, France

† Corresponding author. E-mail: noblet@st-gilles.rennes.inra.fr

Abstract
Four experimental diets differing in the level and the origin of dietary fibre (DF) were studied : a control, low DF
diet (diet C, 100 g total dietary fibre (TDF) per kg dry matter (DM)) and three fibre-rich diets (200 g TDF per kg
DM) which corresponded to a combination of diet C and maize bran (diet MB), or wheat bran (diet WB), or sugar-
beet pulp (diet SBP). During two successive experimental periods, each diet was offered to five pigs at a growing
stage (35 kg body weight (BW)) and at a finishing stage (75 kg BW). In addition, four adult ovariectomized sows
received successively one of the four diets according to a 4 ✕ 4 Latin-square design. Digestive utilization of energy
and nutrients of diets and rate of passage parameters were determined using a pulse dose of ytterbium oxide
followed by total faecal collection. Faecal marker excretion was quantified using an age-dependent, one-
compartment model, from which the mean retention time in the gastro-intestinal tract of pigs (MRT) was obtained.
The digestibility of dietary energy and nutrients, especially the DF fraction, increased with the increase in BW from
growing to finishing pigs (P < 0·01) and was still higher in adult sows; the difference between pig stages was more
pronounced for diet MB. At each stage, the digestibility of energy or nutrients was lower (P < 0·01) for diets MB or
WB than for diet SBP. Accordingly, the energy and DF digestibility of sugar-beet pulp was higher and increased
much less with BW. The MRT was shorter for diets MB and WB in growing pigs and in sows. Sows had a longer
MRT (81 h) than finishing pigs (37 h) and growing pigs (33 h); however, MRT was highly variable between sows.
It is concluded that the degree to which different types of DF are digested depends, in part, on the botanical origin,
and it may be improved by a longer MRT in the gastro-intestinal tract of pigs. Some fibrous foodstuffs (such as
maize-by products) will benefit more from a longer MRT than others.

Keywords: digestibility, fibre, pigs, sows.

and energy decreases due to the low digestibility of

Introduction
Many fibrous, relatively inexpensive foodstuffs such
as cereal by-products are available for animal foods.
In addition, the use of a fibre-rich diet instead of a
conventional sow diet has been shown to lead to
increased gut fill and satiety and improved
behaviour of pregnant sows (Ramonet et al., 2000).
Therefore, there has been increasing interest in
feeding diets high in dietary fibre (DF) to pigs both
from an economic and animal welfare perspective. A
number of studies have shown that as DF increases
in the diet of pigs, total tract digestibility of nutrients
503
the DF fraction and possible interactions with
nutrients (Kuan et al., 1983; Stanogias and Pearce,
1985). However, studies on the capacity of the pig to
digest the DF fraction have given conflicting results,
which can be mainly explained by differences in the
botanical origin of the fibre used. For instance, the
DF fraction of sugar-beet pulp, which contains
mainly soluble carbohydrates, is more digestible
than the DF fraction of cereal bran with more
structural polysaccharides (Graham et al., 1986;
Chabeauti et al., 1991; Yan et al., 1995). In addition,
considerable variation between pigs in the ability to

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 504 Le Goff, van Milgen and Noblet

utilize fibre has been reported. It has been
demonstrated that digestibility of the DF fraction
increases with body weight (BW) of the pig with a
particularly important difference between growing
pigs and adult sows (Noblet and Shi, 1993b and
1994). The improvement appears dependent on the
botanical origin of fibre (Noblet and Bach Knudsen,
1997).
The difference between pig stages is due to a greater
ability of heavier animals to digest DF; it can be
explained by several factors including an increased
intrinsic ability of bacterial flora to digest fibre, a
greater number of bacteria, a reduction of relative
feeding level, and an increased transit time (Dierick
et al., 1989; Noblet and Shi, 1993a; Varel and Yen,
1997). The existence of a relationship between the
improvement of digestibility of DF with BW of the
animal and the ability of its flora to degrade DF has
not been demonstrated (G. Le Goff et al.,
unpublished data). It can therefore be hypothesized
that when the transit time increases, the microbial
flora in the large intestine has more time to ferment
the DF fraction, resulting in a greater digestibility of
fibre.
The aim of this study was to determine the effect of
level and botanical origin of DF (maize, wheat, sugar
beet) on the digestibility of energy and nutrients and
on the rate of passage of digesta through the total
gastro-intestinal tract at three pig stages (growing
pigs, finishing pigs and adult sows).
Material and methods
Experimental diets
Four experimental diets differing in level and origin
of DF were formulated. The control diet (diet C)
contained a low level of DF (100 g/kg dry matter
(DM) of total dietary fibre, TDF) and was prepared
from wheat and isolated soya-bean protein. The
other diets corresponded to a combination of the
control diet and either maize bran (diet MB), wheat
bran (diet WB), or sugar-beet pulp (diet SBP) and
were formulated to contain the same level of DF
(200 g/kg DM of TDF). Ingredients and chemical
characteristics of the diets are given in Table 1 and
chemical characteristics of the fibre-rich ingredients
in Table 2. During non-experimental periods, pigs
received a standard diet containing (g/kg diet):
wheat, 243·9; barley, 250; corn, 160; wheat bran, 50;

Table 1 Composition of experimental diets

Diets† C MB WB SBP

Components (g/kg diet)

