J. Dairy Sci.
85:1482–1490
 American Dairy Science Association, 2002.
Digestion, Milk Production, Milk Composition, and Blood Composition
of Dairy Cows Fed Whole Flaxseed1
Helene V. Petit2,3
Dairy and Swine Research and Development Centre,
Agriculture and Agri-Food Canada,
Lennoxville, QC, Canada J1M 1Z3
ABSTRACT
A total of 90 lactating Holstein cows averaging 628
kg (SE = 8) of body weight (BW) were allotted at calving
to 30 groups of three cows blocked for similar calving
dates to determine the effects of feeding whole un-
treated flaxseed on milk production and composition,
fatty acid composition of blood and milk, and digestibil-
ity, and to determine whether flaxseed could substitute
for other sources of fat such as Megalac and micronized
soybeans. Cows were fed a total mixed diet based on
grass and corn silage and fat supplements for ad libitum
intake. The experiment was carried out from calving
up to wk 16 of lactation. Cows within each block were
assigned to one of the three isonitrogenous, isoener-
getic, and isolipidic supplements based on either whole
flaxseed (FLA), Megalac (MEG), or micronized soybeans
(SOY). Intake of dry matter and change in BW were
similar among diets. Cows fed FLA had greater milk
yield than those fed MEG (35.7 vs. 33.5 kg/d) and there
was no difference between cows fed FLA and those fed
SOY (34.4 kg/d). Fat percentage was higher in the milk
of cows fed MEG (4.14%) than in the milk of those fed
FLA (3.81%) or SOY (3.70%), but milk protein percent-
age was higher for cows fed FLA (2.98%) than for those
fed MEG (2.86%) and SOY (2.87%). Digestibilities of
acid detergent fiber, neutral detergent fiber, and ether
extract were lower for cows fed FLA than for those fed
SOY and MEG. Retention of N was similar among diets.
Feeding FLA resulted in the lowest omega-6-to-omega-
3-fatty-acids ratio, which would improve the nutritive
value of milk from a human health point of view. The
data suggest that micronized soybeans and Megalac
can be completely substituted by whole untreated flax-
seed as the fat source in the diet of early lactating
cows without any adverse effect on production and that
Received November 15, 2001.
Accepted January 10, 2002.
Corresponding author: H. V. Petit; e-mail: petith@em.agr.ca.
1
Contribution Number 726 from the Dairy and Swine Research
and Development Centre, Agriculture and Agri-Food Canada, PO
Box 90, Lennoxville QC, Canada J1M 1Z3.
1482
flaxseed increased milk protein percentage and its
omega-6-to-omega-3-fatty-acids ratio.
(Key words: flaxseed, milk production, reproduction,
fatty acids)
Abbreviation key: ADFI = Atlantic Dairy and Forage
Institute, FLA = fat supplement based on whole flax-
seed, MEG = fat supplement based on Megalac, PUFA
= polyunsaturated fatty acids, SOY = fat supplement
based on micronized soybeans.
INTRODUCTION
Dietary polyunsaturated fatty acids (PUFA) are per-
ceived to be healthier than saturated fatty acids. As a
result, there has been a great deal of interest in manipu-
lating the fatty acid profile of milk fat to respond to
consumer demand. Therefore, nutritionists began to
feed cows PUFA, protected against ruminal biohydro-
genation by microbes, to modify milk fatty acid composi-
tion. For example, feeding PUFA such as flaxseed (rich
in linolenic acid) has increased the proportion of omega-
3 fatty acids in milk fat (see review of Kennelly, 1996),
to meet the guidelines for human health. Flaxseed is
an excellent source of omega-3 fatty acids, which are
known to be anticarcinogenic, to prevent cardiovascular
diseases, and to increase visual activity (Wright et al.,
1998). Feeding up to 15% of the total DM as flaxseed
has no effect on DMI of midlactating dairy cows (Ken-
nelly and Khorasani, 1992). Flaxseed is also known to
increase concentrations of PUFA in milk, but usually
they do not exceed 3 to 4% of total fatty acids (Kennelly,
1996). Moreover, formaldehyde-treated flaxseed fed at
10% of DM has no effect on DMI of dairy cows compared
with Ca salts of palmitic acid, but it increases protein
and linolenic acid concentrations in milk (Petit et al.,
2001), suggesting a better N utilization on the flax diet.
Oilseeds have generally high concentrations of oleic
and linoleic acids, which are effective antiprotozoal
agents. Dietary supplementation of sunflower seed oil
(6% of DM) to sheep reduced protozoa numbers in ru-
men fluid dramatically within 5 d, from approximately
1 million to fewer than 200,000/ml (Ivan et al., 2001).
Partial defaunation has been reported to increase milk
 FLAXSEED AND MILK PRODUCTION
yield by 13.5% and the protein-to-fat ratio by 13.3%
(Moate, 1989). Therefore, oilseeds and oil-rich products
such as soybeans appear to be potential feed ingredients
to control protozoa populations in ruminants, and so
to increase the efficiency of dietary protein utilization.
Flaxseed contains approximately 32% oil (Petit et al.,
2001), which would make it a suitable feedstuff to im-
prove utilization of dietary N. There is, however, very
little information on feed utilization of whole flaxseed
by early lactating dairy cows. Therefore, the objectives
of this experiment were to determine feed utilization
and milk production and composition of dairy cows fed
whole untreated flaxseed and to determine if flaxseed
could substitute for other sources of fat, such as Megalac
and micronized soybeans.
MATERIALS AND METHODS
Dairy A
The experiment was conducted at the Dairy and
Swine Research and Development Centre, Lennoxville,
QC, from November 1998 to May 1999 using 33 multip-
arous and 15 primiparous lactating Holstein cows.
Cows were blocked within parity for similar calving
dates, and there were 11 blocks of multiparous and five
blocks of primiparous cows. The experiment was carried
out from calving up to wk 16 of lactation. Cows were
housed in tie stalls, fed individually, and milked twice
daily at 0545 and 1645 h. Milk production was recorded
at every milking. Milk samples were obtained weekly
from each cow for two consecutive milkings and were
analyzed separately to determine milk composition.
Yield of 4% FCM was calculated according to the equa-
tion of Tyrrell and Reid (1965). Weight and body condi-
tion using a five-point scale (where 1 = emaciated and
5 = fat [Edmonson et al., 1989]) were determined weekly
for each cow. Cows within groups were assigned ran-
domly to one of three treatments. The three total mixed
diets (Table 1) consisted of fat supplements (Table 2)
based on either whole flaxseed (FLA), Megalac (MEG),
or micronized soybeans (SOY). The three treatments
were designed at the beginning of the experiment to
yield similar CP, ether extract, and NEL concentrations
and were formulated to meet requirements for cows
that were a mean 580 kg of BW and produced 40 kg/d
of milk with 3.5% fat (NRC, 1989). Feed consumption
was recorded daily. Diets were fed twice daily for 10%
orts. Total mixed diets were sampled weekly, frozen,
and composited on a 4-wk basis. Composited samples
were mixed thoroughly and subsampled for chemical
analyses. Silage DM was analyzed weekly for adjust-
ment of the total mixed diets.
Blood was collected from the first 10 blocks of cows
on wk 10 postpartum at 1 h before the morning feeding
1483
to give a basal value and at 3 h postfeeding. Blood was
withdrawn from the jugular vein into vacutainer tubes
(Becton Dickinson and Cie, Rutherford, NJ) containing
EDTA for fatty acids, NEFA, and cholesterol analyses
in plasma. The plasma were separated and frozen at
−20°C for subsequent analysis. Milk samples were col-
lected on wk 4 and 8 of lactation from the first 10 blocks
of cows to determine milk fatty acid composition. Total
feces, urine, and milk were collected from the first ten
blocks of cows during wk 12 of lactation for 7 d. Feces
were collected from a rubber mat placed behind the
animals and stored in plastic containers. Daily feces
were weighed and mixed thoroughly. A 10% subsample
was taken and stored at −15°C for subsequent drying
at 55°C. Total urine was collected in stainless steel
containers via Gooch tube (BF Goodrich Co., Kitchener,
ON, Canada) attached to the cow with a nylon netting
covered with neoprene (Spall Bowan Ltd., Guelph, ON,
Canada) affixed to the vulva. A 1% daily subsample
was taken and kept frozen until analysis. Urine was
acidified daily with 100 ml of 10 NⴢH2SO4. Milk samples
were obtained from each cow for 14 consecutive milk-
ings and were analyzed for N to calculate N balance.
Dairy B
The experiment was conducted at the Atlantic Dairy
and Forage Institute (ADFI) of Fredericton Junction,
NB, Canada, from September 1998 to April 2001, with
39 multiparous and three primiparous lactating Hol-
stein cows. Cows were blocked as described for dairy
A. The three dietary treatments (Table 1) were similar
to those fed at Lennoxville, and they were formulated
to meet requirements for cows that were a mean 603
kg of BW and produced 40 kg/d of milk with 3.7% fat
(NRC, 1989). The experiment was carried out from calv-
ing up to wk 16 of lactation. Cows were housed in tie
stalls, fed individually, and milked twice daily at 0615
and 1530 h. Milk production, milk sampling and analy-
sis, and measurements of BW and DMI were performed
as described for dairy A.
Chemical Analysis
Dry matter of total mixed diets was determined by
drying at 100°C for 48 h. Protein N of silage was ana-
lyzed using an acidified extract (20 g of fresh sample
in 200 ml of 0.01 N HCl, agitated at 21°C for 22 h)
and deproteinized with TCA (Novozamsky et al., 1974).
Nitrogen determinations (N and TCA insoluble N) were
done by the Kjeldahl method (AOAC, 1990). Neutral
and acid detergent fiber components were measured
according to the nonsequential procedures of Van Soest
et al. (1991). Dietary values of NEL were predicted using
Journal of Dairy Science Vol. 85, No. 6, 2002
1484 PETIT

