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J. Dairy Sci. 92:55235533 doi:10.3168/jds.2008-1867 American Dairy Science Association, 2009.

Isolipidic additions of fat from corn germ, corn distillers grains, or corn oil in dairy cow diets1
M. M. Abdelqader, A. R. Hippen,2 K. F. Kalscheur, D. J. Schingoethe, and A. D. Garcia
Dairy Science Department, South Dakota State University, Brookings 57007

ABSTRACT

Key words: corn germ, distillers grain, milk fat, cow


INTRODUCTION

Eight multiparous and 8 primiparous Holstein cows were used in a replicated 4 4 Latin square design with 4-wk periods to determine the effects on dairy cow performance of feeding corn germ (CG) compared with dried distillers grains with solubles (DDGS) or corn oil (CO). Four isolipidic dietary treatments were formulated: a control diet, a 14% corn germ diet (CGD), a 30% dry distillers grains with solubles diet (DGD), and a 2.5% corn oil diet (COD). All diets were formulated to contain 6.0% fat, with the fat in the control diet provided by a ruminally inert fat source. Dry matter intake was decreased by feeding the COD compared with the CGD; however, no difference in dry matter intake was observed among the control diet, the DGD, and the COD. Dietary treatments had no effect on milk yield, energy-corrected milk, or 4% fat-corrected milk. Feeding CG had no effect on milk fat percentage when compared with the control diet; however, milk fat percentage tended to decrease with DDGS and decreased with CO when compared with the CGD. Milk protein percentage decreased when cows were fed the COD compared with the control diet. Feeding CO tended to decrease milk fat yield compared with CG; however, dietary treatments had no effect on milk protein and lactose yield. Feed efficiency was not affected by dietary treatments and averaged 1.55 kg of energy-corrected milk/kg of dry matter intake. Feeding DDGS and CO increased the concentration of vaccenic and conjugated linoleic acid in milk fat. Concentrations of monounsaturated and polyunsaturated fatty acids in milk were increased in response to feeding the 3 corn coproducts. Fat from CG appears to be relatively protected in the rumen when compared with that from DDGS and CO and therefore will not affect the production of milk fat to the degree of the more available fat in DDGS and CO.
Received November 4, 2008. Accepted July 29, 2009. 1 Published with the approval of the director of the South Dakota Agricultural Experiment Station as Publication Number 3629 of the Journal Series. 2 Corresponding author: arnold.hippen@sdstate.edu

With the rapid expansion of the ethanol industry, alternative dietary fat sources are becoming available in the form of corn coproducts such as corn germ (CG), wet distillers grains, and dry distillers grains with solubles (DDGS). Corn coproducts are attractive feeds to dairy and beef cattle producers because of their energy, protein, and fiber concentrations. Starch removal during ethanol production concentrates other nutrients in distillers grains 3-fold compared with corn grain; therefore, replacing soybean meal or corn with DDGS or CG increases the ether extract (EE) content of the diets. The general perception is that feeding high concentrations of DDGS can cause milk fat depression. Previous research has demonstrated that feeding DDGS or wet distillers grains at concentrations up to 20% of diet DM had no effect on milk fat percentage (Powers et al., 1995; Anderson et al., 2006; Kleinschmit et al., 2006), particularly when diets contained sufficient amounts of forage fiber. Feeding DDGS at increasing concentrations in diets that contained only 28% total NDF resulted in a linear decrease in milk fat percentage with no adverse effect on milk fat yield (Leonardi et al., 2005). The role of forage fiber in minimizing the effect of oil from DDGS on milk fat content was reported by Cyriac et al. (2005), where DDGS replaced corn silage at increasing concentrations of DM, thus decreasing concentrations of forage fiber. Cyriac et al. (2005) reported a linear decrease in milk fat content as concentrations of DDGS increased in the diet. Griinari et al. (1998) demonstrated that milk fat percentage and yield were decreased when 4% corn oil (CO) was included in a diet containing 14.8% NDF but not when CO was added to a diet containing 32.1% NDF. These data lead to the conclusion that milk fat depression can be prevented when feeding high concentrations of DDGS by providing sufficient physically effective fiber in the diet from forage sources. Recently, Abdelqader et al. (2009) reported that feeding CG at 14% of diet DM had no effect on milk fat

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percentage or yield. Fat content in CG is approximately 20% of DM; therefore, when feeding CG at 14% of diet DM, the amount of fat supplemented from the germ is 40% greater than that provided from DDGS at 20% of diet DM. The form of fat, rather than the concentration of fat, found in CG or DDGS is an important factor in modifying ruminal biohydrogenation processes that could result in differing responses in milk fat synthesis. Therefore, CG, although containing greater concentrations of fat compared with DDGS, may not have as great of a negative effect on milk fat production. The hypothesis for the proposed research was that, because germ is not exposed to the milling processes, the fat contained in the germ would be relatively unavailable in the rumen and would not affect the synthesis and production of milk fat to the degree of the more ruminally available fat in DDGS and CO. The objective of this experiment was to determine the relative effects on lactation performance of feeding equal amounts of fat from CG, DDGS, and CO to lactating dairy cows, including the effect on milk fat and fatty acid composition.
MATERIALS AND METHODS Animals and Diets

corded daily and adjusted to allow for 5 to 10% refusal. Cows were milked 3 times daily at 0600, 1400, and 2100 h and milk yield was recorded at each milking. Cows BW and BCS were recorded at the beginning of the first period and during the last 3 d of each period.
Sampling and Chemical Analysis

