The effect of insoluble fiber and intermittent feeding on gizzard development,

gut motility, and performance of broiler chickens

A. Sacranie,*† B. Svihus,‡1 V. Denstadli,‡ B. Moen,§ P. A. Iji,*† and M. Choct†

*School of Environmental and Rural Sciences, The University of New England, Armidale NSW 2351, Australia;
†Poultry Cooperative Research Centre, Armidale, NSW 2351, Australia; ‡Department of Animal and Aquacultural
Sciences, Norwegian University of Life Sciences, PO Box 5003, N-1432 Aas, Norway;
and §Norwegian Institute of Food, Fisheries and Aquaculture Research, Oslovn. 1, N-1430 Aas, Norway

ABSTRACT Two experiments were conducted to test
the following hypothesis: exposing broiler chickens to
coarse insoluble fiber in the diet will result in enhanced
gizzard function and performance, improved adaptabil-
ity to an intermittent feeding program, and an increase
in the occurrence of reverse peristalsis. In experiment 1,
102 Ross 308 broiler chickens were either intermittently
or ad libitum fed a basal diet, the basal diet diluted
with 15% coarse hulls (consisting of equal weights of
hulls from oats and barley), or the basal diet diluted
with 15% of the same hulls finely ground in a 2 × 3
factorial arrangement with 17 individually caged birds
per treatment. Birds fed ad libitum had access to feed
continuously for 18 h/d, whereas those on intermit-
tent feeding had restricted access to feed from 7 d of
age. From 18 d of age, the restrictive-feeding program
consisted of four 1-h meals and one 2-h meal per day.
In experiment 2, 156 broiler chickens in 12 pen cages
Key words: intermittent feeding, reverse peristalsis, insoluble fiber, oat hulls, broiler
with wood shaving-lined floors were exposed to 1 of 4
treatment groups with 3 pens/treatment: intermittent
or ad libitum feeding of a basal diet and intermittent
or ad libitum feeding of a coarse hull diet, as described
above. At 31 and 32 d of age, birds in experiment 1
were inoculated with chromium EDTA via the cloaca.
There was no interaction between diet and feeding regi-
men. The addition of hulls increased gizzard weight and
content and lowered (P < 0.001) gizzard pH, but it
had no effect on the ability of the birds to handle inter-
mittent feeding. Despite the dilution with coarse hulls,
weight gain and the gain:feed ratio were not affected,
which could partly be explained by an increased (P <
0.001) starch digestibility. Dietary reflux was confirmed
by the presence of chromium in all intestinal tract sec-
tions. Broilers exhibited reverse peristaltic contractions
of sufficient magnitude to propel the marker from the
cloaca to the gizzard.
2012 Poultry Science 91:693–700
http://dx.doi.org/10.3382/ps.2011-01790

INTRODUCTION and duodenum is increased by access to insoluble fiber.

Many feed ingredients, such as barley, oats, and soy-
bean meal contain a considerable amount of insoluble
fiber (Bach Knudsen, 1997). The inclusion of 10% oat
hulls in broiler diets has been found to increase wheat
starch digestibility and stimulate gizzard activity (Ro-
gel et al., 1987; Hetland and Svihus, 2001). In addition,
Hetland et al. (2003) observed a higher total amount of
bile acids in the gizzards of birds with access to wood
shavings, indicating gastro-duodenal reflux proportion-
al to the amount of material in the gizzard, supporting
their hypothesis that digesta reflux between the gizzard
©2012 Poultry Science Association Inc.
Received August 8, 2011.
Accepted October 14, 2011.
1 Corresponding author: birger.svihus@umb.no
This appears to be a logical reaction by the bird to
the presence of structural components, prolonging the
exposure of food to both the mechanical and chemical
components of digestion.
Broiler production relies on fast growth rates and ef-
ficient feed conversion. In view of this, feed is available
to birds ad libitum often under nearly continuous or
continuous lighting programs. This approach to hus-
bandry is associated with potential overconsumption
(Svihus, 2001; Svihus et al., 2010) and physiological
growth-related problems, such as leg weakness and asci-
tes. Restrictive feeding, by the use of intermittent light-
ing programs, has been shown to reduce such problems
and improve feed efficiency (Julian, 1993; Decuypere et
al., 1994; Buyse et al., 1996). The majority of studies
in this area have employed light/dark programs rather
than meal feeding to manipulate feed intake. Provid-
ing birds with access to feed 4 times in a day did not

