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Body composition and muscle glycogen contents of piglets of sows fed diets differing in fatty acids profile and contents
Livestock Science 123 (2009) 329–334
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Body composition and muscle glycogen contents of piglets of sows fed diets differing in fatty acids profile and contents G. Pastorelli a,⁎, M. Neil b, I. Wigren b a b
Department of Veterinary Sciences and Technologies for Food Safety University of Milan, 20133 Milan, Italy Department of Animal Nutrition and Management, Swedish University of Agricultural Sciences, Box 7024, S-750 07 Uppsala, Sweden
a r t i c l e
i n f o
Article history: Received 7 March 2008 Received in revised form 13 November 2008 Accepted 21 November 2008 Keywords: Dietary fatty acids Piglet Body composition Glycogen
a b s t r a c t To study the influence of sow dietary fat on piglet body characteristics, multiparous sows were allocated to one of four different dietary treatments: a conventional low fat (3%) diet (LF) and three high fat (6%) diets; high fat saturated (HFS), high fat oats (HFO), and high fat linseed (HFL). All sows were fed the allocated diet from weaning of the preceding litter until the day after farrowing. At farrowing, one liveborn piglet per litter (NB), was sacrificed and dissected immediately after birth. The heaviest (H) and the lightest (L) piglets in the litter were killed and dissected in the same manner at one day of age. Measurement of body length and circumference, organ weight, body chemical composition and muscle glycogen content were determined. Body measurements were adjusted to the mean body weight (1.67 kg). Dietary treatment did not have any significant influence on body components or carcass traits except for lung weight, being lower in HFO and HFL than in LF piglets. Piglet category affected almost all parameters considered, showing the lowest values for NB piglets, except for lung and circumference that were higher in NB than in L and H piglets; and length which was lower in NB than in L piglets. NB piglets had the highest amount of muscle glycogen content, no difference was found between H and L piglets. Dietary treatments influenced piglet chemical composition, showing the highest overall values of dry matter (DM), protein, and fat for the HFL piglets' carcasses. The present data provide additional information on the depletion of energy reserves; it would appear that sow dietary fat has relatively little effect on progeny since only body chemical composition was significantly influenced by HFL diet. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Dietary fat is important as a concentrated energy source. Feeding fat to sows during late gestation and into lactation may improve piglet survival (Seerley et al., 1974; Pettigrew, 1981). Fat is also important as a source of specific compounds, such as medium-chain triglycerides (MCT) and essential fatty acids (EFAs). MCT have fatty acids 6 to 12 carbon atoms long (Bach and Babayan, 1982) and can be easily utilized by pigs (Benevenga et al., 1989; Chiang et al., 1990; Odle et al., 1991). Improved survival or improved indices of survival have been observed in litters from sows fed ketogenic compounds such as 1,3⁎ Corresponding author. Tel.: +39 0250315758; fax: +39 02 50317898. E-mail address: [email protected] (G. Pastorelli). 1871-1413/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.livsci.2008.11.023
butanediol (Stahly et al., 1985, 1986) and MCT (Rosebrough et al., 1981). Because of the rapid metabolism of MCT to ketone bodies and the ability of ketone bodies to both stimulate lipogenesis and spare carbohydrate use for nonoxidative functions (Allee et al., 1972; Seccombe et al., 1977; Shambaugh, 1985) MCT increased the survival of piglet (Azain, 1983). EFAs are fatty acids which are required by an organism for the maintenance of normal growth and reproduction, are not able to be synthesized by the organism, and are therefore required in appropriate amounts in the diet (Leskanich and Noble, 1999). EFAs and their derivatives have varied biological actions that may have relevance to their involvement in several physiological and pathological processes (reviewed by Das, 2006). Linoleic acid and α-linolenic acid are classified as EFAs.
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Table 1 Composition of experimental diets (as-fed). Diet1
Ingredient, % Barley Wheat Oats High fat oats Soya bean meal Wheat middlings Wheat bran Vitamin and mineral premixe Fat (Akofeed Standard)f Fat (Akofeed Cattle 45)g Monocalcium phosphate Limestone Sodium chloride L-Lysine HCl DL-Methionine L-Threonine Linseed oile Calculated composition, % Dry matter Crude protein Lysine Methionine + Cystine Threonine Crude fat Crude fibre Ca P Metabolizable energy, MJ Analysed composition, % Dry matter Crude protein Crude fat Crude fibre Ash
LFa
HFSb
HFOc
HFLd
14.06 42.25 20.0 – 10.0 5.0 5.0 1.0 0.19 – 1.03 0.95 0.40 0.08 0.01 0.04
46.03 19.17 5.0 – 12.0 5.0 5.0 1.0 – 3.65 1.51 1.02 0.40 0.07 0.07 0.08
1.74 33.48 – 40.0 10.1 5.0 5.0 1.0 0.94 – 1.05 1.19 0.40 0.04 0.04 0.02
– 33.8 20.0 20.0 10.5 5.0 5.0 1.0 0.85 – 1.32 1.00 0.40 0.05 0.07 0.02 1.00
88.6 15.3 0.70 0.56 0.52 3.0 4.8 0.80 0.65 12.3
88.4 15.0 0.72 0.59 0.60 6.0 4.6 0.90 0.75 12.8
89.6 15.4 0.71 0.64 0.53 6.0 6.4 0.90 0.65 12.6
89.4 15.4 0.71 0.63 0.52 6.0 6.0 0.87 0.71 12.6
89.0 15.5 3.2 4.6 5.5
88.7 16.1 5.5 4.5 6.0
90.9 16.0 5.9 5.3 5.8
89.3 15.3 4.9 6.5 6.2
1
Diets were calculated to meet nutrient requirements according to Swedish recommendations (Simonsson, 1994) of breeding sows. aLF = low fat, bHFS = high fat saturated, cHFO = high fat oats, dHFL = high fat linseed oil (without linseed oil). e Contributed per kilogram of complete feed; 10,000 IU vitamin A, 1,000 IU vitamin D3, 60 mg vitamin E, 2 mg vitamin K3, 2 mg vitamin B1, 4 mg vitamin B2, 3 mg vitamin B6, 0.02 mg vitamin B12, 0.25 mg biotin, 1.5 mg folic acid, 20 mg niacin, 15 mg d-pantothenic acid, 0.30 mg I, 0.30 mg Se, 40 mg Fe, 0.01 mg Co, 10 mg Cu, 0.41 mg Mg, 20 mg Mn, 70 mg Zn. f Source of unsaturated fatty acids. g Source of saturated/monounsaturated fatty acids.
