International Dairy Journal 20 (2010) 433e448

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International Dairy Journal

journal homepage: www.elsevier.com/locate/idairyj

Review

Grass-based ruminant production methods and human bioconversion of vaccenic

acid with estimations of maximal dietary intake of conjugated linoleic acids
Remo P. Jutzeler van Wijlen a, *, Paolo C. Colombani b
a
Sponser AG, CH-8832 Wollerau, Switzerland
b
Department of Agricultural and Food Sciences, ETH Zurich, CH-8092 Zurich, Switzerland

a r t i c l e i n f o a b s t r a c t

Article history: Reports on human conjugated linoleic acids (CLAs) intake vary, and usually don't consider food
Received 29 July 2009 production methods and vaccenic acid (VA)-induced CLA biosynthesis in humans. The aim of this review
Received in revised form was to estimate the achievable CLA intake considering both food sources produced by feeding patterns
30 December 2009
natural to animals and human VA bioconversion. Exogenous CLA supply from milk, cheese, lamb, and
Accepted 25 January 2010
beef from grass-based ruminant production methods, was calculated to be about double that of estimates
based on modern production methods (respectively, from the four sources: 1.25, 1.50, 1.44, and 0.69% of
total fatty acids). Using available human consumption data this resulted in an estimated achievable CLA
intake (including VA bioconversion) of between 711 and 1107 mg d 1. Intake of products from natural
grass-fed ruminants leads to a several-fold higher CLA food content and, along with consideration of
endogenously formed CLA from VA, results in a substantially higher CLA availability than previously
estimated.
Ó 2010 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
2. Literature data and selection criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
3. Assimilation of data on CLA food content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
3.1. Fat content of food sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 434
3.2. Distinction of feeding strategy and determination of CLA content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
4. Achievable CLA availability for humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
5. CLA and VA content in food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
5.1. Dietary sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
5.2. Factors influencing CLA and VA content in food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 436
5.3. Correlation between CLA and VA in food sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
5.4. CLA and VA content in non-domesticated animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 437
5.5. CLA and VA content in meat and dairy foods from animals with natural feeding practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438
6. Achievable CLA availability for humans with regard to foods from natural animal food production methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438
6.1. Bioconversion of VA to CLA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438
6.2. CLA intake studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 440
6.3. CLA supply from fat and food disappearance data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 440
6.4. CLA intake from dietary guidelines and a selective diet aiming to maximize CLA availability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 442
6.5. Influence of nutritional behaviour on total available CLA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 443
7. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444

* Corresponding author. Tel.: þ41 43 888 1818; fax: þ41 43 888 1800.
E-mail address: r.jutzeler@sponser.ch (R.P. Jutzeler van Wijlen).