693
694 Sacranie et al.
affect feed efficiency but depressed weight gain com-
pared with birds with ad libitum access to feed (Petek
et al., 2005). Fewer meals also reduced weight gain but
improved feed efficiency compared with ad libitum-fed
birds (Su et al., 1999). Svihus et al. (2010) found that
feed efficiency improved for birds on intermittent feed-
ing.
Intermittent feeding requires the bird to retain in-
gested feed for greater lengths of time than ad libitum
feeding by storing ingesta in the crop and proventricu-
lus or gizzard (Buyse et al., 1993). By stimulating giz-
zard development with the dietary inclusion of barley
and oat hulls, it is possible that the resulting increase in
holding capacity and heightened gizzard function will
enable the broiler to thrive under intermittent feeding.
The purpose of this study was to investigate the ef-
fect of a dietary dilution of broiler diets with insoluble
fiber in the form of hulls on performance, reverse peri-
stalsis in the digestive tract, and a broiler chick’s ability
to adapt to an intermittent feeding regimen.
MATERIALS AND METHODS
Feed, Birds, and Experimental Design
In experiment 1, one-day-old broiler chicks (Ross 308)
were raised on a commercial pelleted starter diet in a
multilevel brooder. At 11 d of age, the birds were moved
to 12 group cages with plastic mat-covered floors, and 2
cages were assigned to each of 6 treatments in a 2-feed-
ing regimen × 3-diet experimental design. At 18 d of
age, 17 birds were randomly selected from each of the
6 groups (102 in total) and transferred to individual
cages (22 cm wide × 38 cm deep × 38 cm high) where
they remained until the termination of the study. The
temperature was gradually reduced from 31 to 28°C at
7 d of age and to 25°C at 16 d of age.
The experimental diets were steam-pelleted through
a 3-mm die and consisted of a basal control diet, a basal
diet diluted with 15% unground coarse hulls (consisting
of equal weights of hulls from oats and barley), and a
basal diet diluted with 15% finely ground hulls of the
aforementioned material (Table 1). The fine hulls were
produced by grinding the coarse hulls through a ham-
mer-mill fitted with a 1-mm sieve. Dry sieving through
a series of sieves while vortexing (amplitude 1.5 mm) on
a Retsch AS 200 (F. Kurt Retsch GmbH & Co., Haan,
Germany) showed that the amount of particles larger
than 1 mm in size was reduced from 77 to 2% as a con-
sequence of grinding. Irrespective of feeding regimen,
ad libitum (AL) or intermittent feeding (IF), birds ex-
perienced complete darkness between the hours of 0200
and 0800. From 11 to 18 d of age, the IF groups had
access to feed from 0800 to 0900 h, 1200 to 1300 h, 1630
to 1730 h, and from 2100 h until the lights went off.
From 18 d of age, these birds had access to feed from
0800 to 0900 h, 1200 to 1300 h, 1600 to 1700 h, 2000 to
2100 h, and from 2400 h until the lights went off.
In experiment 2, 156 broiler chicks from the same
flock as in experiment 1 were divided among 12 pens
with wood shaving-lined floors at 11 d of age. Three
cages per treatment were assigned to each of 4 treat-
ments consisting of a control diet or a coarse hulls-
diluted diet fed either ad libitum or intermittently in
a 2 × 2 experimental design. The diets and feeding
regimens were identical to those in experiment 1. At 25
d of age, the number of birds in each cage was reduced
from 13 to 8.
Data and Sample Collection
In experiment 1, feed intake and weights for indi-
vidual birds were recorded twice a week from 18 d of
age until the end of the study. At 26 d of age, feed was
removed from all chickens at 0700 h and weighed. At
1300 h, all birds were weighed, feed replaced (AL birds
only), and a clean tray placed under each individual
cage for excreta collection. Quantitative excreta collec-
tion was carried out at 27 and 28 d of age. At 29 d of
age, feed was again removed and weighed at 0700 h. At
1300 h the birds were returned to their normal feeding
regimen and excreta trays were removed for the collec-
tion of excreta.
The termination of experiment 1 was carried out over
2 d, 9 birds from each of the 6 groups were sampled
on d 32 and the remaining 8 from each group on d 33.
The sampling days were carefully planned to maintain
the feeding regimens right up until the birds were eu-
thanized and to ensure consistency in exposure to the
marker chromium EDTA. The first 5 birds to be ad-
ministered the marker at 0800 h on d 32 were treated
in the following way: on d 31 the IF birds out of the 5
total birds were given access to feed from 2200 to 2400
h. The feed was then removed from all 5 birds, returned
at 0600 h on d 32, and then removed from all 5 birds
at 0700 h. After 1 h off the feed (at 0800 h), 2 mL of
chromium EDTA (5.32 mg of chromium) marker was
administered via a syringe fitted with a blunt-tipped
crop needle, inserted 6 cm into the cloaca of the bird.
Care was taken to ensure that the crop needle was in-
serted beyond the colonic sphincter, and cages were
checked regularly during the next 2 h to remove excreta
adhering to the wire floor so that oral consumption of
the marker was avoided. If the bird expelled the marker
immediately after or during administration, another 2
mL was given. The birds were then returned to their
cages and birds were given access to feed as determined
by their normal feeding program. The next 5 birds fol-
lowed the same protocol, starting the night before at
2220 h and receiving the marker at 0820 h on d 32. In
this way, all 102 birds were sampled over 2 d in 20-min
batches.
After 2 h of exposure to the marker, the 5 birds were
killed by a cranial blow followed by cervical disloca-
tion and then dissection. Weights were recorded for the
empty gizzard and the full and empty small intestine
 FIBER, INTERMITTENT FEEDING, AND GUT MOTILITY 695
Table 1. Composition of diets
Item Control Hulls

