DOI: 10.1111/j.1439-0396.2010.01049.

ORIGINAL ARTICLE

Comparison of grass haylage digestibility and metabolic plasma

profile in Icelandic and Standardbred horses
S. Ragnarsson1 and A. Jansson2
1 Department of Equine Science, Hólar University College, Sauðárkrókur, Iceland, and
2 Department of Animal Nutrition and Management, Swedish University of Agricultural Sciences, Uppsala, Sweden

Keywords
body weight, breed, body condition score, Summary
forage, insulin, urea
The aim of the present study was to compare digestibility and metabolic
Correspondence
response in Icelandic and Standardbred horses fed two grass haylages
S. Ragnarsson, Department of Equine Science, harvested at different stages of maturity. Six horses of each breed were
Hólar University College, 551 Sauðárkrókur, used in a 24-day change-over design. A total collection of faeces was
Iceland. Tel: +354 4556300; Fax: +354 made on days 15–17 and 22–24. Blood samples were collected on day
4556301; E-mail: [email protected] 24 of each period and analysed for total plasma protein (TPP), plasma
urea, non-esterified fatty acids, cortisol and insulin concentration. There
Received: 6 November 2009;
were no differences in digestibility coefficients of crude protein, neutral
accepted: 8 June 2010
detergent fibre or energy between breeds but organic matter digestibility
was higher in the Standardbred horses. On both haylages, the Icelandic
horses gained weight whereas the Standardbred horses lost weight. The
Icelandic horses had higher TPP, plasma insulin and lower plasma urea
concentrations. Our results indicate that the Icelandic horse may be
more prone to maintain positive energy balance in relation to the Stan-
dardbred horse, but there were no indication of a better digestive capac-
ity in the Icelandic horses.

supply is widespread (Þórhallsdóttir and Eirı́ksson,

Introduction
1996; Vervuert et al., 2001). This is also supported
It is widely accepted in horse management that some by anecdotal information that the Icelandic horse
horses are ‘easy keepers’, suggesting that they has been isolated from other breeds for more than
require less feed than other individuals to maintain 1000 years (Eldjárn, 1981; Pálsson, 1996) and analy-
condition. Convincing differences in digestibility sis of maternal DNA indicating no close relation to
between equid species, such as horses and donkeys, modern breeds such as the Thoroughbred and Stan-
have been shown and might be due to differences in dardbred horse (Vilá et al., 2001). However, scien-
the function of the digestive tract (Pearson and tific data on the digestive capacity and metabolic
Merritt, 1991; Cuddeford et al., 1995; Pearson et al., profile of the Icelandic horse is lacking and most
2001). In a study by Sverrisdóttir et al. (1994), the studies investigating digestibility in different types of
size of the digestive tract in Icelandic horses was equines, including potential ‘easy keepers’, have
investigated and in relation to literature data on been unable to show any significant differences
other breeds. They suggested that Icelandic horses (Slade and Hintz, 1969; Martin-Rosset et al., 1990;
might have a bigger large intestine than other breeds Vermorel et al., 1997).
and that this is because of adaptation to highly The aim of the present study was to compare for-
fibrous feeds. The opinion that the Icelandic horse age digestibility and the concentration of plasma
has special characteristics and is adapted to poor feed urea, non-esterified fatty acids, insulin and cortisol

Journal of Animal Physiology and Animal Nutrition 95 (2011) 273–279 ª 2010 Blackwell Verlag GmbH 273
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Digestibility in Standardbred and Icelandic horses S. Ragnarsson and A. Jansson

in Icelandic and Standardbred horses. The hypothesis a mineral mixture (KRAFFT Miner Röd, Falkenberg,
was that forage digestibility and metabolic response Sweden) was fed, amounting to 1% of the individ-
do not differ between the breeds. ual feed allowance.