0958-6946/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.idairyj.2010.01.008
434 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
1. Introduction
Current dietary recommendations generally call for a moderate
fat intake and are often particularly restrictive with respect to
saturated fatty acids and/or fat of animal origin. However, it is
undisputed that dietary fats contain essential fatty acids and
transport essential fat-soluble vitamins, making dietary fat as
a whole an essential constituent of a balanced diet. Furthermore, it
has become evident during recent decades that a more differenti-
ated view on these still contemporary recommendations is defi-
nitely warranted. One topic related to dietary fat recommendations
of particular interest for dairy science and the dairy industry is the
health impact of naturally occurring trans fatty acids (TFAs), which
are found mainly in fats from ruminant animals. Although a range
of diverse TFA isomers occur in these fats, only a few have been
investigated thoroughly and the conjugated linoleic acids (CLAs)
are among them.
The term CLA was coined to describe linoleic acid isomers with
conjugated double bonds (Ha, Grimm, & Pariza, 1987), thus
including isomers in trans configuration (from 7,9; 8,10; 9,11; etc.,
up to carbon 12,14 in cis/cis, trans/cis, cis/trans, trans/trans
configuration: Martin & Valeille, 2002). In contrast to the purely
chemical nomenclature, where some CLAs are clearly defined as
TFAs, opinions about CLAs in respect to food product labelling are
conflicting. Although according to the Codex Alimentarius
Commission (an FAO/WHO body that aims to develop food stan-
dards, guidelines and related texts) TFAs with conjugated double
bonds such as CLA are excluded from the TFA definition for legis-
lative and declaration purposes (Codex Alimentarius Commission,
2006), some organizations (e.g., the French Food Safety Agency)
propose not distinguishing between industrial or ruminant origin
of TFAs for labelling purposes, thereby including all trans isomers of
CLA (Ledoux, Juanéda, & Sébédio 2007; Léger, Razanamahefa, &
Margaritis, 2007).
The predominant isomer of naturally occurring CLA in food
sources is the c9,t11-18:2 isomer (>80% of total CLAs) (McLeod,
LeBlanc, Langille, Mitchell, & Currie, 2004; Troegeler-Meynadier &
Enjalbert, 2005), also called rumenic acid (RA) due to its forma-
tion in ruminants by microbial hydrogenation of linoleic and lino-
lenic acid. Though the name RA suggests that the rumen is the
major production site, CLA actually originates mainly from endog-
enous desaturation of TFAs in the adipose tissue (Gruffat, Rémond,
Durand, Loreau, & Bauchart, 2008) and mammary gland (Schmid,
Collomb, Sieber, & Bee, 2006) and not from microbial hydrogena-
tion in the rumen (Bauman, Baumgard, Corl, & Griinari, 2007; Kay,
Mackle, Auldist, Thomson, & Bauman, 2004; Palmquist, Lock,
Shingfield, & Bauman, 2005). Next to RA, considerable amounts
(2% of total fatty acids) of other CLA isomers are found in beef and
dairy products (Fritsche et al., 1999; Khanal & Olson, 2004;
Ponnampalam, Mann, & Sinclair, 2006). When discussing the
potential health impact of dietary CLA intake in humans, one
should also consider the dietary intake of the t11-18:1 fatty acid,
named vaccenic acid (VA) after the Latin name ‘vacca’ for cow. VA is
the main trans-18:1 isomer in milk fat from cows fed a natural diet
and it can be converted to RA in humans (Mosley, McGuire,
Williams, & McGuire, 2006; Turpeinen et al., 2002).
CLAs possibly influence a vast range of health aspects such as
body composition, blood lipid profile, insulin sensitivity and dia-
betes, atherosclerosis, oxidative and inflammatory status, carcino-
genesis, bone formation and maintenance, and the immune system
(Belury, 2002; Bhattacharya, Banu, Rahman, Causey, & Fernandes,
2006; Terpstra, 2004; Wahle, Heys, & Rotondo, 2004; Whigham,
Cook, & Atkinson, 2000). Besides, some concomitant safety
concerns (Belury, 2002; Bhattacharya et al., 2006; Wahle et al.,
2004), especially the effect of CLA on body composition, have
gained attention (Close, Schoeller, Watras, & Nora, 2007; Gaullier
et al., 2005; Whigham, Watras, & Schoeller, 2007). However,
because of the large number of possible isomers it is difficult to
assign an observed influence of CLA on human metabolism to
a particular isomer. For example, alone eight geometrical and
positional isomers of 9,11 and 10,12 CLA are possible. The situation
becomes more complicated due to the fact that the different and
multiple effects on health appear to be related to the variety of CLA
isomers (Tricon & Yaqoob, 2006).
In many previous studies, the content of CLA in different food
sources and its variation depending on animal feeding patterns
were examined. However, until now, studies on human intake of
CLA have mainly concentrated on simply recording human diets
and estimating CLA intake by analyzing market foods or referring to
nutrient databases. In Europe, the actual intake of CLA from food
sources is estimated to be about 300 mg d 1 (European Food Safety
Authority, 2004). Yet, up to now, it was not often considered that