Wheat 899·0 680·9 647·7 762·4
Isolated soya-bean proteins 68·5 51·9 49·3 58·1
Maize bran 234·7
Wheat bran 270·5
Sugar-beet pulp 147·0
Dibasic calcium phosphate 12·0 12·0 12·0 12·0
Calcium carbonate 11·0 11·0 11·0 11·0
Salt 4·5 4·5 4·5 4·5
Vitamins and minerals mixture‡ 5·0 5·0 5·0 5·0
Chemical composition (g/kg dry matter)
Ash 49 51 60 58
Crude protein (N ✕ 6·25) 171 168 170 160
Ether extract 16 23 23 15
Crude fibre 27 46 46 51
Neutral-detergent fibre 107 197 196 158
Acid-detergent fibre 28 51 56 55
Total dietary fibre 103 197 197 183
Water-insoluble cell walls 106 195 199 171
Starch 596 516 481 541
Sugars 28 22 39 30
Gross energy (MJ/kg dry matter) 17·97 18·19 18·04 17·71

† C, control diet; MB, maize-bran diet; WB, wheat-bran diet; SBP, sugar-beet pulp diet.
‡ The vitamins and minerals mixture provided the following (per kg diet): 2·7 mg retinyl palmitate; 25 µg cholecalciferol;
20·0 mg DL-a-tocopherol acetate; 2·0 mg thiamin; 4·0 mg riboflavin; 1·0 mg pyridoxine; 20 µg cobalamin; 15 mg niacin; 9·9 mg
D-pantothenate; 200 µg biotin; 1 mg folic acid; 2·0 mg menadione; 500 mg choline chloride; 100·2 mg Zn; 10·0 mg Cu; 37·0 mg
Mn; 80·0 mg Fe; 202 µg I; 100 µg Co; 150 µg Se.

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 Dietary fibre and digestion in pigs at different weights 505

Table 2 Chemical composition of the fibrous ingredients (g/kg dry diets during four successive 21-day periods. The
matter (DM)) feeding level was maintained at 2·4 kg food per day.
During each period, sows were adapted to the diet
Maize Wheat Sugar-beet for 11 days and subsequently moved to metabolism
Ingredients bran bran pulp
cages for collection of faeces for 10 days.
Ash 28 65 82 Measurement of the rate of passage occurred during
Crude protein (N ✕ 6·25) 162 170 100 the first 7 days of the collection period. During the
Ether extract 45 37 12 last 3 days of the collection period, faeces were
Crude fibre 110 107 202 collected daily and pooled (digestibility sample).
Neutral-detergent fibre 495 425 447
Acid-detergent fibre 126 127 215
Acid-detergent lignin 18 42 30 Total tract digestibility measurements
Total dietary fibre 482 432 705 All animals received the diet twice daily (08:00 h and
Water-insoluble cell walls 509 461 601 16:00 h) as mash food mixed with water (1 : 1·5; w/v)
Starch 243 168 † and had free access to water. Pigs were weighed at
Sugars 3 68 67 the beginning and at the end of the collection period.
Gross energy Food refusals and spillage were collected daily and
(MJ/kg DM) 19·59 18·80 16·87
analysed for DM content. For each diet, a sample of
† Not determined. food was collected and measured for its DM content
and subsequently used for chemical analysis. Faeces
were collected daily, stored at 4ºC and weighed,
homogenized and subsampled (digestibility sample)
at the end of the period. One faeces subsample was
soya-bean meal, 230; cane molasses, 30; lysine HCl,
heat-dried for DM determination and a second one
0·6; calcium carbonate, 14; dibasic calcium
was freeze-dried for further chemical analyses. All
phosphate, 12; salt, 4·5 and a vitamins and minerals
animals came from the herd of the Institut National
mixture, 5. The same vitamins and minerals mixture
de la Recherche Agronomique (Saint-Gilles, France).
was used in the experimental and standard diets
During the study, pigs were individually housed in
(Table 1).
metabolism crates located in a temperature-
controlled room (23 + 1ºC). Care and use of animals
Experimental design
were performed according to the Certificate of
Experiment 1: growing and finishing pigs. Five blocks of
Authorisation to Experiment on Living Animals, no.
four Piétrain ✕ (Large White ✕ Landrace) littermate
04739 (delivered by the French Ministry of
barrows, initially weighing 30 kg, were used. Each
Agriculture to J. Noblet).
animal was used during two successive experimental
periods (i.e. the growing (G) and finishing (F) period)
separated by a 28-day non-experimental period. Measurement of the rate of passage
Within each litter, one pig was allotted, based on BW, The first meal (08:00 h) of the measurement period
to one of the four dietary treatments. The pigs was marked with 1 g ytterbium oxide (Yb2O3)
received the same diet during both experimental incorporated directly in the mash food of each
periods. The feeding level was fixed at animal. The pulse dose administered in this way
proportionately about 0·9 of the ad libitum intake provided 878 mg ytterbium (Yb). The same diet not
recorded in previous trials with similar pigs kept in containing the marker (i.e. regular meal) was offered
the same conditions (Quiniou et al., 1996) and it was twice daily throughout the remaining period. The
progressively increased during the study. The end of the meal containing ytterbium oxide was
animals were adapted to the diet for 11 days before a considered as time 0. For each animal, a faecal
10-day total collection of faeces according to the sample was collected prior to ytterbium oxide
classical procedure used in the laboratory (Noblet et administration for determination of the zero level of
al., 1989). During the first 6 days of the collection marker (i.e. blank). Subsequently, faeces were
period, faeces were collected daily and pooled collected regularly and totally during separate
(digestibility sample). Measurement of the rate of intervals (transit samples) 3, 5, 8, 11, 14, 19, 24, 27, 29,
passage occurred during the last 4 days of the 32, 35, 38, 43, 48, 51, 56, 62, 72, 75, 80, 86 and 96 h
collection period. after feeding the marked meal in experiment 1, and
6, 11, 24, 30, 35, 48, 54, 61, 72, 78, 83, 96, 107, 120, 131,
Experiment 2: adult sows. Four Large-White ✕ 144, 155 and 168 h after feeding the marked meal in
Landrace ovariectomized sows (averaging 250 kg experiment 2. For each pig, each transit sample was
live weight) were assigned to the four dietary dried separately for 48 h at 105ºC, weighed, finely
treatments in a 4 ✕ 4 Latin square design. Each sow ground through a 0·5-mm screen (Retsch, Haan,
received alternatively one of the four experimental Germany) and stored at 4ºC.