Eight multiparous and 8 primiparous Holstein cows (206 80 DIM) were used in a replicated 4 4 Latin square design with 4-wk experimental periods. Cows were blocked according to DIM and dietary treatments were randomly assigned to cows within each block. Four isolipidic dietary treatments were formulated: 1) a control diet that contained 2.5% commercial inert fat (Energy Booster 100; Milk Specialties Co., Dundee, IL) consisting of mostly (85%) saturated fatty acids (SFA); 2) a CG diet (CGD) in which CG (Dakota Germ; Poet Nutrition, Sioux Falls, SD) was included at 14% of diet DM; 3) a DDGS diet (DGD) in which DDGS (Glacial Lakes Energy LLC, Watertown, SD) was included at 30% of diet DM, and 4) a CO diet (COD) that contained 2.5% CO (Glacial Lakes Energy LLC) on DM basis (Table 1). All diets were formulated to be similar in CP (18.2%), NDF (33.2%), and EE (6.0%) contents (Table 1). To minimize the effects of varying protein sources, a high-protein, low-fat distillers grains (DakotaGold HP; Poet Nutrition) was included in the control diet, the CGD, and the COD. Diets were formulated to meet or exceed NRC (2001) nutrient requirements for cows producing 50 kg of milk per d at 29 kg/d of DMI. Cows were housed in a free-stall barn and individually fed once daily (0800 h) for ad libitum intake using Calan Broadbent feeder doors (American Calan Inc., Northwood, NH). Amounts fed and refused were reJournal of Dairy Science Vol. 92 No. 11, 2009

Samples of alfalfa hay, corn silage, concentrates, and complete diets were collected weekly. Corn silage and diet samples were stored at 20C until analysis. Alfalfa hay and concentrates samples were stored at room temperature. Feed samples were made into composites by period and dried at 55C in a forced-air oven (style V-23; Despatch Oven Co., Minneapolis, MN) for 48 h and then ground through a 2-mm screen using a Wiley Mill (model 3; Arthur H. Thomas Co., Philadelphia, PA). Samples were reground through a 1-mm screen (Brinkman ultracentrifuge mill; Brinkman Instruments Co., Westbury, NY). Subsamples of the feed composites were dried at 105C for 4 h to determine DM. Composites were analyzed according to the AOAC methods for CP (AOAC, 2002; method 2001.11), EE (AOAC, 2002; method 920.39), and ash (AOAC, 2002; method 942.05). Composites were analyzed sequentially for NDF using heat-stable -amylase and sodium sulfite (Van Soest et al., 1991), and ADF (Robertson and Van Soest, 1981) using an Ankom200 fiber analyzer (Ankom Technology, Fairport, NY). Energy values were calculated according to NRC (2001) as described by Abdelqader et al. (2009). Milk samples from each of the 3 daily milkings were collected for 3 consecutive d at the end of each feeding period. Samples were mixed by gentle inversion and composited in volumes corresponding to the respective milking for each cow on sampling day. Two milk aliquots were obtained from each milk sample; one aliquot was sent to Heart of America DHI laboratory (Manhattan, KS) for compositional analysis during which fat, true protein, and lactose were determined using mid-infrared spectroscopy (Bentley 2000 Infrared Milk Analyzer; Bentley Instruments, Chaska, MN; method 972.16; AOAC, 2002). Concentrations of MUN were determined using chemical methodology based on a modified Berthelot reaction (ChemSpec 150 Analyzer; Bentley Instruments) and SCC was determined using a flow cytometer laser (Somacount 500; Bentley Instruments; method 975.16; AOAC, 2002). The second milk aliquot was stored at 20C for fatty acid analysis. Samples of blood from the coccygeal vein were collected into evacuated tubes (Becton Dickinson and Co., Franklin Lakes, NJ) containing K-EDTA approximately 3 to 4 h after feeding on 3 consecutive days at the end of each period. Samples were immediately placed on

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Table 1. Ingredient and chemical composition (% of DM, unless otherwise noted) of dietary treatments Diet1 Item Ingredient Alfalfa hay Corn silage Ground corn Soybean meal, 44% solvent Soybean hulls DDGHP2 Ruminally inert fat3 Corn germ DDGS Corn oil Vitamin and mineral mix4 Di-calcium phosphate Limestone Salt Vit-E5 Chemical composition6 DM, % CP NDF Forage NDF ADF NFC7 Ether extract Ca8 P8 Mg8 NEL,9 Mcal/kg
1

Control 25.0 30.0 12.0 1.2 11.0 16.3 2.5 0.0 0.0 0.0 0.30 0.70 0.40 0.50 0.10 47.7 (2.54) 17.6 (0.62) 34.6 (1.71) 22.7 20.5 (0.74) 35.1 (1.48) 6.0 (0.18) 0.84 0.38 0.30 1.62

CGD 25.0 30.0 6.8 1.2 7.5 13.5 0.0 14.0 0.0 0.0 0.30 0.70 0.40 0.50 0.10 48.2 (1.54) 17.8 (0.56) 34.4 (1.18) 22.7 19.7 (0.72) 34.8 (1.38) 5.9 (0.14) 0.86 0.45 0.30 1.61