Material and methods Collection of faeces

Animals, experimental design and management For total collection of faeces, the horses were fitted
Six Icelandic and six Standardbred geldings were with modified Stablemaid harnesses. The harnesses
used. The age was 10.4 (SD 2.7) and 9.0 years (SD were emptied into plastic bags every 3–4 h during
2.6), the height at the withers 139 (SD 3.0) and the collection periods. Each 24-h collection was
161 cm (SD 4.2) and the initial body weight (BW) weighed, homogenized and a 5% sub-sample was
423 (SD 28) and 507 kg (SD 49) for the Icelandic bagged and stored in airtight buckets at )24C until
and Standardbred horses, respectively. analysed.
The experiment was arranged as a change-over The haylages were sampled for 3 days during the
design with two forages (treatments), two periods collection periods, starting 2 days before faecal col-
and two groups of six horses (three of each breed). lection began. The samples were taken using a hay-
Each experimental period was 24 days with a 4-day driller and stored in plastic bags placed in airtight
transition to another feed between periods. Within buckets at )24C until analysed. Leftovers were
each period, a two 3-day collections were performed collected over the same 3-day period as the haylage
at days 15 to 17 and 22 to 24. The horses were kept and stored in plastic bags at )24C until analysed.
individually in boxes with concrete floors. Saw dust On the last day of each period, a haylage sample
was used as bedding until 4 days before collection of was taken for microbial analyses.
faeces started when it was switched to sand. The
horses were exercised by walking (15 min/day) dur-
Chemical analysis
ing collection periods and were also subjected to
some non-intensive exercise on adaptation days. All The haylage, leftover and faecal samples were pre-
horses had been kept on pasture for 12 weeks during dried at 60C for 24 h and then left over night at
the summer and were housed 2 weeks before the room temperature to acclimatize before weighing
study began. The experiment was performed at the (dry matter 1, DM1), then samples were milled
National Trotting School, Wången, Nälden, Sweden through a 1 mm sieve. For determination of dry
during the autumn of 2007 and was approved by matter, the samples were dried again at 105C for
the The Umeå local ethics committee. Average air 20 h (DM2). Dry matter content was calculated by
humidity in the stable was 80% and average tem- multiplying DM1 and DM2. Ash was determined by
perature 6C, which is within the thermoneutral incineration at 550C for 3 h. Neutral detergent fibre
zone for acclimatized horses (McBride et al., 1985; (NDF) was analysed in feed and faeces according to
Cymbaluk, 1990). Chai and Udén (1998), acid detergent fibre (ADF)
and acid detergent lignin (ADL) was analysed in feed
according to Goering and Van Soest (1970). Water-
Forage and feeding
soluble carbohydrates (WSC) were analysed accord-
The forages consisted of 60% timothy, 35% mea- ing to Larsson and Bengtsson (1983). The Kjeldahl
dow fescue and 5% rye grass and were harvested method was used to determine nitrogen (N) content
from a 3-year ley fertilized with 77 kg N per hect- in feed and faeces and the value multiplied by 6.25
are. The forages were harvested at two maturity for estimating crude protein (CP). Forage microbial
stages, forage 1 after heading and forage 2 after content was analysed according to Nordic Committee
seeding. The forages were preserved as haylage in on Food Analysis (2005) after modification by using
plastic wrapped square bales. The feed ration was peptone salt water instead of peptone water.
0.023 kg feed per kg BW and based on prior experi-
ence of the requirements of the experimental
Measurements of body weight, body condition scores
horses. The horses were fed four times every 24 h at
and rump fat
6:00 (10% of allowance), 12:00 (19%), 17:00 (25%)
and 21:00 hours (46%). Water was available ad libi- The body weight was recorded in the beginning and
tum from automatic drinking troughs and the horses in the end of the last 10 days in each period (at
had access to salt lick stones. During days 1–12, 8:00 hours) using an electronic scale (E2000S,

274 Journal of Animal Physiology and Animal Nutrition. ª 2010 Blackwell Verlag GmbH
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S. Ragnarsson and A. Jansson Digestibility in Standardbred and Icelandic horses