Table 1. Ingredient and chemical composition of experimental diets (DM basis except DM)1at Lennoxville
and Atlantic Dairy and Forage Institute (ADFI) farms.
Lennoxville ADFI
2
FLA MEG SOY FLA MEG SOY SE
Ingredient
Corn silage 13.1 19.5 14.5 13.9 17.3 13.6
Grass silage 29.0 43.3 32.3 31.4 39.0 30.5
Megalac 0 3.8 0 0 4.0 0
Micronized soybean3 0 0 17.7 0 0 18.4
Whole flaxseed 10.4 0 0 10.8 0 0
Soybean meal 1.8 2.6 0 1.9 2.8 0
Corn gluten meal 6.2 8.1 0 6.7 9.0 0
Barley 35.9 18.9 32.0 31.0 23.9 33.2
Mineral and vitamin premix 3.64,5 3.84 3.54,5 4.36 4.07 4.36
Chemical
DM, % 49.1 44.3 48.7 49.8 45.9 50.7 4.9
NEL, Mcal/kg8 1.69 1.73 1.69 1.68 1.72 1.68 ...
CP, % 19.5 19.6 17.9 15.8 15.3 15.3 2.1
TCA insoluble N, % of N 13.2 13.9 12.4 ND9 ND ND 1.8
Ether extract, % of DM 8.1 6.6 7.0 6.4 5.9 6.0 0.8
NDF, % of DM 37.8 37.6 38.8 29.4 31.5 29.0 4.6
ADF, % of DM 22.2 23.1 21.7 16.9 18.2 15.8 3.0
RUP, % of N8 35 33 39 37 38 43
NFC, % of DM8 37.3 30.3 35.6 34.6 27.1 32.7
Fatty acids, % of total fatty acids
C12:0 0 0.9 0 ND ND ND
C14:0 0 3.5 0 ND ND ND
C16:010.0 35.5 14.4 ND ND ND
C16:1 0 0.4 0 ND ND ND
C18:02.8 3.2 2.7 ND ND ND
C18:1 17.7 26.5 17.8 ND ND ND
C18:225.6 21.7 52.1 ND ND ND
C18:3 43.9 8.3 13.0 ND ND ND
1
Mean of 18 fortnightly and nine monthly samples that were prepared by compositing weekly samples
at ADFI and at Lennoxville, respectively.
2
FLA = Fat supplement based on whole flaxseed, MEG = fat supplement based on Megalac, and SOY =
fat supplement based on micronized soybeans.
3
MICRO-SOYA Elite 40% CP, Semences Prograin Inc. St-Césaire, QC J0L 1T0, Canada.
4
Contained 10.0% Ca, 4.3% P, 4.3% Mg, 13.8% Na, 1.8% S, 1.4% K, 7.5% NaCl, 1760 mg/kg of Zn, 54 mg/
kg of I, 32 mg/kg of Co, 1914 mg/kg of Mn, 436 mg/kg of Cu, 2154 mg/kg of Fe, 15 mg/kg of Se, 197,200 IU/
kg of vitamin A, 66,600 IU/kg of vitamin D, and 1030 IU/kg of vitamin E.
5
250 g of calcium carbonate (38.0% Ca) were fed per cow/d.
6
Contained 19.3% Ca, 3.0% P, 3.4% Mg, 9.1% Na, 1.3% S, 1.0% K, 9.0% NaCl, 1228 mg/kg of Zn, 38 mg/
kg of I, 14.2 mg/kg of Co, 1285 mg/kg of Mn, 305 mg/kg of Cu, 2184 mg/kg of Fe, 15 mg/kg of Se, 167,750
IU/kg of vitamin A, 41,050 IU/kg of vitamin D, and 803 IU/kg of vitamin E.
7
Contained 12.5% Ca, 5.8% P, 4.3% Mg, 11.0% Na, 1.6% S, 1.1% K, 10.5% NaCl, 1431 mg/kg of Zn, 44
mg/kg of I, 19.0 mg/kg of Co, 1527 mg/kg of Mn, 355 mg/kg of Cu, 4182 mg/kg of Fe, 17 mg/kg of Se, 189,500
IU/kg of vitamin A, 48,400 IU/kg of vitamin D, and 900 IU/kg of vitamin E.
8
Calculated using published values of feed ingredients (NRC, 1989).
9
Not determined.