DGD 25.0 30.0 11.3 1.2 0.0 0.0 0.0 0.0 30.0 0.0 0.30 0.70 0.40 0.50 0.10 47.7 (1.57) 17.8 (0.73) 33.8 (0.85) 22.7 17.4 (0.47) 35.3 (1.78) 5.9 (0.21) 0.83 0.45 0.30 1.61

COD 25.0 30.0 12.0 1.2 11.0 16.3 0.0 0.0 0.0 2.5 0.30 0.70 0.40 0.50 0.10 47.1 (1.93) 17.5 (0.63) 34.4 (0.86) 22.7 20.2 (0.25) 35.8 (1.47) 6.0 (0.16) 0.84 0.38 0.30 1.62

Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet. 2 DDGHP = high-protein dried distillers grains (Dakota GoldHP; Poet Nutrition, Sioux Falls, SD). 3 Energy Booster 100 (Milk Specialties Co., Dundee, IL). 4 Dairy Micro Premix (Land O Lakes, Inc., St. Paul, MN). Contained 10% Mg, 4,783 mg/kg of Fe, 4,857 mg/kg of Cu, 122 mg/kg of Co, 17,793 mg/kg of Mn, 26,556 mg/kg of Zn, 408 mg/kg of I, 144 mg/kg of Se, 2,645,520 IU/kg of vitamin A, 529,104 IU/kg of vitamin D, and 10,582 IU/kg of vitamin E. 5 Vitamin E 20,000 IU. 6 Four samples were used for each analysis; each sample was analyzed in duplicate (SD in parentheses). 7 NFC = 100 (% NDF + % CP + % ether extract + % ash). 8 Values are derived from analysis of a single composite of samples collected from all feeding periods. 9 Calculated using NRC (2001) with chemically determined values of feedstuffs (see Table 2).

ice and transported to the laboratory. Samples were centrifuged (2,700 g) and plasma was collected and stored at 20C until analysis. Plasma was thawed and analyzed for glucose using glucose oxidase (glucose kit, cat. no. G7521; Pointe Scientific, Canton, MI) according to Trinder (1969). The concentration of BHBA in plasma was determined (BHBA kit, cat. no. H758758; Pointe Scientific) following the methods of Williamson et al. (1962). Plasma triglyceride concentration was determined (TG kit, cat. no. T7532; Pointe Scientific) according to the procedure of Fossati and Lorenzo (1982), and total cholesterol was determined (cholesterol kit, cat. no. C7510; Pointe Scientific,) following the procedure of Allain et al. (1974).

Milk, plasma, and feed fatty acids were prepared as butyl esters, and fatty acid esters were analyzed by gas chromatography in a chromatograph equipped with a flame ionization detector and an autoinjector (Hewlett Packard model 6890; Hewlett Packard, Palo Alto, CA) as described by Abdelqader et al. (2009).
Statistical Analysis

Data were analyzed as a replicated 4 4 Latin square using the MIXED procedure of SAS (SAS Institute, 2003). Sources of variation in the model included effects of dietary treatment, experimental period, parity, cows within parity, and residual error. Cow within par-

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Table 2. Chemical composition of corn coproducts and forage components (% of DM, unless otherwise noted) Corn coproduct1 Item DM, % CP NDF ADF NFC2 Ether extract Ca P Mg NEL,3 Mcal/kg
1

Forage DDGHP 92.0 45.2 25.8 6.9 36.0 3.3 0.01 0.40 0.10 1.88 Alfalfa hay 90.9 20.7 31.2 22.1 27.2 2.7 1.39 0.30 0.36 1.38 Corn silage 28.3 7.4 49.5 26.1 35.7 4.6 0.34 0.13 0.17 1.39

CG 94.0 15.8 24.5 6.5 36.0 20.0 0.02 1.13 0.49 2.39

DDGS 91.0 30.9 32.7 8.2 36.0 9.9 0.04 0.70 0.32 1.94

CG = corn germ; DDGS = dried distillers grains with solubles; DDGHP = high-protein dried distillers grains. 2 NFC = 100 (% NDF + % CP + % ether extract + % ash). 3 Calculated using NRC (2001).

ity was designated as a random effect in the model. The statistical model was Yijkl = + Ti + Perj + Ckl + Pl + eijkl, where Yijkl = dependent variable for the ith treatment and jth period on the kth cow within the lth parity; = mean; Ti = treatment effect (i = 1, 2, 3, and 4); Perj = period effect (j = 1, 2, 3, or 4); Ckl = cow effect within parity; Pl = parity effect (primiparous or multiparous); and eijkl = residual (error). The interactions (treatment parity, treatment period, parity period, and treatment parity period) were tested but were not included in the model unless found significant. Data were deleted from one primiparous cow because of poor adaptation to Calan feeding doors and foot problems. Significance was declared at (P 0.05) and a trend was reported if 0.05 < P 0.10. Separation of treatment means was conducted using the Tukey test (Steel et al., 1997) for mean comparison.
RESULTS AND DISCUSSION Dietary Treatments

with the other dietary treatments (Table 3). Diets containing corn coproducts had similar fatty acid profiles with greater concentrations of unsaturated fatty acids (UFA), represented in the greater concentrations of C18:2n-6 and total C18:1 fatty acid.
DMI, Milk Yield, and Milk Composition

