2005 Poultry Science Association, Inc.

Role of Insoluble Fiber on Gizzard

Activity in Layers
H. Hetland,*,1 B. Svihus,* and M. Choct†

*Department of Animal and Aquacultrual Sciences, Agricultural University of Norway,

Ås, Norway; and †School of Rural Science and Agriculture,
University of New England, Armidale, Australia

Primary Audience: Egg Producers, Researchers, Nutritionists, Feed Manufacturers

SUMMARY
Effect of access to wood shavings on gizzard activity was examined for birds fed pelleted wheat
and oat diets with or without whole cereal inclusion. Layers on litter floor fed wheat diets with
access to coarse wood shavings showed up to 60% higher weight of the gizzard and its content
than caged layers fed the same diets without access to wood shavings. No such effect was found
by feeding oat diets. Thus, a significant interaction was found, indicating that the appetite for
wood shavings may be dependent of fiber level of the diet. Appetite for wood shavings and paper
was examined for birds fed wheat and oat diets in 2 other experiments. The intake of wood shavings
and paper by birds fed a wheat-based diet was numerically twice as high as in birds fed an oat-
based diet. However, the individual variation was remarkably high. Neutral detergent fiber (NDF)
concentration in the gizzard of birds fed the oat diet was up to twice (P < 0.05) the fiber concentration
in the gizzard of birds fed the wheat diet. Passage of structural fibers through the gizzard and the
appetite for feathers in the absence of structural fibers were also studied in an experiment using
diets based on rice and casein. Coarse fiber structures were observed to accumulate in the gizzard.
In the absence of fiber, birds ate feathers, indicating that birds may eat feathers to compensate
for the lack of structural components in the feed.

Key words: insoluble fiber, gizzard, digestion, layer

2005 J. Appl. Poult. Res. 14:38–46

structures of whole grains. The major chemical

DESCRIPTION OF PROBLEM
During the previous decade, use of whole
wheat (WW) mixed with a pelleted protein con-
centrate has become a common practice in Euro-
pean broiler farming because the inclusion of
whole grains in poultry diets can have beneficial
effect on nutrient digestion [1, 2] and feed use
[3, 4, 5, 6]. One of the important roles of whole
grains has been associated with their ability to
stimulate the activities of the gizzard [1, 2]. This,
in turn, is believed to be related to physical
component important to the structural integrity
of grains is the insoluble fiber, which makes up
the main part of the cell wall architecture. In-
deed, insoluble fiber itself has shown beneficial
effects on nutrient digestion [2, 7, 8, 9] and
gizzard activities [8, 9, 10]. Recent research has
shown that digesta passing through the gizzard
have a remarkably consistent particle size distri-
bution with the majority of particles being
smaller than 40 µm in size regardless of the
original feed structure [1]. The fact that gizzard

1
To whom correspondence should be addressed: harald.hetland@iha.nlh.no.
HETLAND ET AL.: INSOLUBLE FIBER AND GIZZARD ACTIVITY 39

TABLE 1. Composition of the diets (g/kg) used in experiments 1 to 4

Ingredient Wheat diet Oat diet Wheat diet Rice diet Commercial
Experiment 11 1 and 32 23 4 44