our present Western diet and lifestyle, as well as modern cattle
feeding practices, might lead to much lower levels of CLA in our
diets than in ancestral times. Therefore, the objective of this review
was to estimate the potentially achievable CLA availability for
humans considering on the one hand exogenous CLA supply
derived only from foods produced with feeding patterns natural to
animals (and using Swiss food consumption data as an example)
and on the other hand taking into account the endogenous CLA
formation from VA.
2. Literature data and selection criteria
Data for this review were obtained by searching the regularly
updated, web-based reference listing on CLA (Pariza, 2009), which
has been keeping track of published studies on CLA since at least
1987, the year of coinage of the term CLA. The reference listing was
searched against certain inclusion criteria for references published
during the past decade. The criteria were: (i) human studies and
reviews related to CLA and food intake; (ii) studies and reviews on
CLA content in animals and animal-derived food; and (iii) feeding
pattern studies of ruminants and non-ruminants if diet was based
on natural feeding practices of the respective animal species (i.e.,
<50% grain-based diets with ruminants, not enriched with plant
oils, fish oils nor directly supplemented with CLA).
In addition, for completion reasons and to uncover older studies
of interest on human CLA intake, a search by term (CLA, conjugated
linoleic, humans, intake, content, review) was undertaken using
PubMed (National Library of Medicine, 2009) and Google scholar
(Google, 2008). The search considered articles published until the
end of September 2008.
3. Assimilation of data on CLA food content
3.1. Fat content of food sources
The CLA and other fatty acid values reported in the retrieved
articles were not always unambiguously described, i.e., it was,
unfortunately, not always clear if total fatty acids or rather the
analyzed fatty acid methyl esters (FAMEs) were reported. It would
be helpful if a standardized way of reporting fatty acid analyses was
defined by an appropriate body. For this review, unclear data have
been considered as FAMEs.
Furthermore, the total fat content was not always given when
CLA values were reported. This is also somewhat unfortunate,
because without information on absolute values any discussion on
the potential effects of a substance is hardly possible. Thus, the total
fat content needed to be estimated if it was missing. These missing
fat values were either derived from other publications retrieved for
 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
this review that did contain both CLA and total fat values or from
standard national food composition tables/databases.
Cheese data originating from grass-based feeding strategies
were pooled together irrespective of the milk origin, i.e., cow or
sheep (no data for goat), and 32% fat content taken from an earlier
review (Dhiman, Nam, & Ure, 2005a) was taken as the cheese fat
percentage when fat percentage was not indicated.
3.2. Distinction of feeding strategy and determination of CLA
content
The median CLA content of foods derived from animals on grass-
based feeding strategies was calculated and compared with the
median CLA values of foods derived from animals on standard
feeding strategies reported by Dhiman et al. (2005a). However, it
should be noted that grass-based feeding strategies were defined as
those strategies including up to 50% grain-based feed (termed
“optimal modern feeding practice”), because a year-round all-
pasture access (i.e., a 0% grain-based diet) is not a realistic method
in most of Western Europe. Thus, the estimation of a naturally high
CLA content reported below likely represents an underestimation
of the highest possible natural CLA content.
Ovine and caprine milk were disregarded for the calculation of
the median CLA content of milk due to lack of data and their only
minor importance in most Western culinary cultures. Moreover,
a distinction related to the feeding strategy was not possible either
for fermented dairy products or for butter due to insufficient data.
In many studies, only c9,t11 CLA was analyzed. However, consid-
ering that this isomer normally constitutes >80% of total CLA, those
values were taken as total CLA for the purpose of the present
review, thus tolerating a slight underestimation of total CLA
content.
For bovine meat, the CLA median value resulted from intra-
muscular (IM), intermuscular (IT) and subcutaneous (SF) fat, where
the overall content of CLA in dissectible beef fat was formed using
an average fat distribution ratio of 14% IM, 68% IT, and 18% SF,
reflecting the analyzed mean fat contribution in three cattle
genotypes (Aldai, Najera, Dugan, Celaya, & Osoro, 2007). For goat
and sheep meat, however, only lamb IM values were used for
determination of a grass-based median CLA content due to there
being only two available values for CLA content in SF and no value
in IT. To deal with the variety of wild animals and their lesser
culinary importance, only dissectible fat from red deer was taken