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 506 Le Goff, van Milgen and Noblet
Chemical analyses
Dry matter content of each transit sample was
determined. A 0·5 g subsample of each transit sample
was ashed (550ºC, 8 h), digested in a solution of nitric
acid (1·5 mol/l) and analysed for Yb using an atomic
absorption spectrophotometer (Varian 220 FS,
Springvale, Australia); Yb was expressed as mg/g
DM (Mambrini and Peyraud, 1997). The digestibility
sample collected in experiment 2 over the last 3 days
of the collection period was also analysed for its
marker concentration. For food samples, the
methods of Association of Official Analytical
Chemists (1990) were used for measuring moisture,
ash, crude protein (N ✕ 6·25), crude fibre, and ether
extract. Gross energy content was measured using an
adiabatic bomb calorimeter (IKA C5000, Staufen,
Germany). Cell wall fractions (neutral-detergent fibre
(NDF), acid-detergent fibre (ADF) and acid-detergent
lignin (ADL)) were determined according to the
methods of Van Soest and Wine (1967) by using a
sequential procedure with prior amylolytic
treatment. TDF contents were quantified according
to the method of Prosky et al. (1988), and water-
insoluble cell walls (WICW), according to the
method of Carré and Brillouet (1986). Starch content
was determined using the Ewers polarimetric
method (European Economic Community, 1972), and
the content of sugars corresponded to alcohol-
soluble carbohydrates obtained by the method of
Luff-Schoorl (Bureau Interprofessionel d’Etudes
Analytiques, 1976). Moisture, ash, CP, cell wall
fractions and gross energy analyses were carried out
on each digestibility faecal sample; ether extract was
measured after hydrochloric acid hydrolysis on
pooled samples (one per diet and per pig stage).
Calculations and statistical analysis
Digestibility coefficients and energy values. The
digestibility coefficients were calculated for the 10-
day collection periods. For that purpose, the DM
excreted during the 10 days of the collection period
in experiments 1 and 2 were each considered as the
sum of DM excreted during the days of the
cumulated collection period (digestibility sample)
and during each faecal collection of the rate of
passage measurements. It was also assumed that the
so-called faeces digestibility sample (6 days in
experiment 1 and 3 days in experiment 2) was
representative of the faeces excreted over the 10-day
period. Apparent digestibility coefficients of organic
matter, nutrients and energy, and digestible energy
(DE) contents were calculated using routine
procedures (Noblet et al., 1989). According to
Graham et al. (1986) and Bach Knudsen and Hansen
(1991), faecal digestibility coefficients of starch and
sugars can be assumed 1·0. Results obtained on the
four diets were used to calculate total tract
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digestibility coefficients of nutrients and energy
values of the fibre-rich ingredients according to the
difference method (Noblet and Shi, 1994).
Estimation of rate of passage parameters. The quantity of
marker excreted during each faecal collection
interval was calculated as the product of the
concentration of the marker and the DM excreted
during that interval. The cumulated amount of
marker excreted over consecutive collection intervals
was calculated for each pig. The single-compartment
model proposed by Pond et al. (1986) has been used
to fit digesta passage data for pigs in the present
experiment. This model combines a gamma-two
distribution of residence times in a single age-
dependent mixing compartment with a displacement
compartment:
for t < τ, M(t) = 0
for t ≥ τ, M(t) = Co {1 – [exp–λ (t – τ) (1 + λ (t – τ))]},
where: M(t) = cumulative marker excretion at time t
(mg); Co = quantity of marker excreted during the
interval from 0 to ∞ h (mg); τ = time-dependent rate
parameter for compartment turn-over; t = time lapse
between pulse dosing and the end of the respective
collection interval (h); τ = time delay between pulse
dosing and first appearance of marker in faeces (or
residence time due to displacement flow) (h).
According to Pond et al. (1986), the parameters λ and
τ can be used to calculate mean residence time (MRT,
h) of marker in the gastro-intestinal tract (MRT = τ +
(2/λ)).
Therefore, the model was reparameterized as:
for t < τ, M(t) = 0
for t ≥ τ, M(t) = Co {1 – [exp(–2 (t – τ)/(MRT– τ))
(1 + (2 (t – τ)/(MRT – τ)))]}.
For each animal, the parameters τ and MRT and Co
were estimated using the NLIN procedure of
Statistical Analysis Systems Institute (SAS, 1990).
Conceptually, the model considers the MRT as the
sum of the residence times of the marker in a
displacement compartment (τ) and a mixing
compartment (T), calculated as T = MRT – τ. These
compartments are defined in mathematical terms
and they do not necessarily represent anatomical
compartments. Nevertheless, it can be anticipated
that digesta passage through the small intestine is
mostly represented by the displacement
compartment. Most of the passage models based on
mixing compartments originate from ruminant
 Table 3 Digestive utilization of diets in growing pigs and finishing pigs†

Significance‡
Growing pig Finishing pig
Residual Stage ✕
C MB WB SBP C MB WB SBP s.d. Animal Diet Stage diet

No. of observations 5 5 5 5 6 5 5 5
Body weight (kg) 32·1c 33·2c 33·8c 33·8c 77·7b 77·5b 77·1b 80·3a 1·3 * * **
Dry matter (DM) intake (g/day) 1262 1260 1269 1268 2135 2142 2142 2135 NA NA NA NA NA
Average daily gain (g) 622d 568d 556d 616d 949a 812c 836b,c 916a,b 76 * **
Digestibility coefficients
Dry matter 0·877b 0·804e 0·791f 0·863c 0·893a 0·826d 0·810e 0·872b 0·005 ** ** ** *

