GROWTH, CARCASS TRAITS, AND FAlTY ACID

Growth, carcass traits, and fatty acid profiles of adipose tissues from steers fed whole cottonseed N. O. Huerta-Leidenz, H. R. Cross, D. K. Lunt, L. S. Pelton, J. W. Savell and S. B. Smith J ANIM SCI 1991, 69:3665-3672.

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GROWTH, CARCASS TRAITS, AND FAlTY ACID PROFILES OF ADIPOSE TISSUES FROM STEERS FED WHOLE COlTONSEED’s2 N. 0. Huerta-Leidenz3, H. R. Cross4, D. K. Lunt*, L. S. Pelton4, J. W. Savel14 and S. B. Smith4 Texas Agricultural Experiment Station, Texas A&M University, College Station 77843-2471 ABSTRACT

To investigate the impact of dietary whole cottonseed (WCS) level on fatty acid composition, growth, and carcass traits, 45 Hereford steers were assigned to diets containing 0,15, or 30% dietary WCS. The 15 and 30% WCS contributed an estimated 3.3 and 6.6% additional lipid, respectively, to the diets. After being fed for 54 d, all animals were weighed and slaughtered, and carcass measurements were obtained. There we= no differences (P > .05) among dietary treatment groups in live weight or ADG for the 54d feeding period. Control steers had larger (P < -05) longissimus muscle areas than steers fed 30% WCS,which accounted for the advantage in yield grade (P < .05) exhibited by the control group. Feeding of 30% WCS resulted in minor increases in linoleic and total polyunsaturated fatty acid content of perinephric fat expressed in both normalized (area percentage) and gravimetn’c @/lo0 g of fresh tissue) formats. There were no significant differences in the monounsaturated or saturated fatty acid content of adipose tissues from animals fed the different diets. Subcutaneous adipose tissue samples were higher (P < .01) in total unsaturates but had lower (P e .OS) proportions of C18:O and C18:l than perinephric samples. Feeding WCS at the levels repofled herein only had minor effects on fatty acid composition of beef adipose tissues. Kei Words: Fitty Acids, Growth, Carcass Traits, Cottonseed, Steers, Adipose Tissue J. Anim. Sci. 1991. 693665-3672

ruminal microorganisms. However, it has been

lntroductlon

suggested that UFA in oilseeds are protected from ruminal biohydrogenation by the seed coat (Baldwin and Allison, 1983). Feeding dairy cows whole cottonseed (WCS) resulted in milk fat with elevated levels of oleate and stearate (Smith et al., 1981). Preston et al. (1989a,b) indicated that the stearic acid concentration of beef subcutaneous adipose tissue lTecbnical Article 25662 from the Texas Agric. Exp. increased as the result of WCS feeding. Sta. This study was funded partially by Kine; Ranch, Inc., Bonanome and Grundy (1988) provided eviKingsville, TX.Refto a company or a trade name does not imply approval or endorsement by the Texas dence that strongly suggests that stearic acid, as well as other UFA, has a cholesterolAgric. Exp. Sta. 2Appreciation is extcnded to Mrtrcos Gomez-Meza for lowering effect in humans. Given the implicaassistance i n statistical analysis. tions of these findings from a diet and health ~CUITULIaddress: ~ a ~ a l t dc a dAgronomia, universidad standpoint, it is imperative to reexamine del Zulia, M a r a c a i b o - ~Venezuela. “rkpt.~nim. sci., TCX~S MM univ., college station. current information regarding the effects of ’Texas A&M Univ. A@. Res. Cmter, McGregor. WCS feeding on the efficiency of UFA and(or) Received Novcmk 29, 1990. stearate incorporation into bovine adipose Acccpttd March 21, 1991. Producing meat with increased levels of unsaturated fatty acids (UFA) is more difficult in ruminants than in nonruminants because of the extensive hydrogenation of dietary UFA by