ADF concentration of each feed ingredient and the
equations of the NRC (1989). Plasma NEFA (kit
99075401; Wako Pure Chemical Industries, Osaka, Ja-
pan) and total (kit no. 401-25P; Sigma Diagnostic, Inc.,
St. Louis, MO) and HDL (kit no. 352-4; Sigma Diagnos-
tic, Inc., St. Louis, MO) cholesterol concentrations were
analyzed by colorimetric methods. Concentration of
LDL cholesterol was calculated as the difference be-
tween total and HDL cholesterol concentrations. Nitro-
gen, fat, and lactose in milk were determined by infra-
red spectroscopy (Bentley 2000; Bentley Instrument,
Inc. Chaska, MN), except for N in milk samples col-
lected during the digestion trial, which were analyzed
by the Kjeldahl method (AOAC, 1990). Milk fat was
determined by the method of Roese-Goettlib (AOAC,
1990). Ether extraction in feed ingredients, diets, and
feces was conducted with a Soxlec system HT6 appara-
tus (Tecator, Fisher Scientific, Montreal, QC, Canada)
according to method no. 7.060 (AOAC, 1990). Lipid ex-
traction was conducted with a Soxtec system HT 1043
extraction unit (Tecator, Herndon, VA) according to
method no. 7.060 of the AOAC (1990). Concentration