Ingredient, %
 Corn 25.36 21.50
 Wheat 42.00 35.70
  Oat and barley hulls mixture1   15.00
  Soybean meal, 440 g of CP/kg 17.35 14.75
  Fish meal, 700 g of CP/kg 7.00 5.95
  Soybean oil 4.00 3.40
 Limestone 1.30 1.11
  Monocalcium phosphate 1.00 0.85
 l-Lysine, 780 g/kg 0.50 0.43
 dl-Methionine, 990 g/kg 0.30 0.26
 l-Threonine, 985 g/kg 0.10 0.09
 Salt 0.20 0.17
  Sodium bicarbonate 0.30 0.26
  Choline chlorate 0.15 0.13
  Mineral premix2 0.15 0.13
  Vitamin premix3 0.29 0.26
  Calculated lysine 1.46 1.23
  Calculated methionine + cysteine 1.03 0.87
  Calculated threonine 0.85 0.73
  Calculated AMEn4 3,012 2,619
Chemical analysis   Coarse hulls diet Fine hulls diet

  DM, g/kg 896 900 904

  CP, g/kg 212 192 192
  Crude fat, g/kg 62 59 60
  Gross energy, kcal/kg 4,111 4,135 4,159
  Starch, g/kg 399 369 365
1Particle size distribution based on dry sieving was the following for coarse and finely ground hulls, respectively:
>2 mm, 32 and 1%; 1 to 2 mm, 45 and 1%; 0.5 to 1 mm, 15 and 50%; 0.2 to 0.5 mm, 8 and 33%; <0.2 mm, <1
and 15%. The hulls contained 9.6% starch and 5.7% protein and had a negligible fat content.
2Mineral premix provided the following per kilogram of diet: Fe, 75 mg; Mn, 60 mg; Zn, 105 mg; Cu, 15 mg; I,
0.75 mg; and Se, 0.3 mg.
3Vitamin premix provided the following per kilogram of diet: vitamin A (retinyl acetate), 9,000 IU; vitamin D
(cholecalciferol), 5,400 IU; vitamin E (dl-α-tocopheryl acetate), 66 IU; menadione, 9.0 mg; pyridoxine, 6.0 mg;
riboflavin, 24.0 mg; Ca-pantothenate, 26.3 mg; biotin, 0.39 mg; thiamine, 3.75 mg; niacin, 75 mg; cobalamin, 0.03
mg; and folic acid, 3.75 mg.
4As recommended by the NRC (1994).