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Organic matter 0·896b 0·822f 0·814g 0·886c 0·914a 0·846d 0·837e 0·896b 0·004 ** ** ** *
Crude protein 0·835b 0·793c 0·772d 0·799c 0·902a 0·851b 0·843b 0·849b 0·013 * ** **
Ether extract 0·22 0·38 0·29 0·09 0·34 0·45 0·28 0·15 NA NA NA NA NA
Crude fibre 0·41e 0·36f 0·21h 0·63b 0·47c 0·44d 0·29g 0·61a 0·02 ** ** **
Neutral-detergent fibre 0·57c 0·46f 0·49e 0·64b 0·59c 0·54d 0·53d 0·70a 0·02 * ** ** *
Acid-detergent fibre 0·30e 0·35d 0·23f 0·59b 0·33e 0·43c 0·29e 0·63a 0·02 ** ** **
Total dietary fibre 0·48c 0·43d 0·44d 0·65b 0·47c 0·48c 0·44d 0·68a 0·02 * ** * *
Energy 0·867b 0·796f 0·779g 0·855c 0·894a 0·825d 0·808e 0·872b 0·005 ** ** ** *
Digestible energy (MJ/kg DM) 15·58b 14·48e 14·05f 15·15c 16·08a 15·00d 14·58e 15·44b 0·10 ** ** **
a,b,c,d,e,f,g
Diet effects; mean values within a row with unlike superscript letters were significantly different (P < 0·05).
† C, control diet; MB, maize-bran diet; WB, wheat-bran diet; SBP, sugar-beet pulp diet.
‡ From analysis of variance where main effects were animal within diet (no. = 23), diet (no. = 4), pig stage (no. = 2) and the interaction between diet and pig stage;
Dietary fibre and digestion in pigs at different weights

NA, not applicable. The same animals were used at growing and finishing stages and were given the same diet. Ether extract of faeces was measured on samples
pooled per diet and per pig stage and digestibility coefficients could not be calculated for each pig.

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507
 508 Le Goff, van Milgen and Noblet
studies. Although mixing of digesta occurs in the
large intestine in pigs, it has a more tubular structure
than the rumen. The presence of a mixing
compartment results in a gradual excretion of the
marker. The standard deviation of the residence
times (RT) (for both the total tract and the mixing
compartment) can be calculated as σRT = √(2/λ2)
(Matis et al., 1989).
Statistical analyses. Data obtained in experiment 1
were submitted to an analysis of variance with
animal within diet (no. = 23), diet (no. = 4),
physiological stage (no. = 2) and the interaction
between diet and pig stage as the main effects. The
effect of diet was tested against the animal within
diet error term, while the effects of animal within
diet, physiological stage, and the interaction between
diet and pig stage were tested using the residual
variance of the model as error term. Data obtained in
experiment 2 were submitted to a separate analysis
of variance with animal (no. = 4), diet (no. = 4) and
period (no. = 4) as the main effects. SAS (1990) was
used for all statistical analyses.
Results
Experiment 1: growing and finishing pigs
General aspects. Two pigs given diet C were removed
during the non-experimental period because of leg
problems for one pig, and very low food
consumption for the other one. They were replaced
by three finishing pigs that were adapted to diet C
for 11 days before total tract digestibility and rate of
passage measurements. The experimental conditions
for the three replacement pigs were similar to those
for the other pigs. Consequently, a total of 41
measurements were conducted (20 for the growing
stage and 21 for the finishing stage). Due to the
experimental design, BW differed between stages
and averaged 33 and 78 kg for growing and finishing
pigs, respectively (Table 3).
Total tract digestibility of diets. Data presented in Table
3 indicate a marked effect (P < 0·01) of pig stage on
the mean digestibility coefficients of organic matter,
crude protein, dietary fibre (i.e. crude fibre, NDF,
ADF and TDF) and energy. For the four diets, values
were higher (P < 0·01) in finishing pigs than in
growing pigs. The digestibility coefficients of ether
extract were low and values should be considered
with caution because of the low ether extract levels
in diets. Due to the improved digestibility of
nutrients, the DE content of diets was 0·5 MJ/kg
higher on average for the four diets (P < 0·01) in
finishing pigs than in growing pigs.
As indicated in Table 3, the total tract digestibility for
DM, organic matter, crude protein and energy were
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greater (P < 0·01) for diet C than for diets MB, WB or
SBP at both stages. Among the three fibrous diets (i.e.
diets MB, WB and SBP), the digestibility coefficients
of DM, organic matter, and energy were greater
(P < 0·01) for diet SBP, than for diets MB and WB
(Table 3). Furthermore, the difference in total tract
digestibility for organic matter or energy of diet MB
or WB between growing and finishing pigs was more
important than for diets C and SBP, which resulted in
an interaction (P < 0·05) between physiological stage
and diet characteristics. In fact, this result is mainly
due to an interaction between diet and physiological
stage (P < 0·05) on the DF fraction (i.e. NDF and TDF)
digestibility. For instance, the digestibility
coefficients of NDF were similar at both stages for
diet C while they increased markedly with BW for
diets MB, WB and SBP (Table 3). Finally, in
agreement with differences in total tract digestibility
of energy and in connection with differences in the
gross energy content (Table 1) of diets, the DE values
differed (P < 0·01) between diets at each pig stage.
Rate of passage of digesta. The results obtained during
the rate of passage measurements are reported in
Table 4. A lower number of samples (P < 0·05) was
collected when animals received diet C compared
with the other three diets. The total recovery of
marker in faeces was not affected (P > 0·05) by the
pig stage or by diet composition. Over the total
collection period, the proportion of marker recovered
from the 41 measurements averaged 0·92 (ranging
from 0·82 to 1·00). It should be noted that the mean
recovery of marker was quite low in growing pigs
fed the MB diet (0·89), mainly due to the low
recovery in one pig (0·56). As the kinetics of marker
excretion followed a pattern similar to those of the
other pigs, the lower recovery may be due to
incomplete marker intake. In addition, the mean
recovery of marker was also rather low in finishing
pigs given diet WB (0·82) but all values were of
similar magnitude.
Data reported in Table 4 show a longer MRT
(P < 0·01) in finishing (37 h, on average for the four
diets) than in growing pigs (33 h). This difference in
MRT was mainly due to differences in first marker
appearance in the faeces; the displacement flow
residence time (τ) was longer (P < 0·01) in finishing
pigs (25 h, on average for the four diets) than in
growing pigs (20 h). Finally, the mean residence time
in the mixing compartment (T) and the standard
deviation of the distribution of marker excretion
(σRT) were not different (P > 0·05) between pig
stages (Table 4).
Rate of passage of digesta through the gastro-
intestinal tract was not significantly affected by diet
 Table 4 Effect of diet composition on rate of passage estimates for growing and finishing pigs†