Experimental diets contained similar concentrations of nutrients (Table 1), which averaged 17.7 0.73%, 34.3 1.16%, and 6.0 0.17% for CP, NDF, and EE, respectively. Chemical compositions of the CG, DDGS, and high-protein dried distillers grains used in this experiment are presented in Table 2. All dietary treatments contained similar concentrations of total fatty acids, which averaged 5.2 0.24 g/100g of DM. As a result of the experimental design, the control diet contained a greater concentration of SFA compared
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Fats are typically fed to dairy cattle to increase the energy density of the diet and increase milk production (NRC, 2001; Litherland et al., 2005). Despite increasing the energy value of dairy cattle diets, supplemental fat often decreases DMI when total dietary concentrations exceed 6 to 7% of DM (NRC, 2001). In the present study, the EE and total fatty acid concentrations were similar across dietary treatments, which averaged 5.9 and 5.2% of DM, respectively. The DMI of cows fed the DGD was similar to those fed the CGD and the COD; however, cows fed the COD had the least DMI, which was 9% lower than that of cows fed the CGD (Table 4). These results suggest that feeding CO in the free form had a negative effect on feed intake. This decrease in DMI could have been related, in part, to a negative effect of dietary unsaturated fat on rumen fermentation (Jenkins, 1993). Mohamed et al. (1988) reported a negative effect of feeding free oil at 4% of diet DM on feed intake and attributed the response to decreased DM digestibility. On the contrary, Griinari et al. (1998) reported that feeding CO at 4% of DM in a high-forage diet had no effect on DMI when compared with a control diet that contained a saturated fat source. Similarly, Dhiman et al. (2000) reported that feeding soybean oil at 4% of diet DM had no effect on DMI. Consistent with our findings, previous studies have reported that feeding DDGS at approximately 20% of DM had no effect

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Table 3. Fatty acid composition1 of diets (g/100 g of total fatty acid, unless otherwise noted) Diet3 Fatty acid2 Total fatty acid, g/100 g of DM C14:0 C14:1 cis-9 C15:0 C15:1 cis-14 C16:0 C16:1 trans-9 C16:1 cis-9 C17:0 C18:0 C18:1 total C18:2 trans-9, trans-12 C18:2 cis-9, cis-12 C18:3n-6 C18:3n-3 C20:0 C20:1 C20:2 SFA UFA
1 2

Control 5.3 1.66 0.93 0.28 0.01 25.80 0.24 0.85 0.79 20.20 15.99 0.11 23.90 0.93 5.24 0.65 0.21 0.18 48.1 51.8

CGD 5.1 1.14 0.42 0.10 0.01 16.27 0.15 1.14 0.16 3.14 23.51 0.15 42.43 1.04 5.74 0.59 0.25 0.18 21.8 78.3

DGD 5.0 1.11 0.35 0.08 0.01 16.47 0.14 1.02 0.15 2.23 23.88 0.14 44.01 0.96 5.39 0.57 0.24 0.14 21.1 79.1

COD 5.3 1.09 0.35 0.09 0.02 16.99 0.15 1.10 0.13 2.62 24.06 0.16 42.39 1.01 5.47 0.57 0.28 0.18 22.0 78.2

Fatty acids composition of the diet was conducted on TMR samples. SFA = total concentration of saturated fatty acid; UFA = total concentration of unsaturated fatty acid. 3 Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet.

on DMI (Leonardi et al., 2005; Anderson et al., 2006; Kleinschmit et al., 2006). Abdelqader et al. (2009) reported that feeding CG at increasing concentrations of DM resulted in a quadratic response on DMI and that cows fed a 14% CG diet had, numerically, the greatest DMI. The DMI of multiparous cows was greater (P < 0.01) than that of primiparous cows and averaged 25.3 versus 21.5 kg, respectively. Similarly, the intake of CP,

NDF, EE, and total fatty acid were greater for multiparous cows compared with primiparous cows. Feeding CGD, DGD, and COD resulted in an increase in the intake of UFA at the expense of SFA when compared with the control diet. The milk production response to dietary fat supplementation depends on the composition of the basal diet, stage of lactation, energy balance, amount and

Table 4. Least squares means for nutrient intake of dairy cows fed experimental diets Diet2 Item1 Intake, kg/d DM CP NDF Ether extract Total fatty acid SFA, g/d UFA, g/d Total C16, g/d Total C18, g/d C18:0, g/d C18:1, g/d C18:2, g/d C18:3, g/d
a,b c,d

Control 23.2ab 4.0b 8.1cd 1.38 1.24 597a 645b 332a 821b 239a 199b 303b 79

CGD 24.3a 4.3a 8.3d 1.42 1.20 263b 947a 212b 921a 38b 283a 518a 89

DGD 23.7ab 4.2ab 8.0cd 1.40 1.21 256b 960a 213b 930a 27b 290a 537a 76

COD 22.3b 3.9b 7.7d 1.35 1.19 260b 933a 218b 904a 30b 287a 509a 76

SEM 0.72 0.16 0.26 0.04 0.04 12.1 31.3 7.9 29.7 5.3 9.1 16.8 3.2

Means within a row with different superscripts differ (P 0.05). Means within a row with different superscripts tended to be different (0.10 P > 0.05). 1 SFA = saturated fatty acid; UFA = unsaturated fatty acid. 2 Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet.
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Table 5. Least squares means for milk yield, milk composition, and feed efficiency of dairy cows fed experimental diets Diet1 Item Milk, kg/d Fat, % Fat, kg/d Protein, % Protein, kg/d Lactose, % Lactose, kg/d SCS2 MUN, mg/dL ECM,3 kg/d FCM, kg/d ECM/DMI
ac d,e