into account for CLA median formation in game (mixed feeding
strategy) (Phillip, Oresanya, & Jacques, 2007). For non-ruminant
meat and meat products (e.g., chicken, pork, sausages, and ham),
data from two previous reviews of market products (Dhiman et al.,
2005a; Fritsche & Steinhart, 1998) and a single all-pasture study
with turkey (Troegeler-Meynadier & Enjalbert, 2005) were
considered. The CLA values obtained for chicken, pork and meat
products were used for the evaluation of both standard and grass-
based feeding strategies. Meat samples were mostly taken from
a rib section. Where indicated, the muscle samples analyzed were
mainly longissimus dorsi, further semitendinosus, supraspinatus,
semimembranosus, biceps femoris, and thoracis, or mixtures of
several muscles. Differentiation according to muscles pieces was
disregarded in this review.
4. Achievable CLA availability for humans
Three generally different approaches were chosen to estimate
achievable CLA availability for humans to derive an estimate that
was as robust as possible. These approaches were based on food
intake data (i.e., food amounts reported to have been eaten), food
supply data (i.e., food amounts available but not necessarily eaten),
435
and food intake recommendations (i.e., food amounts suggested to
be eaten). For the two approaches of supply and recommendation,
two different models were used. This resulted in a total of five
models.
Model 1 was based on CLA intake studies. All retrieved CLA
intake studies were reviewed and briefly commented on. For the
calculation of the potentially achievable CLA availability with this
model, the retrieved CLA intakes were multiplied by a factor of two
to allow for a higher CLA content in food resulting from natural
grass.
Model 2 was based on CLA supply from fat disappearance data
(using Switzerland as example), with dietary fat supply data from
the Swiss Nutrition Report (Jacob, 2005) providing the basis from
which to work. The CLA supply was derived by multiplying the per
capita fat supply from milk, meat and meat products with the
corresponding CLA content estimated according to Section 3
described above.
Model 3 was based on CLA supply from food disappearance data
(again using Switzerland as example). Swiss per capita food
disappearance retrieved from the web-based FAO Statistics Divi-
sion, section “core consumption data” (FAOSTAT, 2005), and
national Swiss data (Gremaud, Schmid, & Sieber, 2005;
Schweizerischer Bauernverband, 2005) served as the basis for
this model. The difference in the “CLA supply from fat disappear-
ance data” model was that we estimated the fat content of the foods
in this model based on fat content data of foods retrieved during
this study and/or the Swiss Food Composition Database
(Bundesamt für Gesundheit and ETH Zürich, 2003). The CLA supply
was then derived as with the “CLA supply from fat disappearance
data” model.
Model 4 was based on CLA intake from the Swiss food based
dietary guideline. The recommendations for foods of animal origin
as defined by the Swiss Food Pyramid (Walter, Infanger, &
Mühlemann, 2007) are: “Each day, alternate between 1 portion of
meat, fish, eggs, cheese or other sources of protein e.g., tofu
(1 portion ¼ 100e120 g of meat/fish [fresh weight] or 2e3 eggs or
200 g of fresh/cottage cheese or 60 g of hard cheese or 100e120 g of
tofu). In addition, consume 3 portions of milk or dairy products
a day, preferably reduced fat varieties (1 portion ¼ 200 mL of milk
or 150e180 g of yoghurt or 200 g of fresh/cottage cheese or
30e60 g of cheese) .. Use 2 teaspoons (10 g) a day of butter or
margarine made from high-quality oils to spread on bread as
required”. The CLA intake was derived from the food intake
resulting from the pyramid guidelines (i.e., 10 g butter, 120 g beef,
200 mL partly skimmed milk, 180 g yoghurt and 60 g cheese)
multiplied by the CLA content of the respective food source and as
estimated as in Section 3.
Model 5 was based on CLA intake from a selective diet aiming to
“maximize” CLA availability. The CLA intake derived in this model
considered a high, but not the highest possible, CLA intake with
a diet consisting of food products naturally high in CLA (as esti-
mated in Section 3) and the endogenous CLA formation from VA.
The reason why this model does not reflect the real maximal CLA
availability is mainly because with natural grass-based feeding
practices up to 50% grain-based feed was tolerated.
Finally, the CLA quantities from standard and grass-based
feeding strategies resulting from the described five models were
compared. Moreover, the endogenous CLA contribution to be
expected from the bioconversion of VA was added to the median
exogenous CLA values of all models by multiplication with the
corresponding factor as described in Section 6.1.
The literature search revealed numerous articles containing
information on the CLA and/or VA content of food (more than 60
articles, see Tables 2 and 3), which provided the basis for estimating
a maximally achievable CLA intake for humans.
436 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448

