Effects of supplementing fish oil in the drinking water of dairy cows on production performance and milk fatty acid

composition V. R. Osborne, S. Radhakrishnan, N. E. Odongo, A. R. Hill and B. W. McBride J Anim Sci 2008.86:720-729. doi: 10.2527/jas.2007-0342 originally published online Nov 27, 2007;

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Effects of supplementing sh oil in the drinking water of dairy cows on production performance and milk fatty acid composition1
V. R. Osborne,*2 S. Radhakrishnan,* N. E. Odongo,*3 A. R. Hill, and B. W. McBride*
*Department of Animal and Poultry Science, and Department of Food Science, University of Guelph, Guelph, Ontario, Canada N1G 2W1

ABSTRACT: The objective of this study was to determine the effects of supplementing sh oil (FO) in the drinking water of dairy cows on production performance and milk fatty acid composition. Sixteen multiparous Holstein dairy cows (741 84 kg of BW; 60 2.3 d in milk, mean SD) housed in a tie-stall facility were used in the study. The study was conducted as a completely randomized design with repeated measurements. The cows were blocked by days in milk and allocated to 1 of 2 treatments: 10 g of menhaden FO/kg of DM topdressed on the total mixed ration (FOT), and 2 g of menhaden FO/L delivered in the drinking water (FOW). The trial lasted for 5 wk: a 1-wk pretreatment adjustment period and 4 wk of treatment. The animals were fed and milked twice daily (feeding at 0830 and 1300; milking at 0500 and 1500) and had unlimited access to water. Dry matter intake (21.3 kg/d for FOT vs. 22.7 0.74 kg/d for FOW), milk yield (38.2 kg/d for FOT vs. 39.5 1.9 kg/d for FOW), and water intake (101 L/d for

FOT vs. 107 4.4 L/d for FOW) were not affected by treatment. The mode of delivery of FO had no effect on milk fat percentage, but milk fat percentage declined linearly with time. The fatty acid contents of 7:0; 8:0; 9:0; 10:0; 12:0 in the milk of FOT cows were lower than for FOW cows, whereas 18:1 trans-12; 18:1 trans-13 and 14; 18:1 trans-16; and trans-9, trans-11 plus trans10, trans-12 CLA were greater for FOT than for FOW. The contents of 24:1 in the milk of FOW cows were 48% greater than for FOT cows, although the concentrations were low in both groups. There was a tendency for the contents of 14:0 and 22:5n-6 to be greater in FOW cows than FOT cows and for the contents of iso-18:0 to be lower for FOW cows than for FOT cows. Although it appears that the amount of FO added in the study did not bypass the rumen as hypothesized, these results suggest that drinking water can be an alternative for supplementing FO to dairy cows without decreasing feed or water intake relative to cows fed FO in the diet.

Key words: sh oil, drinking water, milk fatty acid, dairy cow 2008 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2008. 86:720729 doi:10.2527/jas.2007-0342

INTRODUCTION
Water is the most important nutrient for dairy cattle. It is required for all biological processes including nutrient transport, digestion and metabolism of nutrients, elimination of waste materials (urine, feces, and respiration), release of excess heat (perspiration), mainte-

1 The authors thank the staff at the Elora Dairy Research Centre, University of Guelph for their technical assistance, Dairy Farmers of Ontario and Ontario Ministry of Agriculture and Food for nancial support, and John K. G. Kramer, Agriculture and Agri-Food Canada, for fatty acid analysis. Fish oil used in this study was generously donated by Omega Protein (Hammond, LA). 2 Corresponding author: vosborne@uoguelph.ca 3 Current address: Agriculture and Agri-Food Canada, Lethbridge Research Centre, Lethbridge, Alberta, Canada T1J 4B1. Received June 11, 2007. Accepted November 19, 2007.

nance of proper uid and acid-base balance, and provision of a uid environment for the developing fetus (Houpt, 1984; Murphy, 1992). Yet, intake of water by lactating dairy cows has been largely overlooked. Ingested water is assumed to equilibrate with ruminal uid (Cafe and Poppi, 1994) although 18 to 80% of ingested water bypassed the rumen (Woodford et al., 1984; Zorrilla-Rios et al., 1990). The transition period can be divided into 2 phases: 5 to 7 d prepartum, characterized by a 30% reduction in DMI (Grummer, 1995), and 0 to 21 d postpartum, when intake increases rapidly. Typically, fats are fed to increase dietary energy density (NRC, 2001), but fat supplementation has other potential benets [e.g., increasing the unsaturated fatty acid (FA) content of milk fat (Middaugh et al., 1988; Stegeman et al., 1992)]. Considering the prepartum reduction in DMI, the rumen bypass potential of water, and advantages of using water as a delivery vehicle for nutrients (Osborne et 720

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al., 2002), use of water as a delivery vehicle for enrichment of n-3 PUFA such as eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) in milk may be warranted. However, there is no report on whether sh oil (FO) supplemented in the drinking water would bypass the rumen and increase the rate of transfer of dietary n-3 PUFA into milk. Conventionally, FO is top-dressed on the total mixed ration (TMR) resulting in its FA being extensively biohydrogenated in the rumen (Griinari and Bauman, 1999; Bauman and Griinari, 2003). We hypothesized that FO in drinking water of cows would bypass the rumen and increase n-3 PUFA in milk. Our objective was to determine effects of supplementing FO in drinking water of dairy cows on production performance and milk FA composition compared with top-dressing FO on the TMR.