Although linolenic acid and linoleic acid require the same enzymes for conversion into longer-chain essential fatty acids, they belong to the omega-3 and omega-6 fatty acids series, respectively, which cannot be inter-converted. The fatty acid elongation pathways are not present in the foetus (Li et al., 2000), implying that the unborn piglet is dependent on maternal supplies. Lipid supply, particularly EFA and long chain polyunsaturated is shown to affect neural development and function (Uauy and Castillo, 2003). Moreover, EFAs are also precursors for the prostaglandins and eicosanoid mediators, which play an important role in reproduction (Ziecik et al., 2000) and immunity (Calder, 2002). Therefore dietary supplementation with fat to enhance energy and EFA supply during the late gestation and lactation may be an effective way of increasing the EFA content in piglet tissue and the EFA level of colostrum and milk, thereby improving the viability of piglets (Gu and Li, 2003) and the
immune status of the neonate (Leskanich and Noble, 1999). Differences in placenta development may cause differences in piglet storage of glycogen (Wu et al., 2006). The stored glycogen provides energy during the birth process and after birth, especially in the period before first colostrum intake. In this way, the piglet becomes more able to respond to the stressful extra-uterine environment and has a bigger chance to survive (Le Dividich et al., 2005). Body glycogen is the major energy reserve of the newborn pig, ranging between 30 and 38 g/kg BW. However, glycogen stores are rapidly depleted after birth (Elliot and Lodge, 1977). The total amount of fat in the newborn pig is very low, ranging from 10 to 20 g/kg BW. Selection of pigs for reduced carcass fatness has resulted in pigs that are leaner at birth (Herpin et al., 1993) and have lighter weight livers and less liver glycogen (Canario et al., 2005). Another factor of great importance for piglet survival is the body weight, since mortality is higher in lighter piglets (Rydhmer, 1992). We hypothesised that the glycogen stores in newborn piglets will be depleted faster in light piglets than in heavy piglets and that there might be differences regarding composition or relative organ sizes. The purpose of this study was to evaluate the effect of level and type of fat in diets for pregnant sows on the typical newborn piglet body composition, proportional organ size
Table 2 Fatty acid composition of feed. Fatty acids, g/ 100 g fatty acids C10:0 C12:0 C14:0 C16:0 C18:0 C18:1 C18:2, n − 6 C18:3, n − 3 C20:0 C20:1 C20:2 C22:0 C22:1 C24:0 C20:4, n − 6(Aa) SFAc MUFAc PUFAc n − 6/n − 3 a
Dieta LF
HFS
HFO
HFL
0.04 0.29 0.30 18.40 1.72 20.76 47.07 4.05 0.22 0.66 0.09 0.23 0.15 0.19 Ndb 21.16 21.57 51.21 11.6
0.21 1.68 1.20 32.02 4.14 24.91 27.75 2.38 0.29 0.36 0.07 0.19 0.08 0.16 0.18 39.89 25.35 30.20 11.6
0.05 0.34 0.29 16.29 3.10 36.29 35.47 2.67 0.29 0.69 0.06 0.20 0.10 0.16 0.02 20.72 37.08 38.52 13.3
0.0 0.0 0.2 13.1 2.2 32.2 34.8 13.2 0.3 0.7 0.1 0.0 0.2 0.1 0.2 15.9 33.1 48.3 2.7
LF: a conventional Low Fat 3%. HFS: High Fat Saturated. HFO: High Fat Oats. HFL: High Fat Linseed. The diets LF, HFO and HFL contained a fat blend intended for pigs (Akofeed Standard), with a mono-/polyunsaturated profile, whereas the HFS diet contained a fat blend intended for cattle (Akofeed Cattle 45). The HFO and HFL diets also included the high-fat oats cultivar Matilda (10% fat, C18:1 dominating) as a fat source. To the HFL diet linseed oil (3.9% C16:0, 3.4% C18:0, 11.8% C18:1, 15.3% C18:2, 63.7% C18:3) was added by hand at feeding time, as a top-dress at a rate of 1% by weight. b Nd: Not detected. c SFA: Saturated fatty acids. MUFA: Monounsaturated fatty acids. PUFA: Polyunsaturated fatty acids.
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and muscle glycogen, and to compare with the extremes regarding body weight in the same litter at one day of age. 2. Materials and methods The piglets in the present study were collected from 38 litters. Sows were allocated to one of four experimental diets (Table 1) The fatty acid composition of the diets is shown in Table 2. The experimental procedures were carried out under the Swedish Animal Welfare Act 1988:534 and were approved by the Uppsala Ethics Board for Animal Experimentation, Sweden (No. C 81/1). At farrowing, one representative liveborn piglet per litter, usually the 3rd piglet born (NB), was selected immediately after birth for organ and tissue sampling, and body measures. This piglet was not allowed to suckle. The piglet was rendered unconscious by carbon dioxide gas, then weighed and bled to death. Body length was measured from snout to the tail base; circumference was measured around the chest immediately behind the fore legs. The heart, liver, lungs, stomach with content, spleen, kidneys (left and right weighed together with the respective adrenals), and the large and small intestines (weighed together), were anatomically dissected and weighed. Samples of longissimus muscle were collected and frozen immediately in liquid nitrogen following dissection and stored at −80 °C. Carcasses were stored in − 20 °C until chemical analysis. At the age of 20–36 h, the piglets that were heaviest (H) and lightest (L) in the litter at first handling after birth were killed and dissected in the same manner as the newborns. 2.1. Analytical procedures Samples of feed from each batch and homogenized samples of piglet carcasses were analysed in duplicates. The contents of dry matter (DM) and ash was determined by drying the samples in 103 °C for 16 h, and by ashing in 550 °C for 3 h, respectively. The content of nitrogen was analysed according to Kjeldahl (Nordic Committee on Food Analysis, 1976) using a Kjeltec (Tecator 2460) and crude protein (CP) was calculated as
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N× 6.25. The content of crude fat (CF) was analysed according to the Official Journal of European Communities (1984) using a Tecator-equipment (hydrolyse unit 1047 and extraction by Soxtec HT6). In feed samples, crude fibre was analysed by the short method as described by Jennische and Larsson (1990). Dietary fatty acids were analysed by extracting the lipids using hexane and isopropanol (Nourooz-Zadeh and Appelqvist, 1988). Methylation of fatty acids were according to Sukhija and Palmquist (1988) and separation was made with a temperature-programmed gas chromatograph (Chrompack CP9001 GC, Chrompack, The Netherlands) equipped with a split injection system and capillary column (Chrompack CP-sil 88, 50 m ×0.25 i.d.). Identification of individual fatty acids was based on standards (Larodan Fine Chemicals, Sigma Chemical Co, Malmö, Sweden) and published data (Precht and Molkentin, 1996). Glycogen concentrations were assayed on freeze-dried material from piglets selected from LF, HFS and HFO treatments. Before analysis the freeze-dried muscle pieces were dissected free from connective tissue, blood and fat. After boiling 1–2 mg of the sample for 2 h in 1 ml of 1 M hydrochloric acid to hydrolyze the glycogen, the glucose concentration was determined using a fluorometric method (Lowry and Passoneau, 1973). 2.2. Statistical analysis Data were analysed as a completely randomised design using the GLM procedure of SPSS (2001). The model included dietary treatment, parity and category as fixed effect. All data relative to carcass composition were studied with body weight as a covariate. Parity was classified as old (O) at order of parity 5 or higher, and young (Y) at order of 2 to 4. Data in tables are presented as least square means. Differences among treatment means were compared by least significant difference. 3. Results The effect of diet, sow parity and piglet category on organ weight and body measurements are presented in Table 3.
