Published December 5, 2014
Absorption of carbohydrate-derived nutrients in sows as influenced
by types and contents of dietary fiber1
A. Serena,2 H. Jørgensen, and K. E. Bach Knudsen3
Department of Animal Health, Welfare and Nutrition, Faculty of Agricultural Sciences,
University of Aarhus, DK-8830 Tjele, Denmark
ABSTRACT: The current investigation was under-
taken to study the absorption and plasma concentration
of carbohydrate-derived nutrients [glucose, short-chain
fatty acids (SCFA), and lactate] and the apparent insu-
lin production in sows fed diets containing contrasting
types and contents of dietary fiber. Six sows were fed
3 experimental diets, low fiber (LF; 177 g of dietary
fiber and 44 g of soluble fiber/kg of DM), high soluble
fiber (HF-S; 429 g of dietary fiber and 111 g of soluble
fiber/kg of DM), and high insoluble fiber (HF-I; 455
g of dietary fiber and 74 g of soluble fiber/kg of DM),
in a repeated crossover design. Variations in dietary
concentration and solubility of dietary fiber were ob-
tained by substituting starch-rich wheat and barley in
the LF diet with dietary fiber-rich co-products (sugar
beet pulp, potato pulp, pectin residue, brewers spent
grain, pea hulls, and seed residue, which have distinct
physicochemical properties). The main carbohydrate
Key words: absorption, catheterized sow, glucose, insulin, short-chain fatty acid
©2009 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2009. 87:136–147
INTRODUCTION
Sugars, oligosaccharides, starch, and nonstarch poly-
saccharides (NSP) are the main organic constituents
of feeds for pigs. Because various carbohydrate compo-
nents are digested and absorbed at different sites and
rates in the gastrointestinal tract (Bach Knudsen, 2005;
Bach Knudsen et al., 2006), differences in the dietary
carbohydrate composition can be used proactively to
modulate the uptake of carbohydrate-derived nutrients,
1
Fiber-rich co-products were donated by the following Danish
companies: KMC, Kartoffelmelcentralen amba (Brande); Danisco
Sugar A/S (Assens); CPKelco ApS (Lille Skensved); Agro-korn A/S
(Videbæk); DLF Trifolium A/S (Roskilde); and Prodana Seeds A/S
(Odense). Technical personnel at the laboratory and the animal unit
are acknowledged for excellent technical assistance.
2
Present address: Arla Foods Amba, Skanderborgvej 277, 8260
Viby J, Denmark.
3
Corresponding author: knuderik.bachknudsen@agrsci.dk
Received November 7, 2007.
Accepted July 28, 2008.
136
component of the LF diet was starch and nonstarch
polysaccharides (cellulose and noncellulosic polysaccha-
rides) for the 2 high dietary fiber diets. Consumption
of the LF diet resulted in increased and rapid glucose
absorption at 0 to 4 h postfeeding. With the HF-I diet,
the glucose absorption pattern was similar but at a
decreased rate, whereas it was decreased and delayed
with the HF-S diet (diet, P < 0.001; time, P < 0.001).
These differences were also reflected in the insulin re-
sponse. The quantitative absorption of SCFA at 0 to
10 h postfeeding was greater when feeding the HF-S
diet compared with the LF diet (P < 0.001) and in-
termediate when feeding the HF-I diet (P < 0.001).
The study showed that feeding the high dietary fiber
diets resulted in a increased and more uniform uptake
of SCFA than when feeding the LF control. Moreover,
the HF-S diet reduced diurnal variation in glucose and
insulin concentrations.
doi:10.2527/jas.2007-0714
glucose, short-chain fatty acids (SCFA), and lactate
(LA), and thereby affect the feeling of satiety and be-
havior of the animals (Brouns et al., 1997; Danielsen
and Vestergaard, 2001; de Leeuw and Ekkel, 2004; de
Leeuw et al., 2005b).
The chemical composition, glycosidic linkages, and
cross-linkages of polysaccharides in the dietary fiber
(DF; NSP + lignin) matrix have a profound effect
on the physicochemical properties of the feed (Bach
Knudsen, 2001; Serena and Bach Knudsen, 2007a). For
instance, soluble DF may raise luminal viscosity and in-
crease the water-binding capacity (WBC) of digesta in
the small intestine (Canibe and Bach Knudsen, 2002),
thereby slowing the movement of digesta and the rate
of glucose absorption (Holt et al., 1979; Ellis et al.,
1995). Insoluble DF, on the other hand, has relatively
little influence on events in the stomach and small in-
testine (Low et al., 1986; Rainbird and Low, 1986), but
the chemical and structural composition and degree of
lignification of the DF will have a profound influence on
the fermentation in the large intestine (Bach Knudsen,
 Absorption of nutrients in sows 137
2001) and the rate of SCFA absorption (Bach Knudsen, Table 1. Composition of experimental diets, low fiber
2005). (LF), high soluble fiber (HF-S), and high insoluble fi-
It is hypothesized that substitution of starch for ber (HF-I)
DF-rich feedstuffs will result in a shift in the nature
Ingredient, g/kg (as-fed basis) LF HF-S HF-I
and diurnal variation of energy that is provided to the
animal. The main purpose of the present investigation Barley 420 145 145
was to study the qualitative and quantitative aspects Wheat 420 145 145
Sugar beet pulp — 140 50
of carbohydrate assimilation in catheterized sows fed
Pectin residue — 140 50
3 experimental diets varying in types and contents of Potato pulp — 140 50
DF. Variations in DF contents and physicochemical Seed residue — — 135
properties were obtained by substituting starch-rich Pea hulls — — 135
wheat and barley in the control diet with DF-rich sugar Brewers spent grain — — 135
Soybean oil 50 50 50
beet pulp, potato pulp, pectin residue, brewers spent
Soybean meal (toasted) 75 216 84
grain, pea hulls, and seed residue, which have distinct Marker (chromic oxide) 2 2 2
physicochemical properties (Serena and Bach Knudsen, Ca(H2PO4)2 22 11 11
2007a). CaCO3 6 6 3
NaCl 3 3 3
Vitamin-mineral mix1 2 2 2
MATERIALS AND METHODS 1
Provided per kilogram of final diet: 8,800 IU of vitamin A as retinol
The experiment protocol complied with the guide- acetate; 1,000 IU if vitamin D3 as cholecalciferol; 60 mg of all rac dl-
α-tocopherol acetate; 2.2 mg of menadione; 2.2 mg of thiamine; 5.5 mg
lines of The Danish Animal Experiments Inspectorate, of riboflavin; 3.3 mg of pyridoxine; 16.5 mg of d-pantothenic acid; 22
Ministry of Justice, Copenhagen, Denmark, concerning mg of niacin; 1.65 mg of folic acid; 220 µg of biotin; 22 µg of cyanoco-
animal experimentation and care of the animals under balamin; 60 mg of BHT; 100 mg of Fe as FeSO4∙7H2O; 150 mg of Zn as
ZnO; 28 mg of Mn as MnO; 20 mg of Cu as CuSO4∙5H2O; 304 µg of I
study. as KI; and 300 µg of Se as Na2SeO3.