Control 34.0 3.88a 1.31de 3.24a 1.10 4.78 1.63 3.22 13.70b 35.9 33.2 1.56

CGD 35.2 3.80ab 1.33d 3.19ab 1.12 4.81 1.70 3.53 13.87b 36.8 34.0 1.52

DGD 35.8 3.59bc 1.30de 3.21ab 1.14 4.83 1.73 3.13 14.97a 36.7 33.8 1.58

COD 34.7 3.50c 1.20e 3.15b 1.08 4.81 1.67 3.36 13.15b 34.7 31.9 1.57

SEM 1.50 0.12 0.06 0.06 0.04 0.06 0.08 0.37 0.55 1.45 1.38 0.06

Means within a row with different superscripts differ (P 0.05). Means within a row with different superscripts tended to be different (0.10 P > 0.05). 1 Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet. 2 SCS = log2(SCC/100,000) + 3. 3 ECM = {[0.327 milk yield (kg)] + [12.95 fat yield (kg)] + [7.2 protein yield (kg)]}; Orth (1992).

source of dietary fat, and DMI (Coppock and Wilks, 1991; NRC, 2001). In the present study, the addition of CG, DDGS, and CO had no effect on daily milk production, ECM, feed efficiency, and yield of 4% FCM (Table 5). Previous research indicated that inclusion of DDGS in dairy cow diets at approximately 20% of DM increased or had no effect on milk yield when DDGS was fed at 27% of diet DM (Hippen et al., 2004; Leonardi et al., 2005; Anderson et al., 2006; Kleinschmit et al., 2006). Inclusion of CG at increasing concentrations of diet DM resulted in a curvilinear response in milk yield, and cows fed a 14% CG diet produced the greatest milk yield (Abdelqader et al., 2009). Our results showed that feeding CO caused a slight decrease in milk protein percentage with no effect on milk protein yield when compared with the control diet; however, no difference was observed between the CGD and the DGD. Although milk protein percentage often decreases when supplemental fat is fed to lactating dairy cows (DePeters and Cant, 1992; Wu and Huber, 1994), protein yield generally remains constant or increases (Knapp et al., 1991; Schingoethe and Casper, 1991). In the current study, the decrease in milk protein percentage when cows were fed the COD could be a function of the numeric increase in milk yield when compared with those fed the control diet. Milk from cows fed the DGD had a greater concentration of MUN compared with that from those fed the other treatments. The mean BW and BCS of cows was similar among dietary treatments (Table 6). Feeding the 3 corn coproducts resulted in similar milk NEL and NEM. Although the estimated diet NEL was similar across treatments and
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averaged 1.73 0.01 Mcal/d, it was 6.2% greater than the predicted average of dietary NEL from the NRC (2001) model. Depression in milk fat percentage is a major concern when feeding DDGS at greater concentrations of diet DM; therefore, DDGS is recommended to be included in dairy cow diets up to 20% of diet DM (Schingoethe, 2006). Previous research has suggested that feeding DDGS or wet distillers grains at concentrations up to 20% of DM does not affect milk fat percentage (Powers et al., 1995; Anderson et al., 2006; Kleinschmit et al., 2006). In contrast, Leonardi et al. (2005) reported that milk fat percentage decreased linearly when DDGS were fed at increasing concentrations up to 15% of DM. Abdelqader et al. (2009) reported that feeding CG at 14% of DM had no effect on milk fat percentage and yield and that milk fat content was maximized when cows were fed a 14% CG diet. The amount of fat supplemented from CG at 14% is equal to 2.8% of diet DM, which is 40% greater than the amount of fat provided by feeding DDGS at 20% of diet DM. This led to the conclusion that CG and DDGS may be different in the physical form of their fat content, such that fat from CG might be less available in the rumen with minimal effect on ruminal fermentation and ultimately milk fat percentage. In the current study, cows fed the diets containing corn coproducts consumed similar amounts of UFA. Consequently, changes in milk fat content could be attributed only to differences in the form of fat and effects on the biohydrogenation process in the rumen rather than to the amount or degree of saturation of fatty acids. Our findings indicated that feeding

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Table 6. Least squares means for BW, BCS, and net energy estimations of dairy cows fed experimental diets Diet1 Item BW, kg BW change, kg BCS BCS change Energy Milk NEL,2 Mcal/d Maintenance NEM,3 Mcal/d NEL empty BW change,4 Mcal/d Total NEL,5 Mcal/d Estimated diet NEL,6 Mcal/d Control 707 20.7 3.28 0.09 25.1 11.0 2.0 39.9 1.72 CGD 712 45.4 3.30 0.06 25.6 11.0 3.1 41.5 1.72 DGD 712 60.3 3.26 0.10 25.5 11.0 4.2 38.9 1.74 COD 703 4.0 3.26 0.01 23.9 10.9 0.2 38.8 1.73 SEM 17.2 18.7 0.06 0.07 1.0 0.2 1.3 1.8 0.10