Table 3 Effect of diet, parity and piglet category on organ weight and body measurements a. Diet
Carcass weight, kg Liver, g Intestinal package, g LKAG, g RKAG, g Stomach, g Spleen, g Heart, g Lungs, g Length, cm Circumference, cm
Parity
Piglet category
SE
LF (N = 27)
HFS (N = 26)
HFO (N = 27)
HFL (N = 27)
Y (N = 62)
O (N = 45)
NB 36
L 35
H 36
1.27a 46.7
1.27a 49.0
1.32a 49.1
1.20b 52.2
1.25 50.4
1.28 48.1
1.29 51.1
1.25 47.4
1.25 49.2
93.7 7.58 7.26 43.0 1.67 12.7 26.7a 38. 7 24.4
94.3 7.25 7.11 46.2 1.57 12.2 25.2ab 38.4 24.0
90.6 7.12 6.98 45.2 1.51 12.3 24.0b 38.6 24.2
83.5 7.21 7.00 44.6 1.44 12.4 23.6b 38.4 25.2
91.4 7.16 6.92 45.7 1.58 12.0 26.0 39.0 24.7
89.6 7.42 7.25 43.9 1.51 12.7 23.7 38.0 24.2
71.9a 6.29a 6.03a 26.5a 1.27a 12.0a 28.6a 37.8a 25.2a
98.1b 7.50b 7.27b 55.9b 1.58b 12.3ab 22.5b 38.9b 24.1b
101.5b 8.08b 7.96b 52.0b 1.78b 12.9b 23.6b 38.8ab 24.1b
Significance b D
P
C
0.01 0.78
† NS
NS NS
NS NS
1.48 0.11 0.11 1.87 0.03 0.11 0.37 0.18 0.16
† NS NS NS NS NS * NS NS
NS NS NS NS NS ** ** ** NS
*** *** *** *** *** * *** * *
Diets: Low fat (LF), High fat saturated (HFS), High fat oats (HFO), High fat linseed oil (HFL). Parity: Young i.e. parity 2–4 (Y), Old i.e. parity 5 or more (O). Piglet category: Newborn (NB), one day old lightest in litter (L), heaviest in litter (H). SE — standard error; LKAG — Left kidney and adrenal gland; RKAG — Right kidney and adrenal gland. abc Means with different superscripts within a row differ at significance indicated. a Values adjusted to the same body weight (1.67 kg). b Significance: NS = P N 0.05; † = P b 0.10; *P b 0.05; **P b 0.01; ***P b 0.001.
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Fig. 1. Glycogen content in muscle from newborn (NB), and one day old piglets born heaviest (H), and lightest (L) in their respective litters.
Except for lung weight there were no significant differences between dietary treatments for any measurement made on body components or carcass traits. Parity had influence on relative lung and heart weight, and body length; piglets of young sows had lighter hearts, heavier lungs and were longer. Piglet category had a significant effect on almost all parameters considered, showing the lowest values for NB piglets. Exceptions were lungs and circumference that were significantly larger in NB than in L or H piglets; and length, which was similar in H and L piglets, with L piglets significantly different from NB piglets. Concerning the muscle glycogen content it was found that NB piglets had the highest amount, and no difference was found between H and L piglets (Fig. 1). Dietary treatments influenced carcass chemical composition (Table 4), showing highest overall values for HFL piglets. Parity had a significant effect on DM only which was higher in piglets of young than of old sows; category showed a significant effect on CF and CP reporting the smallest, intermediate and highest values for the NB, L and H piglets, respectively. 4. Discussion Gestational supplementation with dietary fat did not influence organ weights, except for lung weight being lower in piglets of sows fed diets HFO and HFL, these diets were supplemented with PUFA. This is partly in accordance with Farnworth and Kramer (1988), who fed sows diets supple-
mented with soybean oil (rich in PUFA) or tallow with a cornbased diet as a control, and found no difference between diets in relative weights of heart, liver or lungs in the offspring. These data, however, were from 110 days old fetuses i.e. before birth. Poulos et al. (2004) found that maternal CLA consumption decreased relative heart weight in newborn male piglet but no difference was found regarding liver, kidney or lung. No explanation for this response is apparent. We speculate that our results are due to the supply of fatty acids not being the limiting factor during gestation in development and growth of these organs. Glycogen content is not affected by dietary treatments (data not shown). This is in accordance with results of Newcomb et al. (1991), where no significant difference in piglet glycogen content was found due to sow dietary energy source (starch, MCT (medium chain triglycerides) or soybean oil). Differently, Jean and Chiang (1999) found that sow dietary treatments with MCT and coconut oil resulted in higher (P b 0.10) muscle glycogen content in neonatal piglets than did soybean oil. Anyway, coconut oil and MCT especially, have fatty acids with 6 to 12 carbon atoms length, which can readily cross the placenta (Jean and Chiang, 1999; Rosebrough et al., 1981). Furthermore, the soybean oil dietary treatment (Jean and Chiang, 1999) showed glycogen values of 65 mg/g wet muscle which is similar to those found in the present experiment (51 mg/g wet muscle in NB provided a molar weight of 180 g and 20% DM in muscle). The carcass fat content did not differ between sow dietary energy sources (starch, MCT, or soybean oil) in the study by Newcomb et al. (1991), and was similar to our finding, less than 2%. Such a low content implies the lipid to be primarily structural and as such, may be unavailable as an energy source. The reserves of neonatal piglets are depleted quickly if piglets must metabolize glycogen in an attempt to stay warm (Noblet and Le Dividich, 1981; Herpin et al., 2002). Muscle glycogen content mainly supplies energy for muscle movements (English and Morrison,1984) and this explains why newborn piglets, sacrificed immediately after birth, showed significantly higher values than light and heavy piglets. Values obtained indicate that the rate of mobilization of muscle glycogen was similar in heavy and light piglets; in which the content was 11% lower than in newborns. The depletion after 24 h was similar to the finding of Herpin et al. (2002) in thermoneutral conditions, but lower than in the study by Elliot and Lodge (1977), who found a significant (P b 0.05) decrease of about 40% during the first 12 h post partum establishing a pattern of progressive decline over the full 96 h.
