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Oxidation of nutrients in bull calves

treated with ß‐adrenergic agonists
a a a
A. Chwalibog , K. Jensen & G. Thorbek
a
Department of Animal Science and Animal Health , The
Royal Veterinary and Agricultural University , Copenhagen,
Denmark
Published online: 10 Jan 2009.

To cite this article: A. Chwalibog , K. Jensen & G. Thorbek (1996) Oxidation of nutrients in
bull calves treated with ß‐adrenergic agonists, Archiv für Tierernaehrung, 49:4, 255-261, DOI:
10.1080/17450399609381888

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 Arch. Anim. Nutr., 1996, Vol.49, pp. 255-261 © 1996 OPA (Overseas Publishers Association)
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OXIDATION OF NUTRIENTS IN BULL CALVES

TREATED WITH ß-ADRENERGIC AGONISTS

A. CHWALIBOG, K. JENSEN and G. THORBEK

Department of Animal Science and Animal Health The Royal Veterinary and
Agricultural University, Copenhagen, Denmark
(Received 30 January 1996)
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Oxidation of protein (OXP), carbohydrate (OXCHO) and fat (OXF) was investigated with 12 growing
bulls treated with ß-agonist (L-644, 969) during two 6 weeks trials (Section A and B) at a mean live weight
of 195 and 335 kg. Heat production and nutrient oxidation was calculated from gas exchange, with CO2
reduced for CO2 from fermention processes, and nitrogen excretion in urine.
The ß-agonist had no effect on the level of rumen fermentation as indicated by the same methane pro-
duction for control and treated animals.
Heat Production (HE, RQx) increased by the treatment of ß-agonist corresponding to the increment in
the protein retention.
OXP/HE,RQx was reduced to about 10% in treated animals, indicating that in order to supply amino
acids for an increased protein deposition oxidation of protein is decreased.
OXF/HE,RQx were markedly higher in treated animals, but as indicated by the same CH4 production
the level of the short chain fatty acids (SCFA) production was the same. Therefore, it was concluded that
the increase in OXF was not caused by an increase in SCFA but by a direct influence of ß-agoinst on
mobilization and oxidation of body fat.

KEY WORDS: Bulls: ß-agoinst: Protein oxidation: Carbohydrate oxidation: Fat oxidation: Heat
production

1. INTRODUCTION

The influence of ß-agonist (L-644,969) on protein and fat gain in growing calves was
demostrated in a previous paper (Chwalibog et al, 1995). It was concluded that the
ß-agonist increased protein retention and decreased fat retention. The present paper
deals with the influence of j3-agonist on the oxidation of nutrients.
The total heat production (HE,RQ), calculated from 24 h measurements of O2
consumption, CO2 production and nitrogen excretion in urine, is the sum of heat
produced from oxidation of protein (OXP), carbohydrate (OXCHO) and fat (OXF).
Equations for calculating nutrient oxidation were developed by Brouwer (1957,
1958) and later validated for pigs by Chwalibog et al. (1992). It has been demon-
strated that it is possible to calculate nutrient oxidation from gas exchange measure-
ments when the non-protein respiratory quotient (RQnp) is below or above one.

Prof.Dr.A.Chwalibog, Department of Animal Science and Animal Health, The Royal Veterinary and
Agricultural University, Bülowsvej 13, 1870 Frederiksberg, Denmark.
Supported by the Danish Veterinary and Agricultural Research Council (13-4504) and Merck & Co., Inc.
255
 256 A. CHWALIBOG et al.

However, in ruminants, where a certain part of the measured CO2 is caused by the
fermentative processes and therefore the total amount of CO2 is partly composed
from CO2 produced from nutrient oxidation and partly from nutrient fermentation
in the rumen, the total CO2 production cannot be directly applied in calculations of
OXCHO and OXF. Therefore, in the present paper an attempt has been made to
separate between CO2 from oxidation and fermentation processes in order to evalu-
ate nutrient oxidation in ruminants.
No references concerning an effect of j3-agonist on nutrient oxidation can be
found.