Tru Test, New Zealand) and the weight change was Yij = l + ai + ej + bk + eijk
calculated. The body condition score (BCS) was where Yijk is the observation, l the mean value, ai
assessed on the last day of each period according to the effect of period, ej the effects of collection period
the Icelandic body condition scoring system using (within period), bk the effect of breed and eijk the
scale ranging from 1 to 5, score 3 is moderate condi- residuals; eijk  IND (0, d2).
tion, score 2.5 moderately thin and score 3.5 moder- For comparison of feeds within breed the follow-
ately fleshy according to Stefánsdóttir and ing model was used:
Björndóttir (2001) and the Henneke Body Condition Yijk = l + ai + bj + ck + eijk
Scoring System ranging from 1 to 9 (Henneke et al., where Yijk is the observation, l the mean value, ai
1983). At the end of the experiment, an ultrasonic the effect of period, bj the effect of feed, ck the effect
measurement was performed to estimate rump fat of horse and eijk the residuals; eijk  IND (0, d2).
thickness according to Westervelt et al. (1976).
Results
Plasma analysis
The chemical and microbial compositions of the hay-
Blood samples were taken from the vena jugularis lages are given in Table 1. There were no differences
by vacutainer in heparinized tubes (10 ml) at the between the breeds in the coefficients of total tract
last day of the collection periods at 13:00 to apparent digestibility (CTTAD) for energy, CP and
13:30 hours. They were centrifuged for 10 min NDF but the Standardbred horses had higher
(950 g) and stored at )18C until analysed. For (p > 0.05) CTTAD for OM (Table 2). The measured
quantitative determination of non-esterified fatty DE content (MJ/kg DM) of the haylages were 9.8
acids (NEFA) in plasma an enzymatic colorimetric and 7.5 for haylage 1 and 2, respectively and the
method was used (ACS-ACOD method, Wako Standardbred horses had higher energy intake on
Chemicals GmbH, Neuss, Germany) and calculations haylage 1 (Table 3). The Icelandic horses had higher
were made from a standard curve with a linear BCS than the Standardbred horses (Table 3) and on
curve fit. Total plasma protein concentration (TPP) both haylages, the Icelandic horses gained BW and
was analysed using a refractometer (Atago, Tokyo, the Standardbred horses lost BW (Table 3).
Japan). The plasma urea concentration was analysed The plasma urea concentrations were lower and
using an enzymatic method (Konelab, Thermo Clini- the TPP concentrations were higher in the Icelandic
cal Labsystem, Vantaa, Finland). Plasma cortisol was horses in relation to the Standardbred horses on
analysed using radioimmunoassay (Immulite 2000, both haylages (Table 4). There were no effects of
Siemens, Eschborn, Germany) and plasma insulin by feed on TPP but plasma urea concentration was
ELISA (Mercodia equine insulin kit, Mercodia, higher on haylage 1 in both breeds. The urea/TPP
Uppsala, Sweden). ratio was lower in Icelandic horses in relation to

Statistical analysis
Table 1 Dry matter content (DM, g/kg), chemical compositions (g/kg
Data were subjected to a general linear model analy- DM), gross energy (GE, MJ/kg DM), pH and microbial composition (cfu/
sis, using SAS System for Windows, Release 9.1 g fresh matter) of the haylages used
(SAS, 2002). Values are presented as least square
Haylage 1 Haylage 2
means (LSM) with their standard errors (SE). Differ-
ences were considered statistically significant when DM 788 778
p < 0.05. Organic matter 928 952
Crude protein 111 61
Overall breed effects (both diets included) were
Neutral detergent fibre 611 627
analysed by using the following model:
Acid detergent fibre 362 377
Yijkl = l + ai + ej + bk + cl + (bc)kl + ei j kl Water-soluble carbohydrates 79 147
where Yijkl is the observation, l the mean value, ai Lignin 51 72
the effect of period, ej the effect of collection period GE 18.6 18.6
(within period), bk the effect of breed, cl the effect of pH 6.0 6.0
feed, (bc)kl the effect of interaction between breed Yeast <log 2.0 <log 3.0
Mould <log 2.0 <log 2.0
and feed and eijkl the residuals; eijkl  IND (0, d2).
Enterobacteria log 2.4 log 2.7
For comparison of breeds within feed the follow-
Clostridia <log 2.0 <log 2.0
ing model was used:

Journal of Animal Physiology and Animal Nutrition. ª 2010 Blackwell Verlag GmbH 275
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Digestibility in Standardbred and Icelandic horses S. Ragnarsson and A. Jansson

Table 2 Coefficients of total tract apparent digestibility in Icelandic Table 4 Total plasma protein concentration (TPP), plasma urea (Urea),
and Standardbred horses on two haylage diets non-esterfied fatty acids (NEFA), insulin, cortisol concentrations and
urea, NEFA, insulin, cortisol ratio on TPP in Icelandic and Standardbred
Icelandic Standardbred Breed effect horses fed different haylage diets
Haylage 1 Haylage 2 Haylage 1 Haylage 2 SE p-value Icelandic Standardbred Breed effect
Plasma
OM 0.536A 0.431 0.565B 0.427 0.006 0.0432 variables Haylage 1 Haylage 2 Haylage 1 Haylage 2 SE p-value
CP 0.636 0.478 0.660 0.479 0.012 0.3047
NDF 0.517 0.322 0.536 0.319 0.008 0.3226 TPP (g/l) 71A 70A 65B 64B 1.4 0.0004
Energy 0.517A 0.407 0.540B 0.400 0.007 0.2205 Urea 5.0Aa 3.5Ab 7.0Ba 5.0Bb 0.34 0.0001
(mmol/l)
OM, organic matter; CP, crude protein; NDF, neutral detergent fibre. NEFA 58.3 64.9A 49.9 52.0B 3.97 0.0145
p-values show effect of breed (both forages included). Values in the (lmol/l)
same row with different superscript show significant differences Insulin 38.4A 54.4A 15.5Ba 23.3Bb 6.77 0.0008
(p < 0.05) between breeds within each forage. (lU/ml)
Cortisol 73.7 70.5 85.0 75.7 7.03 0.2553
(¢mol/l)
Table 3 Organic matter intake (OMI, g/kg body weight(BW)/day), Urea/TPP 0.07A 0.05A 0.11Ba 0.08Bb 0.01 0.0001
digestible energy intake (DEI, MJ/kg BW/day), crude protein intake (CPI, NEFA/TPP 0.83 0.92 0.77 0.82 0.05 0.1238
g/kg BW/day), body weight changes (BWC, kg) and body condition Insulin/TPP 0.55A 0.78A 0.24Ba 0.37Bb 0.10 0.0016
score (BCSH and BCSI) in Icelandic and Standardbred horses on two Cortisol/TPP 1.04 1.01 1.30 1.20 0.11 0.0596
haylage diets
p-values show effect of breed (both forages included). Values with
Icelandic Standardbred Breed effect upper case superscript show significant differences (p < 0.05) between
breeds within each forage and values with lower case superscript
Haylage 1 Haylage 2 Haylage 1 Haylage 2 SE p-value show significant differences (p < 0.05) between forages within breed.

OMI 14.9 16.3 15.6 16.5 0.3 0.2330

DEI 0.152A 0.129 0.168B 0.128 0.005 0.1110 positive correlation between BCS and plasma insulin
CPI 1.79 1.04 1.85 1.04 0.04 0.4344 (Fig. 1). The rump fat thickness was 8.8 ± 0.4 mm in
BWC 1.7 ± 2.3 1.7 ± 2.3 )4.4 ± 2.5 )4.4 ± 2.5 –* 0.0235 the Icelandic horses and 4.9 ± 0.4 mm in the Stan-
BCSH 7.5A 7.2A 4.6B 4.4B 0.2 0.0001 dardbred horses (p = 0.0002) corresponding to a
BCSIà 3.7A 3.6A 2.7B
2.7B 0.1 0.0001 total body fat content of 12.8% and 10.5% (Wester-
p-values show effect of breed (both forages included). Values in the velt et al., 1976). There was a correlation between
same row with different superscript show significant differences rump fat thickness (mm) and plasma insulin (lU/
(p < 0.05) between breeds within each forage. ml) at the end of the experiment (Y = 7.063x )
*Due to missing value; Henneke et al. (1983); àStefánsdóttir and 13.87, R2 = 0.41). There was also a correlation
Björndóttir (2001).
90

