The Effect of Added Sucrose on the Digestibility of

Protein and Fiber by Swine

C. N. Skipitaris, R. G. Warner and J. K. Loosli
J ANIM SCI 1957, 16:55-61.

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T H E E F F E C T OF ADDED SUCROSE ON T H E D I G E S T I B I L I T Y
OF P R O T E I N AND F I B E R BY SWINE
C. N. SKIPI~A~!S, 1 R. G. WAgNeR AND J. K. LOOSLI

Cornell University, Ithaca, N. Y.

has been used with success in swine rations, especially in
S UGAR
starters for young pigs (Diaz et al., 1956). Because of its high
solubility, sugar might be expected to be completely digested and
absorbed, but the problem has not been studied with swine. The addition of 15% or more of sugar or molasses to the ration of ruminants
results in a depression of digestibility of the crude fiber and crude
protein (Hamilton, 1942). In 1953, Skipitaris, at the University of
Thessaloniki in Greece, observed that the addition of carob pod meal to
a barley ration depressed the digestibility of the crude protein in the
ration by swine. Since the carob pod meal contained 35 to 40=~o sucrose
and reducing sugars, this result suggested that excess sugar may
decrease the digestibility of certain nutrients in feeds for swine, as has
been observed with sheep and cattle. Mitchell (1942) reported, however,
that such pronounced associative effects have been noted only with
ruminants.
Digestion experiments were carried out to study the influence of added
sugar on the digestibility of crude protein and crude fiber by swine.
Experimental

Procedure

Four barrows, two Yorkshires weighing 215 and 208 lb. and two
Berkshires weighing 164 and 154 lb. were used in 16 digestion trials.
Table 1 shows the plan of the study and the feed intakes during the
four experimental periods. The design was a modified latin square suggested by Henderson (1955) and allowed for an estimation of the
carry over effects.
During the basal periods the larger pigs were fed a constant amount
of 1800 gin. daily of barley, ground to a fine meal, supplemented with
vitamins and minerals, and the smaller pigs received 1600 gin. daily.
Table grade sucrose was added at a level of 16% during the other
periods without lowering the barley allowances. These levels of intake
1 Present address: Department of Animal Husbandry, University of Thessalonike, Greece.
2 The assistance of H. W. Seeley is acknowledged for helpful suggestions in carrying out the
experiment and in reading the manuscript.

55

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56

S K I P I T A R I S , W A R N E R AND

LOOSLI

were established during a pre-experimental period of two weeks. There

were no feed refusals during the trials. The basal barley ration supplied
approximately 60% of Morrison's T D N standard for growing and
fattening pigs of these body weights. The pigs gained 0.64 to 0.67 lb.
daily during the experimental periods. A 10-day preliminary and a
7-day collection period made up each trial. The feed for each period
was carefully mixed and the allowance for each feeding was weighed
into a paper bag at the start of each period, samples being saved at the
same time for chemical analyses.
The pigs were kept in metabolism crates similar to those used by
Crampton and Whiting (1943). They were fed twice daily at 8.30 a.m.
TABLE

1. T H E F E E D I N G S C H E D U L E O F P I G S R E C E I V I N G G R O U N D
BARLEY WITH AND WITHOUT ADDED SUCROSE
Feed intake, grams per day
Period I

Pig No. 9
1u
2B
3Y
4B

Barley b Sucrose
1800
1600
I800
1600

288
256

a y is for Vorkshire and B for

b The barley was ground and
recommended by Beeson et al.
final barley meal fed contained

Period I[

Period I I I

Barley

Sucrose

Barley

1800
1600
1800
1600

288

1800
1600
1800
1600

256

Period IV

Sucrose

Barley

Sucrose

288
256

1800
1600
1800
1600

256
288

Berkshire.
supplemented with vitamins and minerals to furnish the amounts
(1953). Zinc was added at a level of 4 ppm of the barley, the
10.90% and 4.91% of crude protein and crude fiber, respectively.

and 5:30 p.m. The feed was mixed into a thick slop and water was
allowed ad lib. after feeding. The cages permitted the complete collection of feces without contamination with urine. The feces were collected
twice daily, weighed and a 7% aliquot placed in a tightly closed glass
jar with 1% of thymol and stored at approximately 40 ~ F. At the end
of the collection period, the daily aliquots were composited, mixed and
a subsample used for chemical analyses. Nitrogen was determined on
the fresh feces by the Kieldahl method. Crude fiber was determined by
the A.O.A.C. method.
Bacterial protein in the feces was determined by a modification of
the method of Mattill and Hawk (1911) using a series of fractional
sedimentations in a high-speed centrifuge. Dilute HC1 solution (0.2%)
was used to free the bacteria of fecal residues as described in detail
by Skipitaris (1956). The crude protein fractions studied are described
in table 2.
Counts were made of the total fecal bacteria by the nigrosine slide

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SUGAR AND DIGESTIBILITY OF PROTEIN AND FIBER

57

TABLE 2. THE RESULTS OF THE MICROSCOPIC EXAMINATION OF

VARIOUS COMPONENTS OF FECAL PROTEIN OF SWINE
SEPARATED BY FRACTIO'NAL CENTRIFUGATION
Fractionafion
designation ~
1. Residue protein
2. Bacterial protein
3. Soluble protein

Approx.
No. of gs?