3665


3666

HUERTA-LEJDENZ ET AL.

tissue. Accordingly, the objectives of this study were to determine the effects of WCS on 1) growth and carcass traits of beef cattle and 2) fatty acid composition of subcutaneous and perinephric adipose tissue. Materials and Methods

Animals, Feeding, and Sample Collection. Forty-five Hereford steers were stratified by live weight and hip height and allotted randomly to receive either 0, 15, or 30% levels of WCS in diets with concentrate formulations adjusted to balance major nutrients, except energy (Table 1). The WCS contained 22.3% ether extractable lipid by weight (as fed); thus, the 15 and 30% levels contributed an additional 3.3 and 6.6% lipid, respectively, to the diets. Diets were formulated to meet all NRC (1984) requirements. The animals started treatments at an approximate live weight of 340 kg. Cattle were housed in pens by treatment group (15 per pen). During the treatment period, two control steers and three steers fed 15% WCS lost weight continually and consequently were removed from the study. After being fed for 54 d, all remaining animals were slaughtered at a commercial packing plant and carcass measurements were obtained. Immediately after slaughter, 8- to 10-g duplicate samples of subcutaneous fat over the 13th rib and fat from the perinephric fat were obtained. Individual samples were placed in small plastic vials, cap-sealed, and kept frozen on dry ice for shipping. On arrival, samples were kept in frozen storage (-20°C) until they were analyzed. Fatry Acid Composirion. Total lipids from adipose tissue samples were extracted from homogenates of 1-g samples using chlorofommethanol (21, vol/vol), according to the procedure of Folch et al. (1957). An aliquot of the total lipid extract from adipose tissue samples was freed of solvent under nitrogen using an analytical evaporate$. Approximately 100 mg of the total lipid extract from adipose tissue samples was mixed with the internal standard (C12:O methyl ester) and methylated

%eyer N-Evap, Organomation Associates Inc.. Ber-

lin,

m.

7Model 437A.Packard, Chrompack Inc., Raritan, NJ. 'Model SP 4290, Spectraphysics, San Jose, CA. 9Nu-Check-Rep Inc., Elysian, MN.

with boron trifluoride-methanol following the procedure described by Morrison and Smith (1964) after saponification. To saponify, 2 ml of .5 N KOH in methanol was added to the lipid extracts and the mixture was heated at 70'C for 15 min. This step was included before the transesterification procedure of Morrison and Smith (1964) to ensure complete saponification and methylation of the samples. Fatty acid methyl esters (FAME) were analyzed using a flame ionization gas chromatograph7 equipped with a 2-m x 3.175-mm stainless steel column packed with 15% cyanopropyl: phenylpolysiloxane (9: 1 wt/ wt) on a 100-120 mesh solid support. The column was run isothermally at a temperature of 185'C. The injection port and detector were maintained at 250'C and 275'C, respectively. Nitrogen was the carrier gas with a flow rate of 25 d m i n . Chromatograms were recorded with a computing integrators. The gas chromatograph system was calibrated with standard FAME mixtures9. Identification of sample fatty acids were made by comparing the relative retention times of FAME peaks from samples with those of standards. For quantification of sample FAME, reference standards of pure triglycerides of C14:0, C16:0, C18:0, and C2O:O fatty acids9 were saponified, mixed with the internal standard, and methylated, and the methyl esters were chromatographed to obtain response factors for each external standard. Fatty acid response factors were entered in the equation used for gravimetric quantitation of sample FAME as grams of individual fatty acid triglycerides per 100 g of wet tissue (Slover and Lanza, 1979). Data also were calculated as normalized area percentages of fatty acids. Sruriszical Analysis. Data were analyzed using the SAS GLM procedure for unequal subclass numbers (SAS, 1985). With the exception of ADG and hot carcass weight, growth or carcass traits were compared using one-way ANOVA. When F-tests for main effects indicated a difference among treatments, simple means for carcass traits were separated using the Tukey-Kramer procedure for unequal cell sizes (SAS, 1985). Despite the intent of balancing initial (0 d) weight among treatments, initial weight contributed significantly to the variation of ADG and hot carcass weight. To compare ADG and hot carcass weights across dietary treatments, means were adjusted to a common initial live weight.