Journal of Dairy Science Vol. 85, No. 6, 2002

 FLAXSEED AND MILK PRODUCTION 1485
Table 2. Chemical composition of feed ingredients (DM basis).

Micronized Corn gluten

Megalac Flaxseed soybeans Barley meal
CP, % of DM 0 25.2 42.5 13.2 64.7
Ether extract, % of DM 76.4 31.4 24.4 2.3 2.4
ADF, % of DM 0 21.7 5.6 10.8 4.9
NDF, % of DM 0 34.3 ND1 28.8 ND
Fatty acids, % of total fatty acids
C10:0 0.6 0 0 ND ND
C12:0 1.6 0 0 ND ND
C14:0 6.2 0 0 ND ND
C14:1 0.3 0 0 ND ND
C16:0 47.1 5.2 11.9 ND ND
C16:1 0.3 0 0 ND ND
C18:0 3.5 3.4 3.3 ND ND
C18:1 32.3 18.5 20.8 ND ND
C18:2 7.8 16.1 56.9 ND ND
C18:3 0.3 56.8 7.1 ND ND
1
Not determined.

of lipid in Megalac was determined by combustion of RESULTS AND DISCUSSION

OM at 550°C overnight in a muffle furnace (AOAC,
1990).
Fatty acids were extracted and methylated according
to the method described by Chouinard et al. (1997),
whereas in situ transesterification was performed on
feed ingredients according to Park and Goins (1994).
Plasma fatty acids were extracted according to the pro-
cedures outlined by Delbecchi et al. (2001), and prepara-
tion of plasma fatty acid methyl esters was carried out
as described by Folch et al. (1957). Fatty acid methyl
ester profiles were measured by GLC on a Hewlett-
Packard 6890 chromatograph (Hewlett-Packard Ltée,
Montreal, QC, Canada) with a G1315A autosampler
equipped with a flame ionization detector and a split-
splitless injector, as described by Delbecchi et al. (2001).
Statistical Analysis
All results were subjected to ANOVA using the gen-
eral linear models procedure of SAS (1985). Data re-
corded during the digestibility trial were analyzed as
a randomized block design, and block and treatment
were the main sources of variation. Data on blood and
milk fatty acid composition were analyzed as a split-
plot design with treatment, sampling time (before or
after feeding for blood and wk 4 or 8 of lactation for
milk), and the interaction treatment by sampling time
as main sources of variation. Data on production were
analyzed as repeated measurements using PROC
MIXED of SAS and as mean values for the 16-wk experi-
ment when there was no interaction between week and
treatment (P > 0.10). Probability values greater than
0.10 were considered nonsignificant.
Diets were designed to be similar in CP, NEL, and
EE concentrations, but there was some variation among
treatments (Table 1). The MEG diet had greater concen-
tration of NEL than the FLA and SOY diets, and this
was true for both sites. Concentration of CP was lower
on the SOY diet than the FLA and MEG at dairy A but
not at dairy B. Ether extract concentration was more
variable at dairy A than at dairy B. In general, chemical
composition of diets fed at ADFI was more similar than
that of diets fed at Lennoxville. However, the response
to dietary treatments was similar for both sites, as
shown by the lack of interaction (P > 0.10) between
treatment and site. Therefore, one can presume that
although there was a confounding effect of diet composi-
tion (e.g., CP and ether extract concentrations) on pro-
duction of dairy cows, such effect would not be too im-
portant.
There was no interaction between week of lactation
and treatment and between treatment and site for DMI,
milk production, milk composition, and BW measure-
ments; therefore, only mean values per treatment for
each site are reported (Table 3). Intake of DM was
similar among treatments, which would agree with the
results of Petit et al. (2001), who compared formalde-
hyde-treated flaxseed and Megalac, and to those of
Abel-Caines et al. (1998), who compared Megalac and
nonenzymatically browned soybeans. Untreated whole
flaxseed is readily accepted by dairy cows, and feeding
up to 15% of the total DM as flaxseed has no effect on
DMI (Kennelly and Khorasani, 1992). Moreover, the
effects of level and type of fat supplement on DMI are
negligible when total fat concentration is below 6% of
the DM (Schingoethe and Casper, 1991; Kennelly, 1996;