Centrifugation
time

Microscopic
observation

min.
1

230
10800

Fecal particles
protozoa, few bacteria

20

Bacteria, traces of
fecal debris

Supernantant
From no. 2

Free of cells and

debris

a NX6.25.
b No. of times gravity.
technique described by Gall et al. (1947, 1949). A 0.01 ml. sample of
the carefully mixed bacterial suspension was transferred by a Breed
pipette to a slide and mixed with a 3 ram. loopful of a s a t u r a t e d methyl
alcohol solution of nigrosine. T h e mixture was spread evenly on the
slide and dried on a hot plate. Counts were made of 20 fields on each
of two slides made from duplicate samples from each period. T h e
bacterial counts, the slide area and volume, the volume of bacterial
suspension and its relation to total feces being known, it was possible
to estimate the total bacteria excreted on each diet.
T h e method used in calculating the analysis of variance is shown
in table 3.

Results and Discussion

T h e average apparent digestion coefficients are shown in table 4. I t
is evident that the added sucrose depressed the digestibility of the crude
TABLE 3. METHOD USED FOR ANALYSIS OF VARIANCE

Total
Pigs
Periods
Treatments, direct
Treatments, residual
Error

d.f.

SS

15
3
3
1
1
7

Computed iu usual way

Computed in usual way
Computed in usual way
( T ~ T b ) 2/16
(R~--R~) 2/12
By difference

Ta=total for treatment A.

T~total for treatment B.
R~=total of the 6 observationspreceded by A.
Rb=total of the 6 observationspreceded by B.

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58

SKIPITARIS, WARNER AND LOOSLI

protein and crude fiber. This depression amounted to approximately

5% and 38% for crude protein and crude fiber, respectively. The differences were highly significant ( P < 0 . 0 1 ) . The marked decrease in
digestibility of crude fiber is probably the result of intestinal bacteria
utilizing the sugar for energy in place of fiber, as occurs in ruminants.
I t is possible that the depression in crude fiber digestibility increased
the loss of protein in the feces because of the less complete digestion of
"fiber-bound" protein. The bacterial population was greater on the
TABLE 4. T H E A V E R A G E A P P A R E N T D I G E S T I O N C O E F F I C I E N T S OF
P R O T E I N AND C R U D E F I B E R OF S W I N E R E C E I V I N G B A R L E Y
W I T H AND W I T H O U T A D D E D SUCROSE
Ration
Barley
Pig. No.

Protein

Crude fiber

1Y
2B
3Y
4B
Av.

76.8
75.6
77.8
76.0
76.6

16.4
15.0
17.8
16.5
16.4

Barley+Sucrose
Protein
73.0
70.8
74.2
72.1
72.5**

Crude fiber
11.0
9,2
10,6
10.0
10,2"*

Corrected
protein ~
74.0
72.4
76.2
74.0
74.2**

a Corrected for lower digestibility of crude fiber.

~ Significantly lower than barley alone (P<O.O1).

sugar ration and these bacteria carried additional nitrogen into the
fecal excretions.
Presented also in table 4 are coefficients for protein digestibility of
the barley plus sucrose ration corrected for the lowered digestibility of
crude fiber. This calculation was based on the assumption that the
increase in fecal crude fiber accompanying sucrose feeding, resulted in
a proportionate increase in fecal protein. The corrected protein digestion
coefficient thus reflects the maximum theoretical amount of fecal protein
which could have resulted from the increased excretion of "fiberbound" protein. The corrected digestion coefficients remain lower than
those for the protein in barmy fed alone, supporting the view that
increased bacterial growth on the barley plus sucrose effected a lowered
apparent digestibility of dietary nitrogen by the pig.
The average crude protein contents of the various fecal fractions are
shown in table 5. For each pig, the feces contained more nitrogen when
the ration contained added sucrose. The differences were highly significant ( P < 0 . 0 1 ) for each fraction of the fecal crude protein as well
as for the total fecal protein. I t is possible that a portion of the residual

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59

SUGAR AND DIGESTIBILITY OF PROTEIN AND FIBER

and soluble protein may have been the result of an increase in metabolic
fecal nitrogen as a result of sucrose feeding. The highly soluble nature
of the sucrose would suggest that this effect does not exist.
Bacterial counts made on the fecal samples by the nigrosine slide
technique gave larger total numbers of bacteria excreted on the diets
containing sucrose than on barley alone. The average values were
156 X 1012 on barley plus sugar and 126 X 101~ on barley. In every
instance the feces contained more bacteria when sugar was fed. Although
the counting techniques may be less accurate than one might desire it
T A B L E 5. A V E R A G E C R U D E P R O T E I N C O N T E N T OF F E C A L F R A C T I O N S
OF PIGS R E C E I V I N G R A T I O N S W I T H A N D W I T H O U T
ADDED SUCROSE
Crude protein fraction ~
Pig No.