Table 1. Ingredients and chemical composition of the diet

Item Ingredient composition Corn silage Alfalfa-grass silage High moisture corn Alfalfa/grass hay Expeller soybean meal1 Soybean meal Corn gluten meal Premix2 Cobalt iodized salt Limestone Sodium bicarbonate Chemical composition DM CP, N 6.25 ADF NDF Calcium Phosphorus Potassium Magnesium Sodium Fat Ash NEl,3 Mcal/kg
1 2

% of DM 24.2 24.2 31.9 3.91 3.95 1.98 6.65 1.37 0.47 0.76 0.61 51.1 17.8 15.4 34.4 1.07 0.53 1.42 0.32 0.41 4.33 7.0 1.65

MATERIALS AND METHODS Animals and Experimental Design

Animals were cared for and handled in accordance with the Canadian Council on Animal Care regulations, and the University of Guelph Animal Care Committee reviewed and approved the experiment and all procedures carried out in the study. Sixteen lactating multiparous Holstein dairy cows (741 84 kg BW; 60 2.3 d in milk, mean SD) housed in a tie-stall facility at the Elora Dairy Research Center, University of Guelph (Guelph, Ontario, Canada) were used in the study. The study was conducted in a completely randomized design with repeated measurements. The cows were offered a TMR (Table 1) for ad libitum intake, allowing for 5 to 10% refusal. The TMR was offered twice daily at 0830 and 1300, and DMI was monitored daily throughout the experiment. The cows were blocked by days in milk and allocated to 1 of 2 treatments: 10 g of menhaden FO [specic gravity (H2O = 1): 0.93; Omega Protein Inc., Reedville, VA] per kg of DM top-dressed on the TMR (FOT), or 2 g of menhaden FO/L delivered in the drinking water (FOW). The trial lasted for 5 wk: a 1-wk pretreatment adjustment period and 4 wk of treatment. The tie-stalls were tted with individual Della stainless steel water bowls (Alfa Laval Inc., Peterborough, Ontario, Canada) equipped with a high-volume valve for regulating water ow, and each bowl was serviced by a positive displacement, volumetric water meter (Model C-700 Bronze, ABB Kent Meters Inc., Mississauga, Ontario, Canada). A CR10X measurement and control module (Campbell Scientic Corporation, Edmonton, Alberta, Canada) was used to collect and store the water meter pulse information, air temperature, relative humidity, and water temperature every 15 min for the entire experimental period. The information stored in the data logger was downloaded daily. A volumetric, medicator delivery system (Dosatron International Ltd., Clearwater, FL) administered the menha-

SoyPlus, West Central, Ralston, IA. Lactating cow premix (as-fed): 11% Ca; 15% P; 12.5% Mg; 2% S; 2% K; 1,200 mg/kg of Fe; 100 mg/kg of I; 1,500 mg/kg of Cu; 3,300 mg/kg of Mn; 5,000 mg/kg of Zn; 36 mg/kg of Co; 22 mg/kg of Se; 550,000 IU/kg of vitamin A; 220,000 IU/kg of vitamin D; and 3,100 IU/kg of vitamin E. 3 Estimated according to the NRC (2001).

den FO in the water. Water meters were calibrated weekly by diverting water to a carboy and measuring the volume of water against the computer pulse count and meter dial. Total water intake from each bowl was also monitored by an additional in-line water meter for validation. The cows had unlimited access to water. The cows were milked in their stalls twice daily at 0500 and 1500. Milk samples were collected from the morning milking and preserved with 2-bromo-2-nitropropane-1-2-diol, and the samples were immediately submitted to the Central Milk Testing Laboratory (Laboratory Services Division, University of Guelph, Ontario, Canada) for compositional analysis. A second set of milk samples without preservative was also collected daily from the morning milking and stored at 70C for milk FA analysis. Blood samples were obtained weekly via puncture of a coccygeal artery or vein before the morning feeding into 10-mL evacuated tubes (Becton Dickinson Vacutainer Systems, Franklin Lakes, NJ). Samples were allowed to stand at room temperature for 30 min and then centrifuged at 2,500 g for 15 min in a refrigerated centrifuge (Beckman, Model TJ-6, Palo Alto, CA) at 4C. Serum was separated and transferred to 7-mL, plastic scintillation vials and stored at 70C until analysis.