Diets
The diets (Table 1) were prepared from wheat and
barley supplemented with different co-products from
the Danish vegetable food and agricultural industries
[potato pulp, KMC (Brande, Denmark); sugar beet
pulp, Danisco Sugar A/S (Assens, Denmark); pectin
residue, CPKelco ApS (Lille Skensved, Denmark);
brewers spent grain, Agro-korn A/S (Videbæk, Den-
mark); pea hulls and seed residue, DLF Trifolium A/S
(Roskilde, Denmark)]. The variation in the chemical
and physicochemical composition of these co-products
collected over a 2-yr period has been published recent-
ly (Serena and Bach Knudsen, 2007a). The diets were
formulated to contain different contents and types of
DF. A low-DF diet (LF; 177 g of DF/kg of DM) was
prepared from wheat and barley as the main carbohy-
drate source, and 2 high-DF diets (approximately 440
g of DF/kg of DM) were prepared by substituting the
wheat and barley with sugar beet pulp, potato pulp,
and pectin residue (HF-S) and with approximately
one-third sugar beet pulp, potato pulp, and pectin resi-
due and two-thirds brewers spent grain, pea hulls, and
seed residue (HF-I). The diets were formulated to meet
the Danish recommendations for essential macro- and
micronutrients (Jørgensen and Tybirk, 2005) and were
milled to pass through a 2-mm screen. In our diet for-
mulation, proper adjustment was made to ensure a suf-
ficient supply of digestible AA, which was the reason
for the greater protein concentration of the 2 high-DF
diets compared with the LF diet (Table 2). The amount
of carbohydrates was greatest in the LF diet (698 g/kg
of DM) and similar in the HF-S and HF-I diets (628
to 622 g/kg of DM). Diet LF had the greatest starch
content, whereas it was much less in the 2 high-DF
diets. The 2 high-DF diets were similar in total NSP
(367 to 369 g/kg of DM) and similar in cellulose and
noncellulosic polysaccharides (NCP), but with vari-
able proportions of soluble-to-insoluble NCP (30:70 in
the HF-S diet and 20:80 in the HF-I diet). The different
chemical compositions translated into differences in the
physicochemical properties of the diets. Swelling and
WBC were greatest in the HF-S diet and least in the
LF diet.
Animal Experiment
Measurement of the quantitative absorption of nu-
trients was carried out as a 3 × 3 repeated crossover
design, with 6 sows fed the 3 different diets (i.e., LF,
HF-S, and HF-I, for 7 d per diet period). Nonpregnant
sows with an initial average BW of 202 ± 28 kg were
selected after weaning their first litter (Peter Bøjlesen,
Vammen, Denmark). After 10 d of adaptation, each
sow was surgically fitted with 2 catheters. The first
was placed in the portal vein (1.2 mm i.d. and 2.3 mm
o.d.; Cole-Parmer, Vernon Hills, IL), and the second
was placed in the mesenteric artery (1.0 mm i.d. and
1.8 mm o.d.; Cole-Parmer), and also with an ultra-
sonic blood flow probe (28A probe, 28 mm; Transonic
System Inc., Ithaca, NY) around the portal vein. A
flowmeter (Transonic T206 flowmeter with P-option;
Transonic System Inc.) was used to measure the blood
flow rate. After a 7-d recovery period from the surgery,
the sows were introduced to 1 of the 3 experimental
diets. During the experimental period, the sows were
fed once daily at 1000 h (leftover feed was removed af-
138 Serena et al.