Wheat 556.25 150 707.51 — 600

Oats 150 547.45 — — —
Rice — — — 700 —
Sorghum — — — — 25.75
Millrun — — — — 50
Soybean meal 59.6 59.6 73.3 — 150
Fish meal 50 50 41.8 — —
Fish ensilage 41 41 41.0 — —
Casein — — — 160 —
Meat and bonemeal — — — — 20
Cottonseed meal — — — — 40
Soybean oil 22.3 40.0 17.7 — —
Canola oil — — — 30 —
Sunflower oil — — — — 3
Monocalcium phosphate 9.3 9.3 4.6 — —
Dicalcium phosphate — — — 10 15
Ground limestone 15 15 15.0 88 90
Shell meal 88.8 79.9 87.0 — —
Sodium chloride — — 0.15 3 2.5
Sodium bicarbonate 0.74 0.74 0.64 — —
Vitamin and mineral premix 2.35 2.35 2.35 26 26
Manganese oxide 0.12 0.12 0.12 — —
Titanium dioxide — — 5.0 — —
DL-Methionine 1.21 1.21 1.1 4 1.2
L-Lysine 0.31 0.31 0.28 1 0.2
L-Threonine 0.36 0.36 0.24 1 —
Choline chloride 0.2 0.2 0.2 1 0.35
Carotenoid mixtures 1.6 1.6 0.45 — —
Avizyme 1200 0.5 0.5 1.5 — —
Avizyme 1210 0.3 0.3 — — —
Betain 0.11 0.11 0.11 — —
1
Calculated nutrient contents per kilogram: AMEn, 11.19 MJ; CP, 153.3 g; lysine, 7.77 g; methionine, 4.23 g;
methionine+cysteine, 6.7 g; threonine, 5.7 g; calcium, 36 g; available phosphorus, 4.8 g. Analyzed natural detergent fiber
content, 107 g.
2
Calculated nutrient contents per kilogram: AMEn, 10.95 MJ; CP, 149.3 g; lysine, 8.1 g; methionine, 4.2 g; methionine+cysteine,
6.8 g; threonine, 5.6 g; calcium, 37.7 g; available phosphorus, 4.1 g. Analyzed natural detergent fiber content, 164 g.
3
Calculated nutrient contents per kilogram: AMEn, 11.4 MJ; CP, 160 g; crude fat, 36 g; lysine, 7.9 g; methionine, 4.2 g;
methionine+cysteine, 6.7 g; threonine, 5.8 g; calcium, 36 g; available phosphourus, 3.2 g. Analyzed natural detergent fiber
content, 86 g.
4
Calculated nutrient contents per kilogram: AMEn, 11.02 MJ; CP, 170.6 g; crude fat, 22.0 g; lysine, 7.8 g; methionine, 3.4
g; methionine+cysteine, 5.9 g; threonine, 5.5 g; calcium, 3.9 g; phosphorus, 5.0 g.
5
Vitamin premix provided the following per kilogram of diet: Retinyl acetate, 2.7 mg; cholecalciferol, 0.056 mg; DL-α-
tocopheryl acetate, 36 mg; menadione, 4.2 mg; pyridoxine, 3.2 mg; riboflavin, 9.8 mg; Ca-pantothenate, 11.8 mg; biotin,
0.15 mg; thiamine, 2 mg; niacin, 34 mg; cobalamin, 0.016 mg; folic acid, 1.55 mg, Fe, 50 mg; Mn, 40 mg; Zn, 70 mg; Cu,
10 mg; I, 0.5 mg; Se, 0.20 mg.
6
Provided per kilogram of control diet: retinol 2.42 mg, cholecalciferol 0.72 mg, D-α-tocopherol 0.08 mg, menadione 1.6
mg, thiamine 1.6 mg, riboflavin 4.8 mg, pyridoxine hydrochloride 5 mg, cobalmin 0.16, biotin 0.08, niacin 40 mg, calcium
pantothenate 9.6 mg, folic acid 1.6 mg, monensin 80 mg, Mn 64 mg, Fe 48 mg, Cu 6.4 mg, I 0.8 mg, Co 0.24 mg, Mo 0.8
mg.