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Table 2. Fatty acid composition of the supplemental menhaden sh oil1

Fatty acid 14:0 15:0 iso-15:0 16:0 16:1 cis-7 16:1 cis-9 16:3 16:4 17:0 iso-17:0 ai-17:0 17:1 18:0 18:1 cis-9 18:1 cis-11 18:2 n-6 18:2 cis-9, trans-11 18:2 trans-11, trans-13 18:3 n-6 18:3 n-3 18:4 n-3 20:0 20:1 cis-9 20:1 cis-11 20:2 n-6 20:3 n-3 20:3 n-6 20:4 n-6 20:4 n-3 20:5 n-3 22:0 22:1 cis-11 22:1 cis-13 22:3 n-3 22:4 n-6 22:5 n-6 22:5 n-3 22:6 n-3 23:0 24:1 cis-15 26:0
1

g/100 g of fatty acid 6.56 0.66 0.16 18.4 0.31 8.92 1.34 0.23 0.62 0.74 0.34 0.23 3.97 9.00 3.24 1.37 0.02 0.24 0.25 1.17 2.46 0.38 0.40 1.49 0.26 0.21 0.28 0.83 1.41 10.44 0.28 0.11 0.21 0.03 0.19 0.14 2.06 10.95 0.12 0.38 0.02

3.002; AOAC, 1990), OM by ashing at 500C for 16 h (method 942.05; AOAC, 1990), and CP by use of Leco FP 428 nitrogen analyzer (Leco Corporation, St. Joseph, MI; method 4.2.08; AOAC, 1990). The samples were also analyzed for ether extract (method 920.39; AOAC, 1990), ADF (method 973.18c; AOAC, 1990), and NDF (Van Soest et al., 1991) using -amylase (Sigma No. A3306, Sigma Chemical Co., St. Louis, MO), sodium sulte, and correction for ash concentration adapted for an Ankom 200 ber analyzer (Ankom Technology, Fairport, NY). The Ca, P, K, Mg, and Na were determined by inductively coupled plasma spectroscopy (method 945.46; AOAC 1990). Milk samples were analyzed for true protein and fat using a near infrared analyzer (MilkoScan 4000, Foss NIR Systems Inc., Hillerod, Denmark).

Analysis of FA Composition
Frozen serum and milk samples were thawed in a water bath at 38C, and the milk samples were pooled weekly according to method 925.21 (AOAC, 1990). Lipids were extracted with chloroform:methanol:water in the ratio of 1:1:0.9 using a modication of the method described by Bligh and Dyer (1959). The total lipids were methylated using NaOCH3 as the catalyst (CruzHernandez et al., 2004) and analyzed directly by GLC. Fatty acid methyl ester analysis was performed using a Hewlett-Packard Model 5890 Series II GLC (Palo Alto, CA) equipped with a split/splitless injector at 250C, a ame ionization detector at 250C, and a CP Sil 88 column (100 m 0.25 mm, 0.2- m lm thickness, Varian Inc., Mississauga, Ontario, Canada). Hydrogen was used as the carrier gas at a constant ow rate of 1 mL/min. The temperature of the GLC oven was set to 45C for 4 min, increased at 13C/min to 175C and held for 27 min, and again increased at the rate of 4C/ min to a nal temperature of 215C and held for 35 min (Kramer et al., 2001; Cruz-Hernandez et al., 2004). Agilent Technologies ChemStation software (Version A.10, Palo Alto, CA) was used for data analysis. A 1L sample was injected using the splitless mode set at 0.3 min. Peaks were identied by comparison of retention times with FA methyl ester standards (GC 463, UC59M, 21:0, 23:0, and 26:0, Nu-Check-Prep Inc., Elysian, MN). The individual isomers of 18:1 were determined as follows: the temperature of the GLC oven was maintained at 45C for 4 min, increased to 163C at a rate of 13C/min and held for 40 min, and again increased at the rate of 4C/min to a nal temperature of 215C and held for 23 min. Peaks of 18:1 isomers were identied by comparison to published data (Kramer et al., 2002; Shingeld et al., 2003; Loor et al., 2004). Fatty acids are reported as g/100 g of total FA.

The data are expressed as the percentage of total fatty acids.

Chemical Analysis
Representative samples of the TMR were collected from the data ranger (American Calan, Northwood, NH) 3 times/wk before top-dressing the FO, pooled weekly and stored at 20C until analyzed. Orts from individual cows were weighed each morning before feeding, and representative samples were collected and stored at 20C until analyzed. The TMR and orts were analyzed for DM by oven-drying at 60C for 48 h (method 930.15; AOAC, 1990). The dried TMR samples were then ground to pass through a 1-mm screen (Wiley Mill, Arthur H. Thomas, Philadelphia, PA), and the chemical composition was determined in duplicate at a commercial laboratory (Agri-Food Laboratories, Guelph, Ontario, Canada). Analytical DM content was determined by oven-drying at 135C for 2 h (method

Calculations and Statistical Analysis

The transfer coefcient of DHA, docosapentanoic acid (DPA), or EPA was calculated as

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Table 3. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on DMI, water intake, and milk yield and composition1
Fish oil Item DMI, kg/d Water intake, L/d Milk yield, kg/d Milk fat, % Milk fat yield, g/d Milk protein, % Milk protein yield, g/d
1

FOT 21.3 101 38.2 2.25 793 2.87 978

FOW 22.7 107 39.5 2.40 965 2.78 1,114

SEM 0.74 4.5 1.93 0.238 128 0.056 75

Treatment P-value 0.19 0.31 0.65 0.68 0.42 0.35 0.29

P-value for time Linear 0.97 0.65 0.04 <0.01 <0.01 0.24 0.27 Quadratic 0.64 0.11 0.06 0.15 0.56 0.22 0.96 Cubic 0.06 0.05 0.06 0.10 0.28 0.44 0.15

Means are based on 8 cows per treatment.