Journal of Dairy Science Vol. 85, No. 6, 2002

1486 PETIT

Table 3. Average DMI, milk production and composition, and BW of Holstein cows fed a concentrate based
on whole flaxseed (FLA), Megalac (MEG), or micronized soybeans (SOY) between calving and wk 16 of
lactation.1
Treatment Site
FLA MEG SOY ADFI Lennoxville SE

DMI, kg/d 19.4 18.3 18.7 19.0 18.2 0.5

Peak milk yield, kg/d 40.1a 37.5b 38.8ab 37.2 36.9 1.1
Milk production, kg/d 35.7a 33.5b 34.4ab 33.4 34.6 1.0
Milk composition, %
Fat 3.81c 4.14b 3.70c 4.02e 3.79f 0.08
Protein 2.98c 2.86d 2.87d 2.88f 2.95e 0.03
Lactose 4.71c 4.57d 4.70c 4.59f 4.73e 0.04
4% FCM, kg/d 34.5 33.7 32.9 34.2 32.8 1.1
Production, kg/d
Fat 1.35 1.36 1.29 1.34 1.28 0.05
Protein 1.05c 0.95d 0.98d 0.96 1.01 0.04
Lactose 1.68c 1.50d 1.63cd 1.53 1.63 0.04
Inital BW, kg 634 629 625 639 628 8
Final BW, kg 636 621 623 607 646 7
BW change, g/d 14 −7 −2 −285f 232e 5
a,b
Means within treatment with different subscripts differ (P ≤ 0.10).
c,d
Means within treatment with different subscripts differ (P < 0.05).
e,f
Means within site with different subscripts differ (P < 0.05).
1
Least squares means with pooled standard error (SE).

Dhiman et al., 2000; Petit et al., in press). Declines in
DMI with fat-supplemented diets appear to be related
to ruminal effects of fats, whereas studies in which
ruminal effects of fat were not observed no depression
in DMI occurred (Petit et al., 2002). Benson et al. (2001)
hypothesized that long-chain fatty acids are utilized
differently in early compared with midlactation, sug-
gesting that the negative effect of lipid supplementation
on DMI is more important as lactation progressed.
Cows fed FLA had higher milk production than those
fed MEG, and there was no difference between cows
fed either FLA or MEG and those fed SOY. Similar
differences among treatments were observed in peak
milk yield. Cows fed FLA (63 d) peaked earlier (data
not shown) than those fed either SOY (70 d) or MEG
(75 d). Similar milk yield was observed for cows fed
Megalac and those fed nonenzymatically browned soy-
beans (Abel-Caines et al., 1998). This would disagree
with the results of Petit et al. (2001), who previously
reported lower milk yield when midlactating dairy cows
were fed formaldehyde-treated flaxseed instead of Meg-
alac. In the present experiment, although FLA and
MEG had similar percentages of CP (Table 1), FLA
contained more ether extract than MEG, and this could
have contributed to increased milk production. Ken-
nelly and Khorasani (1992) found no difference in milk
yield and milk fat when cows were fed no flaxseed or
up to 15% of the total DM as flaxseed, although flaxseed
decreased milk protein. Cows fed supplemental fat sup-
plied by oilseeds, either soybeans or sunflowers, usually
peak later (wk 9 postpartum) than cows fed a control
diet (wk 6 postpartum) and they also have higher milk
production (Schingoethe and Casper, 1991). No study,
however, was found in the literature on the effect of
feeding flaxseed on lactation peak.
Fat percentage in milk was higher for cows fed MEG,
and there was no difference between cows fed FLA and
those fed SOY. Supplementation of dairy cow diets with
fat sometimes (Schingoethe and Casper, 1991), but not
always (Romo et al., 1996), depressed milk fat percent-
ages. Similar percentages of protein, fat, and lactose
were found for cows fed Megalac and those fed nonenzy-
matically browned soybeans (Abel-Caines et al., 1998).
Milk protein percentage was higher for cows fed FLA
than for those fed MEG and SOY, and it was similar for
cows fed MEG and SOY. Greater protein concentration
was previously observed for midlactating dairy cows
fed formaldehyde-treated flaxseed compared to those
fed Megalac (Petit et al., 2001). Protein concentration
is usually lower for cows supplemented with Ca long-
chain fatty acids (Canale et al., 1990), although Schin-
goethe et al. (1996) found a trend (P = 0.06) for higher
protein concentration when cows were fed sunflower
seeds than when they were fed extruded soybeans. Ac-
cording to Schingoethe et al. (1996), the effect of fat
supplement on milk protein depends on the source of
fatty acids being fed.
In general, supplemental fat decreases fat and pro-
tein percentages in milk (Schingoethe and Casper,
1991), but these reductions would seem to be due to
the highly unsaturated fat sources affecting rumen fer-
mentation. Formaldehyde-treated flaxseed has been