from the pancreatic duct to the ileo-ceco-colonic junc-
tion. The pH of the gizzard with contents was recorded
by immersing the electrode into the center of the giz-
zard through the duodenal sphincter. Contents from
the crop, gizzard, duodenum, jejunum, and ileum from
half of the birds were collected quantitatively and im-
mediately frozen in liquid nitrogen. To ensure that the
concentrations of chromium observed in chickens from
this trial were a result of the introduced marker, 6 test
chickens were killed and dissected as described above.
The birds came from the control group of another trial
where they were raised on a similar diet as used in the
present study. They had not been exposed to any ex-
ternal source of chromium. Five birds died and 14 were
discarded at termination because they showed signs of
having ascites, and these birds were equally distributed
among treatments. The birds were cared for accord-
ing to laws and regulations governing experiments with
live animals in Norway (the Animal Protection Act of
December 20, 1974, and the Animal Protection Ordi-
nance concerning experiments with animals of January
15, 1996).
In experiment 2, 4 birds per pen were killed as de-
scribed for experiment 1 and dissected at 31 d of age.
Contents from the crop and gizzard were collected and
weighed after drying, and the weight of the empty giz-
zard was recorded; these data were pooled per pen be-
fore statistical analysis. The IF birds were killed 45 min
after having access to feed.
Chemical Analyses
Contents from the gizzard, duodenum, jejunum, and
ileum were dried at 80°C for 24 h, ground in a coffee
grinder, digested in nitric acid using a closed chamber
microwave, and then analyzed for chromium content as
described by Binnerts et al. (1968), using inductively
coupled plasma emission spectroscopy. The contents
from the crop and gizzard obtained from the litter-
reared birds were dried at 105°C overnight.
Excreta samples were frozen immediately after each
quantitative collection, mixed, dried overnight at 105°C,
and then analyzed for gross energy content on a Parr
1281 isoperibol calorimeter (Parr, Moline, IL) along
with feed samples. Starch content in feed and excreta
was determined enzymatically following the procedure
of McCleary et al. (1994), whereas Kjeldahl-N was de-
termined according to AOAC (2003) on a Kjeltech 2300
696 Sacranie et al.
(Foss Tecator, Höganäs, Sweden). The AME was cal-
culated by correcting to N balance, using the N levels
analyzed for in the feed and excreta, as described by
Bourdillon et al. (1990).
Statistical Analyses
In experiment 1, data were analyzed using a 2 × 3
factorial arrangement followed by pairwise comparisons
using the Duncan procedure with P < 0.05 as the sig-
nificance level (SAS Institute, 2002). In experiment 2,
data were subjected to a 2 × 2 factorial arrangement
followed by pairwise comparisons using the Duncan
procedure with P < 0.05 as the significance level. In
both experiments, variation was represented by the re-
sidual SE (root of mean square error).
RESULTS
Experiment 1
There was no significant interaction between feeding
regimen and diet for any variable measured (Table 2).
The AL-fed birds tended to consume more feed than
the IF birds (P = 0.07). Diet affected feed intake (P <
0.05), which was because of an increased feed intake by
birds fed diets with coarse hulls added. No significant
effect of feeding regimen on weight gain was observed,
but due to a reduced weight gain for the diet with finely
ground hulls, diet affected (P < 0.01) weight gain. Di-
lution of the diet with hulls reduced (P < 0.001) the
gain:feed ratio. Feeding regimen did not affect AME or
starch digestibility. Dilution with hulls, however, im-
proved (P < 0.001) starch digestibility.
Both IF and dilution with hulls reduced (P < 0.001)
gizzard pH (Table 2). The addition of hulls to diets re-
sulted in fuller and larger gizzards, with birds exposed
to coarse hulls yielding the largest (P < 0.001) gizzards.
Diet affected (P < 0.05) the weight of the small intes-
tine due to a reduced weight with finely ground hulls.