2. MATERIALS AND METHODS

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2.1. Animals, Procedure and Diets

The experiment included 12 Red Danish bull calves measured in 58 individual balance
periods (5 d collection) with 24 hours measurements of gas exchange in each period.
The animals were fed ad libitum with identical feed compounds and measured in two 6
weeks trials at a mean LW of 195 and 335 kg in Section A and B respectively. The
calves were divided into three groups, with no treatment in Group 1 and with addition
of 5 and 10 mg/d of /J-agonist (L-644,969) in Group 2 and 3 respectively in Section A,
increased to 10 and 20 mg/d in treated animals (Group 2 and 3) in Section B.
Measurements were performed by means of nitrogen balance and with use of indirect
calorimetry. Further details of the experiment are given in Chwalibog et al. (1995).
The mean initial LW, intake of metabolizable energy and digestible protein together
with production of CH4 per kg of ingested organic matter (IOM) are shown in Table 1.

Table 1 Live weight (LW), intake of metabolizable energy (ME) and digested protein (DP) in relation to
metabolic live weight (kg0•") and production of methane (CH4) in relation to intake of organic matter
(IOM) (Means ± SEM)
Section A Section B
Group 1 Group 2 Group 3 Group 1 Group 2 Group 3

5 10 12 12
)3-agonist mg/d 0 5 10 0 10 20
LW initial, kg 175 + 6.3 205 + 10.3 188 + 5.4 326±12.6 343 + 28.0 357 + 20.2
ME, kJ/kg 075 1248 + 36.9 1216 + 14.7 1082 + 26.6 1082 + 21.7 1109 + 31.6 1001 ±32.0
DP, g/kg075 12.6 + 0.34 12.2 + 0.20 11.3 + 0.30 11.2 + 0.32 11.0 + 0.54 10.5 + 0.38
CH 4 , I/kg IOM 12.7±0.21 12.3+0.32 13.1+0.36 20.9 + 0.82 19.9 + 1.02 22.1+0.74

2.2. Calculations
The following terminology was used:
HE,RQx: heat production by RQ-method with correction for CO2 from fermenta-
tion
O2np: O2 from oxidation of non-protein nutrients
 /3-AGONIST AND OXIDATION OF NUTRIENTS 257

CO2npx: CO 2 from oxidation of np nutrients (corrected for CO 2 from fermenta-

tion)
RQnpx: respiratory quotient of non-protein nutrients
UN: nitrogen excreted with urine
OXP: heat from oxidized protein
OXCHO: heat from oxidized carbohydrate
OXF: heat from oxidized fat
All calculations were performed with constants and factors in accordance to
Brouwer (1965). It was assumed that in maximum three litres of CO2 are released per
one litre CH 4 produced by fermentation of hexoses into short chain fatty acids
(SCFA) (Fahey and Berger 1988).
Units: HE, OXP, OXCHO and OXF in U; O2, CO2 and CH4 in litres ; UN in grams.
HE,RQx = 16.18 • O2+5.02 • (CO2-3 • CH4) - 5.99 • UN-2.17 • CH4
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CO 2 npx = CO 2 - UN • 6.25 • 0.774 - 3 • CH 4

O2np = O 2 -UN-6.25-0.957
RQnpx = CO2npx/O2np
O X P = U N - 6.25 -18.42
OXCHO = [(RQnp*-0.711)/0.289 • O2np]/0.829 • 17.58
OXF =[CO 2 npx-(RQnpx-0.711)/0.289 • O2np]/1.431 • 39.76
Data are expressed as mean values ± standard error of the mean (SEM).

3. RESULTS

The feed intake, being ad libitum, was not influenced by the addition of /?-agonist as
discussed in the previous paper (Chwalibog et ai, 1995) and summarized in Table 1.
The mean intake of DP in Section A and B was 12.0 and 10.9 g/kg 075 , while the CH 4
production was 12.7 and 21.21 I/kg IOM in Section A and B respectively.
The mean values of heat production and oxidation of nutrients in the different
groups are shown in Table 2.
The mean RQnpx was around 0.96 for all groups in Section A, decreasing in
Section B to the lowest value of 0.88 in Group 3 treated with 20 mg/d of L-644,969.
The mean values of HE,RQx were in both sections significantly higher (P < 0.05) in