Standardbred horses on both haylages. There were 80

no differences in the urea/TPP ratio between haylag- 70
es in the Icelandic horses but in the Standardbreds
Plasma insulin (μU/ml)

60
the ratio was higher on haylage 1 in relation to hay-
lage 2 (p = 0.0014). There was a positive correlation 50
between crude protein intake (g/day) and plasma
40
urea (mmol/l) across breeds and feeds
(Y = 145.3 + 79.9x, R2 = 0.51, SD = 114). The NEFA 30

concentration was higher in Icelandic horses in rela- 20

tion to Standardbred horses. However, there were
10
no differences between breeds or feeds in the NEFA/
TPP ratio (Table 4). The Icelandic horses had higher 0
plasma insulin concentrations than the Standardbred 3 4 5 6 7 8 9
BCS
horses on both feeds and the Standardbred horses
had higher plasma insulin concentration on haylage Fig. 1 Correlation between the Henneke et al. (1983) body condition
2 in relation to haylage 1 (Table 4). There were no scores (BCS) and plasma insulin concentration in Icelandic and Stan-
differences in the plasma cortisol concentration, nei- dardbred horses (mean value of two haylage diets).
ther between breeds or feeds (Table 4). There was a Y = 9.8259x ) 26.498, R2 = 0.65.

276 Journal of Animal Physiology and Animal Nutrition. ª 2010 Blackwell Verlag GmbH
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S. Ragnarsson and A. Jansson Digestibility in Standardbred and Icelandic horses