Journal of Dairy Science Vol. 85, No. 6, 2002

 FLAXSEED AND MILK PRODUCTION 1487
Table 4. Average milk fatty acid composition on wk 4 and 8 of lacta- Table 5. Feed intake and digestibility on wk 12 of lactation of Holstein
tion of Holstein cows fed a total mixed diet containing whole flaxseed cows fed a total mixed diet containing whole flaxseed (FLA), Megalac
(FLA), Megalac (MEG), or micronized soybeans (SOY).1 (MEG), or micronized soybeans (SOY).1
Item FLA MEG SOY SE Item FLA MEG SOY SE
(% of total fatty acids) DMI, kg/d 20.6 18.7 19.1 0.6
C10:0 3.4a 2.6b 3.3a 0.1 Digestibility, %
C12:0 3.4a 2.7b 3.3a 0.1 DM 66.1b 66.8ab 68.7a 0.5
C14:0 10.6 9.6 10.4 0.2 CP 71.4 72.7 70.2 1.6
C14:1 0.8 0.7 0.8 0.1 NDF 45.5b 51.5a 49.5a 1.1
C16:0 26.9b 36.9a 28.5b 0.9 ADF 44.1b 52.9a 49.2a 0.9
C16:1 1.8 1.7 1.5 0.1 Ether extract 80.0b 86.7a 88.0a 0.9
C18:0 17.0a 12.0b 17.0a 0.5 Milk yield, kg/d 34.6 32.6 34.8 0.9
C18:1c11 0.77 0.66 0.73 0.02 N intake, g/d 537 465 492 16
C18:1c9 29.8a 27.2ab 25.6b 0.7 Output, g/d
C18:1t9 1.5 1.8 1.7 0.1 N in feces 153c 126d 145cd 5
C18:2c6 2.7c 3.3b 5.7a 0.2 N in urine 180a 145b 143b 6
C18:2t6 0.18 0.07 0.07 0.02 N in milk 157 142 150 4
C18:3n3 0.90b 0.44c 1.04a 0.04 Total 490a 413b 438b 12
C20:3n6 0.08b 0.14a 0.17a 0.01 N retention
C20:4 0.17 0.19 0.19 0.01 % of intake 9.1 11.2 11.0 1.5
C22:6n3 0 0 1.6 0.6 g/d 47 52 54 7.9
Omega-6/omega-3 3.5c 8.5a 5.3b 0.4 Means within a row with different subscripts differ (P < 0.05).
a,b

Means within a row with different subscripts differ (P < 0.05).

a,b,c Means within a row with different subscripts differ (P = 0.07).
c,d

1 1
Least squares means with pooled standard error (SE). Least squares means with pooled standard error (SE).

shown to have little effect on ruminal fermentation
(Petit et al., 2002). Feeding fat through seeds maintains
or increases milk fat content (Dhiman et al., 2000).
However, when oilseeds and calcium salts of fatty acids
are compared, fat concentration is higher (Petit et al.,
2001) or similar (Petit et al., 2002) for cows fed Ca
salts of fatty acids and those fed formaldehyde-treated
flaxseed. There is a negative correlation in vitro be-
tween the acetate-to-propionate ratio and chain length
of unsaturated fatty acids (Jenkins and Palmquist,
1983), which would suggest that diets rich in PUFA,
such as SOY and FLA in the present experiment, could
decrease milk fat percentage as compared to cows fed
a more saturated fat source such as MEG. Feeding un-
protected fat in the form of fish oil is known to depress
milk fat percentage (Cant et al., 1997), which likely
results from the generation of high levels of trans-fatty
acids in the rumen (Baumgard et al., 2000). A greater
dietary supply of C18:2 and C18:3 fatty acids is also
known to increase milk C18:1 concentration through
ruminal biohydrogenation (Dhiman et al., 1995), with
a parallel decrease in milk fat (Dhiman et al., 2000),
which would corroborate the increased proportion of
C18:1 in the milk of cows fed FLA and SOY (Table 4).
Lactose concentration was lower for cows fed MEG
compared with those fed FLA and SOY. Yields of 4%
FCM and of daily fat were similar among treatments.
Protein production was the highest for cows fed FLA,
and there was no difference between those fed MEG,
and SOY. Daily production of lactose was higher for
cows fed FLA than for those fed MEG and there was
no difference between cows fed SOY and those fed either
FLA or MEG. The only difference between sites was in
milk composition: at ADFI, percentage of fat was
higher, and percentages of protein and lactose were
lower than at Lennoxville. The higher milk fat percent-
age at ADFI could have resulted from the dietary ether
extract concentration being lower for ADFI than at Len-
noxville (Table 1). Moreover, CP concentration was
greater in the diets of cows at Lennoxville than in those
of cows at ADFI (Table 1), which could explain the
higher protein percentage in the milk of cows at Len-
noxville.
Initial and final BW and change in BW were similar
among treatments. However, the change in BW was
significantly different between the two sites. Cows at
ADFI lost weight, whereas those at Lennoxville gained
weight during the 16-wk period of the experiment,
which could explain the higher fat percentage in milk
of cows at ADFI. There was no difference among treat-
ments for SCC, initial, final, and change in BCS (data
not shown).
Total DMI measured on wk 12 of lactation was simi-
lar among treatments (Table 5) as observed for the 16-
wk experiment. Digestibility of DM was greater for cows
fed SOY than for those fed FLA. Cows fed MEG had
intermediate values, and they had DM digestibility that
was similar to that of cows fed SOY and FLA. Digestibil-
ity of CP was similar among treatments; lower CP di-
gestibility was reported by Petit et al. (2002) for cows
fed formaldehyde-treated flaxseed than for those fed
Megalac, which may reflect the effects of the formalde-
hyde treatment. Cows fed FLA had lower digestibility
of ADF and NDF than cows fed MEG and SOY, and