Table 2 Non-protein-respiratory quotient (RQnpx), total heat production (HE, RQx) and heat from oxidation
of protein (OXP), carbohydrate (OXCHO) and fat (OXF) in relation to HE, RQx (Means + SEM)
Section A Section B
Group 1 Group 2 Group 3 Group 1 Group 2 Group 3
n 5 10 8 12 6 12
)3-agonist mg/d 0 5 10 0 10 20
RQnpx 0.97 + 0.01 0.95 + 0.01 0.97 + 0.01 0.94 + 0.01 0.90 + 0.02 0.88 + 0.01
HE, RQx, MJ/d 40.5 + 1.86 46.8+1.80 42.1 + 1.15 58.4 + 1.45 66.8 + 3.19 62.7 + 2.01
OXP/HE,RQx,% 10.0 + 0.76 9.0 + 0.42 10.0 + 0.38 14.7+1.11 10.3 + 1.48 11.3 + 0.90
OXCHO/HE,RQ.v,% 82.7 + 0.86 75.6+1.65 80.1 + 1.88 67.4 + 2.06 60.4 + 5.52 53.6 + 4.41
OXE/HE,RQx,% 7.7+1.86 15.6+1.67 10.2 + 2.05 18.4 + 2.05 29.7 + 4.60 35.6 + 3.93
 258 A. CHWALIBOG et al.

Group 2 than in the control group. The heat production from oxidation of protein
in relation to the total heat production (OXP/HE,RQx) was in Section A identical
for all groups (P>0.05) with a mean value of 9.6 + 0.28%. The values in Section B
were identical for the treated calves in Group 2 and 3 (P>0.05), with a mean value
of 11.0 + 0.76% being significantly lower (P< 0.001) than the mean value of
14.7+ 1.11% in the control group.
The heat from the oxidation of carbohydrate in relation to the total heat produc-
tion (OXCHO/HE,RQx) was in Section A between 76-83% and in Section B
between 54-67%, in both sections with the lowest value for the treated calves. The
values of OXF/HE,RQx were in Section A between 8-16% increasing 18-36% in
Section B. The mean value in Section A of 15.6 ± 1.67% in Group 2 was significantly
higher than the control group (P<0.01). In Section B the values were 29.7 + 4.60%
and 35.6 + 3.93% in Group 2 and 3 respectively, the differences being significant
(P < 0.05 and P < 0.001) from the control group.
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4. DISCUSSION

Calculations of nutrient oxidation from gas exchange measurements are based on

the assumption that CO2 produced is caused by oxidation of nutrients. However, in
ruminants the total measured CO2 derives both from nutrient oxidation in the tissue
and nutrient fermentation in the rumen. In an attempt to seperate the origin of CO2
it was assumed that 3 litres of CO2 follwed the production of 1 litre of CH4 by rumen
fermentation, therefore oxidation-CO2 was calculated by reducing the total mea-
sured CO2 with 3 • CH4, 1. This value is a maximum CO2 output per litre of CH4 for
high grain diet, as it was fed in the present investigation, giving an acetate: propio-
nate molar ratio of 1:1, according to the equation (Fahey and Berger, 1988):
3 glucose -> 2 acetate+2 propionate+butyrate+3CO2+CH4+2H2O.
In contrast, a high forage diet might give an acetate : propionate ratio 3:1/5 glu-
cose —> 6 acetate+2 propionate+butyrate+5CO2+3CH4+6H2O, thus with 1.7 mole
CO2 per mole of CH4.
After reducing the total CO2 into CO2x both heat production from nutrient oxida-
tion (HE, RQx) and the partition of nutrient oxidation could be estimated from the
gas exchange and excretion of urinary nitrogen. As heat production is calculated
from protein oxidation and oxidation of non-protein substrates, the classical
assumption that UN corresponds to the amount of oxidized protein can be crucial in
these calculations. Especially in ruminants a part of UN may originate from non-
protein nitrogen (NPN) and thus affecting OXP values and subsequently the propor-
tions OXP,OXCHO and OXF to HE,RQx. However, in the present investigations
the animals were fed only with concentrates and straw and the amount of NPN was
neglible. Furthermore, determination of nitrogen in urine is always attached with
risk of loosing some ammonia during the time of collection. In order to evaluate an
impact of a potential error of UN on the results a hypothetical calculation was
made. In the Table 3 the average values of UN and the corresponding values of
HE,RQx and OXP are compared assuming that UN was underestimated by 10%.
 J3-AGONIST AND OXIDATION OF NUTRIENTS 259