Table 2. Chemical composition and physicochemical properties of experimental diets,

low fiber (LF), high soluble fiber (HF-S), and high insoluble fiber (HF-I)
Item LF HF-S HF-I

Chemical composition, g/kg of DM

  Ash 57 65 76
  Protein (N × 6.25) 138 178 170
  Fat 84 86 94
  Total carbohydrates 698 628 622
   Sugars 19 36 24
   Glucose 2 2 3
   Fructose 1 2 2
   Sucrose 16 26 13
   Fructan 9 7 6
   Starch 517 216 225
   Total nonstarch polysaccharides1 144 (44) 369 (111) 367 (74)
    Cellulose 32 151 160
    Noncellulosic polysaccharides 112 (44) 218 (111) 207 (74)
  Arabinose 23 43 38
  Xylose 39 29 60
  Mannose 4 13 7
  Galactose 7 47 22
  Glucose 30 21 44
  Uronic acid 7 63 35
  Klason lignin 33 60 88
  Dietary fiber (lignin + nonstarch polysaccharides) 177 (44) 429 (111) 455 (74)
Physicochemical properties
  Swelling, L/kg of DM 4.13 6.43 5.44
  Water-binding capacity, kg/kg of DM 1.34 3.14 2.58
1
Values in parentheses are soluble nonstarch polysaccharides.