Journal of Dairy Science Vol. 85, No. 6, 2002

1488 PETIT
there was no difference between cows fed MEG and
those fed SOY. Cows fed FLA had lower ether extract
digestibility than those fed MEG and SOY, which may
reflect the greater accessibility to rumen microbes and
enzymes of fats added in the form of Ca salts or mi-
cronized soybeans compared with lipids that are inti-
mately associated with the fiber-protein matrix in
whole flaxseed. Lower ether extract digestibility for
cows fed formaldehyde-treated flaxseed than fed Mega-
lac has been reported by Petit et al. (2002), whereas
Garcia-Bojalil et al. (1998) reported an improved digest-
ibility of ADF and NDF when feeding calcium salts of
fatty acids compared to a no-fat diet. Improved fiber
digestion could result in an increase in the acetic-to-
propionic-acid ratio when feeding Ca salts of fatty acids
(Schauff and Clark, 1992).
Milk yield measured over the 7-d period on the week
of digestibility (wk 12 of lactation) was similar among
treatments and averaged 34.0 kg/d. Total N intake,
expressed in g/d, was similar among treatments. Out-
put of N in feces (P = 0.07) and in urine (P < 0.05) and
total N output (P < 0.05) were greater for cows fed FLA
compared to cows fed MEG or SOY, although there was
no difference in N output in feces between cows fed
FLA and those fed SOY. Cows fed MEG and SOY had
similar output of N in feces, urine, and in total. Output
of N in milk was similar among treatments. Retention
of N, expressed either as a percentage of intake or in g/
d, was similar for the three diets. Fat supplementation
improves the efficiency of utilization of dietary N for
milk N deposition, and the effect is better when feeding
cis compared to trans-fatty acid isomers (Romo et al.,
1996). However, no difference was found in the present
experiment among fat sources, which all contained
mainly cis-fatty acid isomers.
Milk fatty acids concentrations (Table 5) of C10:0,
C12:0, C18:0, and C18:3 were higher for cows fed FLA
than for those fed MEG, whereas the inverse was ob-
served for concentrations of C16:0 and C18:2n6cis. This
is in general agreement with the results of Petit et al.
(2001), who compared formaldehyde-treated flaxseed
and Megalac, and with those of Kennelly and Khorasani
(1992), who fed up to 15% of flaxseed in the total DM
of dairy cows. Feeding FLA compared to SOY increased
concentrations of C18:1n9cis and decreased those of
C18:2n6cis, C18:3n3, and C20:3n6. Concentration of
C18:1n9trans was similar among diets. Feeding roasted
cracked soybeans, soybean oil, or flaxseed oil has been
shown to increase milk C18:1n9trans concentration
compared with feeding raw cracked soybeans (Dhiman
et al., 2000). This would suggest that the natural protec-
tion of whole flaxseed and heat treatment of micronized
soybeans were adequate to prevent a rapid release of
oil and thus biohydrogenation of long-chain fatty acids
Journal of Dairy Science Vol. 85, No. 6, 2002
into conjugated linoleic acid. Cows fed SOY had the
highest C18:2 concentration in milk. Similarly, Dhiman
et al. (2000) reported that cows fed soybeans compared
with those fed free oil had greater C18:2 concentration
in milk as a result of the oil being released more slowly
from oil seeds than from free oil in the rumen, thereby
decreasing the probability of biohydrogenation of oil-
seeds. Feeding oil through micronized soybeans would
make more unsaturated oil available in the small intes-
tine for absorption, resulting in higher transfer effi-
ciency from the diet to the milk fat (Dhiman et al.,
1995). Feeding SOY increased C18:2 concentration in
milk fat, as previously reported by Abel-Caines et al.
(1998), who compared nonenzymatically browned soy-
beans and Megalac. Feeding FLA resulted in the lowest
omega-6-to-omega-3-fatty-acid ratio in milk, which
would improve the nutritive value of milk from a human
health point of view. According to Sim (1998), the cur-
rent high ratio should be decreased to less than 4-to-1
to reduce the potential risk of coronary heart diseases;
feeding whole untreated flaxseed to dairy cows could
contribute in improving human health by a greater in-
take of omega 3 fatty acids in enriched dairy products.
There was no interaction (P > 0.10) between sampling
time and treatment for plasma composition. However,
cows fed MEG had greater concentrations of NEFA
than those fed either FLA or SOY, and there was no
difference between cows fed FLA and those fed SOY
(Table 6). On average, plasma concentrations of NEFA
increased (P < 0.05), respectively, from 234 to 278 µEq/
L from −1 to 3 h after feeding (data not shown). Greater
NEFA concentration is related to increased body fat
mobilization (Roberts et al., 1981), suggesting that in-
creased fat mobilization could have contributed to milk
yield, as there was a numerical greater loss of BW for
cows fed MEG over the 16-wk experiment. Greater
NEFA concentrations have been previously reported
for cows fed Megalac than for those fed formaldehyde-
treated flaxseed (Petit et al., 2001).
Total cholesterol concentration was greater for cows
fed MEG, intermediate for those fed SOY, and lower
for those fed FLA. Cows fed MEG and SOY had higher
and similar concentrations of HDL cholesterol com-
pared with those fed FLA. Concentrations of LDL cho-
lesterol were similar among treatments, although Petit
et al. (2001) reported greater LDL cholesterol concen-
trations for cows fed Megalac than for those fed formal-
dehyde-treated flaxseed. Fat supplementation is known
to increase blood cholesterol (Garcia-Bojalil et al.,
1998), although the types of fatty acids would seem to
differ. As previously reported by Petit et al. (2001),
feeding FLA decreased blood-cholesterol concentration
compared with feeding MEG.
 FLAXSEED AND MILK PRODUCTION 1489
Table 6. Blood composition of Holstein cows fed a total mixed diet containing whole flaxseed (FLA), Megalac
(MEG), or micronized soybeans (SOY).1
Item FLA MEG SOY SE
b a b
NEFA, µeq/L 235 307 229 12
Total cholesterol, mg/100 mL 246c 319a 278b 13
HDL cholesterol, mg/100 mL 137b 161a 153a 5
LDL cholesterol, mg/100 mL 109 158 125 13
Fatty acid, % of total fatty acids
C14:0 1.2 1.4 1.4 0.1
C16:0 10.9b 14.2a 11.4b 0.2
C16:1 1.7a 1.3b 1.2b 0.1
C18:0 15.4a 12.2b 14.1b 0.2
C18:1t9 0.69b 0.81a 0.85a 0.02
C18:1c9 8.5a 9.1a 5.0b 0.3
C18:1c11 0.69 0.67 0.79 0.03
C18:2 47.6c 51.9b 57.3a 0.6
C18:3n6 1.16 1.12 0.92 0.04
C18:3n3 8.0a 2.8c 3.4b 0.3
C20:3n6 1.82ab 1.99a 1.78b 0.04
C20:4 1.65b 1.89a 1.51b 0.05
C20:5n3 0.79a 0.57b 0.52b 0.03
omega-6/omega-3 5.9b 17.0a 16.7a 0.7