FA transfer coefficient =

FA yield in milk (g) . FA intake (g)

Milk FA yield was estimated as described by Glasser et al. (2007), whereas FA intake was calculated as DMI concentration of FA in the diet (g/kg). The ANOVA for averages of DMI, milk yield, and serum and milk FA and FA transfer coefcients were conducted using the MIXED procedure (SAS Inst. Inc., Cary, NC) using the model Yij = + i + j + ij + ij, where Yij = the dependent variable, = the overall mean, i = the effect of treatment (i = 1, 2), j = the effect of time (j = 1, 2, 3, 4), ij = the effect of the treatment time interaction (ij = 1, 2, ....., 8), and ij = the random residual error. The effects of treatment were considered xed, and cow within treatment was included as a random effect. To determine time-dependent changes, and interactions between time and treatment, the effects of treatments over time were evaluated using orthogonal contrasts. Day in milk and pretreatment DMI, milk yield, and milk fat percentage were used as covariates. Effects were considered signicant at a probability of P < 0.05. Differences among treatment means were tested for signicance using Tukeys multiple range test. Data are expressed as mean SEM, which represents the pooled SEM for the model.

cows fed saturated and unsaturated FA supplements (Harvatine and Allen, 2006). Other studies have shown reduced DMI when FO was fed at 3% of ration DM (Donovan et al., 2000; Whitlock et al., 2002). The milk yield prole showed a positive quadratic (P = 0.03) effect with time, whereas milk fat percentage and yield in both treatments decreased linearly (P < 0.05) with time (Table 3). In general, oils with a high degree of unsaturation disturb rumen fermentation and ber digestibility, leading to lower acetate production and milk fat synthesis (Coppock and Wilks, 1991). Pretreatment milk fat percentages were 3.45 vs. 3.59% 0.28, FOT vs. FOW, respectively, whereas the pretreatment milk fat yields were 1,272 vs. 1,534 g 276, FOT vs. FOW, respectively. This milk fat depression is consistent with other studies where FO has been fed (Cant et al., 1997; Jones et al., 1998; Whitlock et al., 2002). High concentrate diets, addition of oil seeds (Middaugh et al., 1988), and other rumen-active fats cause milk fat depression in lactating cows (Sutton, 1989). The protein content and yield in milk were not affected (P 0.15) by either treatment or over time, consistent with previous studies (Jones et al., 2000; Donovan et al., 2000; Whitlock et al., 2002), but contrary to Cant et al. (1997) who observed a reduction in milk protein content when FO, monensin, or both were added to the diet of dairy cows.

RESULTS AND DISCUSSION Milk and Serum FA Composition DMI, Water Intake, Milk Yield, and Milk Components
The FA composition of the menhaden FO is presented in Table 2. Dry matter intake, water intake, milk yield, and milk components are presented in Table 3. No interaction of treatment sampling time was observed; therefore, treatment means across all sampling times are reported. Supplementing FO in the drinking water of dairy cows had no effect (P 0.19) on DMI, water intake, or milk yield compared with top-dressing it on the TMR. Dry matter and water intakes remained relatively uniform throughout the trial. Similar water intake patterns have been reported for lactating dairy Milk and serum FA, 18:1 isomers, and CLA contents (g/100 g of total FA) are summarized in Tables 4 to 10. The major characteristic of ruminant milk fat is the high proportion of saturated FA (Palmquist et al., 1993; see Table 4). The contents of the short-chain saturated FA 7:0, 8:0, 9:0, 10:0, and 12:0 in the milk of FOT cows were lower (P < 0.05) than for FOW cows. Short-chain FA (4:0 to 12:0) are considered products of de novo synthesis within the mammary gland using acetate as the precursor. As mentioned, milk fat content was reduced linearly over time (Table 3). This was associated with altered rumen biohydrogenation characterized by a shift in major biohydrogenation pathways: decreased