Table 4 Effect of diet, parity group and piglet category on carcass chemical composition. Diet (D)
Dry matter Ash Crude protein Crude fat
Parity (P)
Significance a
Piglet category (C)
LF (N = 27)
HFS (N = 26) HFO (N = 27) HFL (N = 27) Y (N = 62)
O (N = 45) NB (N = 36)
L (N = 35)
H (N = 36)
D
P
C
20.82a 4.42ab 12.88ab 1.34a
20.52a 4.36a 12.74a 1.25a
20.47a 4.43 12.68 1.27
20.38b 4.50 12.98b 1.32b
21.07a 4.40 13.43a 1.64a
** * * †
* NS † †
* NS *** ***
20.15a 4.31a 12.52a 1.27a
21.43b 4.66b 13.15b 1.51b
20.99b 4.44 12.97 1.41
20.75ab 4.42 12.06c 1.07c
Diets: Low fat (LF), High fat saturated (HFS), High fat oats (HFO), High fat linseed oil (HFL). Parity: Young i.e. parity 2–4 (Y), Old i.e. parity 5 or more (O). Piglet category: Newborn (NB), one day old lightest in litter (L), heaviest in litter (H). a,b,c Means in the same row with different superscripts differ at significance indicated. a Significance: NS = P N 0.05; † = P b 0.10; *P b 0.05; **P b 0.01; ***P b 0.001.
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Body energy reserves at birth are present as glycogen and fat because the newborn has a very low capacity to oxidise protein before 5–7 days of life (Marion and Le Dividich, 1999). Total body glycogen stores range from 30 to 35 g per kg, of which 89% are located in the muscle, but after birth the rapid depletion of glycogen stores (Elliot and Lodge, 1977) starts. The overall values for percentage composition of piglets at birth agree well with the values reported in the literature (Elliot and Lodge, 1977; Jean and Chiang, 1999) for the same age. Brooks and Davis (1969) report values for moisture, protein and ether extract (EE) of 78.5%, 12.5% and 2.5% at 48 h that agree in average with those presented herein (Table 4), except for EE being higher. In piglets aged 6 days (Van den Brand et al., 2000) the body composition was 248, 74.1 and 140 g/kg of DM, CF and CP, respectively, and after two weeks these figures were increased, in average to 327.27 kg of DM, 138.6 g/kg CF and 155.1 g/kg CP, corresponding to relative increments of 31.96%, 87.04% and 10.78%. Indeed the inverse relationship between moisture and lipid and the general increase in lipid and decrease in water as body weight increases is well known (Landgraf et al., 2006). Once absorbed, fat is deposited or oxidized. In our study the carcass fat content was increased from NB piglets by 26% in L and by 58% in H piglets in 24 h. Adipose tissue LPL, an enzyme playing a key role in the regulation of fat storage, undoubtedly contributes to this remarkable capacity of pigs to deposit large amounts of fat soon after birth (Le Dividich et al., 1997). Indeed its activity is already high at birth, being similar to that found in 33 kg pigs (Rinaldo and Le Dividich, 1991) and the increase found at 24 h is quite similar to that found during the same period by Le Dividich et al. (1997). The incorporation of fat into the diet during gestation does not affect piglet body measurements; the present data provide additional information on the depletion of energy reserves. The decline in muscle glycogen during the immediate postnatal period emphasizes the importance to the neonatal piglet of an early external energy source. It would appear that sow dietary fat has relatively little effect on progeny although the fatty acid composition in piglets should be determined. References Allee, G.L., Romsos, D.R., Leveille, G.A., Baker, D.H., 1972. Metabolic consequences of dietary medium-chain triglycerides in the pig. Proc. Soc. Exp. Biol. Med. 139, 422. Azain, M.J., 1983. Effects of adding medium-chain triglycerides to sow diets during late gestation and early lactation on litter performance. J. Anim. Sci. 1 (71), 3011–3019. Bach, A.C., Babayan, V.K., 1982. Medium-chain triglyceride: an update. Am. J. Clin. Nutr. 36, 950. Benevenga, N.J., Steinman-Goldsworthy, J.K., Crenshaw, T.D., Odle, J., 1989. Utilization of medium-chain triglycerides by neonatal piglets: I. Effects on milk consumption and body fuel utilization. J. Anim. Sci. 67, 3331. Brooks, C.C., Davis, J.W., 1969. Changes in the perinatal pig. J. Anim. Sci. 29, 325. Canario, L., Tribout, T., Thomas, F., David, C., Cogué, J., Herpin, P., Bidanel, J.P., Père, M.C., Le Dividich, J., 2005. Estimation of the effects of selection using frozen semen, between 1977 and 1998 in Large White population, on body composition and physiological state of the new-born piglet. Journées Rech. Porcine en France 37, 427–434. Calder, P.C., 2002. Dietary modification of inflammation with lipids. Proc. Nutr. Soc. 61, 345–358. Chiang, S.H., Pettigrew, J.E., Clarke, S.D., Cornelius, S.G., 1990. Limits of medium-chain and long-chain triacylglycerol utilization by neonatal piglets. J. Anim. Sci. 68, 1632.