Means within a row with different subscripts differ (P < 0.05).

a,b,c
1
Least squares means with pooled standard error (SE).

Concentrations of C14:0 were similar among diets. ACKNOWLEDGMENTS

Cows fed FLA had lower concentrations of C16:0,
C18:1n9trans, C18:2, and C20:4n6 than those fed MEG,
whereas the inverse was observed for concentrations
of C16:1, C18:0, C18:3n3, and C20:5n3. Feeding SOY
decreased concentrations of C16:1, C18:0, C18:1n9cis,
C18:3n3, and C20:5n3 compared with feeding FLA. In
general, feeding FLA increased the omega-3 fatty acids
compared to feeding MEG and SOY, which resulted in
the lowest omega-6-to-omega-3-fatty-acid ratio in blood.
CONCLUSIONS
Feeding whole flaxseed compared to Megalac or mi-
cronized soybeans increased milk protein percentage.
Moreover, cows fed flaxseed produced more milk and
peaked earlier than those fed Megalac although milk
fat was lower with flaxseed. Dietary fat sources had no
effect on BW change, but cows fed Megalac had greater
NEFA concentrations than those fed flaxseed or mi-
cronized soybeans. Cows fed flaxseed had lower digest-
ibility of DM and fiber than those fed micronized soy-
beans. Output of N and retention of N was similar
among diets. Feeding flaxseed resulted in the lowest
omega-6-to-omega-3-fatty-acid ratio, which would im-
prove the nutritive value of milk from a human health
point of view. The data suggest that micronized soy-
beans and Megalac can be completely substituted by
whole untreated flaxseed as the fat source in the diet
of early lactating cows without any adverse effect on
production and that flaxseed increased milk protein
percentage and its omega-6-to-omega-3-fatty-acid ratio.
The author thanks N. Clark, J. Sweetland, L. Veil-
leux, and I. Sabourin for technical assistance and the
Atlantic Dairy and Forage Institute, Fredericton Junc-
tion, NB E5L 1R1 and Semences Prograin Inc., St-Cés-
aire, QC J0L 1T0 for their financial help. This project
was sponsored in part by the Matching Investment Ini-
tiative of Agriculture and Agri-Food Canada.
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