contents have higher fiber content than the feed
illustrates that fiber is harder to grind than other
nutrients and thus is accumulated in the giz-
zard [7].
For broilers, turkeys, and layers on floor and
in modified cages, litter can be a significant
source of insoluble fiber. In addition, recent stud-
ies indicate that structural components of the
feed or litter ingesta may play a role in pre-
venting cannibalism among layers [11, 12]. If
so, such components as wood shavings may also
be an important enrichment of housing systems.
40
TABLE 2. Relative weight of empty gizzard and gizzard contents in 36-wk-old birds fed pellets with whole or ground
wheat or oats in conventional 3-hen cages or on litter floor (experiment 1)1
Conventional cages
Item GW WW
Empty gizzard, g/kg of live weight 11.7 11.6
Gizzard contents, g/kg of live weight 4.3 5.4
GW = ground wheat; WW = whole wheat; GO = ground oats; WO = whole oats.
1
Thus, the aim of these studies was to investi-
gate the response of birds to coarse wood shav-
ings in gizzard activities and gut weight when
they were fed diets low and high in fiber or
coarse and regular structure. Furthermore, we
examined the effect of fiber with and without
gizzard-stimulating properties on the voluntary
intake of fiber in birds fed low- and high-fiber
diets.
MATERIALS AND METHODS
Experiment 1
Nonbeak trimmed layers [13] to be used in
this experiment were used from another large
scale experiment. The birds were obtained at 16
wk of age and were housed in conventional 3-
bird cages [14] or in a litter floor system [15].
The birds housed in 3-bird cages were reared in
cages, while the birds housed on litter floor were
reared on litter floor. In the floor system, wood
shavings were used as litter cover, and it was
refilled weekly by new wood shavings. Cages
and the litter floor system were located in 2
different rooms, both with temperature and light
control. Light was given for 10 h from 16 wk
of age and incrementally to 15 h at 27 wk of
age by following commercial recommendations.
Diets were either wheat-based or oat-based
(Table 1). Both were fed either as a commonly
ground complete diet [ground wheat (GW) and
ground oats (GO)] or diets with 40% of the
whole cereals incorporated into the pellets [WW
and whole oats (WO)]. In the complete diets,
all cereals were ground through a 3-mm sieve.
The composition of the diets with whole cereal
was the same as in the corresponding control
diets, except 40% of the wheat or oats (of the
total diet) was added whole into the mixer before
pelleting. In the WW and WO diets, the shell
meal was also taken out of the mixture and given
JAPR: Research Report
Litter floor Pooled
standard
GO WO GW WW GO WO deviation
13.3 16.0 18.4 17.9 15.6 16.8 2.43
5.5 6.1 6.5 7.1 6.5 6.1 1.29
in coarse form on top of the feed in the troughs
daily in cages and in separate troughs on litter
floor. All feeds were conditioned at approxi-
mately 75°C, pelleted, and crumbled at a com-
mercial feed mill.
At 36 wk of age, 10 birds per feeding regi-
men and housing system were weighed and
killed by a cranial blow followed by cervical
dislocation. The gizzard was weighed with and
without its contents. Two to three birds were
selected at random from different cages from
the cage system, while birds housed on litter
floor were picked at random. Because different
numbers of bids were taken out of the cages, no
statistical analysis was performed.
Experiments 2 and 3
Birds and Housing System. Pullets [13]
were reared from d 1 in commercial rearing
cages without access to litter. At 21 wk of age,
40 birds were moved to 20, 2-bird cages in a
room with light and temperature control. Each
cage was equipped with 2 separate feed troughs
with 1 opening in the cage front for each trough.
Light was given for 12 h from 21 wk of age and
incrementally up to 16 h at 27 wk of age by
following commercial recommendations.
Feed Composition and Form. The feed in
experiment 2 was based on wheat. Wheat was
ground using a hammer mill [16] through a 3-
mm sieve. The feed mixture was heated to 75°C
in a conditioner [17] and pelleted through a pellet
press [18] fitted with a 3-mm die with 42 mm
effective thickness and crumbled. In experiment
3 the oat-based feed (GO) from experiment 1
was used (Table 1).
Experimental Design, Dissection and Sta-
tistical Analysis. Wheat (experiment 2) or oat-
based diet (experimant 3) was weighed out accu-
rately into 1 trough for each cage. In addition,
wood shavings or paper (processed through a
HETLAND ET AL.: INSOLUBLE FIBER AND GIZZARD ACTIVITY 41

TABLE 3. Performance and characteristics of the gizzard in birds fed wheat diets, 23 to 28 wk of age (experiment
2)

Wheat Wheat diet + Wheat diet +

diet paper wood shavings √MSE1
Feed consumption excluding fiber, g/d 108 113 114 9.48
Egg production, g/d 55.6 58.6 57.7 4.37
Egg to feed, kg/kg 0.51 0.52 0.51 0.033
Empty gizzard, g/kg of live weight 8.9b 9.9ab 11.2a 1.52
Gizzard contents, g/kg of live weight 3.3b 3.6b 5.1a 1.33
Neutral detergent fiber in gizzard, g/kg of DM 250 269 341 100
Bile acids in gizzard, mg/g of DM 8.3 7.3 6.6 2.00
Total bile acids in gizzard, mg 15.1 12.3 20.6 7.39
Live weight, g 1,751 1,739 1,753 95
a,b
Means in a row with different superscript are significantly different (P < 0.05).
1
√MSE = Square root of mean square error in the analysis of variable.