ter 45 min). The daily feeding level was 2,000 g of feed plasma from the EDTA-heparinized tubes was used for
(as-fed basis), and the sows had free access to water analysis of LA. On the days of blood sampling, any feed
(Table 3). Sows were placed in farrowing pens during that remained was collected.
the collection of blood from the portal vein and mesen-
teric artery. Blood samples were collected on the last Analytical Methods
day of each period at −120, −60, 0, 15, 30, 45, 60, 90,
and 120 min after feeding, and at 60-min intervals up All chemical analyses were performed in duplicate
to 600 min. Blood flow in the portal vein was measured on freeze-dried materials. The DM was measured by
at the same time as blood collection as a mean of 4 drying to a constant weight (mostly 20 h) at 103°C,
readings in 1 min, and hematocrit was measured at and ash was analyzed according to the AOAC (1990)
−60 and 360 min after feeding. The blood was collected method. Nitrogen was determined by the Kjeldahl
into 2 Na-heparinized plastic tubes (9 and 4 mL) and method (AOAC, 1990), and protein was calculated as
1 EDTA-heparinized plastic tube (2 mL; Greiner Bio- N × 6.25. Fat was extracted with diethyl ether after
One GmbH, Kremsmünster, Austria) and centrifuged acid hydrolysis and analyzed as described by Stoldt
(1,800 × g for 10 min at 8°C). Plasma was frozen until (1952). The diets were analyzed for sugars (glucose,
further analysis. The plasma from the Na-heparinized fructose, and sucrose) and fructans as described by
tubes was analyzed for glucose, SCFA, and insulin. The Larsson and Bengtsson (1983), and starch and NSP
as described by Bach Knudsen (1997). Total NSP was
divided into cellulose, soluble NCP, and insoluble NCP
Table 3. Daily intake of feed and energy (units/d) by their constituent sugars by GLC for neutral sug-
in sows fed experimental diets, low fiber (LF), high ars and by colorimetry for uronic acid (Bach Knudsen,
soluble fiber (HF-S), or high insoluble fiber (HF-I) 1997). Klason lignin was measured as the sulfuric acid-
Diet LF HF-S HF-I
insoluble residue as described by Theander and Åman
(1979). The diets were analyzed for WBC and viscosity
Feed, g 2,000 2,000 2,000 as described by Canibe and Bach Knudsen (2002) and
DM, g 1,820 1,840 1,840
Johansen et al. (1997), respectively. Briefly, the proce-
GE,1 MJ 31.9b 33.2a 33.1a
DE,1 MJ 26.2a 26.0a 22.3b dure for swelling was as follows: 300 mg of sample was
ME,1 MJ 24.9a 24.0a 20.7b weighed into a 15-mL conical centrifuge tube with 0.1-
a,b
Within a row, means without a common superscript letter differ
mL divisions, dissolved in 10 mL of 0.9% NaCl:0.02%
(P < 0.05). NaN3 and placed in a shaking water bath (150 move-
1
Data from Serena et al. (2008a). ments/min) for 20 h at 39°C. The swelling capacity (L/
 Absorption of nutrients in sows
kg of DM) was measured by reading the volume that
fiber occupied 1 h after removing the water bath.
Plasma was analyzed for SCFA (acetate, propionate,
butyrate, valerate, caprionate, isovalerate, and isobu-
tyrate) essentially as described by Brighenti (1998) by
using 2-ethyl butyrate (Fluka No. 03190; Sigma Al-
drich, St. Louis, MO), rather than isovaleric acid, as an
internal standard. The LA was measured according to
the method described by Noll (1984), and insulin was
analyzed as described by Løvendahl and Purup (2002).
Glucose concentrations in plasma were analyzed by a
glucose-oxidase kit (Trinder, 1969).
Calculations and Statistical Analysis
Absorption of glucose, LA, and SCFA into the portal
vein and apparent production of insulin in the portal
vein were calculated from the porto-arterial differences
and the portal flow measurement as described by Rérat
et al. (1984):
q = (C p - C a ) ´ F (dt ), and Pearson product-moment correlation coefficient by us-
tn
Q = å q,
t0
where q is the amount of glucose, LA, or SCFA ab-
sorbed and the amount of insulin produced within the
time period dt, Cp is the concentration in the portal
vein, Ca is the concentration in the mesenteric artery,
F is blood flow in the portal vein, and Q is the amount
absorbed from t0 to tn or the amount of insulin pro-
duced from t0 to tn. The calculated insulin production
can only be described as apparent because insulin has a
pulsatile secretion that is broken down by the liver and
kidneys and that has a variable half-life value (10 to 30
min). The energy coefficients reported by Weast et al.
(1984) were used when converting millimoles to energy
for glucose, LA, and the SCFA mix.
Differences between diets and time in concentrations
(mesenteric artery and portal vein) of glucose, LA,
SCFA (acetate, propionate, butyrate, valerate, capri-
onate), and branched-chain fatty acids (isobutyrate
and isovalerate), and absorption of glucose, LA, and
SCFA were analyzed as repeated measurements by us-
ing PROC MIXED (SAS Inst. Inc., Cary, NC), with a
level of significance at P ≤ 0.05:
Xijkl = μ + αi+ βj + νk + γl + (αγ)il + εijkl,
where Xijkl is the dependent variable, μ is the overall
mean, αi is the diet (i = 1, 2, or 3), βj is the week (j
= 1, 2, or 3), νk is the sow (k = 1 to 6), γl is time (l
= 1 to 17), (αγ)il is the 2-factorial interaction between
diet and time, and εijk is the error term. If P > 0.05 for
the interaction between diet and time, (αγ)il was taken
out of the model. Data on insulin concentrations in the
139
mesenteric artery and portal vein were transformed to
log(x) before the statistical analysis to obtain variance
homogeneity.
For analyzing the effect of diet and time on the ab-
sorption of glucose, LA, and SCFA and the apparent
production of insulin before feeding in the absorptive
phase and in the postabsorptive phase, the data set
was divided into 3 groups (before feeding, −2 to 0 h;
absorptive phase, 0 to 4 h; and postabsorptive phase, 4
to 10 h) and the 3 data sets were analyzed as repeated
measurements by using PROC MIXED of SAS, with
the level of significance at P ≤ 0.05:
Xijk = μ + αi+ βj + νk + (αβ)ij + εijkl,
where Xijk is the dependent variable, μ is the overall
mean, αi is the diet (i = 1, 2, or 3), βj is the period (j
= 1, 2, or 3), νk is the sow (k = 1 to 6), and (αβ)ij is
the 2-factorial interaction between diet and period. The
correlation between total apparent glucose absorption
and apparent insulin production was calculated as the
ing PROC CORR of SAS.
RESULTS
To obtain a fixed reference point for the measurements
of postprandial changes in nutrients in the blood, the
eating period was initially restricted to 45 min. Only 1
sow fed the HF-I diet did not consume all the allowance
on 1 of the sampling days.
There were no differences in blood flow among sows
fed the 3 diets (P = 0.26; Table 4). Mean blood flow for
all diets was 19.8 mL/(kg of BW∙min). Blood flow was
3.4 L/min before feeding, increasing to 4.0 L/min in the
absorptive phase and to 4.2 L/min in the postabsorp-
tive phase (data not shown).
The concentrations of glucose and insulin in the por-
tal vein and in the mesenteric artery are shown in Table
4. There was a rapid postprandial increase in glucose
concentration in the portal vein when feeding the LF
diet (Figure 1A). Glucose concentration increased from
3.4 mmol/L at feeding (time 0 h) to 7.0 mmol/L after
1 h, and slowly decreased to 4.5 mmol/L in the portal
vein 10 h after feeding. When feeding the HF-I diet,
there was also a rapid response in the portal vein 1 h
after feeding and the same descending pattern in glu-
cose concentration up to 10 h after feeding as observed
with the LF diet. However, when feeding the HF-S diet,
a decreased and delayed glucose response in the portal
vein was observed. The concentration of glucose in the
portal vein throughout the day was also more stable
when feeding the HF-S diet compared with the other
diets. However, in the mesenteric artery, the concen-
tration was similar when sows were fed the 3 different
diets.
The absorption (mmol/h) of glucose was greater
when feeding the LF diet than when feeding the high-
DF diets (Table 4). Before feeding, no differences were
 140

Table 4. Plasma concentrations (0 to 10 h postfeeding) in the mesenteric artery and portal vein, blood flow in the portal vein, and absorption in sows fed
low fiber (LF), high soluble fiber (HF-S), or high insoluble fiber (HF-I)1
Mesenteric artery Portal vein