5. CLA and VA content in food
5.1. Dietary sources
The most common dietary sources of CLA are ruminant milk,
dairy products from ruminants, and their meat and adipose tissues.
An overview on CLA containing food sources (Khanal & Olson,
2004) classified milk as the most important CLA food source. In
ruminant milk, the primary CLA isomer is present in the c9,t11
configuration and mainly synthesized through desaturation of
dietary VA by D9-desaturase (i.e., stearoyl-CoA desaturase, EC
1.14.19.1) in the mammary glands at a conversion rate of z80%
(Mosley, Shafii, Moate, & McGuire, 2006), making VA the main
contributor to milk-fat CLA. However, CLA is also directly synthe-
sized e also from VA e in muscle and adipose tissues both in
ruminants (Bauman et al., 2007; Raes et al., 2004), humans (Adlof,
Duval, & Emken, 2000; Kuhnt, Kraft, Mockel, & Jahreis, 2006;
Mosley, McGuire, et al., 2006; Salminen, Mutanen, Jauhiainen, &
Aro, 1998; Turpeinen et al., 2002), and monogastric animals, for
example, pigs (Gläser, Wenk, & Scheeder, 2002).
Except for the t7,c9 CLA isomer, which is converted from t7 18:1
(Collomb, Sieber, & Butikofer, 2004) and frequently mentioned the
second-most important isomer in ruminant fat (Alfaia et al., 2007a;
Bauman et al., 2007), all other isomers are supposed to arise from
ruminal microflora by biohydrogenation (Collomb et al., 2004).
Although there is also evidence for the formation of CLA or its
precursors by human colon microflora, and the fact that local
effects such as anti-inflammatory and anti-proliferative effects on
colonocytes have been demonstrated, the uptake of CLA from this
source is probably negligible (Devillard, McIntosh, Duncan, &
Wallace, 2007).
5.2. Factors influencing CLA and VA content in food
Multiple factors influence the CLA content of food sources (Table
1). Animals in modern production systems are fed on silage and
crop concentrate rather than pasture and this leads to a great
reduction in CLA content in foods derived from these animals and
consequently to lower CLA availability in human diets. It has been
repeatedly shown that an animal-adapted feeding pattern, i.e.,
grazing for cows, not only increased CLA in meat and dairy prod-
ucts, but also mono- (MUFA), polyunsaturated (PUFA), n 3 fatty
acids, and VA content, while at the same time reduced saturated
fatty acids (SFA) and increased the n 6/n 3 ratio substantially
(Bisig et al., 2008; Couvreur, Hurtaud, Lopez, Delaby, & Peyraud,
2006; Dannenberger et al., 2004; Dewhurst, Shingfield, Lee, &
Scollan, 2006; French et al., 2000; Garcia, Pensel, et al. (2008);
Hollo et al., 2005; Kraft, Kramer, Schoene, Chambers, & Jahreis,
2008; Nuernberg, Nuernberg et al., 2005; Or-Rashid, Odongo,
Subedi, Karki, & McBride, 2008).
Two studies reported a fivefold higher CLA content in milk fat
from grazing cows (2.21% and 2.53% of total fatty acids) compared
with cows fed a 50:50 diet of conserved forage and grain (Dhiman,
Anand, Satter, & Pariza, 1999; Khanal, Dhiman, & Boman, 2008).
Similarly, beef muscle from steers raised on fresh pasture and/or
forage contained up to six times more CLA compared with that from
steers fed a corn grain-based feedlot diet (Poulson, Dhiman,
Cornforth, & Olson, 2001; Poulson, Dhiman, Ure, Cornforth, &