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Short communication
Body composition and muscle glycogen contents of piglets of sows fed diets differing in fatty acids profile and contents G. Pastorelli a,⁎, M. Neil b, I. Wigren b a b
Department of Veterinary Sciences and Technologies for Food Safety University of Milan, 20133 Milan, Italy Department of Animal Nutrition and Management, Swedish University of Agricultural Sciences, Box 7024, S-750 07 Uppsala, Sweden
a r t i c l e
i n f o
Article history: Received 7 March 2008 Received in revised form 13 November 2008 Accepted 21 November 2008 Keywords: Dietary fatty acids Piglet Body composition Glycogen
a b s t r a c t To study the influence of sow dietary fat on piglet body characteristics, multiparous sows were allocated to one of four different dietary treatments: a conventional low fat (3%) diet (LF) and three high fat (6%) diets; high fat saturated (HFS), high fat oats (HFO), and high fat linseed (HFL). All sows were fed the allocated diet from weaning of the preceding litter until the day after farrowing. At farrowing, one liveborn piglet per litter (NB), was sacrificed and dissected immediately after birth. The heaviest (H) and the lightest (L) piglets in the litter were killed and dissected in the same manner at one day of age. Measurement of body length and circumference, organ weight, body chemical composition and muscle glycogen content were determined. Body measurements were adjusted to the mean body weight (1.67 kg). Dietary treatment did not have any significant influence on body components or carcass traits except for lung weight, being lower in HFO and HFL than in LF piglets. Piglet category affected almost all parameters considered, showing the lowest values for NB piglets, except for lung and circumference that were higher in NB than in L and H piglets; and length which was lower in NB than in L piglets. NB piglets had the highest amount of muscle glycogen content, no difference was found between H and L piglets. Dietary treatments influenced piglet chemical composition, showing the highest overall values of dry matter (DM), protein, and fat for the HFL piglets' carcasses. The present data provide additional information on the depletion of energy reserves; it would appear that sow dietary fat has relatively little effect on progeny since only body chemical composition was significantly influenced by HFL diet. © 2008 Elsevier B.V. All rights reserved.
1. Introduction Dietary fat is important as a concentrated energy source. Feeding fat to sows during late gestation and into lactation may improve piglet survival (Seerley et al., 1974; Pettigrew, 1981). Fat is also important as a source of specific compounds, such as medium-chain triglycerides (MCT) and essential fatty acids (EFAs). MCT have fatty acids 6 to 12 carbon atoms long (Bach and Babayan, 1982) and can be easily utilized by pigs (Benevenga et al., 1989; Chiang et al., 1990; Odle et al., 1991). Improved survival or improved indices of survival have been observed in litters from sows fed ketogenic compounds such as 1,3⁎ Corresponding author. Tel.: +39 0250315758; fax: +39 02 50317898. E-mail address: [email protected] (G. Pastorelli). 1871-1413/$ – see front matter © 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.livsci.2008.11.023
butanediol (Stahly et al., 1985, 1986) and MCT (Rosebrough et al., 1981). Because of the rapid metabolism of MCT to ketone bodies and the ability of ketone bodies to both stimulate lipogenesis and spare carbohydrate use for nonoxidative functions (Allee et al., 1972; Seccombe et al., 1977; Shambaugh, 1985) MCT increased the survival of piglet (Azain, 1983). EFAs are fatty acids which are required by an organism for the maintenance of normal growth and reproduction, are not able to be synthesized by the organism, and are therefore required in appropriate amounts in the diet (Leskanich and Noble, 1999). EFAs and their derivatives have varied biological actions that may have relevance to their involvement in several physiological and pathological processes (reviewed by Das, 2006). Linoleic acid and α-linolenic acid are classified as EFAs.
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Table 1 Composition of experimental diets (as-fed). Diet1
Ingredient, % Barley Wheat Oats High fat oats Soya bean meal Wheat middlings Wheat bran Vitamin and mineral premixe Fat (Akofeed Standard)f Fat (Akofeed Cattle 45)g Monocalcium phosphate Limestone Sodium chloride L-Lysine HCl DL-Methionine L-Threonine Linseed oile Calculated composition, % Dry matter Crude protein Lysine Methionine + Cystine Threonine Crude fat Crude fibre Ca P Metabolizable energy, MJ Analysed composition, % Dry matter Crude protein Crude fat Crude fibre Ash
LFa
HFSb
HFOc
HFLd
14.06 42.25 20.0 – 10.0 5.0 5.0 1.0 0.19 – 1.03 0.95 0.40 0.08 0.01 0.04
46.03 19.17 5.0 – 12.0 5.0 5.0 1.0 – 3.65 1.51 1.02 0.40 0.07 0.07 0.08
1.74 33.48 – 40.0 10.1 5.0 5.0 1.0 0.94 – 1.05 1.19 0.40 0.04 0.04 0.02
– 33.8 20.0 20.0 10.5 5.0 5.0 1.0 0.85 – 1.32 1.00 0.40 0.05 0.07 0.02 1.00
88.6 15.3 0.70 0.56 0.52 3.0 4.8 0.80 0.65 12.3
88.4 15.0 0.72 0.59 0.60 6.0 4.6 0.90 0.75 12.8
89.6 15.4 0.71 0.64 0.53 6.0 6.4 0.90 0.65 12.6
89.4 15.4 0.71 0.63 0.52 6.0 6.0 0.87 0.71 12.6
89.0 15.5 3.2 4.6 5.5
88.7 16.1 5.5 4.5 6.0
90.9 16.0 5.9 5.3 5.8
89.3 15.3 4.9 6.5 6.2
1
Diets were calculated to meet nutrient requirements according to Swedish recommendations (Simonsson, 1994) of breeding sows. aLF = low fat, bHFS = high fat saturated, cHFO = high fat oats, dHFL = high fat linseed oil (without linseed oil). e Contributed per kilogram of complete feed; 10,000 IU vitamin A, 1,000 IU vitamin D3, 60 mg vitamin E, 2 mg vitamin K3, 2 mg vitamin B1, 4 mg vitamin B2, 3 mg vitamin B6, 0.02 mg vitamin B12, 0.25 mg biotin, 1.5 mg folic acid, 20 mg niacin, 15 mg d-pantothenic acid, 0.30 mg I, 0.30 mg Se, 40 mg Fe, 0.01 mg Co, 10 mg Cu, 0.41 mg Mg, 20 mg Mn, 70 mg Zn. f Source of unsaturated fatty acids. g Source of saturated/monounsaturated fatty acids.