Significance‡
Growing pig Finishing pig
Residual Stage ✕
C MB WB SBP C MB WB SBP s.d. Animal Diet Stage diet

No. of observations 5 5 5 5 6 5 5 5
Indigestible dry matter (g/day) 160f 247d 265c 173f 228e 371b 407a 273c 11 * ** ** **
Rate of passage parameters
Recovery of marker
Amount ytterbium measured (mg)§ 840a,b 780a,b 834a,b 878a 831a,b 775a,b 727b 874a,b 79 +
Relative to ytterbium

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incorporated (%)§ 95·6a,b 88·8a,b 95·0a,b 100·0a 94·6a,b 88·3a,b 82·8b 94·9a,b 9·0 +
MRT (h) 34·3b,c 31·5c 32·1c 34·4b,c 37·8b 34·3b,c 34·2b,c 42·1a 2·7 ** **
τ (h) 22·7a,b 18·6c 18·4c 19·9b,c 26·0a 24·7a,b 24·6a,b 26·4a 2·6 ** **
T (h) 11·6a,b 12·9a,b 13·7a,b 14·5a 11·7a,b 9·6b 9·6b 15·6a 3·3
σRT (h) 8·2a,b 9·1a,b 9·7a,b 10·2a 8·3a,b 6·8b 6·8b 11·1a 2·3
No. of samples per animal 17a,b,c 19a 19a 18a,b 15c 16b,c 18a,b 16b,c 2 * *
a,b,c,d,e,f,g,h
Diet effects, mean values within a row with unlike superscript letters were significantly different (P < 0·05).
† C, control diet; MB, maize-bran diet; WB, wheat-bran diet; SBP, sugar-beet pulp diet; τ, displacement flow residence time (time delay between pulse dosing and
first appearance of marker in faeces) ; MRT, mean residence time; T corresponds to the mean residence time in the mixing compartment and was determined as the
difference between MRT and τ; σRT corresponds to the standard deviation of mean residence time in the mixing compartment and was calculated as σRT = √(2/λ2).
Indigestible dry matter corresponds to the difference between dry matter intake and digestible dry matter intake.
‡ From analysis of variance where main effects were animal within diet (no. = 23), diet (no. = 4), pig stage (no. = 2) and the interaction between diet and pig
Dietary fibre and digestion in pigs at different weights

stage; +, P ≤ 0·10. The same animals were used at growing and finishing stages and were given the same diet.
§ Total amount of marker incorporated (1 g ytterbium oxide) provided 878 mg ytterbium.

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509
 510 Le Goff, van Milgen and Noblet

composition (P > 0·05) (Table 4). It can be noted that
the MRT was numerically lower in animals given
diets MB or WB than in animals on diet C or SBP.
Similarly, the time of first appearance of the marker
(τ) was numerically greater (P > 0·05) in growing pigs
and finishing pigs on diet C than values obtained in
animals given diet MB or WB. Both the mean
residence time in the mixing compartment (T) and its
standard deviation were numerically greater
(P > 0·05) in pigs given diet SBP than for animals on
the other diets.
Experiment 2: adult sows
Total tract digestibility of diets. The mean BW of the
sows during the balance experiment was 252 kg and
was not significantly different between diets but was
affected (P < 0·01) by the experimental period (Table
5), resulting in a positive average BW gain (230 g/
day) for the four sows. Results show a clear
reduction of the total tract digestibility of DM,
organic matter and energy when the fibre level of the
diet was increased (P < 0·01). Differences between
the basal diet (i.e. diet C) and the high-fibre diets (i.e.
diets SBP, MB and WB) varied according to the type
of fibrous ingredient incorporated in the diets (Table
5). For instance, relative to the diet C, energy
digestibility was decreased by 0·020, 0·048 and 0·071
units (P < 0·01) for diets SBP, MB and WB,
respectively. The digestibility of crude protein was
affected by the composition of the diet (P < 0·01),
resulting in higher nitrogen faecal losses (P < 0·01) in
sows given the high-fibre diets (i.e. diets MB, WB and
SBP) than in animals on the diet C (result not
presented). Digestibility coefficients of crude fat
(ether extract) were low and quite variable between
diets (ranging from 0·07 to 0·42). Furthermore, the
digestibility coefficients of DF differed between diets
(P < 0·01), being the highest for diet SBP, irrespective
of the DF criterion used.
Rate of passage of digesta. The mean number of
samples collected per experimental period during
the rate of passage measurements differed (P < 0·01)
between sows (Table 6; no. = 11 on average). The
number of samples was particularly low for one sow
(no. = 9); the recovery of marker was also the lowest
for this sow (P < 0·05). Moreover, the concentration
of marker in the digestibility sample collected during
the last 3 days in experiment 2 was not negligible for
this sow while it was zero for the other sows.
Similarly, the MRT and the displacement flow
residence time (τ) were greater (P < 0·01) in this sow
(119 and 94 h, respectively) than in the other three
sows (68 and 52 h, on average; data not shown).
The proportion of marker recovered averaged 0·89,
with the lower value (0·85) obtained in sows given
diet MB (Table 6). The rate of passage was affected by
diet composition. First, the MRT was longer
(P < 0·01) in sows on diet C (95 h, on average) than
for animals given diets SBP, MB and WB (82, 78 and
69 h, respectively), mainly due to differences
(P < 0·05) in the time of first marker appearance (τ).
The MRT in the mixing compartment (T) and the