1 Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet. 2 NEL (milk), Mcal/d = Milk yield (kg) (0.0929 fat % + 0.0563 true protein % + 0.0395 lactose %); NRC (2001). 3 NEM (maintenance), Mcal/d = 0.08 BW0.75. 4 NEL (empty BW change) calculated according to NRC (2001). 5 Total NEL (Mcal/d) = [NEL (milk) + NEM (maintenance) + NEL (empty BW change)]. 6 Estimated diet NEL = [Total NEL (Mcal/d)/DMI (kg)].

the CGD had no effect on milk fat percentage and yield when compared with the control diet; however, feeding the COD decreased milk fat percentage and tended to decrease milk fat yield when compared with the CGD. Milk fat percentage and yield from cows fed the DGD were similar to that obtained from cows fed the COD; however, feeding the DGD tended only to decrease milk fat content compared with the CGD. Feeding DDGS resulted in an intermediate response in milk fat percentage and yield when compared with CG and CO. These findings clearly indicate that DDGS may contain small proportions of free oil that interfere with rumen fermentation and alter rumen biohydrogenation, leading to increased concentrations of trans-10 C18:1 and trans-10, cis-12 CLA, which have been demonstrated to be effective inhibitors of mammary gland de novo fatty acid synthesis (Griinari et al., 1998; Baumgard et al., 2000). Similarly, Dhiman et al. (2000) reported that milk fat percentage was decreased when soybean oil was fed in comparison with raw and roasted soybeans. The authors concluded that feeding oilseeds instead of free oil may decrease the potential of inducing milk fat depression. Milk fat percentage was also depressed when cows were fed diets containing soy oil compared with soybeans (Larson and Schultz, 1970; Mohamed et al., 1988). The lack of a depression effect on milk fat percentage when oilseeds are fed can be attributed to the protection of oil in the seed combined with the slow release of encapsulated oil in the rumen (Knapp et al., 1991). The slow release of oil in the rumen can minimize the effect on the ruminal biohydrogenation process, therefore decreasing the accumulation of bio-

hydrogenation intermediates that may cause milk fat depression.


Milk Fatty Acid Profiles

The fatty acid profiles of milk fat are presented in Tables 7 and 8. Feeding the 3 corn coproducts decreased the concentrations of C4:0, C14:0, and C16:0 and increased the concentrations of total C18:1 and cis9, cis-12 C18:2 compared with the control diet (Table 7). Grouping fatty acids based on their origin indicated that the inclusion of CG in dairy cow diets had no effect on the concentrations of de novo synthesized fatty acids when compared with the control diet or the DGD (Table 8). Feeding DDGS and CO, however, decreased concentrations of de novo synthesized fatty acids (<16 carbon) and increased the concentrations of preformed fatty acids (>16 carbon) in milk fat when compared with the control diet. Palmitate in milk fat can be derived either directly from the diet or from de novo synthesis in the mammary gland. The decrease in palmitic acid concentration when cows were fed the 3 corn coproducts can be attributed to lower dietary concentration combined with decrease in de novo synthesis in the mammary gland. Changes in milk fatty acid composition observed in this study are consistent with known effects of unsaturated fat supplements on de novo synthesized fatty acids in the mammary gland (Palmquist et al., 1993). Feeding the 3 corn coproducts increased concentrations of long-chain fatty acids at the expense of both the short- and medium- chain fatty acids and increased concentrations of monounsaturated
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Table 7. Least squares means for relative proportions of fatty acids (g/100 g) in milk fat from dairy cows fed experimental diets Diet1 Fatty acid C4:0 C5:0 C6:0 C7:0 C8:0 C9:0 C10:0 C11:0 C12:0 C12:1 cis-11 C13:0 C14:0 C14:1 trans-9 C14:1 cis-9 C15:0 C15:1 cis-14 C16:0 C16:1 trans-9 C16:1 cis-9 C17:0 C17:1 cis-10 C18:0 C18:1 total C18:2 trans-9, trans-12 C18:2 cis-9, cis-12 C18:3n-6 C18:3n-3 C20:0 C20:1 C20:2 C20:3n-6 C20:3n-3 C20:4n-6 Other fatty acids2
ac 1

Control 3.34 0.31 2.02a 0.09 1.24 0.03ab 2.93 0.06 3.47 0.12 0.09ab 10.60a 0.25a 1.39 0.99a 0.24a 28.79a 0.15 1.42a 0.57a 0.27 10.01b 25.26b 0.03c 3.13c 0.01 0.45a 0.15 0.14b 0.02 0.13 0.04ab 0.19b 1.55b
a

CGD 3.28 0.29 2.02a 0.08 1.25 0.04ab 2.97 0.06 3.46 0.11 0.10ab 10.25ab 0.24a 1.30 0.93ab 0.22ab 24.60b 0.15 1.21b 0.43b 0.20 11.71a 28.63a 0.04bc 3.57b 0.01 0.47a 0.16 0.15b 0.03 0.14 0.05a 0.20a 1.67a
b

DGD 3.24 0.29 1.99ab 0.11 1.23 0.04a 2.85 0.07 3.22 0.11 0.10a 9.83b 0.22b 1.29 0.97a 0.21ab 23.96b 0.16 1.24b 0.39b 0.29 11.37a 29.07a 0.05b 3.53b 0.03 0.42b 0.16 0.21a 0.03 0.14 0.05ab 0.21a 1.61ab
b