Although linolenic acid and linoleic acid require the same enzymes for conversion into longer-chain essential fatty acids, they belong to the omega-3 and omega-6 fatty acids series, respectively, which cannot be inter-converted. The fatty acid elongation pathways are not present in the foetus (Li et al., 2000), implying that the unborn piglet is dependent on maternal supplies. Lipid supply, particularly EFA and long chain polyunsaturated is shown to affect neural development and function (Uauy and Castillo, 2003). Moreover, EFAs are also precursors for the prostaglandins and eicosanoid mediators, which play an important role in reproduction (Ziecik et al., 2000) and immunity (Calder, 2002). Therefore dietary supplementation with fat to enhance energy and EFA supply during the late gestation and lactation may be an effective way of increasing the EFA content in piglet tissue and the EFA level of colostrum and milk, thereby improving the viability of piglets (Gu and Li, 2003) and the
immune status of the neonate (Leskanich and Noble, 1999). Differences in placenta development may cause differences in piglet storage of glycogen (Wu et al., 2006). The stored glycogen provides energy during the birth process and after birth, especially in the period before first colostrum intake. In this way, the piglet becomes more able to respond to the stressful extra-uterine environment and has a bigger chance to survive (Le Dividich et al., 2005). Body glycogen is the major energy reserve of the newborn pig, ranging between 30 and 38 g/kg BW. However, glycogen stores are rapidly depleted after birth (Elliot and Lodge, 1977). The total amount of fat in the newborn pig is very low, ranging from 10 to 20 g/kg BW. Selection of pigs for reduced carcass fatness has resulted in pigs that are leaner at birth (Herpin et al., 1993) and have lighter weight livers and less liver glycogen (Canario et al., 2005). Another factor of great importance for piglet survival is the body weight, since mortality is higher in lighter piglets (Rydhmer, 1992). We hypothesised that the glycogen stores in newborn piglets will be depleted faster in light piglets than in heavy piglets and that there might be differences regarding composition or relative organ sizes. The purpose of this study was to evaluate the effect of level and type of fat in diets for pregnant sows on the typical newborn piglet body composition, proportional organ size
Table 2 Fatty acid composition of feed. Fatty acids, g/ 100 g fatty acids C10:0 C12:0 C14:0 C16:0 C18:0 C18:1 C18:2, n − 6 C18:3, n − 3 C20:0 C20:1 C20:2 C22:0 C22:1 C24:0 C20:4, n − 6(Aa) SFAc MUFAc PUFAc n − 6/n − 3 a
Dieta LF
HFS
HFO
HFL
0.04 0.29 0.30 18.40 1.72 20.76 47.07 4.05 0.22 0.66 0.09 0.23 0.15 0.19 Ndb 21.16 21.57 51.21 11.6
0.21 1.68 1.20 32.02 4.14 24.91 27.75 2.38 0.29 0.36 0.07 0.19 0.08 0.16 0.18 39.89 25.35 30.20 11.6
0.05 0.34 0.29 16.29 3.10 36.29 35.47 2.67 0.29 0.69 0.06 0.20 0.10 0.16 0.02 20.72 37.08 38.52 13.3
0.0 0.0 0.2 13.1 2.2 32.2 34.8 13.2 0.3 0.7 0.1 0.0 0.2 0.1 0.2 15.9 33.1 48.3 2.7
LF: a conventional Low Fat 3%. HFS: High Fat Saturated. HFO: High Fat Oats. HFL: High Fat Linseed. The diets LF, HFO and HFL contained a fat blend intended for pigs (Akofeed Standard), with a mono-/polyunsaturated profile, whereas the HFS diet contained a fat blend intended for cattle (Akofeed Cattle 45). The HFO and HFL diets also included the high-fat oats cultivar Matilda (10% fat, C18:1 dominating) as a fat source. To the HFL diet linseed oil (3.9% C16:0, 3.4% C18:0, 11.8% C18:1, 15.3% C18:2, 63.7% C18:3) was added by hand at feeding time, as a top-dress at a rate of 1% by weight. b Nd: Not detected. c SFA: Saturated fatty acids. MUFA: Monounsaturated fatty acids. PUFA: Polyunsaturated fatty acids.
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and muscle glycogen, and to compare with the extremes regarding body weight in the same litter at one day of age. 2. Materials and methods The piglets in the present study were collected from 38 litters. Sows were allocated to one of four experimental diets (Table 1) The fatty acid composition of the diets is shown in Table 2. The experimental procedures were carried out under the Swedish Animal Welfare Act 1988:534 and were approved by the Uppsala Ethics Board for Animal Experimentation, Sweden (No. C 81/1). At farrowing, one representative liveborn piglet per litter, usually the 3rd piglet born (NB), was selected immediately after birth for organ and tissue sampling, and body measures. This piglet was not allowed to suckle. The piglet was rendered unconscious by carbon dioxide gas, then weighed and bled to death. Body length was measured from snout to the tail base; circumference was measured around the chest immediately behind the fore legs. The heart, liver, lungs, stomach with content, spleen, kidneys (left and right weighed together with the respective adrenals), and the large and small intestines (weighed together), were anatomically dissected and weighed. Samples of longissimus muscle were collected and frozen immediately in liquid nitrogen following dissection and stored at −80 °C. Carcasses were stored in − 20 °C until chemical analysis. At the age of 20–36 h, the piglets that were heaviest (H) and lightest (L) in the litter at first handling after birth were killed and dissected in the same manner as the newborns. 2.1. Analytical procedures Samples of feed from each batch and homogenized samples of piglet carcasses were analysed in duplicates. The contents of dry matter (DM) and ash was determined by drying the samples in 103 °C for 16 h, and by ashing in 550 °C for 3 h, respectively. The content of nitrogen was analysed according to Kjeldahl (Nordic Committee on Food Analysis, 1976) using a Kjeltec (Tecator 2460) and crude protein (CP) was calculated as
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N× 6.25. The content of crude fat (CF) was analysed according to the Official Journal of European Communities (1984) using a Tecator-equipment (hydrolyse unit 1047 and extraction by Soxtec HT6). In feed samples, crude fibre was analysed by the short method as described by Jennische and Larsson (1990). Dietary fatty acids were analysed by extracting the lipids using hexane and isopropanol (Nourooz-Zadeh and Appelqvist, 1988). Methylation of fatty acids were according to Sukhija and Palmquist (1988) and separation was made with a temperature-programmed gas chromatograph (Chrompack CP9001 GC, Chrompack, The Netherlands) equipped with a split injection system and capillary column (Chrompack CP-sil 88, 50 m ×0.25 i.d.). Identification of individual fatty acids was based on standards (Larodan Fine Chemicals, Sigma Chemical Co, Malmö, Sweden) and published data (Precht and Molkentin, 1996). Glycogen concentrations were assayed on freeze-dried material from piglets selected from LF, HFS and HFO treatments. Before analysis the freeze-dried muscle pieces were dissected free from connective tissue, blood and fat. After boiling 1–2 mg of the sample for 2 h in 1 ml of 1 M hydrochloric acid to hydrolyze the glycogen, the glucose concentration was determined using a fluorometric method (Lowry and Passoneau, 1973). 2.2. Statistical analysis Data were analysed as a completely randomised design using the GLM procedure of SPSS (2001). The model included dietary treatment, parity and category as fixed effect. All data relative to carcass composition were studied with body weight as a covariate. Parity was classified as old (O) at order of parity 5 or higher, and young (Y) at order of 2 to 4. Data in tables are presented as least square means. Differences among treatment means were compared by least significant difference. 3. Results The effect of diet, sow parity and piglet category on organ weight and body measurements are presented in Table 3.