between rump fat and BCS across breeds elevated in the Standardbred horses in relation to the
(Y = 0.667x + 1.72; R2 = 0.66). Icelandic horses but the Standardbred horses consis-
tently had higher plasma urea concentrations (even
on haylage 2, low crude protein feed in relation to
Discussion
the Icelandic horses on haylage 1, high crude protein
This study indicates that there are no major differ- feed), perhaps indicating catabolic metabolism.
ences in the digestive capacity of Icelandic horses in The Icelandic horses had higher TPP. This might be
relation to Standardbred horses. This is in accor- a breed effect but might also reflect a smaller plasma
dance with other studies on different types of horses volume. Difference in blood volume between breeds
(Slade and Hintz, 1969; Martin-Rosset et al., 1990; has earlier been reported (Persson, 1967). A larger
Vermorel et al., 1997). plasma volume in the Standardbred horses might be
However, the OMD was lower in Icelandic horses due to the Standardbred horses having been sub-
in relation to Standardbred horses. This was mainly jected to high intensity training earlier (racing) and
caused by lower OMD on haylage 1 because the training has been shown to increase the plasma vol-
OMD on haylage 2 was similar. The reason for the ume (McKeever et al., 1987). For this reason, the
lower OMD in Icelandic horses on haylage 1 is not statistical analysis for all plasma variables was run
clear. Both breeds had higher OMI on haylage 2 in both in absolute values and as a ratio of TPP. With
relation to haylage 1 (p < 0.0001) and no marked this treatment, the breed difference in NEFA disap-
selection was detected. This indicates that the intake peared whereas the breed differences in urea and
could have been regulated on an energy intake basis insulin remained. The plasma insulin concentrations
as suggested by Edouard et al. (2008) because the were higher in the Icelandic horses. This could be a
digestibility was higher on haylage 1. On both hay- breed difference but it could also be breed indepen-
lages, the Icelandic horses were in positive energy dent and influenced by differences in body condition
balance, indicated by increased BW. This is in con- (Pagan et al., 2009). The Icelandic horses had higher
trast to the Standardbred horses who failed to main- BCS and thicker rump fat. The BCS has been shown
tain BW, in spite of a higher DEI on haylage 1. The to be correlated to plasma insulin concentrations in
higher energy requirements of the Standardbred horses and ponies (Carter et al., 2009). In the present
horses in relation to the Icelandic horses might be study, the correlation was even stronger than in the
due to a higher average voluntary activity and/or to study by Carter et al. (2009). The R2 explaining the
a higher maintenance requirement because of a variation in insulin in relation to BCS (R2 = 0.65)
more energy-requiring body tissue composition (less was higher than the R2 for the correlation between
fat, more muscle) (Blaxter, 1989). A higher meta- rump fat and insulin (R2 = 0.41, equation data not
bolic rate in the Standardbred horses could also be shown) suggesting that a BCS measurement could be
supported by the tendency to higher cortisol levels of value as indicator of metabolic changes as sug-
(cortisol/TPP) in this breed. gested by Carter et al. (2009). However, another rea-
The reason for the unbalanced appetite in the son for the higher plasma insulin concentrations in
Standardbred horses (i.e. feed intake not matching the Icelandic horses might be differences in the feed
energy losses) remains unclear and highlights the intake pattern and glucose uptake since the samples
phenomenon of easy keepers vs. not so easy keepers. were taken 1 h after feeding. This explanation seem,
Based on DEI intake and the changes in BW of the however, less likely because the difference between
horses in the present study, the Icelandic horses are breeds in insulin concentrations was very large
in the range ‘normal to easy keepers’ or 0.127– (>200% higher in the Icelandic horses) and the OMI
0.139 MJ DE/kg BW and the Standardbred horses in was similar or even numerically higher in the Stan-
the range ‘normal to hard keepers’ or 0.139– dardbred horses. The reason for the higher plasma
0.152 MJ DE/kg BW according to NRC (2007). The insulin concentration in the Standardbred horses on
‘easy keeper’ suggestion is supported by Ragnarsson haylage 2 is not clear but can be due to the higher
(2009) who estimated the maintenance require- WSC content on this feed.
ments of sedentary Icelandic horses to be 0.123 MJ
DE/kg BW.
Conclusion
It has earlier been shown that the plasma NEFA
concentration increases in horses in negative energy Our results indicate that the Icelandic horse might
balance (Sticker et al., 1995). However, in the pres- be more prone to maintain positive energy balance
ent study the plasma NEFA concentration was not in relation to the Standardbred horses, but that this

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Digestibility in Standardbred and Icelandic horses S. Ragnarsson and A. Jansson

is not because of a better digestive capacity. The measurement, and body fat percentage in mares.
study also showed that the metabolic profile differed Equine Veterinary Journal 15, 371–372.
between the breeds with respect to plasma urea, Larsson, K.; Bengtsson, S., 1983: Bestämning av lätt till-
total plasma proteins and plasma insulin concentra- gängliga kolhydrater i växtmaterial (determination of readily
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Acknowledgements horses. Livestock Production Science 25, 257–264.
McBride, G. E.; Christopherson, R. J.; Sauer, W., 1985:
The authors want to thank the staff at Wången
Metabolic rate and plasma thyroid concentrations of
National Trotting School and Sarah Hamilton for
mature horses in response to changes in ambient temper-
their assistance during the study, Agricultural Pro- atures. Canadian Journal of Animal Science 65, 375–382.
ductivity fund of Iceland, Trioplast AB Sweden for McKeever, K. H.; Schurg, W. A.; Jarrett, S. H.;
funding and HästKraft AB and Lantmännen Box Convertino, V. A., 1987: Exercise training-induced
Sweden, for providing the forages. hypervolemia in the horse. Medicine and Science in Sports
and Exercise 19, 21–27.
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