P-value P-value

Item LF HF-S HF-I SEM Diet Time D × T2 LF HF-S HF-I SEM Diet Time D × T2

Flow, L/min — — — — — — — 3.97 4.09 3.92 0.25 0.26 0.004 —

Glucose, mmol/L 3.70 3.46 3.55 0.27 <0.001 <0.001 0.002 5.42a 3.96b 4.34b 0.36 0.009 <0.001 —
  Absorption, mmol/h — — — — — — — 419 124 189 21 0.001 <0.001 0.02
Insulin, pmol/L 73 42 50 5.01 0.05 <0.001 0.03 138a 87b 98b 9.6 0.01 <0.001 —
  Apparent production, nmol/h — — — — — — — 16.2 11.7 11.7 11 0.46 0.005 —
Lactate, mmol/L 0.56 0.57 0.55 0.05 0.43 0.001 — 0.74a 0.72b 0.74a 0.06 0.02 <0.001 —
  Absorption, mmol/h — — — — — — — 49 44 43 2.8 0.33 <0.001 —
SCFA,3 µmol/L 223 512 432 33 <0.001 <0.001 0.03 798c 1,770a 1,416b 111 <0.001 0.01 —
  Acetate, µmol/L 205 483 409 31 <0.001 <0.001 0.03 517c 1,243a 1,022b 7.7 <0.001 0.01 —
Serena et al.

  Propionate, µmol/L 6.2c 8.4a 7.1b 0.59 <0.001 0.07 — 180c 335a 260b 22 <0.001 0.02 —
  Butyrate, µmol/L 5.3c 12a 9.7b 0.79 <0.001 0.11 — 63b 126a 85b 7.7 <0.001 0.001 —
  Valerate, µmol/L 0.7c 0.9a 0.9b 0.06 <0.001 0.36 — 14b 23a 18b 1.6 <0.001 <0.001 —
  Caprionate, µmol/L 3.3 3.7 3.1 0.28 0.37 <0.001 — 5.8b 11a 8.4b 0.72 0.006 0.13 —
BCFA,4 µmol/L 2.4c 4.0a 3.1b 0.27 <0.001 <0.001 — 17c 31a 22b 2.0 <0.001 0.09 —
  Isobutyrate, µmol/L 1.0c 1.9a 1.5b 0.12 <0.001 0.008 — 6.5b 12a 8.9b 0.75 <0.001 0.09 —
  Isovalerate, µmol/L 1.2c 2.2a 1.6b 0.14 <0.001 0.006 — 11c 19a 13b 1.2 <0.001 0.09 —
  Absorption of SCFA, mmol/h — — — — — — — 133c 321a 218b 13.7 <0.001 0.04 —
a–c
Within a row, means without a common superscript letter differ (P < 0.05).
1
Values are means; n = 6.
2
D × T = diet × time.
3
SCFA = short-chain fatty acids.
4
BCFA = branched-chain fatty acids.
 Absorption of nutrients in sows 141

Figure 1. Portal and arterial concentrations (A) or absorption (B) of glucose in sows fed diets containing low fiber (LF), high soluble fiber
(HF-S), or high insoluble fiber (HF-I).Values are means ± SEM; n = 6. P-values (absorption): (−2 to 0 h) diet, P = 0.07, and time, P = 0.5; (0
to 4 h) diet, P < 0.001, and time, P < 0.001; (4 to 10 h) diet, P < 0.001, and time, P = 0.14. Mesenteric artery (– – –); portal vein (——); LF
(–○–); HF-S (–■–); HF-I (–□–).

observed among the diets, whereas feeding the LF diet
led to a greater absorption of glucose in the absorptive
and postabsorptive phases compared with feeding the 2
high-DF diets (P < 0.001; Figure 1B). In the absorptive
phase, glucose absorption was delayed when feeding the
HF-S diet compared with feeding the LF and HF-I di-
ets (P < 0.001). The portal glucose absorption relative
to the intake of starch and sugars (% of intake) was less
in the absorptive and postabsorptive phases when feed-
ing the HF-S diet (26 and 17%, respectively) compared
with the HF-I (33 and 27%, respectively) and LF (34
and 34%, respectively) diets.
The concentration of insulin in the portal vein mim-
icked glucose to a large extent (Figure 2A): a rapid
response when feeding the LF diet, a decreased and less
attenuated response when feeding the HF-I diet, and
a more decreased and less attenuated response when
feeding the HF-S diet. Four hours after feeding, insulin
concentration in the portal vein was similar for all 3
diets (70 to 80 pmol/L), and this persisted up to 10 h
postfeeding. The apparent insulin production (Figure
2B) at 0 to 10 h postfeeding was not different among
the 3 diets (Table 4). The total apparent insulin produc-
tion (0 to 10 h after feeding) was correlated (r = 0.75,
P = 0.009) with the total apparent glucose absorption
(0 to 10 h after feeding). In the absorptive phase, the
correlation was greater when feeding the HF-S diet (r =
0.96, P < 0.001) than when feeding the other diets (LF:
r = 0.78, P = 0.01; HF-I: r = 0.69, P = 0.04).
Feeding the HF-S diet decreased portal vein con-
centration of LA, whereas there was no difference in
either mesenteric artery concentrations of LA or the
142 Serena et al.