Table 3 Effect of diet, parity and piglet category on organ weight and body measurements a. Diet
Carcass weight, kg Liver, g Intestinal package, g LKAG, g RKAG, g Stomach, g Spleen, g Heart, g Lungs, g Length, cm Circumference, cm
Parity
Piglet category
SE
LF (N = 27)
HFS (N = 26)
HFO (N = 27)
HFL (N = 27)
Y (N = 62)
O (N = 45)
NB 36
L 35
H 36
1.27a 46.7
1.27a 49.0
1.32a 49.1
1.20b 52.2
1.25 50.4
1.28 48.1
1.29 51.1
1.25 47.4
1.25 49.2
93.7 7.58 7.26 43.0 1.67 12.7 26.7a 38. 7 24.4
94.3 7.25 7.11 46.2 1.57 12.2 25.2ab 38.4 24.0
90.6 7.12 6.98 45.2 1.51 12.3 24.0b 38.6 24.2
83.5 7.21 7.00 44.6 1.44 12.4 23.6b 38.4 25.2
91.4 7.16 6.92 45.7 1.58 12.0 26.0 39.0 24.7
89.6 7.42 7.25 43.9 1.51 12.7 23.7 38.0 24.2
71.9a 6.29a 6.03a 26.5a 1.27a 12.0a 28.6a 37.8a 25.2a
98.1b 7.50b 7.27b 55.9b 1.58b 12.3ab 22.5b 38.9b 24.1b
101.5b 8.08b 7.96b 52.0b 1.78b 12.9b 23.6b 38.8ab 24.1b
Significance b D
P
C
0.01 0.78
† NS
NS NS
NS NS
1.48 0.11 0.11 1.87 0.03 0.11 0.37 0.18 0.16
† NS NS NS NS NS * NS NS
NS NS NS NS NS ** ** ** NS
*** *** *** *** *** * *** * *
Diets: Low fat (LF), High fat saturated (HFS), High fat oats (HFO), High fat linseed oil (HFL). Parity: Young i.e. parity 2–4 (Y), Old i.e. parity 5 or more (O). Piglet category: Newborn (NB), one day old lightest in litter (L), heaviest in litter (H). SE — standard error; LKAG — Left kidney and adrenal gland; RKAG — Right kidney and adrenal gland. abc Means with different superscripts within a row differ at significance indicated. a Values adjusted to the same body weight (1.67 kg). b Significance: NS = P N 0.05; † = P b 0.10; *P b 0.05; **P b 0.01; ***P b 0.001.
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Fig. 1. Glycogen content in muscle from newborn (NB), and one day old piglets born heaviest (H), and lightest (L) in their respective litters.
Except for lung weight there were no significant differences between dietary treatments for any measurement made on body components or carcass traits. Parity had influence on relative lung and heart weight, and body length; piglets of young sows had lighter hearts, heavier lungs and were longer. Piglet category had a significant effect on almost all parameters considered, showing the lowest values for NB piglets. Exceptions were lungs and circumference that were significantly larger in NB than in L or H piglets; and length, which was similar in H and L piglets, with L piglets significantly different from NB piglets. Concerning the muscle glycogen content it was found that NB piglets had the highest amount, and no difference was found between H and L piglets (Fig. 1). Dietary treatments influenced carcass chemical composition (Table 4), showing highest overall values for HFL piglets. Parity had a significant effect on DM only which was higher in piglets of young than of old sows; category showed a significant effect on CF and CP reporting the smallest, intermediate and highest values for the NB, L and H piglets, respectively. 4. Discussion Gestational supplementation with dietary fat did not influence organ weights, except for lung weight being lower in piglets of sows fed diets HFO and HFL, these diets were supplemented with PUFA. This is partly in accordance with Farnworth and Kramer (1988), who fed sows diets supple-
mented with soybean oil (rich in PUFA) or tallow with a cornbased diet as a control, and found no difference between diets in relative weights of heart, liver or lungs in the offspring. These data, however, were from 110 days old fetuses i.e. before birth. Poulos et al. (2004) found that maternal CLA consumption decreased relative heart weight in newborn male piglet but no difference was found regarding liver, kidney or lung. No explanation for this response is apparent. We speculate that our results are due to the supply of fatty acids not being the limiting factor during gestation in development and growth of these organs. Glycogen content is not affected by dietary treatments (data not shown). This is in accordance with results of Newcomb et al. (1991), where no significant difference in piglet glycogen content was found due to sow dietary energy source (starch, MCT (medium chain triglycerides) or soybean oil). Differently, Jean and Chiang (1999) found that sow dietary treatments with MCT and coconut oil resulted in higher (P b 0.10) muscle glycogen content in neonatal piglets than did soybean oil. Anyway, coconut oil and MCT especially, have fatty acids with 6 to 12 carbon atoms length, which can readily cross the placenta (Jean and Chiang, 1999; Rosebrough et al., 1981). Furthermore, the soybean oil dietary treatment (Jean and Chiang, 1999) showed glycogen values of 65 mg/g wet muscle which is similar to those found in the present experiment (51 mg/g wet muscle in NB provided a molar weight of 180 g and 20% DM in muscle). The carcass fat content did not differ between sow dietary energy sources (starch, MCT, or soybean oil) in the study by Newcomb et al. (1991), and was similar to our finding, less than 2%. Such a low content implies the lipid to be primarily structural and as such, may be unavailable as an energy source. The reserves of neonatal piglets are depleted quickly if piglets must metabolize glycogen in an attempt to stay warm (Noblet and Le Dividich, 1981; Herpin et al., 2002). Muscle glycogen content mainly supplies energy for muscle movements (English and Morrison,1984) and this explains why newborn piglets, sacrificed immediately after birth, showed significantly higher values than light and heavy piglets. Values obtained indicate that the rate of mobilization of muscle glycogen was similar in heavy and light piglets; in which the content was 11% lower than in newborns. The depletion after 24 h was similar to the finding of Herpin et al. (2002) in thermoneutral conditions, but lower than in the study by Elliot and Lodge (1977), who found a significant (P b 0.05) decrease of about 40% during the first 12 h post partum establishing a pattern of progressive decline over the full 96 h.