Table 5 Digestive utilization of diets in adult sows†

Diet Significance‡
Residual
C MB WB SBP s.d. Diet Animal Period

No. of observations 4 4 4 4
Body weight (kg) 251·6 253·7 251·4 251·8 1·6 ** **
Dry matter intake (g/day) 2106 2110 2116 2115 NA NA NA NA
Average daily gain (g) 232 232 212 260 124
Digestibility coefficients
Dry matter (DM) 0·885a 0·844c 0·819d 0·865b 0·010 **
Organic matter 0·916a 0·873c 0·852d 0·900b 0·009 **
Crude protein 0·913a 0·851d 0·877b 0·870c 0·005 ** *
Ether extract 0·31 0·42 0·32 0·07 NA NA NA NA
Crude fibre 0·39c 0·55b 0·38c 0·67a 0·03 **
Neutral-detergent fibre 0·58b 0·64a,b 0·58b 0·71a 0·04 **
Acid-detergent fibre 0·30c 0·52b 0·35c 0·62a 0·04 **
Total dietary fibre 0·47c 0·57b 0·50b,c 0·69a 0·05 **
Energy 0·896a 0·848c 0·825d 0·876b 0·008 **
Digestible energy (MJ/kg DM) 16·10a 15·42b 14·89c 15·52b 0·15 **
a,b,c,d
Diet effects, mean within a row with unlike superscript letters were significantly different (P < 0·05).
† C, control diet; MB, maize-bran diet; WB, wheat-bran diet; SBP, sugar-beet pulp diet.
‡ From analysis of variance with effects of diet (no. = 4), animal (no. = 4) and period (no. = 4); NA, not applicable. Ether extract
of faeces was measured on samples pooled per diet and per pig stage and corresponding digestibility coefficients could not be
calculated for each pig.

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 Dietary fibre and digestion in pigs at different weights 511

Table 6 Effect of diet composition on rate of passage estimates for adult sows†

Diet Significance‡
Residual
C MB WB SBP s.d. Diet Animal Period

No. of observations 4 4 4 4
Indigestible dry matter (g/day) 241d 329b 383a 285c 22 **
Rate of passage parameters
Recovery of marker
Amount ytterbium measured
(mg)§ 822a 748b 773a,b 777a,b 28 * **
Relative to ytterbium
incorporated (%)§ 93·6a 85·2b 88·8a,b 88·4a,b 3·2 * **
MRT (h) 95·0a 78·1b,c 68·6c 82·2b 6·0 ** **
τ (h) 75·9a 60·3b 54·7b 58·2b 6·6 * **
T (h) 19·1a,b 17·8a,b 13·9b 24·0a 4·6 + **
σRT (h) 13·5a,b 12·6a,b 9·8b 17·0a 3·3 + **
No. of samples per animal 11 11 12 11 1 *
a,b,c,d
Mean values within a row with unlike superscript letters were significantly different between diets (P < 0·05).
† C, control diet; MB, maize-bran diet; WB, wheat-bran diet; SBP, sugar-beet pulp diet; τ, displacement flow residence time
(time delay between pulse dosing and first appearance of marker in faeces) ; MRT, mean residence time; T corresponds to the
mean residence time in the mixing compartment and was determined as the difference between MRT and τ; σRT corresponds
to the standard deviation of mean residence time in the mixing compartment and was calculated as σRT = √(2/λ2). Indigestible
dry matter corresponds to the difference between dry matter intake and digestible dry matter intake.
‡ From analysis of variance with effects of diet (no. = 4), animal (no. = 4) and period (no. = 4); levels of significance : +, P ≤ 0·10.
§ Total amount of marker incorporated (1 g ytterbium oxide) provided 878 mg ytterbium.

standard deviation of residence times (σRT) were
numerically higher (P = 0·10) in sows given diet SBP
than for animals on the other diets.
Digestive utilization of fibrous ingredients in growing and
finishing pigs and in adult sows
Total tract digestibility coefficients of nutrients and
energy and energy values of maize bran, wheat bran
and sugar-beet pulp were numerically determined
according to the difference method; data cannot be
statistically compared (Table 7). Digestibility
coefficients of organic matter, and energy of
ingredients were numerically greater in adult sows
than in finishing pigs or in growing pigs. The
difference between stages was mainly explained by
increased digestibility coefficients of the DF fraction
(i.e. crude fibre, NDF, ADF and TDF). However, the
results indicate that the difference between pig stages
was more marked for maize bran, than for wheat
bran or sugar-beet pulp. For instance, the
digestibility coefficient of NDF determined in adult
sows was 0·32 units greater than in growing pigs for
maize bran, while the difference between these two
stages was 0·18 and 0·14 units for sugar-beet pulp
and wheat bran, respectively. As a consequence, the
difference in DE value between adult sows and

Table 7 Digestive utilization of maize bran, wheat bran and sugar-beet pulp in growing pigs (G), finishing pigs (F) and adult sows (A)†