EWECIS OF WHOLE CO'ITONSEED FEEDING ON STEERS

Adjusted means for ADG or hot carcass weights were compared using the test of equality for a l l pairs of least squares means (SAS, 1985). To account for day-today variation in fatty acid composition analyses, data were analyzed using a randomized complete block design (Montgomery, 1984) with blocks consisting of daily sample batches. When F-tests for dietary treatments were significant, means for fatty acids were separated using the Tukey-Kramer method. The effect of depot site as a main effect was tested in a separate analysis using the same linear model as for dietary treatment. Results and Discussion

Means for growth traits are presented in Table 2. No differences (P > .05) were observed among treatment groups in hip height or body condition, and there were no differences (P > .05) in live weight or adjusted ADG for the 54d feeding period. Keele et al. (1989) reported that WCS can be fed up to 25.3% of the dietary DM without adversely affecting fiber digestibility. &viously, Moore et al. (1986) reported that the addition of 6.3% fat in the form of WCS (fed as 30% of the diet), as cottonseed oil or as

3667

animal fat, significantly reduced mean ADF digestibility. However, all forms of added fat increased apparent digestibility of the lipid. In their study (Moore et al., 1986), digestibilities of DM and 8 were higher for the WCS diet than for the diets containing free fats; their results, however, were based on feeding highroughage diets and cannot be extrapolated to common dietary conditions such as those encountered in most feedlot operations. Our results indicated that the addition of WCS in low-forage diets did not affect animal performance. Unadjusted means for hot carcass weights were similar for all dietary treatment groups (Table 3). However, when adjusted for variation in initial live weight, increased concentrations of WCS in the diet were associated with lighter hot carcass weights. This observation agrees with those reported by Preston et al. (1989a,b). Those workers reported that hot carcass weight decreased progressively with increasing levels of WCS. Control steers had larger (P e .05) longissimus muscle (ribeye) areas than steers fed 30% WCS, which accounted for the lower yield grade (P e .05) exhibited by the control group (Table 3). No other carcass trait was affected by WCS treatment (P > .05).

TABLE 1. INGREDIENT COMPOSITION AND NUTRlENT DENSlTY OP DIETS DieP Item

Ingredient composition.as fed 96 No. 2 yellow corn Whole cottomeed cottonseed meal Cottonseed hulls Molasses Ground limestone Trace m i n e d s ~ l t b Potassium chloride Vitamin mixc Nutrient densityd

m % crude protein, 96 Crude fiber, % Mcavkg mg.Mcavkg

%*

Control

15% wcs

30% WCS

63.00 0 16.00 15.00 4.00 1.00

5850 15.00 950 11.00 4.00 1.00 .25

54.75

.25

.30

30

.45

.45

89.30 12.80 9.70 1.72

89.50 12.80 9.80 1.83 1.15 .43

1.05

30.00 2.75 6.50 4.00 1.00 25 .30 .45 89.70 12.80 9.70 1.94 127 .43 .40

calcium,9% .43 P h o s p ~% , .38 39 WCS = added whole cottonseed. bcomposition of m e mineralizad sale NaCl, 98% ZN, .35% Mn, 28% Fe. .175% Cu, ,03596;and I, .007%. CCompositionof vitamin mix: vitamin A, 2.2 x io6 vitamin D,1.1 x 106 IUW,ami vitamin E, 2,200 nr/ks. dNutrien~data were estimated from table values W C , 1984).