Table 4 Effect of diet, parity group and piglet category on carcass chemical composition. Diet (D)
Dry matter Ash Crude protein Crude fat
Parity (P)
Significance a
Piglet category (C)
LF (N = 27)
HFS (N = 26) HFO (N = 27) HFL (N = 27) Y (N = 62)
O (N = 45) NB (N = 36)
L (N = 35)
H (N = 36)
D
P
C
20.82a 4.42ab 12.88ab 1.34a
20.52a 4.36a 12.74a 1.25a
20.47a 4.43 12.68 1.27
20.38b 4.50 12.98b 1.32b
21.07a 4.40 13.43a 1.64a
** * * †
* NS † †
* NS *** ***
20.15a 4.31a 12.52a 1.27a
21.43b 4.66b 13.15b 1.51b
20.99b 4.44 12.97 1.41
20.75ab 4.42 12.06c 1.07c
Diets: Low fat (LF), High fat saturated (HFS), High fat oats (HFO), High fat linseed oil (HFL). Parity: Young i.e. parity 2–4 (Y), Old i.e. parity 5 or more (O). Piglet category: Newborn (NB), one day old lightest in litter (L), heaviest in litter (H). a,b,c Means in the same row with different superscripts differ at significance indicated. a Significance: NS = P N 0.05; † = P b 0.10; *P b 0.05; **P b 0.01; ***P b 0.001.
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Body energy reserves at birth are present as glycogen and fat because the newborn has a very low capacity to oxidise protein before 5–7 days of life (Marion and Le Dividich, 1999). Total body glycogen stores range from 30 to 35 g per kg, of which 89% are located in the muscle, but after birth the rapid depletion of glycogen stores (Elliot and Lodge, 1977) starts. The overall values for percentage composition of piglets at birth agree well with the values reported in the literature (Elliot and Lodge, 1977; Jean and Chiang, 1999) for the same age. Brooks and Davis (1969) report values for moisture, protein and ether extract (EE) of 78.5%, 12.5% and 2.5% at 48 h that agree in average with those presented herein (Table 4), except for EE being higher. In piglets aged 6 days (Van den Brand et al., 2000) the body composition was 248, 74.1 and 140 g/kg of DM, CF and CP, respectively, and after two weeks these figures were increased, in average to 327.27 kg of DM, 138.6 g/kg CF and 155.1 g/kg CP, corresponding to relative increments of 31.96%, 87.04% and 10.78%. Indeed the inverse relationship between moisture and lipid and the general increase in lipid and decrease in water as body weight increases is well known (Landgraf et al., 2006). Once absorbed, fat is deposited or oxidized. In our study the carcass fat content was increased from NB piglets by 26% in L and by 58% in H piglets in 24 h. Adipose tissue LPL, an enzyme playing a key role in the regulation of fat storage, undoubtedly contributes to this remarkable capacity of pigs to deposit large amounts of fat soon after birth (Le Dividich et al., 1997). Indeed its activity is already high at birth, being similar to that found in 33 kg pigs (Rinaldo and Le Dividich, 1991) and the increase found at 24 h is quite similar to that found during the same period by Le Dividich et al. (1997). The incorporation of fat into the diet during gestation does not affect piglet body measurements; the present data provide additional information on the depletion of energy reserves. The decline in muscle glycogen during the immediate postnatal period emphasizes the importance to the neonatal piglet of an early external energy source. It would appear that sow dietary fat has relatively little effect on progeny although the fatty acid composition in piglets should be determined. References Allee, G.L., Romsos, D.R., Leveille, G.A., Baker, D.H., 1972. Metabolic consequences of dietary medium-chain triglycerides in the pig. Proc. Soc. Exp. Biol. Med. 139, 422. Azain, M.J., 1983. Effects of adding medium-chain triglycerides to sow diets during late gestation and early lactation on litter performance. J. Anim. Sci. 1 (71), 3011–3019. Bach, A.C., Babayan, V.K., 1982. Medium-chain triglyceride: an update. Am. J. Clin. Nutr. 36, 950. Benevenga, N.J., Steinman-Goldsworthy, J.K., Crenshaw, T.D., Odle, J., 1989. Utilization of medium-chain triglycerides by neonatal piglets: I. Effects on milk consumption and body fuel utilization. J. Anim. Sci. 67, 3331. Brooks, C.C., Davis, J.W., 1969. Changes in the perinatal pig. J. Anim. Sci. 29, 325. Canario, L., Tribout, T., Thomas, F., David, C., Cogué, J., Herpin, P., Bidanel, J.P., Père, M.C., Le Dividich, J., 2005. Estimation of the effects of selection using frozen semen, between 1977 and 1998 in Large White population, on body composition and physiological state of the new-born piglet. Journées Rech. Porcine en France 37, 427–434. Calder, P.C., 2002. Dietary modification of inflammation with lipids. Proc. Nutr. Soc. 61, 345–358. Chiang, S.H., Pettigrew, J.E., Clarke, S.D., Cornelius, S.G., 1990. Limits of medium-chain and long-chain triacylglycerol utilization by neonatal piglets. J. Anim. Sci. 68, 1632.
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