Chromium, at levels considerably higher than the
background level, was present in all sections of the
gastrointestinal tract, with no significant difference be-
tween treatments (Table 3). However, it tended (P ≤
0.07) to be higher in the gizzards and the jejunum of
the AL-fed birds.
Experiment 2
There was a tendency for reduced weight gain both
with IF (P = 0.06) and with hull dilution in the diet (P
= 0.11; Table 4). Dilution of the diet with hulls reduced
(P < 0.05) the gain:feed ratio. The weight of the empty
gizzard increased with IF (P < 0.05) and with hull dilu-
tion (P < 0.001; Table 4). In addition, IF tended (P =
0.06) to elevate the amount of DM in the crop.
DISCUSSION
Access to coarse hulls stimulated gizzard develop-
ment. This is in agreement with previous studies high-
lighting the stimulatory effect of structural material on
the gizzard (Preston et al., 2000; Hetland et al., 2002;
Plavnik et al., 2002; Svihus et al., 2004, 2010; Ravin-
dran et al., 2006) and as discussed in a recent review
(Svihus, 2011).
The large particle size of the coarse hulls and their
hardness as a result of the insoluble fiber content ex-
plains why birds consuming the coarse hull diet devel-
oped the heaviest gizzards. The coarse hull particles are
retained in the gizzard until they are ground to a cer-
tain critical size that allows them to pass through the
pyloric sphincter (Clemens et al., 1975; Moore, 1999;
Hetland et al., 2002, 2003). This leads to an increase
in the volume of the organ’s contents and a muscular
adaptation to meet the greater demand for grinding.
Birds fed the fine hull diet also exhibited heavier giz-
zards than those of the control group, although less so
than those of the coarse hull group, suggesting that
the fine particles were retained and partly induced the
same response. Hetland et al. (2003) reported that the
mean duodenal particle size for birds fed a diet con-
taining ground or whole oat hulls was between 120 and
127 μm. This agrees with an earlier report that stated
that 90% of duodenal particles were less than 300 μm
(Hetland et al., 2002) and suggests that hulls ground
through a 1-mm sieve are large enough to be retained
in the gizzard as a grinding medium.
The birds fed the coarse hull diet in experiment 2,
reared on wood shavings, exhibited increases in relative
empty gizzard weights in a similar way to the birds in
experiment 1. Birds have been shown to consume litter
material when it is available, presumably to compen-
sate for a lack of structural material in the diet (Het-
land et al., 2004, 2005). It is interesting to note in this
context that relative gizzard size tended to be larger
for the birds in experiment 2 that had access to litter
material than for the birds in experiment 1. In experi-
ment 2, there was a slight increase in relative gizzard
weights in AL birds fed the control diets compared with
birds on the same diet and feeding conditions in ex-
periment 1. This effect was even more dramatic in the
IF birds. With limited exposure to feed but with full
access to litter, birds possibly developed heavier giz-
zards as a result of consuming the litter between meals.
Although litter consumption was not measured in this
study, presence of litter material was observed during
the dissection of the birds. Access to litter resulted in
a higher relative gizzard weight in the IF birds raised
on the hull diet compared with that of the caged birds
fed the coarse hull diet. This is contrary to the idea
that when wood shavings are available to a bird it will
monitor its intake depending on the level of indigestible
fiber present in the diet (Hetland et al., 2005). These
results indicate that given the opportunity or motiva-
Table 2. Experiment 1 performance and digestive characteristics of birds (17 to 32 or 33 d of age)
Starch Crop Relative Gizzard Relative Relative empty
Feed BW AMEn, digestibility Crop content, Gizzard gizzard content, full intestine intestine
Item intake, g gain, g G:F kcal/kg coefficient content, g g of DM pH weight, %1 g of DM weight, %1 weight, %1