COD 3.17 0.28 1.91b 0.09 1.16 0.03ab 2.70 0.07 3.17 0.11 0.09b 9.90b 0.24a 1.33 0.85b 0.21b 24.13b 0.16 1.32ab 0.38b 0.18 10.63ab 29.98a 0.07a 3.74a 0.01 0.46a 0.15 0.22a 0.02 0.13 0.04b 0.20a 1.81a
b

SEM 0.07 0.01 0.05 0.01 0.04 0.01 0.10 0.01 0.11 0.01 0.01 0.17 0.01 0.08 0.03 0.01 0.58 0.01 0.05 0.01 0.05 0.33 0.59 0.01 0.12 0.01 0.02 0.01 0.01 0.01 0.01 0.01 0.01 0.07

Means within a row with different superscripts differ (P 0.05). Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet. 2 Other fatty acids = sum of (unidentified fatty acid, C19:0, C19:1, C22:1, C23:0, C20:5, C22:2, C22:3, C22:5n-6, and C22:5n-3).

and polyunsaturated fatty acids in milk. The decrease in milk fat concentrations of myristic and palmitic acids may represent an improvement in milk fatty acids profile because these fatty acids have been reported to adversely alter blood lipid in human, thus increasing the risk of coronary heart disease (Tholstrup et al., 1994; Vega-Lpez et al., 2006). The concentrations of total trans- and cis-18:1 isomers in milk fat were increased in cows fed the corn coproducts (Table 8), and pronounced changes in the concentrations of specific trans and cis isomers of octadecanoic acid were observed. Feeding the CGD and the DGD resulted in comparable concentrations of trans-10 C18:1, which was greater than that obtained in milk fat from cows fed the control diet; however, cows fed the COD had the greatest concentration of trans-10 C18:1
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compared with those fed the other dietary treatments. Feeding the DGD and the COD increased milk fat concentrations of trans-11 C18:1 when compared with the control diet and the CGD. Abdelqader et al. (2009) reported that feeding CG at 14% of DM resulted in 42% and 50% increases in the concentration of trans-10 and trans-11 C18:1 fatty acids, respectively, when compared with control diet. In the current study, feeding the CGD resulted in only 16% and 20% increases in the concentration of trans-10 and trans-11 C18:1 fatty acids, respectively. Variation in the results could be attributed to differences in diet composition in addition to differences in the EE content of the control diet in both studies. Milk fat concentrations of the predominant CLA isomer, cis-9, trans-11, as well as trans-10, cis-12 CLA, were increased in response to feeding the

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Table 8. Least squares means for relative proportions of C18:1, conjugated linoleic acid (CLA) isomers, and saturation classes of fatty acids (g/100 g) in milk fat from cows fed corn coproducts Diet1 Fatty acid Total trans C18:1 trans-4 C18:1 trans-6 C18:1 trans-9 C18:1 trans-10 C18:1 trans-11 Total cis C18:1 cis-6 C18:1 cis-9 C18:1 cis-11 C18:1 cis-12 C18:1 cis-13 C18:1 cis-15 Other cis-C18:1 Conjugated C18:2 Trans-10, cis-12 Cis-9, trans-11 Other CLA isomers De novo2 Preformed2 C16 carbons2 SCFA3 MCFA3 LCFA3 SFA4 MUFA4 PUFA4
ac 1

Control 1.98 0.01b 0.07 0.22c 0.57c 1.11b 23.29b 0.22c 21.37b 0.54b 0.41b 0.37c 0.08c 0.27b 0.05b 0.53b 0.12b 27.64a 41.91b 30.65a 7.04a 52.02a 41.08b 64.89a 30.18b 4.61c
c

CGD 2.41 0.02ab 0.08 0.32b 0.66b 1.33b 26.22a 0.33b 23.31a 0.61ab 0.69ab 0.54a 0.10b 0.63a 0.06b 0.60b 0.13b 27.04ab 47.59a 26.21b 6.95a 46.82b 46.96a 62.01b 33.13a 5.19b
b

DGD 3.18 0.02ab 0.06 0.34b 0.66b 2.11a 25.92a 0.35b 23.35a 0.60ab 0.82a 0.48b 0.10bc 0.57a 0.09a 0.91a 0.23a 26.09bc 47.97a 25.59b 6.90ab 45.50b 47.26a 60.26bc 33.69a 5.69a
a

COD 3.40 0.03a 0.08 0.43a 0.79a 2.08a 26.59a 0.42a 22.27a 0.62a 0.92a 0.56a 0.14a 0.65a 0.10a 0.94a 0.25a 25.66c 48.67a 25.87b 6.65b 45.42b 48.11a 59.08c 34.69a 5.93a
a

SEM 0.13 0.01 0.01 0.02 0.02 0.10 0.52 0.02 0.48 0.03 0.09 0.01 0.01 0.05 0.01 0.04 0.01 0.44 0.84 0.59 0.14 0.80 0.93 0.73 0.70 0.17

Means within a row with different superscripts differ (P 0.05). Dietary treatments: CGD = corn germ diet; DGD = dry distillers grains with solubles diet; COD = corn oil diet. 2 Fatty acid based on origin: de novo = fatty acid < C16 carbon; preformed = fatty acid > C16 carbon; C16 carbons = fatty acid from both origins. 3 Fatty acid based on chain length: SCFA = short-chain fatty acid (C4 to C9); MCFA = medium-chain fatty acid (C10 to C16); LCFA = long-chain fatty acid (C17). 4 Fatty acid based on degree of saturation: SFA = saturated fatty acid, MUFA = monounsaturated fatty acid; PUFA = polyunsaturated fatty acid.