rum,


3668

HUWTA-LEIDENZ ET AL. TABLE 2. EFFECT OF DIETARY WHOLE COTTONSEED ON GROWTH "&UT'S DieP

Item No. of steers OdWtkg 54 d Wt, kg Hip height, cm Od 54 d Body condition scoreb Od 54 d Daily Pains', ks

ovaall

15% WCS 12 342.9

15 347.2'

4029

404.6'

115.7d 1185'

115.Sd 118.3'

116.3'

5.8d 5.8' .95d

Peedlgaine. kg/kg overall

30% wcs

Control 13 326.9 377.3'

8.3

119.1'

SE

26.5 30.9

2.7 2.9

5.8' 6.3'

5 9 6.2'

.43 58

1.15'

1. O d

.1

7.8

8.2

-

W C S = added whole cottomeed. bl = very tnin; 9 = extremely fat. 'Vatues adjusted for initial weight as the covariate. 'Means in the same row with saperscrjptS that do not have a common superscript letter differ (P < .05). 'Cattle groupfed, therefore. no measure of within-treatment variation is available.

With the exception of linoleic acid (C18:2) compared with the control. The PUFA of and the total polyunsaturated fatty acid VU- perinephric adipose tissue from cattle fed 30% FA) content of perinephric samples, there were WCS was higher (P< .OS) than for either the no significant differences (P > .05) in fatty control or the 15% WCS group. These acid composition among the dietary treatment differences were less than .5% in magnitude groups (Table 4). The linoleic acid weight and, as in the case of linoleate, were considpercentage in perinephric adipose tissue in- ered to be minor Fable 4). Similar trends were creased .55% by feeding 30% WCS (P < .M) observed when fatty acid variables were

TABLE 3. EPPECTS OF DIETARY WHOLE COTTONSEED ON CARCASS TRAITS

DieP Item

Control

Hot carcass wt, kg Adjustedbhot carcass wt, kg Fat thickness, mm Adjusted fat thickness, mm Ribeye area,cm2 Kidney, pelvic and heart fat, 46 Marbling score' skeletal maturityd Leanmaturityd overallmaturityd Qualitygradec Yield grade