Ration

1Y

Barley
Barley+sugar
Barley
Barley+sugar
Barley
Barley+sugar
Barley
Barley+sugar
Barley
Barley+sugar

2B
3Y
4B
Av.

Residue

Total
protein ~

Bacterial

Soluble

gm.

gm.

gm.

gm.

80
86
74
82
90
102
80
84
81
88

144
166
132
156
136
150
136
154
137
157

94
118
96
120
74
100
79
102
86
110

318
370
302
358
300
352
295
340
303
355

a NX6.25.

is interesting that the total bacterial counts support the other data in
suggesting that the addition of 16 3 of sugar to barley increases the
fecal bacteria and at the same time lowers utilization of both the crude
protein and the crude fiber.
I t is evident from the data that the addition of sugar to the ration of
pigs lowers the digestibility of both the protein and the crude fiber in
about the same way as is true for ruminants. The decreases in the
digestion coefficients, appear to be similar in magnitude to those reported
for ruminants. Briggs and Helter (1943) observed that 25~o of cane
molasses added to a ration for sheep lowered the digestibility of the
crude protein by 6.5%. Hamilton (1942), using sheep, obtained depressions of 12 3 for crude protein and 27 3 for crude fiber in a ration
containing up to 30 3 of glucose.
From a practical standpoint, the depressed digestibility of crude fiber

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60

SI(IPITARIS, WARNER AND LOOSLI

resulting from the addition of sucrose appears to be o.f less importance

in swine rations than in ruminant rations, at least in terms of the
availability of energy, because of the lower content and poorer utilization of the fiber under all conditions. The results suggest that a ration
for growing-fattening swine which contains as much as 16% of sugar
should be higher in protein to compensate for its lowered digestibility.
The depression in digestibility of crude fiber presumably would be of
greatest importance for sows maintained on large amounts of silage
or for pigs on pasture; obviously, the question should be studied.
The statistical analysis of the data showed that the mean square for
carry over effect was negligible. I t can be inferred from this observation
that for a study of digestibility and/or bacterial changes a preliminary
period of 10 days is completely satisfactory and in all probability
longer than is absolutely necessary.
Summary
Studies were conducted with pigs to measure the influence of added
sugar upon the digestibility of the crude protein and the crude fiber of
barley. Adding 16% of sucrose lowered the apparent digestibility of
the crude protein and the crude fiber approximately 5% and 38%,
respectively. The depression of protein digestibility was found to be
greater than could be explained by an increase in the excretion of "fiberbound" protein. Fecal nitrogen fractionation revealed more nitrogen
present as bacteria, as soluble nitrogen and residual nitrogen when
sucrose was added than when barley was fed alone. The depression in
digestibility of protein and crude fiber as the result of feeding sugar
seems to be similar in magnitude to the depressions observed with
ruminants.
L i t e r a t u r e Cited
Beeson, W. M., E. W. Crampton, T. J. Cunha, N. R. Ellis and R. W. Luecke. 1953.
Nutrient requirements for swine. National Research Council Publ. 295.
Washington, D. C.
Briggs, I-L M. and V. G. Heller. 1943. The effect of adding blackstrap molasses,
potassium salts, sucrose and corn syrup to a lamb-fattening ration. J. Agr.
Res. 67:359.
Crampton, E. W. and F. Whiting. 1943. The digestibility of typical eastern Canadian
feeds by market bacon hogs. Sci. Agr. 23:518.
Diaz, F., V. C. Speer, G. C. Ashton, C. H. Liu and D. V. Catron. 1956. Comparison
of refined cane sugar, invert cane sugar and unrefined cane sugar in starter
rations for early weaned pigs. J. Animal Sci. 15:315.

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SUGAR AND DIGESTIBILITY OF PROTEIN AND FIBER

61

Gall, L. S., C. N. Stark and J. K. Loosli. 1947. The isolation and preliminary study
of some physiological characteristics of the predominating flora from the
rumen of cattle and sheep. J. Dairy Sci. 30:891.
Gall, L. S., W. Burroughs, P. Gerlaugh and B. H. Edgington. 1949. Special methods
for rumen bacterial studies in the field. J. Animal Sci. 8:433.
Hamilton, T. S. 1942. The effect of added glucose upon the digestibility of protein
and fiber in rations for sheep. J. Nutr. 23 : 101.
Henderson, C. R. 1955. Personal Communication.
Mattill, H. A. and R. B. Hawk. 1911. A method for the quantitative determination
of fecal bacteria. J. Exp. ivied. 14:433.
Mitchell, H. H. 1942. The evaluation of feeds on the basis of digestible and metabolizable nutrients. J. Animal Sci. 1 : 159.
Skipitaris, C. N. 1953. Digestibility and nutritive value of carob pods in farm
animals. Dept. Animal Hus., University Thessalonike, Greece.
Skipitaris, C. N. 1956. The effect of added soluble carbohydrate upon the digestibility of protein and fiber in rations for swine. Ph.D. Thesis. Cornell University,
Ithaca, N. u

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