absorption of LA at 0 to 10 h postfeeding (Table 4).
The absorption of LA decreased from approximately
57 mmol/h in the absorptive period (diet, P = 0.96) to
approximately 37 mmol/h in the postabsorptive period
(diet, P = 0.54). The statistical analyses revealed that
there was an effect of time (P = 0.002) only in the ab-
sorptive phase.
The 3 diets established 3 different concentrations of
SCFA in the portal vein: least for the LF diet, interme-
diate for the HF-I diet, and greatest for the HF-S diet
(Figure 3A). These concentration differences were also
observed in the mesenteric artery for all acids except
caprionate (Table 4). The absorption of SCFA (Figure
3B) was consistently greater before and after feeding
when sows were consuming the 2 high-DF diets (diet,
P < 0.001, 0.001, and 0.002 for before feeding, in the
absorptive phase, and in the postabsorptive phase, re-
spectively). However, there was an effect of time only
in the absorptive phase (time, P = 0.04). The total ab-
sorption of SCFA at 0 to 10 h was greater when feeding
the HF-S diet compared with feeding the LF diet, with
the HF-I diet being between the 2 diets (Table 4).
The apparent uptake of energy (MJ/h) from glucose
and SCFA varied considerably among diets in the 3

Figure 2. Portal and arterial concentrations (A) or production (B) of insulin in sows fed diets containing low fiber (LF), high soluble fiber
(HF-S), or high insoluble fiber (HF-I).Values are means ± SEM; n = 6. P-values (apparent production): (−2 to 0 h) diet, P = 0.49, and time,
P = 0.55; (0 to 4 h) diet, P = 0.09, and time, P < 0.001; (4 to 10 h) diet, P = 0.66, and time, P = 0.66. Mesenteric artery (– – –); portal vein
(——); LF (–○–); HF-S (–■–); HF-I (–□–).
 Absorption of nutrients in sows 143

Figure 3. Portal and arterial concentrations (A) or absorption (B) of total short-chain fatty acids in sows fed diets containing low fiber (LF),
high soluble fiber (HF-S), or high insoluble fiber (HF-I). Values are means ± SEM; n = 6. P-values (absorption): (−2 to 0 h) diet, P < 0.001, and
time, P = 0.88; (0 to 4 h) diet, P < 0.001, and time, P = 0.04; (4 to 10 h) diet, P = 0.002, and time, P = 0.24. Mesenteric artery (– – –); portal
vein (——); LF (–○–); HF-S (–■–); HF-I (–□–).

periods (before feeding, absorptive phase, and postab-
sorptive phase), whereas for LA, there was no diet ef-
fect (Table 5). A period effect was further observed for
glucose and LA, but not for SCFA, which was relatively
constant at all sampling periods. Whereas the uptake
of total energy varied by a factor of 2.3 from before
feeding to the absorption phase for the HF-S diet, the
ratio was 7.3 for the LF diet, with the HF-I diet being
between the 2 diets (P < 0.001). Collectively, the ap-
parent uptake of energy (0 to 10 h postfeeding) from
carbohydrate-derived nutrients was 14.5, 8.2, and 8.3
MJ in the LF, HF-S, and HF-I diets, respectively (Ta-
ble 6). Although the intake of ME was similar for sows
fed the LF and HF-S diets, the accumulated uptake of
144 Serena et al.

glucose was only one-third when feeding the HF-S diet

Table 5. Apparent absorption of energy (MJ/h) derived from glucose, lactate, and short-chain fatty acids (SCFA) in the different periods (before feeding,

D×P
compared with the LF diet, whereas the energy from

0.32
0.98
0.75
0.75
SCFA was more than twice as high with the HF-S com-
pared with the LF diet.

<0.001
<0.001

<0.001
P-value2

Period

0.09
DISCUSSION

The current investigation was designed to study the

<0.001

<0.001
<0.001
0.53
Diet
qualitative and quantitative absorption of products de-

absorptive phase, and postabsorptive phase) in sows fed low fiber (LF), high soluble fiber (HF-S), or high insoluble fiber (HF-I)1
riving from carbohydrate assimilation in sows fed di-
ets that varied in types and contents of DF. To our
knowledge, this kind of study has never been carried

 
 
out with sows before; therefore, we do not have com-

0.151
0.006
0.048
0.135
SEM
parable literature data regarding the flow rate in the
portal vein. In the present study, the portal flow rate in

Postabsorptive phase
sows [19.8 mL/(kg of BW∙min)] was greater in absolute

0.288b
0.718b
0.044
0.386
HF-I
terms, but less than that shown for growing pigs [33.0
to 38.8 mL/(kg of BW∙min)] when using ultrasonic flow
probe techniques as in the current study (Ellis et al.,

0.713b
0.456a
HF-S

0.046
0.233
1995; Bach Knudsen et al., 2000, 2005), the electro-
magnetic flow probe technique [40 to 42.5 mL/(kg of
BW∙min); Rérat et al., 1984], or dye [31.7 to 36.5 mL/
(kg of BW∙min); van der Meulen et al., 1997a,b] when