Table 1
Factors influencing conjugated linoleic acids (CLA) content in food sources.

Factor Leading to CLA reduction Leading to CLA increase References

Season Winter, autumn Spring, summer Addis et al. (2007); Bisig et al. (2008); Dunshea et al. (2008); Ferlay et al.
(2008); Nudda, McGuire, Battacone, and Pulina (2005); Rego et al.
(2008); Thorsdottir, Hill, and Ramel (2004)
Feeding regimen Silage, concentrate Grazing, forage Chilliard and Ferlay (2004); Collomb, Schmid, Sieber, Wechsler, and
Ryhanen (2006); D'Urso et al. (2008); Dhiman et al. (1999); Dunshea
et al. (2008); Ferlay et al. (2008); Ferruzzi et al. (2007); French et al.
(2000); Kay et al. (2004); Kelly et al. (1998); Kraft et al. (2008); Leiber,
Kreuzer, Nigg, Wettstein, and Scheeder (2005); Lucas et al. (2006);
Nuernberg, Nuernberg et al. (2005); Ponnampalam et al. (2006);
Poulson et al. (2001, 2004); Realini et al. (2004); Rego et al. (2004);
Stockdale et al. (2003); Tyagi et al. (2007)
Botanical Mono-cultural, species-poor Species-rich, clover Addis et al. (2005); Couvreur et al. (2006); De Noni and Battelli (2008);
composition Lorenzen et al. (2007); Mel'uchová et al. (2008); Or-Rashid et al. (2008)
Freshness of grass Maturity, conserved, grass Freshness Dhiman et al. (1999); French et al. (2000); van Dorland, Wettstein,
silage Aeschlimann, Leuenberger, and Kreuzer, 2007; Wyss and Collomb
(2008)
Supplementation Barley grain CLA, fish oil, plant oil and seeds AbuGhazaleh, Schingoethe, Hippen, and Kalscheur (2004); Dhiman et al.
(soy, linseed, rapeseed, sunflower, (2005b); Gonthier et al. (2005); Nudda et al. (2006); Song, Kang, Lee,
flaxseed) Jeung, and Yang (2007); Stockdale et al. (2003); Zheng et al. (2005)
Altitude Lowland pastures Highland, mountain pastures Collomb, Butikofer, Sieber, Bosset, and Jeangros (2001); De Noni and
Battelli (2008); Leiber, Scheeder, Wettstein, and Kreuzer (2004); Leiber,
Kreuzer, Nigg, Wettstein, and Scheeder (2005) ;Or-Rashid et al. (2008)
Animal suborder Non-ruminants Ruminants Rule et al. (2002)
Species (meat) Cattle Buffalo, lamb, bison, kangaroo Cordain et al. (2002); Engelke et al. (2004); Giuffrida, Arenas de Moreno,
Beriain, Huerta-Leidenz, and Smith (2005); Patkowska-Sokola, Jamroz,
Bodkowski, Cwikla, and Wertelecki (2002); Rule et al. (2002)
Breed Cows' milk: Brown Swiss > Holstein-Friesian > Jersey White et al. (2001); Hollo et al. (2005)
Cows' meat/fat: Hungarian Grey > Holstein-Friesian Wachira et al. (2002)
Lambs' meat/fat: Soay > Friesland > Suffolk Tsiplakou, Mountzouris, and Zervas, (2006); Tsiplakou, Kominakis, and
Sheeps' milk: no influence of breed Zervas (2008)