Table 3 Comparison of average values of UN and corre-

sponding values of HE.RQx and OXP

UN, g/d HE.RQx.kJId OXP.kJId OXP/HE.RQx, %

Section A

36 43801 4162 9.6

40(4-10%) 43779 4578 10.5

Section B

68 61798 7785 12.4

74(4-10%) 61758 8563 13.7

It is demonstrated that 10% error in UN values would have neglible effect on

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HE,RQ* . Although OXP would increase by 10%, the effect on the proportion
OXP/HE,RQx would not change the biological interpretation of the results.
In the previous paper (Chwalibog et al, 1995) it was demonstrated that treatment
with the j3-agonist L-644,969 had no significant effect on feed consumption but in-
creased daily live weight gain. The ß-agonist had no effect on metabolizability of
energy and protein digestibility, while utilization of digested protein and consequently
protein retention (RP) increased. Furthermore it was shown that higher levels of ß-
agonist used in Group 3 did not further increase RP, but reduce fat retention (RF).
In the present paper it was demonstrated that the yS-agonist had no effect on CH4
production, being an indicator of fermentation processes in the rumen. However,
caused by the diet applied, the CH4 production was lower than measured by
Thorbek (1980) for a high forage diet.
The values of HE,RQx Were highest for Group 2 in both sections, corresponding
with higher RP and lower RF (Chwalibog et al., 1995), hence heat increment by pro-
tein retention is higher than by fat retention.
A simplified model of nutrient partition between oxidation and retention process-
es in ruminants is demonstrated in Figure 1.
In this investigation protein oxidation was of the same magnitude in Section A
and accounted for about 10% of HE,RQx, while in Section B the control animals
increased OXP to 14.7%, but the treated bulls still had relatively low values about
11%. These values are much lower than 15-18% recalculated experiments with grow-
ing calves (Thorbek, 1980) and pigs (Chwalibog et al., 1992) and may indicate too
low protein concentration in the diet. In order to obtain the highest possible protein
retention the animals had^to reduced protein oxidation. This is evident for younger
bulls (Section A) which had the same OXP independent on treatment. However,
for older bulls (Section B) where the control animals had higher OXP than Group 2
and 3, it indicates a direct effect of ß-agonist on OXP reduction in order to save
amino acids for an increased protein retention.
The RQnpjc was not different between the control and the treated groups in
Section A, while it was reduced in Section B at 20 mg/d of /3-agonist. This reduction
is consistent with the switch between OXCHO and OXF as less carbohydrate and
more fat was oxidized supressing RQ values.
 260 A. CHWALIBOG et al.

CHO^Glucose-OXCHO
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Figure 1 A model of nutrient oxidation and retention in ruminants. Retained protein (RP), oxidized
protein (OXP), dietary carbohydrate (CHO), oxidized carbohydrate (OXCHO), short chain fatty acids
(SCFA), oxidized fat (OXF) and retained fat (RF)

As it is demonstrated by the model, a higher proportion of carbohydrate will be

oxidized as fat and not as carbohydrate by increased fermentation causing a higher
production of SCFA which partly are oxidized. In the present experiment
OXF/HE,RQx was markedly higher in animals treated with ß-agonist, especially
with 20 mg/d. However, as CH4 production was not influenced by the treatments,
indicating no changes in fermentation of dietary carbohydrate to fatty acids, it can
be postulated that the increase in OXF was caused by a direct influence of ß-agonist
on lipolysis (Miller et al., 1988) and oxidation of body fat.
In the conclusion, these findings, in concurrence with the model, demonstrate an
influence of ß-agonist on catabolic processes by reducing protein oxidation and
enhancing fat oxidation.

References
Brouwer, E.: Acta Physiol. Pharmacol. Nederlandia, 6,795 (1957)
Brouwer, E.: Proc.1st Symp. on Energy Metabolism, EAAP Publ., 8,182 (1958)
Brouwer, E.: Proc. 3rd Symp. on Energy Metabolism, EAAP Publ., 11, 441 (1965)
 0-AGONIST AND OXIDATION OF NUTRIENTS 261

Chwalibog, A., K.Jakobsen, S.Henckel and G.Thorbek:J Anim. Physiol. a. Anim. Nutr. 68,123 (1992)
Chwalibog, A., K.Jensen and G.Thorbek: Arch.Anim.Nutr., 49, 159 (1996)
Fahey Jr. G.C. and L.L.Berger: Carbohydrate nutrition of ruminants. In: The Ruminant Animal. Ed:
D.C.Church, Prentice-Hall, Englewood Clinks, N.J. pp 269-297 (1988)
Miller, M.F., D.K.Garcia, M.F.Coleman, P.A.Ekern, D.K.Lunt, K.A.Wagner, Procknor, M.,
T.H.Welsh, and S.B.Smith: J.Anim. Sci., 66,12 (1988)
Thorbek, G.: Report from the National Institute of Animal Science, Copenhagen, 498,193 pp. (1980)
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