DGD and the COD compared with the control diet and the CGD. In the present study, feeding the DGD and the COD resulted in 80% and 100% increases, respectively, in the concentrations of trans-10, cis-12 CLA in milk fat compared with feeding the control diet. The decrease in milk fat percentage reported in the current study when cows were fed the COD could be attributed to increased concentrations of trans-10, cis-12 CLA; however, other biohydrogenation intermediates may also be involved in causing milk fat depression (Peterson et al., 2003; Loor et al., 2005). An increase in the concentration of both trans-10, cis-12 CLA and trans-10 C18:1 fatty acids indicates that ruminal lipid metabolism had been modified, which caused a shift in the biohydrogenation pathway toward the formation of trans-10 C18:1 fatty acid. Shingfield et al. (2006) reported that milk fat content was inversely correlated (r = 0.808) with con-

centrations of trans-9, cis-11 CLA in milk and that the changes in this CLA isomer could account for approximately 80% of the variation in milk fat content when cows were fed diets containing fish oil and sunflower oil. Recently, Perfield et al. (2007) demonstrated that the trans-9, cis-11 CLA isomer can decrease concentrations of fat in milk. In the current study, inclusion of DDGS and CO in dairy cow diets resulted in approximately 92% and 107% increases, respectively, in the concentrations of total unidentified CLA isomers compared with the control diet. Furthermore, a significant increase (P < 0.01) in the concentration of a specific unidentified CLA isomer was observed. Feeding the control diet and the CGD, DGD, and COD resulted in milk fat concentrations of this unidentified CLA isomer of 0.04, 0.04, 0.08, and 0.09%, respectively. Previously, we speculated that this unidentified CLA isomer could be the trans-9, cis-11 CLA (Abdelqader et al., 2009).
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Plasma Metabolites and Fatty Acids Composition

Dietary treatment had no effects on plasma concentrations of glucose, triglycerides, and BHBA, which averaged 69.2 1.2, 11.8 1.5, and 6.7 0.5 mg/ dL, respectively. Our results demonstrated that plasma cholesterol concentration (180, 203, 192, and 213 mg/ dL for the control diet and the CGD, DGD, and COD, respectively) was increased (P = 0.02) when cows were fed the COD compared with those fed the control diet; however, no statistical difference was observed among cows fed the 3 corn coproducts. These findings are in agreement with previous studies that reported a positive relationship between serum concentrations of cholesterol and dietary fat content (Palmquist and Conrad, 1978; Drackley et al., 1992). Recently, Bu et al. (2007) reported an increase in plasma cholesterol concentration when cows were fed 4% soybean oil, 4% flaxseed oil, or a mixture of both compared with those fed the control diet. Feeding the CGD, DGD, and COD had no effect on plasma concentrations of C14:0, C16:0, C18:0, total C18:1, and C18:2 fatty acids, which averaged 16 1.5, 314 11.9, 404 15.0, 200 10.4, and 1,428 68.7 g/mL, respectively. Plasma concentrations of trans-11 C18:1 (20, 19, 27, and 30 g/mL for the control diet and the CGD, DGD, and COD, respectively) increased by feeding the DGD and the COD, and cows fed the COD had the greatest plasma concentration of trans-11 C18:1. Trans-11 C18:1 is the precursor for endogenously synthesized cis-9, trans-11 C18:2 CLA isomer via the activity of 9-desaturase in the mammary gland (Griinari et al., 1998). Dietary treatments had no detectable effect on plasma concentrations of trans-10 C18:1, cis-9, trans-11 CLA, and trans-10, cis-12 CLA, which averaged 0.7 0.4, 7.2 1.1, and 2.6 0.29 g/mL, respectively.
CONCLUSIONS

the milk fat concentration of trans-11 C18:1 and cis9, trans-11 CLA. Overall, CG provides an alternative source of fat for energy in diets of lactating dairy cows when fed up to 14% of dietary DM; furthermore, fat in CG is relatively protected and has minimal effects on milk fat production.
ACKNOWLEDGMENTS

The authors express their gratitude to the farm crew at the South Dakota State University Dairy Research and Training Facility (Brookings) for care of the animals. This research was partially supported by funding from the South Dakota Corn Utilization Council (Sioux Falls).
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Our results indicate that fat from CG appears to have minimal effect on concentrations of milk fat when compared with fat from DDGS or CO. Our findings suggest that oil from CG may be relatively protected and slowly released in the rumen when compared with that from DDGS, which may contain small concentrations of free oil that are more readily available during rumen digestion, thus interfering with fermentation and biohydrogenation. Inclusion of corn coproducts in dairy cow diets increased the concentration of monounsaturated and polyunsaturated fatty acids in milk fat, which have been associated with several health benefits to consumers. Moreover, feeding corn coproducts, particularly DDGS and CO, in dairy cow diets increased
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