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Table 4. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on milk fatty acid contents1,2
Fish oil Fatty acid 4:0 5:0 6:0 7:0 8:0 9:0 10:0 11:0 12:0 iso-13:0 ai-13:0 13:0 iso-14:0 14:0 iso-15:0 ai-15:0 15:0 iso-16:0 16:0 16:1n-9 16:1n-7 iso-17:0 ai-17:0 17:0 iso-18:0 18:0 18:2n-6 18:3n-6 18:3n-3 18:4n-3 19:0 20:0 20:1 20:1 cis-9 20:1 cis-11 20:2n-6 20:3n-6 20:4n-6 20:4n-3 20:5n-3 21:0 22:0 22:1 22:1 cis-11 22:1 cis-13 22:4n-6 22:5n-6 22:5n-3 22:6n-3 23:0 24:1 26:0 Unidentied FA Total CLA3 Total SFA4 Total C4:0 to C12:0 Total C13:0 to C16:0 Total C17:0 to C26:0 Total MUFA5 Total n-6 PUFA6 Total n-3 PUFA7 n-6:n-3 ratio FOT 2.53 0.018 1.28 0.018 0.89 0.023 2.04 0.22 2.49 0.034 0.069 0.15 0.087 8.6 0.20 0.40 0.96 0.28 24.6 0.16 1.35 0.23 0.58 0.57 0.074 8.6 2.37 0.029 0.43 0.10 0.110 0.180 0.037 0.23 0.12 0.051 0.11 0.12 0.10 0.08 0.096 0.052 0.01 0.003 0.07 0.025 0.07 0.10 0.044 0.022 0.012 0.03 8.3 1.49 55.4 9.4 35.4 10.5 31.3 2.77 0.86 3.3 FOW 2.40 0.019 1.28 0.030 1.08 0.039 2.58 0.26 2.97 0.027 0.070 0.19 0.086 10.7 0.21 0.41 1.09 0.27 24.5 0.17 1.31 0.24 0.52 0.58 0.041 7.9 2.46 0.023 0.44 0.07 0.104 0.139 0.024 0.16 0.13 0.058 0.12 0.14 0.14 0.10 0.068 0.053 0.17 0.067 0.11 0.022 0.11 0.16 0.049 0.023 0.017 0.09 7.9 1.00 58.0 10.7 37.6 9.8 29.1 2.93 0.97 3.0 SEM 0.12 0.0020 0.083 0.0020 0.062 0.0029 0.13 0.017 0.12 0.0015 0.0038 0.015 0.0080 0.74 0.017 0.039 0.098 0.021 1.10 0.0079 0.072 0.025 0.040 0.037 0.012 0.48 0.12 0.0034 0.033 0.015 0.0033 0.014 0.0025 0.025 0.011 0.0056 0.012 0.014 0.031 0.010 0.0068 0.0031 0.120 0.045 0.045 0.0028 0.014 0.026 0.0094 0.0015 0.0015 0.049 1.98 0.13 2.07 0.06 0.58 0.18 1.27 0.14 0.11 0.32 P-value 0.46 0.70 0.98 <0.01 0.05 <0.01 0.01 0.11 0.01 <0.01 0.84 0.07 0.95 0.06 0.73 0.83 0.37 0.78 0.95 0.78 0.70 0.83 0.27 0.88 0.08 0.33 0.61 0.24 0.75 0.43 0.26 0.05 <0.01 0.06 0.44 0.40 0.39 0.42 0.31 0.16 0.01 0.86 0.37 0.33 0.07 0.34 0.06 0.17 0.70 0.70 0.03 0.40 0.84 0.13 0.38 0.03 0.34 0.77 0.35 0.44 0.50 0.50 Continued

Table 4 (Continued). Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on milk fatty acid contents1,2
Fish oil Fatty acid Total 18:1 trans Total 18:1 cis
1 2

FOT 10.1 19.2

FOW 10.2 16.9

SEM 0.89 0.34

P-value 0.85 0.21

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment. 3 Total CLA = cis-9, trans-11 CLA; cis-10, trans-12 CLA; trans-9, cis-11 CLA; trans-10, cis-12 CLA; trans-11, trans-13 CLA; trans-9, trans-11 CLA and trans-10, trans-12 CLA. 4 Total SFA = all fatty acids without any double bond (9:0 to 26:0). 5 Total MUFA = all fatty acids with a single double bond (16:1 to 24:1). 6 Total n-6 PUFA = trans-9, trans-12 18:2; cis-9, trans-12 18:2; trans9, cis-12 18:2; 18:2n-6; 18:3n-6; 20:2n-6; 20:3n-6; 22:2n-6; and 22:5n6. 7 Total n-3 PUFA = 18:3n-3; 20:3n-3; 20:4n-3; and 22:4n-3.

formation of 18:1 trans-11 (vaccenic acid) and increased formation of 18:1 trans-10 in the rumen (Griinari and Bauman, 1999) via trans-10, cis-12 CLA as a direct inhibitor of milk fat synthesis in the mammary gland (Bauman and Griinari, 2003). The concentration of total medium-chain FA (13:0 to 16:0) was not (P = 0.34) affected by treatments. However, the concentration of 14:0 was much greater in milk than in serum (10 vs. 0.5 g/100 g in milk and serum, respectively), whereas 16:0 in milk was double that in serum. This was not surprising because half of the 16:0 in milk is synthesized de novo in the mammary tissue, whereas 14:0 arises almost exclusively from de novo synthesis (Mansbridge and Blake, 1997; Wattiaux and Grummer, 2004). Irrespective of the treatments, the concentration of 18:0 in serum was 66% higher than in milk (Tables 4 and 5). Under normal conditions 18:0 is one of the major FA that are transferred without alteration into milk lipid (Duncan and Garton, 1963; Mansbridge and Blake, 1997). The contents of 20:0 and 20:1 in the milk of FOW cows were lower (P < 0.05) than for FOT cows. There was no treatment difference (P 0.60) in the content of 20:0 or 20:1 in serum (Table 5). The contents of nervonic acid (24:1n-9) in the milk of FOW cows were higher (P < 0.05) than for FOT cows although the concentrations were very low in both groups. The concentration of 24:1 in serum was 10 times higher than in milk, but there was no difference (P = 0.33) between treatments. The concentrations of the highly unsaturated FA arachidonic acid (20:4n-6), EPA (20:5n-3), DPA (22:5n-3), and DHA (22:6n-3) in milk were not (P 0.16) affected by treatment (Table 4). Normally, DHA occurs only in trace amounts ( 0.01%) in milk fat of cows fed conventional diets, whereas EPA and DPA are present at about 0.05 and 0.08%, respectively (Wright et al., 2007). By supplementing 2 g of FO/L in the drinking water of dairy cows in the current study, the yields of EPA (P =