paper cutter) was weighed out to 7 cages each
in experiments 2 and 3.
At 28 wk of age, all birds per feeding regi-
men were weighed and killed by a cranial blow
followed by cervical dislocation. The gizzard
was weighed with and without its contents. Giz-
zard contents were analyzed for neutral deter-
gent fiber (NDF) and bile acids. Bile acid con-
centration of the gizzard contents was deter-
mined at the Norwegian School of Veterinary
Science, Oslo, Norway, using the Enzabile kit
[19]. Data from experiments 3 and 4 were ana-
lyzed separately using the general linear model
procedure of SAS software [20]. Differences be-
tween treatments were determined using the
least significant difference test. Residual stan-
dard deviation was used as a measure of ran-
domness.
Experiment 4
Birds and Housing System. A rice-based
control diet (as described in Table 1) with a very
low fiber level (analyzed nonstarch-polysaccha-
ride contents = 8 g/kg), the control diet with
coarse oat hulls (OH-Coarse), the control diet
with finely ground oat hulls (OH-Fine), and a
commercial diet were manufactured. The diets
with oat hulls were made by mixing the control
diet with fine and coarse oat hulls, respectively,
in the ratio 1:10 prepelleting. Analyzed content
of nonstarch-polysaccharides of the oat hulls
was 582 g/kg. The coarse oat hulls consisted of
large flakes of hulls from commercial dehulling
of oats, without further grinding except in the
pelleting process. The fine oat hulls were ground
to a powder on a laboratory hammer mill through
a 0.5-mm sieve.
Birds, Housing, and Dissection. One hun-
dred and fifty commercial layers [21], 68 wk old,
were placed in single-bird cages with individual
feeders. Prior to the experiment, the birds were
fed a commercial diet in similar cages.
To study the passage of structural fiber
through the gizzard, the OH-Fine and OH-
Coarse diets were fed to 36 birds each for 1
wk to get accustomed to the respective fiber-
enriched diets. During the next 10 d, all these
birds were fed the control diet to empty the
digestive tract of fiber.
Prior to the trial day, the birds were fasted
overnight. The birds were given access to excess
amounts of OH-Fine and OH-Coarse diets, re-
spectively, for 30 min, and the consumption was
measured. Thereafter, the OH-Fine and OH-
Coarse diets were replaced by the control diet,
and consumption was again measured. Prior to
feeding (t = 0) and after 2, 4, 8, 24, and 48 h,
6 birds for each feed were killed and dissected.
All the contents of the crop and gizzard were
collected and stored in a freezer until freeze-
drying.
The remaining 78 birds were used to study
feather pecking. Prior to the start of the experi-
ment (d 0), 6 birds were examined to determine
whether the population of birds had pecked and
swallowed feathers. All 4 feeds were weighed
out to 18 birds each. At d 3, 7, and 14, 6 birds
from each feed were killed and dissected. The
gizzard was opened, and its contents were col-
lected in jars for freeze-drying. Freeze-dried ma-
42 JAPR: Research Report

TABLE 4. Performance and characteristics of the gizzard for birds fed oat diets, 23 to 28 wk of age (experiment
3)

Oat Oat diet + Oat diet +

diet paper wood shavings √MSE1
Feed consumption excluding fiber, g/d 107 105 107 6.13
Egg production, g/d 53.9 51.5 52.5 4.87
Egg to feed, kg/kg 0.51 0.49 0.49 0.033
Empty gizzard, g/kg of live weight 16.2 15.5 15.2 1.57
Gizzard contents, g/kg of live weight 7.7 7.1 7.4 1.30
Neutral detergent fiber in gizzard, g/kg of DM 465 533 420 77
Bile acids in gizzard, mg/g of DM 6.9 5.2 5.4 2.11
Total bile acids in gizzard, mg 42.2a 26.5b 29.0b 10.47
Live weight, g 1,615 1,611 1,564 128
a,b
Means in a row with different superscript are significantly different (P < 0.05).
1
√MSE = Square root of mean square error in the analysis of variable.