Maize bran Wheat bran Sugar-beet pulp

G F A G F A G F A

Digestibility coefficients
Organic matter 0·578 0·637 0·738 0·588 0·635 0·682 0·804 0·800 0·804
Crude protein 0·616 0·697 0·643 0·583 0·704 0·780 0·368 0·397 0·462
Ether extract 0·53 0·58 0·54 0·33 0·24 0·33 NA NA NA
Crude fibre 0·36 0·38 0·67 0·10 0·21 0·38 0·77 0·79 0·89
Neutral-detergent fibre 0·38 0·50 0·70 0·44 0·48 0·58 0·71 0·85 0·89
Acid-detergent fibre 0·38 0·50 0·67 0·19 0·26 0·37 0·80 0·87 0·86
Total dietary fibre 0·41 0·46 0·64 0·43 0·44 0·51 0·74 0·86 0·89
Energy 0·565 0·624 0·707 0·541 0·597 0·648 0·744 0·747 0·756
Digestible energy (MJ/kg
dry matter) 11·03 12·22 13·86 10·13 11·23 12·19 12·66 12·61 12·75

† Determined by the difference method (Noblet and Shi, 1994). NA, not applicable.

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 512 Le Goff, van Milgen and Noblet
growing pigs was quite important for maize bran
and wheat bran (+2·8 and 2·0 MJ/kg DM,
respectively) and close to zero for sugar-beet pulp
(Table 7).
Discussion
Effect of body weight of pigs on total tract digestibility of
dietary energy and nutrients and energy values
In experiment 1, the digestibility of organic matter,
energy and nutrients of all diets increased between
35 kg BW growing pigs and 80 kg BW finishing pigs.
A larger effect of BW can be observed when adult
sows (experiment 2) and growing pigs (experiment
1) are compared. These observations are in
agreement with previous results (Roth and
Kirchgessner, 1984; Fernandez and Jørgensen, 1986;
Fernandez et al., 1986) and are mainly explained by
the improved digestibility of the DF fractions (Tables
3 and 5). This is well illustrated when digestibility
coefficients of maize bran or wheat bran are
compared between pig stages (Table 7). However, in
the case of sugar-beet pulp, the digestibility of
energy or organic matter was reasonably stable
between pig stages, despite a positive increase in the
digestibility of the DF fraction with BW of pigs. This
discrepancy can be related to the limits of the
difference method in terms of accuracy of
calculations and the assumption that the digestive
utilization of the basal diet is not affected by the
presence of sugar-beet pulp. Finally, it can be
hypothesized that the difference in digestibility
values, especially in digestibility of DF, would have
been less pronounced if the duration of adaptation to
the diets before digestibility measurements in
experiment 1 in growing pigs (11 days) had been
similar to that used in sows (18 days in experiment
2). Indeed, Longland et al. (1993) demonstrated that 3
to 5 weeks are necessary before stability of
measurement of digestibility of DF is reached in
growing pigs.
The first practical application of this study concerns
the estimation of the digestible energy values of
diets, especially for the fibre-rich diets, for different
physiological stages (Tables 3 and 5). According to
conclusions given previously (Noblet and Shi, 1993b;
Le Goff and Noblet, 2001), foods should be given
energy values depending on BW of pigs. Similarly,
the energy values measured in adult sows for wheat
bran in the present study or in the studies of Noblet
and Bourdon (1997) or Noblet and Le Goff (2000) are
greater than those proposed in the Centraal
Veevoeder Bureau (2000) feeding tables. It can then
be concluded that energy values given in feeding
tables are adapted to growing animals and new
values applicable to adult pigs are required (Le Goff
and Noblet, 2001).
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Rate of passage parameters in the pig
The model of Pond et al. (1986) provided a
reasonably good statistical fit of the pattern of
marker appearance in faeces. However, in some
cases, especially in sows, the statistical fit was not
excellent due to a low number of samples (5 to 9) per
animal. In the present study, the marker was not
totally recovered in faeces (0·92 and 0·89, on average
for experiments 1 and 2, respectively). The lower
results obtained in the present study, especially in
sows, may be due to (1) incomplete total collection,
(2) an overestimation of the theoretical ingested
marker and/or (3) cumulative errors during marker
analysis in the faeces. However, according to the
model used, the recovery rate has no influence on the
estimates of rate of passage parameters.
The rate of passage varied between animals given
the same diet (Tables 4 and 6); this phenomenon was
particularly important in adult sows. The sow with
the greater MRT defaecated relatively less frequently
than the other sows. Similar variability in digesta
passage in pigs has been previously reported
(Stanogias and Pearce, 1985). This is a short-coming
of the current technique which essentially measures
defaecation patterns. In addition, it has been
suggested that variations in MRT or in frequency of
defaecation are related to the activity of the pigs
(Latymer et al., 1990). Independently of the diet and
feeding level, the retention of digesta in the gastro-
intestinal tract was twice as long for one sow, and
thus leaving more time for digestion; one would
therefore expect a greater digestibility of energy and
nutrients in this sow (Dierick et al., 1989). In fact, the
digestibility coefficient of energy was not different
for this sow when compared to the sows with lower
MRT (data not shown). This suggests that the digesta
retention time in the sows is sufficiently long for its
duration not to affect faecal energy digestibility.
Effect of body weight on digesta passage
The results show clearly that the MRT and τ
increased between growing and finishing pigs
(experiment 1, Table 4); this effect is more
pronounced when these values are compared with
those obtained in adult sows (experiment 2, Table 6).
However, results obtained in experiments 1 and 2
must be compared with caution since growing and
finishing pigs were adapted to the diet for a longer
period (17 days) than adult sows (11 days) before
transit measurements. The MRT values measured in
pigs given the basal diet in the present study are of
similar magnitude to those measured in previous
studies in pigs given low DF diets. Firstly, Potkins et
al. (1991) reported a MRT, defined as the excretion of
0·5 of the marker, equivalent to 32 h in 28 kg BW
growing pigs. Secondly, Pond et al. (1986) using the