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Table 5. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on serum fatty acid contents1,2
Fish oil Fatty acid 14:0 iso-15:0 ai-15:0 15:0 iso-16:0 16:0 16:1n-9 16:1n-7 iso-17:0 ai-17:0 17:0 iso-18:0 18:0 19:0 18:2n-6 18:3n-6 18:3n-3 18:4n-3 20:0 20:1 20:1 cis-7 20:1 cis-9 20:1 cis-11 20:2n-6 20:3n-6 20:3n-3 20:4n-3 20:4n-6 20:5n-3 22:0 22:1 22:1 cis-11 22:1 cis-13 22:4n-6 22:5n-6 22:5n-3 22:6n-3 23:0 24:0 24:1 26:0 Unidentied FA Total CLA3 Total SFA4 Total C14:0 to C16:0 Total C17:0 to C26:0 Total MUFA5 Total n-6 PUFA6 Total n-3 PUFA7 n-6:n-3 ratio Total 18:1 trans Total 18:1 cis
1 2

Table 6. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on eicosapentaenoic acid (EPA), docosahexaenoic acid (DHA), and docosapentanoic acid (DPA) intakes, milk yields, and transfer coefcients1
Fish oil Item Intake EPA, g/d DHA, g/d DPA, g/d Yield EPA, g/d DHA, g/d DPA, g/d Transfer coefcient EPA, mg/g DHA, mg/g DPA, mg/g
1

FOT 0.46 0.25 0.32 0.54 0.28 11.7 0.25 1.10 0.44 2.05 0.67 0.17 14.0 0.154 38.2 0.35 2.84 0.071 0.130 0.079 0.057 0.13 0.07 0.123 1.08 0.034 0.51 1.42 1.02 0.32 0.008 0.028 0.050 0.077 0.084 0.32 0.17 0.54 1.02 0.14 0.034 5.0 0.41 30.4 13.6 16.9 16.3 41.4 5.0 8.3 5.8 8.6

FOW 0.51 0.28 0.35 0.58 0.29 12.6 0.27 1.18 0.43 1.98 0.63 0.18 13.4 0.147 37.6 0.38 2.65 0.068 0.136 0.079 0.055 0.11 0.07 0.120 1.15 0.038 0.52 1.26 0.95 0.32 0.011 0.021 0.042 0.082 0.084 0.33 0.15 0.55 0.95 0.12 0.031 5.1 0.41 30.8 14.6 16.2 16.6 40.7 4.8 8.5 5.9 8.8

SEM 0.048 0.021 0.031 0.033 0.023 0.45 0.052 0.014 0.023 0.14 0.092 0.073 0.57 0.0044 0.90 0.038 0.20 0.0075 0.0084 0.0059 0.0030 0.010 0.0030 0.0064 0.075 0.0047 0.079 0.12 0.11 0.014 0.0030 0.0040 0.0053 0.0082 0.0099 0.050 0.027 0.030 0.11 0.018 0.0031 0.35 0.018 0.12 0.54 0.73 0.57 0.97 0.43 0.70 0.33 0.10

P-value 0.46 0.41 0.60 0.44 0.68 0.16 0.43 0.32 0.95 0.74 0.80 0.94 0.50 0.25 0.64 0.49 0.52 0.80 0.60 0.99 0.58 0.28 0.70 0.71 0.52 0.54 0.89 0.36 0.48 0.75 0.52 0.27 0.33 0.68 0.99 0.94 0.68 0.80 0.66 0.33 0.46 0.67 0.87 0.52 0.27 0.33 0.65 0.64 0.68 0.66 0.81 0.86

FOT 22.3 23.3 4.4 0.67 0.36 0.83 30 16 191

FOW 22.4 23.5 4.4 0.75 0.41 0.94 34 17 214

SEM 0.68 0.72 0.14 0.037 0.019 0.046 1.7 0.9 11.2

P-value 0.89 0.89 0.89 0.12 0.11 0.12 0.16 0.17 0.16

Means are based on 8 cows per treatment.

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment. 3 Total CLA = cis-9, trans-11 CLA; cis-10, trans-12 CLA; trans-9, cis-11 CLA; trans-10, cis-12 CLA; trans-11, trans-13 CLA; trans-9, trans-11 CLA; and trans-10, trans-12 CLA. 4 Total SFA = all fatty acids without any double bond (9:0 to 26:0). 5 Total MUFA = all fatty acids with a single double bond (16:1 to 24:1). 6 Total n-6 PUFA = trans-9, trans-12 18:2; cis-9, trans-12 18:2; trans9, cis-12 18:2; 18:2n-6; 18:3n-6; 20:2n-6; 20:3n-6; 22:2n-6 and 22:5n6. 7 Total n-3 PUFA = 18:3n-3; 20:3n-3; 20:4n-3; and 22:4n-3.