0.182b
a

1.338a
0.057
1.006
LF
expressed per kilogram of BW. The most likely cause
for the reduced blood flow is the decreased proportion
of visceral organs in mature and almost fully grown ani-

 
 
mals compared with young growing animals (Doornen-

0.012
0.244

0.022
0.246
SEM
bal and Tong, 1981).
The most noticeable differences among diets were the
contents of starch and the types, contents, and degrees

0.227ab
of lignification of the polysaccharides that made up the
b

0.999b
Absorptive phase

HF-I

0.067
0.705
DF fraction. Although the concentrations of NSP, cel-
lulose, and NCP were practically the same in the 2
high-DF diets, the proportions of soluble and insoluble
b

0.935b
0.303a
HF-S

0.084
0.548

NCP varied substantially, which was related to the dif-

ferences in the DF composition of the raw materials in Within a row, means without a common superscript letter differ (P < 0.05).
the high-DF diets (Serena and Bach Knudsen, 2007a).
The greater solubility of DF in the HF-S diet was re-
0.148b
a

1.751a
1.519
0.084
LF

sponsible for the greater WBC and swelling compared

with the DF in the HF-I diet, whereas the high de-
gree of lignification of the HF-I diet was responsible
 
 

for the decreased total tract digestibility of energy for

0.024
0.004

0.031
0.024
SEM

this diet compared with the other 2 diets (Serena et al.,

2008b).
The contrasting starch contents and chemical and
0.274a

physicochemical properties of DF in the experimental

0.011
0.007

0.292
HF-I
Before feeding

diets were key factors regulating the rate at which glu-

cose was absorbed into the portal vein. The glucose
absorption pattern in the absorptive phase when feed-
0.326a
HF-S

0.068
0.014

0.408

ing the LF diet was increased and was similar to that

Values are means; n = 6.

shown for growing pigs fed a concentrated diet (Ellis

D × P = diet × period.

et al., 1995; van der Meulen et al., 1997b; Bach Knud-

0.130b

sen et al., 2000, 2005). This was caused by the open

0.106
0.003

0.239
LF

structure of cereal starch, which enabled easy access

for the salivary and pancreatic α-amylases in the small
intestine (Englyst et al., 1992; Gallant et al., 1992;
Glucose
Lactate