Feeding                        
  Ad libitum control 1,724 1,076 0.62 3,143 0.98 4.1 1.1 3.60 1.2 0.3 3.9 2.3
  Ad libitum coarse 1,803 1,113 0.62 2,881 0.99 2.3 0.6 2.27 2.4 4.6 3.7 2.4
  Ad libitum fine 1,707 1,021 0.60 2,902 0.99 3.3 1.4 2.28 1.8 4.6 3.2 2.1
  Intermittent control 1,646 1,050 0.64 3,065 0.97 3.7 1.8 3.01 1.1 1.1 3.5 2.2
  Intermittent coarse 1,716 1,058 0.62 2,871 0.99 2.3 1.2 1.96 2.3 3.8 3.3 2.3
  Intermittent fine 1,649 989 0.60 2,960 0.99 4.9 1.5 1.97 1.7 4.0 3.1 2.1
Root mean square error (n = 83) 46.6 104.7 0.034 108.7 0.012 6.14 2.43 0.580 0.29 1.70 7.02 0.37
Feeding regimen                      
  Ad libitum 1,745 1,070 0.61 2,923 0.99 3.2 1.0 2.72 1.8 3.2 3.6 2.3
 Intermittent 1,671 1,033 0.62 2,947 0.99 3.6 1.5 2.31 1.7 2.9 3.3 2.2
Diet                        
 Control 1,685b 1,063ab 0.63a 3,023a 0.97b 3.9 1.5 3.31a 1.2c 0.7b 3.7a 2.3a
 Coarse 1,760a 1,086a 0.62a 2,844c 0.99a 2.3 0.9 2.12b 2.4a 4.2a 3.5a 2.4a
 Fine 1,678b 1,005b 0.60b 2,940b 0.99a 4.1 1.5 2.13b 1.8b 4.3a 3.1b 2.1b
Main effect                      
  Feeding regimen 0.07 NS NS NS NS NS NS *** NS NS 0.09 NS
 Diet * ** *** *** *** NS NS *** *** *** * *
a–cMeans within a column not sharing a common superscript differ significantly at P < 0.05.
1Relativeweights are calculated as a percentage of BW.
*P < 0.05; **P < 0.01; ***P < 0.001.
FIBER, INTERMITTENT FEEDING, AND GUT MOTILITY
697
698 Sacranie et al.
Table 3. Amounts of chromium detected in the digestive tracts of experiment 1 birds1
Total Cr, μg