0.16) and DPA (P = 0.17) in milk were 13 and 12% higher, respectively, for FOW compared with FOT (Table 6). Offer et al. (1999) reported transfer coefcients of less than 0.03 for PUFA of chain length C20. Cant et al. (1997) reported transfer efciencies of 0.09 and 0.16 for EPA and DHA, respectively, whereas Wright et al. (1999) reported that the transfer of EPA and DHA declined from 0.27 to 0.07, and 0.34 to 0.11, respectively, as intakes increased from 1.6 to 16 g/d. Palmquist and Griinari (2006) reported that only 10 mg of DHA/g of intake, 57 mg of EPA/g of intake, and 110 mg of DPA/ g of intake appeared in milk compared with the 17, 34, and 214 mg/g of DHA, EPA, and DPA intake, respectively, in the drinking water of the cows in the current study. We had hypothesized that FO supplemented in the drinking water of cows would bypass the rumen and increase the milk FA contents of n-3 PUFA such as EPA and DHA. The transfer coefcients for EPA, DPA, and DHA from diet to milk fat were not different (P 0.16) between treatments (Table 6). It appears the amount of FO added in the current study did not bypass the rumen as hypothesized, and based on milk fat data, it appears both modes of delivery of FO affected rumen function in a similar manner resulting in similar transfer coefcients between treatments for EPA, DPA, and DHA. We however concur with Palmquist and Griinari (2006) that a more precise approach for determining transfer efciencies is to use different levels of FO in the diet and determine the slope of the regression of the amount of FO FA in milk vs. intake instead of using single concentrations of the individual FA in diet and milk. The contents of 18:1 trans-12, 18:1 trans-13 and -14, and 18:1 trans-16 in the milk of FOW cows were lower (P < 0.05) than for FOT cows (Table 7). Fish oils in lactation diets have been shown to modify ruminal bio-

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Table 7. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on milk 18:1 isomers1,2
Fish oil Fatty acid 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1
1 2

FOT 0.036 0.042 0.90 0.74 2.45 3.62 0.78 1.13 0.40 17.8 0.85 0.36 0.142

FOW 0.043 0.043 0.75 0.85 2.82 3.83 0.64 0.95 0.28 15.5 0.96 0.32 0.132

SEM 0.0033 0.0030 0.099 0.088 0.67 0.58 0.034 0.046 0.019 1.11 0.065 0.023 0.0071

P-value 0.18 0.84 0.30 0.37 0.70 0.81 <0.01 0.02 <0.01 0.16 0.27 0.28 0.25

trans-4 trans-5 trans-6 and trans-8 trans-9 trans-10 trans-11 trans-12 trans-13 and trans-14 trans-16 cis-9 cis-11 cis-12 cis-13

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment.

hydrogenation and increase 18:1 trans isomers in milk fat (Pennington and Davis, 1975; Kalscheur et al., 1997). It has been suggested that the very long-chain FA of FO inhibit the nal biohydrogenation step to 18:0, thereby maximizing yield of 18:1 trans intermediates (Wonsil et al., 1994; Shingeld et al., 2003; Lee et al., 2005). The concentration of 18:1 cis-9 in milk was highest among the MUFA in both groups. In milk, vaccenic acid was the main 18:1 trans isomer in both treatment groups (Table 7). The concentration of 18:1 trans-11 was higher in milk than in serum (Tables 7 and 8), which was surprising because 70 to 80% of 18:1 trans11 in blood is converted to cis-9, trans-11 CLA in the mammary tissue, which should lead to a depletion of 18:1 trans-11 in milk (Griinari et al., 1998). The higher concentration of 18:1 trans-11 in milk may reect differential esterication or availability of other FA for esterication. Similarly, the concentration of 18:1 trans-10

(Tables 7 and 8) and total 18:1 trans (Tables 4 and 5) in milk were higher than in serum. Differences in saturated and unsaturated FA concentrations between serum and milk (Tables 4 and 5) indicate the existence of desaturation activity in the mammary gland (Chilliard et al., 2000; Voigt and Hagemeister, 2001). The concentration of 18:1 trans-10, 18:1 trans-11, and total 18:1 trans increased quadratically (P < 0.001) with time (data not shown). Several CLA isomers and their precursors were also identied. Generally, the concentration of cis-9, trans11 CLA; trans-7, cis-9 CLA; and total CLA was higher in milk than in serum (Tables 9 and 10). The contents in milk of trans-9, trans-11 plus trans-10, trans-12 CLA in FOW cows were lower (P < 0.05) than for FOT cows (Table 9). All the other CLA identied were not (P 0.08) affected by treatments. The trans-10, cis-12 CLA was present in milk in low concentrations in both

Table 8. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of dairy cows on serum 18:1 isomers1,2
Fish oil Fatty acid 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1 18:1
1 2

FOT 0.042 0.055 0.34 0.39 0.97 2.36 0.56 0.84 0.20 6.70 0.29 0.87 0.60

FOW 0.049 0.062 0.38 0.44 1.05 2.29 0.55 0.86 0.18 6.91 0.28 0.89 0.59

SEM 0.0022 0.0031 0.026 0.026 0.193 0.24 0.034 0.037 0.011 0.37 0.011 0.045 0.048

P-value 0.03 0.11 0.30 0.21 0.78 0.83 0.79 0.74 0.31 0.70 0.78 0.80 0.83

trans-4 trans-5 trans-6 and trans-8 trans-9 trans-10 trans-11 trans-12 trans-13 and trans-14 trans-16 cis-9 cis-11 cis-12 cis-13

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment.