Lang et al., 1999). For the HF-I diet, the absorption

Total 
SCFA
Item

a,b

profile resembled that of the LF diet, but in quantita-

1
2
 Absorption of nutrients in sows
Table 6. Apparent energy absorption 0 to 10 h postfeeding in sows fed low fiber (LF),
high soluble fiber (HF-S), or high insoluble fiber (HF-I)1
Item LF
a 2
Glucose, MJ 12.11 (84)
Lactate, MJ 0.68 (5)
SCFA,3 MJ 1.68c (11)
Total absorbed energy, MJ 14.5a
Absorbed energy, % of ME 58
a–c
Within a row, means without a common superscript letter differ (P < 0.05).
1
Values are means; n = 6.
2
Values in parentheses are percentages of total absorbed energy.
3
SCFA = short-chain fatty acids.
tive terms, the uptake was decreased because of the
decreased starch load. We believe that this was because
the starch in the 2 diets came from the same source
(the cereals) and that insoluble DF present in the HF-I
diet had little, if any, influence on either gastric empty-
ing or the movement of digestion products in the lumen
of the small intestine (Low et al., 1986; Rainbird and
Low, 1986). However, when feeding the HF-S diet, the
portal glucose concentration was decreased and peak
glucose was reached at a later stage than with the 2
other diets. We presume that this was due to delayed
gastric emptying (Rainbird and Low, 1986; Flourié,
1992) and a bulky luminal environment (Serena et al.,
2008a), which may have slowed down amylolysis and
the movement of digestion products (Rainbird et al.,
1984; Low et al., 1986; Rainbird and Low, 1986). How-
ever, a contributing factor could be the potato starch
in the HF-S diet (16% of total starch), with a reduced
prececal digestibility (Sun et al., 2006) and reduced ab-
sorption rate (van der Meulen et al., 1997a).
Our quantitative and qualitative direct measure-
ments of glucose absorption with the HF-S diet con-
firmed the measurements in peripheral blood (jugular
vein) observed by Vestergaard (1997), who compared a
control diet similar to the LF diet with a high-DF diet
with added sugar beet pulp. Whereas other studies fed
growing pigs a concentrated diet supplemented with
highly viscous guar gum (Nunes and Malmlöf, 1992; El-
lis et al., 1995) and documented the importance of the
viscosity of digesta (Flourié, 1992) in regulating glucose
absorption, our study with the HF-S diet pointed more
toward WBC and swelling as regulatory factors in the
HF-S diet.
We showed a greater attenuated insulin response
compared with glucose in the immediate postprandial
period, but otherwise a pattern that was very similar
to that of glucose. This was a reflection of absorbed
glucose being the prime factor regulating the pancre-
atic secretion of insulin (Shils et al., 1994; Ellis et al.,
1995) in the experimental setting. After a large meal,
increased serum insulin concentrations stimulate fuel
and energy storage, followed by a decreased systemic
glucose concentration. A substantial portion of total
insulin secreted from the pancreas is lost during the
first passage through the liver via hepatic extraction.
145
HF-S HF-I SEM
b b
3.59 (44) 5.14 (62) 1.69 (3.5)
0.61 (7) 0.53 (6) 0.07 (0.4)
3.95a (49) 2.64b (32) 0.32 (3.3)
8.2b 8.3b 3.07
34 40  
However, our results concerning the insulin response in
the portal vein and mesenteric artery confirm the jugu-
lar vein insulin responses in sows fed a low-DF diet,
and diets high in soluble and insoluble DF, respectively
(Vestergaard, 1997). It is also clear from the studies of
de Leeuw et al. (2004, 2005a) that fermentable DF has
a stabilizing effect on insulin concentration in sows.
The diets resulted in 3 levels of carbohydrate fer-
mentation in the large intestine (Serena et al., 2008b),
which were directly reflected in the portal concentra-
tions of all SCFA as well as in total SCFA absorption.
For the LF diet, the SCFA concentrations in the portal
vein were comparable with values found in growing-
finishing pigs fed diets with approximately the same
DF level (Giusi-Perier et al., 1989; Bach Knudsen et
al., 2000), whereas the concentration was much greater
when feeding the 2 high-DF diets. For these diets, por-
tal concentrations were far greater than those reported
for growing pigs and were similar to those observed
for sheep (approximately 1.6 mmol/L; Bergman, 1990),
but were less than those observed for cows (approxi-
mately 2.9 mmol/L; Huntington and Reynolds, 1983).
Therefore, these data point to the importance of NSP
as a measure of the fraction of dietary carbohydrates
fermented in the large intestine and the portal and sys-
temic concentrations of SCFA (Bach Knudsen, 2005).
It is also clear from this comparison that the SCFA
concentrations in the portal vein of sows more closely
resembled those of herbivores, which rely primarily on
SCFA as an energy source, than those of pigs fed con-
centrated diets (Bergman, 1990). In spite of the oxida-
tion of SCFA in the various body tissues, the SCFA
concentration ranking in the portal vein would be re-
flected in the peripheral tissues.
Although the polysaccharides in the high-DF diets had
different physiochemical properties and fermentabilities
(Serena et al., 2008a,b), they did not lead to differences
in the SCFA profile. In this aspect, the results of this
study were different from other studies in which one
type of DF polysaccharide was the predominant source
for fermentation. For instance, the fermentation of cel-
lulose stimulates acetate formation (Giusi-Perier et al.,
1989), whereas arabinoxylans and resistant starches are
substrates that stimulate butyrate formation (van der
Meulen et al., 1997a; Bach Knudsen et al., 2005).
146 Serena et al.
Absorbed LA contributed only 5 to 7% of absorbed
carbohydrate-derived nutrients, with virtually no influ-
ence from the dietary composition. As was discussed by
Serena and Bach Knudsen (2007b), this is a reflection
of the absorption profile of LA being better synchro-
nized to that of glucose than that of SCFA.
Because of the substantial oxidation of energy in
splanchnic tissues (Brundin and Wahren, 1991), the
portal appearance of energy could only be considered as
apparent. However, it is evident from this study that a
decreased proportion of the absorbed energy was deliv-
ered to the body during the first 10 h postfeeding when
feeding the high-DF diets compared with the low-DF
diet. It should also be noted that, whereas the portal
flux of starch declined gradually during the late absorp-
tive phase to reach nonsignificant levels in the last 2 h
before feeding, the portal flux of SCFA in the last 2 h
before feeding was at approximately the same level as
10 h after feeding when feeding the 2 high-DF diets.
The 2 high-DF diets altered several factors that are
thought to influence the feeling of satiety. First, substi-
tution of NSP for starch results in a shift in the nature
of absorbed energy from readily absorbed glycogenic
energy to more slowly released SCFA. Second, a direct
consequence of the reduced diurnal variation in ab-
sorbed energy is less fluctuation in insulin. Third, DF-
rich diets resulted in more materials in the stomach and
the remaining gut system (Serena et al., 2008a). Thus,
sows consuming high-DF diets can be expected to be
satiated for a longer period of time, which is regulated
in the short-term by the physical presence of digesta in
the gut, which influences the stretch and chemorecep-
tors in the stomach and duodenum, and in the long run
by metabolic means obtained through the decreased
diurnal variation in energy uptake (Read et al., 1994).
These conditions can be expected to reduce the inci-
dence of aggressiveness, stress, and stereotype behavior
in sows induced by hunger (Brouns et al., 1997; Ramo-
net et al., 1999; Bergeron et al., 2000; Danielsen and
Vestergaard, 2001; Grieshop et al., 2001).
In conclusion, we showed a clear influence of dietary
composition on the rate and types of carbohydrate-de-
rived nutrients to the body. The study showed that DF-
rich diets provided a greater and more uniform SCFA
uptake than did the low-DF diet, and that the diet high
in soluble DF provided a more regulated glucose and
insulin response. It is expected that decreased diurnal
variation in portal energy appearance may counteract
the aggressiveness, stress, and stereotypic behavior
caused by hunger in sows.
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