223.9 233.1f 8.5 9.2 67.1f 2.0

15%wcs

30% wcs

SE

234.3 2316 8 99 10.2 65.4'8 2.4 s189

23 1.2 225.28 10.2 10.6 61.68 2.3

A36

A4' A33 A3? se76 2.98

21.6 2.4 4.3 3.5 5.6 .5 0 43.1 8.3 5.5 5.7 34.5 .55

~~~

s1@ A36 A3' A3' SeM 2.3'

A33 A3' seS3 2.7%

sP3

WCS = added whole cottonseed. bvalues for hot carcass weight adjusted for initial live weight as the covariatc. 'SI = slight; degrees = 0 to 99. dmcarcasses were of A maturity; de= 0 to 99. 'Se = Select; degrees = 0 to 99. &Means in the same row with superscripts that do not have a common superscript letter differ (P < .05).


EFFECTS

expressed as grams/lOO g of fresh tissue pable 5). As previously indicated by the normalized data, only the perinephric depot site was altered by treatment with WCS. There are numerous studies dealing with the dietary manipulation of fatty acid composition of beef adipose tissues (Willey et al., 1952; Roberts and McKirdy, 1964, Johnson and McClure, 1972; Dryden and Marchello, 1973; Dryden et al., 1973; Clemens et al. 1974; Kimoto et al., 1974; Garrett et al., 1976; Hood and Thomton, 1976; St. John et al., 1987; Preston et al., 1989a,b). Most of these reports have been restricted to the evaluation of oils instead of oilseeds. In an earlier report (Willey et al., 1952), feeding cottonseed oil to steers at 5 % of the diet produced a more saturated body fat with a concomitant reduction in oleic acid. These effects were contrary to those encoun-

TABLE 4.

3669

OF WHOLE COTTONSEED FEEDING ON STEERS

MEAN" NO-

tered with the use of other free vegetable oils (Roberts and McKirdy, 1964). The latter researchers demonstrated the feeding rapeseed and sunflower seed oil slightly increased the percentage of monounsaturated fatty acids in beef perinephric fat. Fats compose approximately 20% of the DM in WCS (Smith et al., 1981). Little is hown about the effects of feeding WCS on the fatty acid composition of beef depot fats. Preston et al. (1989a,b) indicated that the stearic acid concentration of beef subcutaneous fat increased as the result of WCS feeding. The increase in stearic acid was accompanied by a reduced concentration of linolenic acid and some monounsaturated fatty acids &e., C16:l and (201). This would support the increase in beef fat saturation reported previously by Willey et al. (1952). Feeding WCS to dairy

PERCENTAGES OP MAJOR FAlTY ACIDS IN DEPOT PATS AFpEcI13D BY DIETARY WHOLE COTTONSEED

OP EIERJ3FORD STEERS AS

Did CQllkOl

Item

Depot site' No. of obsavations Fatty acidd c140 c141 C160 C16:l C18:O C18:l C182 C18:3 SPA UFA MUFA PUPA DFA OFA Total fatty acids Ratios UFABFA MUFMFA DFAIOFA PUPAISPA

15%WCS

SE

30% WCS

PN

sc

PN

sc

PN

sc

13

13

12

12

15

15

4.06 1.31 30.82 5.49 17.63 36.10 4.w 31 52.50 47.47 42.90 4.5F 65.10 34.87 99.97

3.58 1.99 26.33 6.77 10.45 44.81 326 .48 40.38 57.32 53.57 3.75 67.77 29.92 97.69

4.55 1.22 31.95 5.41 1826 34.00 4.14ef .45 54.76 45.23 40.64 4.M 63.49

4.33 1.11 33.33 5.82 15.84 34.5 1 4.61f .44 53.50 46.49 41.44 5.05f 62.33 37.70 99.99

3.94 2.12 27.13 5.80 9.6 1 44.00 3.61 .45 40.67 56.00 51.92 4.06 65.58 31.06

99.99

3.99 2.23 27.63 6.77 9.30 43.97 3.14 .48 40.91 56.59 52.97 3.62 65.88 31.62 97.50

96.64

.03

.91 .83 1.97 .09

1.46 1.37 2.82 .09

.83 .74 1.80 .08

1.39 1.30 2.10 .09

.87 .77 1.72 .09

1.39 128 2.13 .10

.12 .ll

36.50

PN 40 .7 .4 .1 1.7 7.6 2.1 5

.1 2.8 2.8 2.7 .6 6.7 6.7

sc 40 .7 .4 22 3.3 2.8 3.4 .7 .1 4.3 3.8 3.9 .7 4.6 2.6 4.5

22

51

21 27

.01

.02