 Dietary fibre and digestion in pigs at different weights
same method of measurement as in the present study
obtained a 44 h MRT value in 80 to 85 kg BW pigs.
Finally, MRT values equivalent to 82 and 95 h were
estimated by Holzgraefe et al. (1985) and Münchow
et al. (1986), respectively, when mature sows were
given a control low DF meal. As illustrated by Pekas
(1991), the size and weight of the digestive system
increase with age and BW, with a longer hindgut in
270 kg BW pigs (7·5 m) than in 70 kg BW pigs (5·4 m)
or in 30 kg BW pigs (4·3 m). Moreover, the digesta
residence time in the hindgut has been shown to be
the main component of the total residence time
(Latymer et al., 1990) and represents 0·60 to 0·80 of
the total residence time in the gastro-intestinal tract
(Keys and DeBarthe, 1974; Kass et al., 1980). From
these observations, it appears that the greater
retention time of digesta in the gastro-intestinal tract
of sows is mainly related to the greater size of the
hindgut. The σRT is greater in sows than in growing
and finishing pigs. The σRT is indicative of the size
of the mixing compartment (relative to the flow of
digesta), which supports the idea that the greater
MRT in sows is due to the larger size of the hindgut.
Moreover, the feeding level may also affect passage
in the gastro-intestinal tract (Roth and Kirchgessner,
1985). Accordingly, the lower feeding level used for
adult sows in the present experiment partly explains
the increased MRT in adult animals.
Effect of botanical origin of fibre on rate of passage
parameters
At each pig stage, the utilization of maize bran or
wheat bran (relative to the basal diet) resulted in a
numerical decrease of MRT (Tables 4 and 6), which is
in agreement with previous results (Fioramnoti and
Bueno, 1980; Stanogias and Pearce, 1985). In fact, at
the current level of DF, adding maize or wheat bran
increased the quantity of indigestible DM 1·4- to 1·8-
fold (Tables 4 and 6). Therefore, cereal bran
supplementation exerts a direct physical action in the
hindgut of pigs which stimulates propulsive colonic
motility due to a greater bulk of digesta (Bardon and
Fioramonti, 1983; Cherbut et al., 1988). Since sugar-
beet pulp is degraded to a greater extent by the
bacterial flora (McBurney and Thompson, 1990;
Salvador et al., 1993) with little increase of the
undigested organic matter (Tables 4 and 6), it does
not act physically on the transit of digesta
throughout the hindgut. However, the effect of
adding sugar-beet pulp to the basal diet on passage
is less clear since there was no effect in growing pigs,
a positive effect in finishing pigs and a negative
effect in adult sows (Tables 4 and 6) when compared
with the basal diet. Literature data on the effect of
soluble DF sources such as sugar-beet pulp on MRT
are rather contradictory with an increase when
offered to 30 to 85 kg BW pigs (Latymer et al., 1990),
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513
no effect on MRT in 28 kg BW growing pigs (Potkins
et al., 1991) and an acceleration of the transit in adult
humans (Salvador and Cherbut, 1992). In this latter
study, it was proposed that products of degradation
of DF, especially volatile fatty acids, may stimulate
colonic wall contractions with a subsequent
increased rate of passage of digesta in the gastro-
intestinal tract. Nevertheless, variations in the effect
of sugar-beet pulp on MRT between pig stages are
not clearly explained.
Comparative aspects
The physico-chemical properties of sugar-beet pulp
DF favours digestive degradation (Salvador et al.,
1993; Guillon et al., 1998). Subsequently, little
improvement with increased BW is expected with an
increased MRT in adult animals. The insoluble
lignified structure of wheat bran DF resists
degradation in the digestive tract, irrespective of the
duration of fermentation (McBurney and Thompson,
1990; Bach Knudsen, 1997). Consequently, the
digestibility of wheat bran DF is only slightly
improved when MRT is increased. Maize bran is
rather specific since its DF fraction is poorly
digestible in the growing pig but highly digestible in
the adult sow (Table 7). Such an interaction has
rarely been described in the literature (Noblet and
Bach Knudsen, 1997). These authors hypothesized
that the digestibility of maize bran could be greatly
improved when the retention time in the hindgut of
animals is increased. Results of the present study
support this hypothesis since the increased MRT
between finishing pigs and adult sows was
considerable for diet MB (+40 h, on average). In
conclusion, the degree to which different types of DF
are digested depends primarily on the physico-
chemical characteristics, which differ according to
botanical origin (Longland and Low, 1991). However,
for maize by-products, the utilization of the DF can
be improved with the greater retention time in the
hindgut of adult animals.
Conclusion
The results of the present study show that the
digestibility of energy and nutrients (especially the
DF fraction) of fibre-rich diets or fibrous ingredients
improved with increasing BW of the pig. In parallel,
the MRT of digesta in the gastro-intestinal tract
increased with BW, probably due to a larger size of
the hindgut and a lower relative feeding level. It is
hypothesized that improved digestibility of the DF
fraction of diets with BW can be mainly explained by
a greater retention time of digesta. However, the
degree to which different types of DF are digested
also depends on the botanical origin. From a
practical point of view, these results emphasize that
the determination of energy values of fibre-rich foods
 514 Le Goff, van Milgen and Noblet
should be carried out at different BW and the
botanical origin of fibrous ingredients has to be
considered.
Acknowledgements
This work was partly supported by the GERNA (Paris). The
authors gratefully acknowledge R. Vilboux for preparing
experimental diets, J. Delanoë, Y. Lebreton, F. Le Gouevec,
A. Roger and J. F. Rouaud for their technical assistance in
animal management and P. Bodinier, Y. Jaguelin and S.
Hillion for their technical help in laboratory analyses. We
also thank J. L. Peyraud for his assistance in the preparation
of the protocol.
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(Received 7 June 2001—Accepted 24 December 2001)