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Table 9. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of lactating dairy cows on milk CLA content1,2
Fish oil Fatty acid cis-9, trans-11 CLA trans-9, cis-11 CLA trans-10, cis-12 CLA trans-11, trans-13 CLA trans-9, trans-11 plus trans-10, trans-12 CLA
1 2

FOT 1.33 0.058 0.022 0.030 0.045

FOW 0.88 0.047 0.016 0.025 0.035

SEM 0.17 0.0069 0.0029 0.0069 0.0017

P-value 0.08 0.30 0.13 0.11 <0.01

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment.

treatments (Table 9). This CLA isomer has been reported to depress milk fat (Peterson et al., 2002). In the current study, milk fat percentage and yield were numerically lower for FOT cows, although not signicantly different from FOW cows (Table 3). The concentration of trans-10, cis-12 CLA was also lower during diet-induced milk fat depression than when comparable depression of milk fat was induced by postruminal trans-10, cis-12 CLA infusions (Bauman and Griinari, 2003), suggesting that other biohydrogenation intermediates may also have contributed to the reduction in milk fat secretion. Shingeld et al. (2003) reported that FO may reduce the ow of trans-10, cis-12 CLA leaving the rumen and suggested that the decrease in milk fat content in response to FO was associated with increased milk fat 18:1 trans-10 concentrations that arose from increased ruminal formation of this biohydrogenation intermediate. However, in an experiment to examine the effect of 18:1 trans-10 on milk fat synthesis, Lock et al. (2007) infused pure 18:1 trans-10 in the abomasum of dairy cows and reported that although 18:1 trans-10 was taken up by the mammary gland and transferred to milk fat, it had no effect on milk fat synthesis even when provided at a dose 10 times greater that the effective dose of trans-10, cis-12 CLA. Pereld et al. (2007) suggested that an increase in milk fat content of trans-9, cis-11 CLA was associated with dietinduced milk fat depression. These authors provided evidence of a role for this isomer in milk fat depression based on the 15% reduction in milk fat yield with abomasal infusion of a CLA enrichment that supplied 5 g/ d of trans-9, cis-11 CLA.

The concentration of vaccenic acid in milk was not different between FOT and FOW (P = 0.81; Table 7). It has been suggested that FO reduces the nal biohydrogenation step where vaccenic acid is converted to stearic acid (Griinari and Bauman, 1999). Similarly, DHA has been shown to promote vaccenic acid accumulation in mixed ruminal cultures when incubated with linoleic acid (AbuGhazaleh and Jenkins, 2004). The effect of FO on the concentration of cis-9, trans11 CLA in milk for both treatments increased quadratically (P = 0.003) with time, consistent with previous studies. Whitlock et al. (2002) reported that the level of cis-9, trans-11 CLA in milk decreased after 14 d when FO and extruded soybeans were fed, whereas AbuGhazaleh et al. (2004) reported that the concentration of cis-9, trans-11 CLA in milk increased until d 21 and then declined thereafter. In the rumen, cis-9, trans-11 CLA is formed primarily from isomerization of dietary linoleic acid (18:2n-6) during the rst step of biohydrogenation (Harfoot and Hazlewood, 1988). In conclusion, there were no differences between the 2 modes of administration of FO (10 g FO per kg of DM top-dressed on the TMR vs. 2 g of FO/L metered in the drinking water) on DMI, water intake, or milk yields. Although it appears that the amount of FO added in the study did not bypass the rumen as hypothesized, these results suggest that drinking water can be an effective alternative for supplementing FO in the diet of dairy cows. Further research is warranted to explore the use of drinking water as a vehicle for supplying nutrients to dairy cows, especially during the transition period when DMI is normally reduced.

Table 10. Effects of top-dressing sh oil on the total mixed ration (FOT) or supplementing it in the drinking water (FOW) of lactating dairy cows on serum CLA content1,2
Fish oil Fatty acid cis-9, trans-11 CLA trans-9, cis-11 CLA trans-10, cis-12 CLA trans-11, trans-13 CLA trans-9, trans-11 plus trans-10, trans-12 CLA
1 2

FOT 0.098 0.025 0.066 0.030 0.168

FOW 0.096 0.031 0.079 0.029 0.161

SEM 0.0075 0.0024 0.0054 0.0072 0.0059

P-value 0.87 0.07 0.12 0.61 0.66

The data are expressed as the percentage of total fatty acids. Means are based on 8 cows per treatment.

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