Item Gizzard Duodenum Jejunum Ileum

Feeding
  Ad libitum control 4.3 6.1 41.8 11.9
  Ad libitum coarse 32.5 11.7 14.9 17.9
  Ad libitum fine 34.8 5.1 30.7 25.0
  Intermittent control 3.3 22.0 9.4 16.2
  Intermittent coarse 8.5 4.4 3.2 25.2
  Intermittent fine 11.4 8.9 4.2 7.0
Root mean square error (n = 83) 38.57 26.46 58.66 39.68
Feeding regimen        
  Ad libitum 23.9 7.6 29.1 18.3
 Intermittent 7.7 11.8 5.6 16.1
Diet        
 Control 3.8 14.1 25.6 14.1
 Coarse 20.5 8.1 9.1 21.6
 Fine 23.1 7.0 17.5 16.0
Main effect        
  Feeding regimen 0.07 NS 0.06 NS
 Diet NS NS NS NS
1For chickens that were not exposed to the chromium EDTA marker, the levels of Cr detected in the gizzard, du-
odenum, jejunum, and ileum were 1.3 ± 0.48 μg, 0.17 ± 0.046 μg, 0.64 ± 0.21 μg, and 2.66 ± 0.41 μg, respectively.

tion by way of feed restriction, birds will display forag-
ing behavior independent of the level and particle size
of insoluble fiber in the diet.
In accordance with previous findings, the increase in
volume and weight of gizzards in birds exposed to insol-
uble fiber in the diet was accompanied by a reduction
in pH in the caged bird (Nir et al., 1994; Gabriel et al.,
2003; Svihus, 2011). The strong effect of feeding regi-
men on pH in the control birds is especially interest-
ing. This may be explained by the increase in the DM
content of the crop and gizzard in the same birds. The
significant rise in DM content in the control birds, as a
result of IF, could lead to an increase in fermentation,
resulting in more acidic digesta flowing into the proven-
triculus and gizzard. The significant effect of feeding on
gizzard DM contents alludes to a longer retention time,
which leads to an increase in hydrochloric secretions
from the proventriculus (Nir et al., 1994).
These gains in mechanical and chemical gizzard func-
tion coupled with a longer retention time have been
shown to result in improvements in nutrient digest-
ibility and feed utilization (Svihus, 2011). This was
confirmed by starch digestibility data from the current
experiment and by the considerably higher AMEn of
the hulls-diluted diets than what would be expected
based on the calculated energy value of the control diet
and the hulls (NRC, 1994). Starch granules in wheat
are embedded in a protein matrix. In order for starch
digestion to take place, the granules need to be released
by proteolysis (Barlow et al., 1973; Simmonds et al.,
1973). Péron et al. (2007) showed that proventricular
pepsin activity was highest between pH 2 and 3. In
addition to pH, a prolonged retention time of gizzard
contents, as observed in birds fed the hull diets, would
provide improved conditions for pepsin activity expos-
ing the starch granules more readily than in birds with

Table 4. Performance and digestive characteristics of experiment 2 birds (11–31 d of age)

Feed BW Crop content, Relative gizzard Gizzard content,
Item intake, g gain, g G:F g of DM weight, %1 g of DM