terial was weighed before gentle grinding. Feath-
ers were separated and weighed for each bird.
Chemical Analyses, Calculations, and Sta-
tistical Analysis. Total fiber analysis was per-
formed using the AOAC alditol acetate proce-
dure [22].
Amount of oat hulls passed gizzard (% of
entered gizzard) was calculated as follows:
100 − [(oat hulls in gizzard × 100)/
(ingested oat hulls − oat hulls in crop)]
The experiment was designed to be com-
pletely balanced. However, 7 of the birds used
to study feather pecking, distributed among the
rice-based diet treatments (only d 7 and 14),
were excluded from the analysis because of their
very low feed intake (from 13 to 50 g/d). Data
for passage of oat hulls and feather contents in
the gizzard were not normally distributed, and
thus no statistical analysis is presented for these
data. The data for feed consumption and gizzard
weights were analyzed in a 2-way ANOVA us-
ing the general linear model procedure of SAS
software [20].
RESULTS AND DISCUSSION
The weight of the empty gizzard did not
differ with feeding pellets with ground or whole
cereals (Table 2). However, a tendency (P =
0.066) for increased weight of gizzard contents
was found with whole cereals in the feed. Birds
on litter floor had up to 60% higher (P < 0.05)
weight of empty gizzard and gizzard content
weights than birds in cages. However, the differ-
ence in gizzard weight and gizzard contents
weight due to housing system was greater for
birds fed wheat diets than those for birds fed
oat diets. Thus, significant (P < 0.05) interac-
tions were found between cereals and housing
systems. The significant (P < 0.05) interaction
between cereal and feed structure was due to an
increase in the empty gizzard weight when GO
were substituted for WO, while no difference
was found for the wheat diets.
Neither paper nor wood shavings affected
performance (Table 3 and 4). While carrying
out experiment 3, a warm weather period caused
difficulties for maintaining recommended tem-
perature. This may explain the lower egg produc-
tion for the oat diet compared with the wheat
diet.
Weights of the empty gizzard and gizzard
contents of birds fed the wheat diet were in-
creased due to their access to wood shavings,
but no such effect was apparent in birds fed the
oat diet. Access to paper did not affect gizzard
weight in either experiment. The NDF concen-
tration was considerably higher in gizzard con-
tents than in the feed in all treatments. The NDF
concentration in gizzard was twice as high (P <
0.05) in birds fed the oat diet as those fed the
wheat diet.
Bile acid concentration of gizzard contents
was similar for wheat and oat diets and also
similar with and without access to wood shav-
ings or paper. However, total bile acid content
was more than doubled in birds fed oats than
wheat diet, proportional to the difference in giz-
zard contents.
HETLAND ET AL.: INSOLUBLE FIBER AND GIZZARD ACTIVITY 43

TABLE 5. Daily consumption of paper and wood were normally distributed, statistical analysis of
shavings for birds fed wheat and oat diet (experiment the data from experiment 4 is presented. A nu-
2 and 3)
merically higher amount of oat hulls in the giz-
Wheat diet Oat diet zard was found for the OH-Coarse diet than OH-
Wood Wood Fine diet. Approximately 50% of the ingested
Paper shavings Paper shavings oat hulls from the OH-Coarse diet and 90% of
the ingested oat hulls from OH-Fine diet had
Mean, g/bird 1.5 0.5 0.2 0.3
Minimum, g/bird 0.0 0.2 0.0 0.0 passed the gizzard after 2 h. No oat hulls were
Maximum, g/bird 5.9 1.4 0.9 1.1 found in birds fed OH-Fine diet after 48 h,
whereas as much as 30% of the ingested oat
hulls were found in the gizzard of birds fed OH-
Numerically, birds fed the wheat diet con- Coarse diet.
sumed almost double the amount of wood shav- No significant amount of feathers was found
ings than those fed the oat diet, while consump- in birds prior to the start of the experiment (0.00
tion of paper was 6 times higher for birds fed to 0.02 g). A clearly lesser amount of feathers
the wheat diet than for birds fed the oat diet was found in the gizzard contents of birds fed
(Table 5). The numerical distance between the the commercial and OH-Coarse diets than those
mean and extreme values was observed to be fed control and OH-Fine diets, despite high indi-
up to 3 times the mean. vidual variation within a treatment (Table 6).
Approximately 6 and 3% of ingested oat Birds fed the OH-Coarse and commercial diets
hulls were found in the crop after 2 and 4 h, showed higher (P < 0.05) amounts of gizzard
respectively. Thereafter, no significant amount contents than those fed the control and OH-Fine
of oat hulls was found in the crop. diets. The gizzard contents in birds fed OH-
The amount of oat hulls that had passed the Coarse were observed to consist mainly of oat
gizzard (percentage of entered gizzard) at differ- hulls.
ent times is illustrated in Figure 1. Since only the The gizzard is known as the pacemaker or-
data for feed consumption and gizzard contents gan in birds, regulating the particle size of food