*ercentage of the total peak area of lfre fatty acids listed. bwCS = added whole cottonseed. 'PN = perinephric site, sc = subcutaneous site. d~~~ = total saturated fatty acids; UFA = total unsaturated fatty acids; MUFA = total m 0 - m ~ ~ fatty acids; PUPA = total polyunsaturated fatty acids; DFA = those d m i fatty acids having desirable (either neutral or hypocholesteroldc) effect in hmnans (is., UFA + C18:O); OFA = those dietary fatty acids haviug objectionable (hypercholesterolemic) effect in humans (i.e., C140 + C160). in the same row within depot site with superscripts that do not have a common mperscript letta differ (P c

.os).


3670

HUWTA-LEIDENZ ET AL.

cows also has resulted in a twofold increase in The present study demonstrated a small but yield of stearic acid in milk fat (Smith et al., significant increase in PUFA levels. Similarly, 1981). Results of the present study do not increases in linoleic acid of bovine adipose support those findings. The reasons for the tissues as the result of feeding formaldehydediscrepancy between our results and those of treated sunflower seed have been reported Preston et al. (1989a,b) are not known but may previously (Hood and Thornton, 1976). Howbe related to the length of the feeding period ever, several reports have demonstrated that or, alternatively, to the method of feeding. It is the fatty acid composition of bovine depot fat possible that the WCS feeding time required is unchanged by the consumption of vegetable for achieving an increased deposition of stearic oils or seeds (Willey et al., 1952; Roberts and acid in depot fats exceeds 56 d. The twofold McKirdy, 1964; St. John et al., 1987). Some increase in PUFA levels of adipose increase of oleic acid in milk from cows fed WCS, as reported by Smith et al. (1981), tissue of cattle fed WCS is not unexpected probably reflected the fact that different because, on a weight basis, slightly more than physiological mechanisms of dietary fat trans- half (52 dl00 g) of edible cottonseed oil in fer and(or) modification exist between secre- PUFA, 99% of which is linoleic acid (USDA, 1978). It is also probable that dietary linoleic tory and depot fats. Changing the PUFA content of bovine acid that has escaped ruminal biohydrogenaadipose tissue has met with limited success. tion would be more easily absorbed than other

TABLE 5. MEAN GRAVIMETRIC CO-

OF MAJOR FATI” ACIDS IN DEPOT FATS OF HEREFORD S T E W S AS AFFECED BY DIETARY WHOLE COlTONSEED Dietb

Depot site‘ No. of observations Fatty acidd c140 c141 C16:O C161 C18:O

C18:l C18:2 C18:3 SFA UFA PUPA MUFA DFA OFA Totalfattyacids Ratios UPNSFA PUPNSFA DFNOFA M”FA/SFA

15%wcs

Control

Item

m

sc

PN

sc

PN

13

13

12

12

15

3.91 1.05 27.00 4.60 16.68 31.33 3.8V .42 47.60 4120 4.22e 37.00 57.88 30.90 88.78

2.10 1.14 13.31 4.41

3.80 .97 25.78 4.96 14.99 32.60 4.33f .4 1 47.32 4323 4.74f 38.50 58.22 32.33 90.60

3.44 1.10 25.78 4.52 16.02 33.01 3.72= .47 45.24 42.82 4.1p 38.63 58.84 29.22 88.06

1.63 lo.% 3.52 5.11 19.60 1.48 .2 1 17.69 25.65 1.69 23.96 30.76 1258 43.34

.% .09 2.15 .87

1.61 .10 2.56 1.51

.85

.87

.09 1.96 .78

SE

30% WCS

5.25

23.80 1.76 25 20.64 31.35 2.02 29.33 36.60 15.39 52.00 1.54 .09 2.36 1.44

.92 .10 1.88 .82

SC 15

1.99 .99 12.63 3.96 5.32 22.32 1.89 23 19.94 29.40 2.13 2727 34.72 14.62 49.34 1.55 .ll 2.40 1.44

PN 40 .6 .3 6.0 1A 7.1 2.4 .5

.1 4.8 2.9 5

2.8 6.9 6.5 6.0 .12 .01 .56 .ll

sc 40 1.1 .5 5.6 1.5

3.4 9.5 1.0 .1 9.9 12.3 1.1 11.3 15.4 6.6 21.9 .30

.m

.31 .30