Feeding            
  Ad libitum control 2,408 1,675 0.70 7.0 1.4 2.1
  Ad libitum coarse 2,444 1,601 0.66 9.4 2.3 7.0
  Intermittent control 2,333 1,612 0.69 19.9 1.7 2.6
  Intermittent coarse 2,416 1,563 0.65 20.5 2.7 6.1
Root mean square error (n = 12) 87.7 49.2 0.032 9.44 0.26 0.73
Feeding regimen            
  Ad libitum 2,426 1,638 0.68 8.2 1.8 4.6
 Intermittent 2,375 1,588 0.67 20.2 2.2 4.4
Diet            
 Control 2,371 1,644 0.69 13.5 1.5 2.4
 Coarse 2,430 1,582 0.65 15.0 2.5 6.6
Main effect            
  Feeding regimen NS 0.11 NS 0.06 * NS
 Diet NS 0.06 * NS *** ***
1Relativeweights are calculated as a percentage of BW.
*P < 0.05; ***P < 0.001.
 FIBER, INTERMITTENT FEEDING, AND GUT MOTILITY
passive-functioning gizzards. This is a possible mecha-
nism for the recorded improvement in starch digestibil-
ity in the hull diet groups. Starch digestibility may also
be improved by a more active gizzard, resulting in a
finer grinding and acting as a regulator for digesta flow
into the small intestine (Hetland and Svihus, 2001).
Heightened gizzard activity has also been linked to a
stimulation of pancreatic secretions, resulting in an in-
crease in amylase (Hetland et al., 2003).
The lack of interaction between feeding system and
diet structure indicates that the increase in volume
of the gizzard does not influence the bird’s ability to
adapt to meal feeding. Svihus et al. (2010) recorded the
same trend and suggested that if the frequency of feed-
ing intervals is too great, as in this study, the variation
in gizzard volume will have less effect.
Despite the very strong limitations on feed availabil-
ity through the 5 feeding times and only 6 h with feed
available compared with 18 h for the AL-fed birds, the
IF birds were able to adapt quite quickly to IF, as indi-
cated by the similar weight gain as the AL birds after
an adaptation period and no significant reduction in
bird weight at the termination of the experiment. This
is due to an extensive use of the crop as an intermedi-
ary storage organ for feed, as shown here and previ-
ously (Svihus et al., 2010).
Although some research has shown the prevalence
of reverse peristalsis in turkeys (Duke, 1992), the cur-
rent observations are the first to indicate the presence
of reflux through the whole gastrointestinal tract of
chickens. In the current study, stunning the bird fol-
lowed by cervical dislocation was chosen over other
methods to euthanize birds. In previous studies carried
out by the same research group, CO2 asphyxiation has
been employed due to the assumed low level of spastic
movements by using this method (Coenen et al., 2000).
However, a considerable loss of intestinal contents via
regurgitation, far from ideal when measuring gut motil-
ity, was experienced by using this method (Svihus et
al., 2010), and thus stunning, followed by cervical dis-
location, was chosen in the current experiment. It was
observed in this study that birds died without excessive
movement after cervical dislocation and without regur-
gitating. Thus, although some spastic movements af-
fecting digesta reflux may have taken place, it is unlike-
ly to have had a large effect in the current experiment.
Perhaps the most suitable method for gastric motility
studies in terms of minimizing convulsions would be
intravenous pentobarbital, which is known to inhibit
gastric motility (Duke et al., 1977). However, due to
the time-limiting factors in the current protocol and
potential difficulties in administering the injection in
a timely fashion, this method was deemed unsuitable.
The tendency for higher levels of chromium to be
present in the gizzards and jejunal contents of AL birds
fed the hull diets is contrary to what was expected.
Reflux has been characterized as a highly organized
motility response to feed shortage, resulting in the re-
699
establishment of motility patterns along the digestive
tract and satiety for a short period of time (Clench and
Mathias, 1992; Jiménez et al., 1994). It was assumed
that reverse peristalsis, especially the gastroduodenal
reflux, would be one method by which the IF birds
would improve feed utilization. In view of this, the inter-
mittent feeding system was perhaps not severe enough
to induce an antiperistaltic response. Conversely, the
theory that reflux is induced by feed shortages may not
be sound and that reflux is rather a characteristic of
optimal mixing of the lumen contents of satiated birds.
However, chromium EDTA is a liquid phase marker
and retention time in the gizzard is dependent on par-
ticle size (Vergara et al., 1989). If digesta were refluxed
back into the gizzard, the marker would be the part of
the fraction to pass first through the pylorus sphincter.
Consequently, the chromium levels measured in the giz-
zard may not be wholly representative of the extent
of the gastroduodenal reflux. Previous research has
suggested the stimulating effect of coarse fiber on giz-
zard function, in particular more frequent and powerful
contractions and the subsequent intraluminal pressure
changes that they induce, will lead to an increase in the
occurrence of gastric refluxes (Duke, 1982; Hetland et
al., 2003). The aforementioned limitation of the liquid
phase marker and increase in gizzard contents in birds
exposed to the hull treatments, suggest that further
work should be carried out to investigate the potential
for reflux of solid feed particles and possible interaction
with recorded improvements in starch digestibility.
The difference in detected ranges of the chromium
level in the birds unexposed to the marker and experi-
mental birds is adequate to conclude that chromium
recovered from digesta samples from experimental birds
originated from the introduced marker. It is remark-
able that the intestinal motility of a bird is capable
of propelling the marker from just above the colonic
sphincter to the gizzard with an average recovery of
0.30%. This equates to one-fourth of the total recovered
chromium from the gastrointestinal tract. Samples were
taken 2 h after infusion of the marker and it is more
than likely that a large proportion of chromium was
passed out in the excreta; but previous unpublished
results point to a large flow of the marker into the ceca
where it is held and then released slowly according to
cecal contractions. The results from this study clearly
demonstrate reflux of the liquid fraction of intestinal
contents. However, additional studies are required to
quantify the amount and frequency in which material
is relocated by reverse peristalsis.
In conclusion, this study demonstrates that birds
raised on diets diluted with hulls exhibit improved per-
formance due to improved gizzard function and hold-
ing capacity. This did not improve the ability of birds
to deal with restricted feeding. Furthermore, the study
clearly illustrates the prevalence of reverse peristalsis,
or reflux, along the entire length of the gastrointestinal
tract in broiler chickens.
700 Sacranie et al.
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