FIGURE 1. Passage of oat hulls through the gizzard.

44 JAPR: Research Report

TABLE 6. Feed consumption, gizzard contents, and feathers in the gizzard (experiment 4)

Control Commercial Standard

Item Day diet OH-Fine1 OH-Coarse1 diet deviation

Daily feed consumption, g 3 135 134 137 131 26.11

7 113 110 102 132
14 84 124 109 119
Gizzard contents excluding feathers, g DM 3 0.53 0.69 2.64 1.48 1.44
7 0.67 1.55 3.58 3.31
14 0.39 1.42 4.45 4.63
Feathers in gizzard, g 3 0.52 0.10 0.02 0.04 0.57
7 0.79 1.00 0.02 0.06
14 0.11 0.28 0.02 0.00

OH-Fine = the control diet with finely ground oat hulls; OH-Coarse = the control diet with coarse oat hulls.
1

entering the small intestine for downstream di-
gestion. Feeding studies with broilers have
shown that the gizzard becomes stimulated when
whole cereals replace ground cereals [1, 3, 8].
Replacing 50% GW with the same quantity of
GO gives the same increase in gizzard weight
as replacing 50% GW with WW [1]. A relatively
small amount of insoluble fiber (e.g., 10% oat
hulls coming from 50% of oats) stimulates the
gizzard to the same extent as 50% WW. The
current data indicate that the gizzard weight is
less affected by cereal structure in layers. How-
ever, insoluble fiber seems to play an even more
profound role for gizzard activity in the more
mature digestive tract of the layer. This corres-
ponds with an earlier experiment with layers
in which consumption of 4% of feed as wood
shavings resulted in 50% heavier gizzard,
whereas including 40% WW in the wheat-based
diet increased the gizzard weight by only 10%
[7].
In the current study, over 50% heavier giz-
zards in birds on litter floor than in cages ap-
peared to be a result of the birds consuming the
litter materials. Visual observations of gizzard
contents showed that the birds housed in a litter
floor system had consumed a considerable
amount of wood shavings. The interaction be-
tween cereal-grain type and system may indicate
that the voluntary intake of insoluble fiber from
the floor varies with dietary insoluble fiber level.
The numerical difference in gizzard weight and
its contents due to housing system was small for
birds fed oat diets and large for those given
wheat diets. Since oats contain considerably
more fiber than wheat, the appetite for wood
shavings seems to be related to the fiber content
of the diet. This was also supported by the results
of experiments 2 and 3 in which the intake of
wood shavings by birds fed a wheat-based diet
was almost twice that of birds fed an oat-based
diet. However, the very high individual variation
makes these results less conclusive. In contrast
to digestible nutrients, insoluble fiber plays a
physiological role in the digestive tract by inter-
acting with gut functions, nutrients and enzymes
[23]. Thus, the birds may perceive the physiolog-
ical response less sensitively than for digestible
nutrients, leading to vast variation in the intake
of wood shavings and paper. Individual pecking
and searching behavior may also influence the
consumption of indigestible fiber components.
In a previous study in which wood shavings
were sprinkled on top of the feed, a considerably
higher consumption was observed than in the
current experiment. This indicates that how and
where structural components, such as wood
shavings, are presented may influence the bird’s
ability to consume these materials [7].
In poultry, insoluble dietary fiber has shown
beneficial effects on starch digestion [7, 8, 9,
10]. It has been speculated whether the beneficial
effect of insoluble fiber on digestion is a result
of increased gizzard activity. The higher level
of total bile acids in the gizzard by feeding oats
compared with wheat suggests that structural
components increase digesta reflux, possibly
through increased gizzard activity. This result
agrees with a previous study [7] in which insolu-
ble fiber increased total bile acids in the gizzard.
That insoluble fiber decreases the nutrient con-
centration may also play a role by increasing
digestive juices and substrate relationship,
which is supported by the fact that fine cellulose
HETLAND ET AL.: INSOLUBLE FIBER AND GIZZARD ACTIVITY 45