%lo0 g of fresh tissue. bwCS = added whole cottonseed. ‘PN = perinephric site, SC = snbcutane~ussite. dSFA = total saturated fatty acids; UPA = total unsaturated fatty acids; MUFA = total monounsaturated fatty acids; PUPA = total polyunsaturated fatty acids; DFA = those dietary fatty acids having desirable (either neutral or hypocholesterolemic) effect in humans (Le., UFA + C180); OFA = those dietary fatty acids having objectionable (hyparholesterolemic) effect in humans (Le., C140 + C160). e*fMeansin the same row within depot site with superscriptStbat do not have a common superscript letter differ (P < .05).


EFPECTS OF WHOLE COlTONSEED FEEDING ON STEERS

individual fatty acids. In the work of Keele et al. (1989), estimates of true digestibility (assuming that there was no biohydrogenation in the colon) as measured by fatty acid disappearance between duodenum and feces were 71% for palmitic acid, 45% for stearic acid, 79% for oleic acid, and 100% for linoleic acid. From a compositional standpoint, an increased absorption of WCS monounsaturates to depot fats is less probable. Compared with PUFA, the monounsaturate content of cottonseed oil is relatively low (17.8 g/lOO g of edible oil portion), 96% of this amount being oleic acid (USDA, 1978). Our results, in general, agree with those of Clemens et al. (1974). who concluded that ruminant adipose tissues are relatively insensitive to changes in type or amount of dietary fatty acids. In the present study, the addition of WCS at relatively high concentrations in the diet (30%) could not effectively overwhelm the biohydrogenating capacity of ruminal

3671

microbes. Recent data, presented by Keele et al. (1989), have challenged the supposition that unsaturated fatty acids in WCS are protected from ruminal biohydrogenation. Presumably, the protective effect of the seed coat is lost during mastication (Keele et al., 1989). Overall fatty acid composition of the perinephric and subcutaneous depot sites is depicted in Table 6. Relative proportions of most fatty acids differed significantly (P e .OS). There were no significant differences in the percentages of palmitoleic (C16:l) or linolenic (C18:3) acids between the subcutane ous and perinephric depot sites. However, when expressed as grams/lOO g of fresh tissue, small differences (P< .05) between depot sites in palmitoleic and linolenic acids were noted. Gravimetric values (Table 6) indicated that only myristoleic acid (C14:l) content was not different between depot sites. In general, the subcutaneous depot was more unsaturated (P< .05) than the perinephric depot. Consequently,

TABLE 6. FAlTY ACID COMPOSITION OF SUBCUTANEOUS AND PWINEPHRIC FAT DEP(YTS OF W O R D STEERS m o t site

perinephric (n = 40)

Subcutaneous (n= 40)

Fatty acida

Area%

g/100 8

Area 96

c140 c141 C160 C161 C18:O C18:l Cl8:2 C18:3

DFA

4.31b 121b 32.10b 559b 17.15b 34.9ob 4.2gb .47b 53.59 46.43b 41.67b 4.76b 6358b

OFA Total fatty acids

99.98b

3.71d 1.@Id 27.1 8d 4.71d 15.83d 32.34d 3.97d ,43d 46.72d 42.d 38.08d 4.d 58.32d 30.89 89.21d

3.89 2.11' 27.u 6.4Ib 9.8@ 44.26f 3.39 .47b 40.65' 56.M 52.77 3.83' 66.3sc 30.86f 97.24'

.87b .7gb

.92d .82d

1.41' 1.32b

SFA UFA MUFA

PUFA

3 6 . d

g

1.w

.99d 12.3@ 3.95e 5.23e 2 1.8? 1.72e

.ae 19.42e 28.76e 26.81e 1.95e 34.w 14.1p 48.18e

Ratios 1.5? 1.4? .09b .d .wb .lcF 1.83b 1.d 2.17 2.44d %FA = total sahuatcd fatty acids; UFA = total unsataratcd fatty acids; MUFA = total mononnsaturated fatty acids, PUFA = total polyunsaturated fatty acids; DFA = those dietary fatty acids haviug desirable (either neutral or hypocholesterolemic) effect in humans (i.e., UFA + C18:O); OFA = those dietary fatty acids having objectiomble (hypercholesterolemic) effect in hnmans (Le.. C140 + C16:O). b.CAreaperccntagc values in the same row with superscripts that do not have a common superscript lctta differ (P < UFA/SFA MUFA/SFA PUFNSFA DFA/OFA

.W). 4eGravimetri~

(P