powder has also shown beneficial effects on
starch digestion in wheat diets [2].
Coarse feed particles need to be ground to
a certain critical size before they can leave the
gizzard [24, 25], causing the volume of gizzard
contents to increase when diets with whole cere-
als or insoluble fiber are fed [7]. The current
data and previous experiments show that the
fiber level of the gizzard contents is about twice
that of the feed [7]. This means that fiber struc-
tures accumulate in the gizzard and that retention
time of insoluble fiber is longer than for other
nutrients. Fibers, such as cereal hulls, are very
solid and can probably be retained for a long
time in the gizzard. In contrast, whole cereals
that mainly consist of starch granules and protein
will be dissolved very fast in the acidic gastric
fluid in the gizzard. Thus, the gizzard activity
is more strongly stimulated by fiber structures
compared with whole cereal structures. Accu-
mulation of fiber in the gizzard and thus its
slower passage out of the gizzard are contrary
to the conventional theory that insoluble fiber
speeds up feed passage. However, it must be
stressed that this phenomenon may only be true
for the coarse, insoluble fiber fraction. Recent
work [10, 26] indicates that the passage time of
fine particles decreases when coarse fiber is fed
to broilers.
Similar to fine cellulose powder [2], paper
materials did not stimulate gizzard activity. The
higher consumption of paper than wood shav-
ings by feeding the wheat diet may indicate that
the appetite for fiber is triggered by a need for
gizzard-stimulating components. Birds with ac-
cess to wood shavings easily get the gizzard
stimulated by the consumption of a small amount
of the material. In contrast, birds with access to
paper get no such response of fiber consumption.
This lack of gizzard stimulation by paper may
be a reason why the birds with access to paper
showed a considerably higher fiber consumption
than those with access to wood shavings. The
similar consumption levels of paper and wood
shavings by birds fed the oat diet may be due
to the satisfaction of their fiber needs for gizzard
stimulation by the high fiber content of the diet.
An empty gizzard in birds fed the rice and
OH-Fine diets was probably due to the low-fiber
diet and the finely processed oat hulls passing
through the gizzard without stimulating it. This
reinforces the hypothesis that the holding of food
in the gizzard depends on the particle size of
the food and its insoluble fiber contents. In other
words, the gizzard will not retain food if it has
no need for grinding. Possibly, the bird has a
need for fibrous or structural feed components to
maintain a normal gizzard. In the current study,
birds fed a high-fiber commercial diet and a
diet containing coarsely ground oat hulls had a
negligible amount of feathers in their gizzards
compared with those fed a low-fiber diet and
a diet containing finely ground oat hulls. This
indicates that feather-pecking behavior may be
partly related to the feed structure and consis-
tency. Since the birds were housed in single-
bird cages, this phenomenon could not be caused
by cannibalism and aggressive pecks. Thus,
feather pecking and swallowing may be driven
by a need for gizzard stimulation. The gizzard
has been found to play a major role for gastrodu-
odenal reflux of digesta [27]. An empty gizzard
will not have feed stimuli and as such will not be
able to regulate downstream digestive processes.
This, once again, supports the hypothesis that
birds may have a requirement for fiber for stimu-
lation of the anterior digestive tract and that a
functional gizzard needs contents with struc-
tural components.

CONCLUSIONS AND APPLICATIONS

1. The gizzard has an excellent ability to grind coarse components. Coarse particles are selectively
retained in the gizzard until they are ground to a certain critical size.
2. Structural components in the form of whole cereals and coarse water-insoluble fiber can improve
feed use in birds fed highly concentrated diets. Improved feed use is at least partly caused by
increased digestibility of starch and may be due to increased gizzard activity.
46
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