Inter-individual variation and breed-specific fatness De Smet, Raes, and Demeyer (2004); Lock, Bauman, and Garnsworthy,
probably more important than breed effects (2005)
Food processing Negligible or no influence Badiani et al. (2004); Buccioni, Rapaccini, Minieri, and Antongiovanni
(2007); Herzallah, Humeid, and Al-Ismail (2005); Knight et al. (2004);
Lock et al. (2005); Shantha, Crum, and Decker (1994)
 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
Olson, 2004) or concentrate (Realini, Duckett, Brito, la Rizza, & De
Mattos, 2004; Steen & Porter, 2003). Such increases of CLA in
milk and meat were similarly demonstrated in other studies (Kelly,
Kolver, Bauman, Van Amburgh, & Muller, 1998; Schroeder et al.,
2003; White et al., 2001). This outcome is related to a higher
PUFA concentration in grass compared with concentrates (French
et al., 2000) and might also be related to the high a-linolenic
(ALA):linoleic acid ratio of the animals' diet (Mel'uchová, Blasko,
Kubinec, & Górová, 2008). Both fatty acids contribute either
directly or indirectly in pasture for c9,t11 CLA biosynthesis. Grazing,
compared with conserved forage and grain-based diets, increases
ruminal pH and thereby favours the microbial production of CLA
and VA, while the addition of grain and forage conservation tech-
niques may decrease ruminal pH, which in turn negatively affects
ruminal fermentation in respect to formation of CLA and VA
(Dhiman et al., 1999; French et al., 2000).
In moderate climates like in central Europe, where cattle cannot
feed on fresh grass the whole year round, important seasonal
differences with lower CLA content during wintertime have been
demonstrated, as well as a greater CLA content at higher altitudes
(Table 1). High altitude was not only associated with an increased
total CLA level in milk fat, but also with a shift in the relation
between the different isomers, for example, t11,c13 CLA increased
by 310% thus becoming the second-most important isomer
(Collomb et al., 2004). Such a shift in the distribution pattern of CLA
isomers in muscle tissues and subcutaneous fat was also observed
in another study comparing bulls on a pasture or a concentrate diet,
respectively. t11,c13 CLA again became the second-most important
isomer with a very high accumulation rate of up to a factor of 14.5,
and the second- and third-highest accumulation rates were
observed for the t11,t13 and t12,t14 isomers, with a factor of up to
7.1 (Dannenberger et al., 2005). Of some concern is that milk fat and
milk yield depression were reported and correlated with increased
proportions of t10,c12 CLA (Baumgard, Corl, Dwyer, Saebo, &
Bauman, 2000; Peterson, Matitashvili, & Bauman, 2003), caused
by energy-dense diets high in starch and C18:2 n 6 fatty acids as
are present in concentrates or maize silage (Nielsen, Straarup,
Vestergaard, & Sejrsen, 2006). However, other studies found no
(Rego et al., 2004) or only transient (Khanal et al., 2008) milk yield
and milk fat depression (yield declined after changing from pre-
pasture to pasture diet and returned to near pre-pasture levels after
the pasture, fat content increased initially when cows were turned
on pasture and then declined and stabilised).
Regarding meat fat, a crucial factor in divergent CLA and VA
values between breeds is obviously the varying inbred fat accu-
mulation. Furthermore, the sampling site of a piece of meat, a lean
versus a fatty steak and the type of tissue fat, results in considerable
differences in fatty acid profile. SF and IT adipose tissues are not
only richer in total CLA than IM fat, but also contain more VA (Hollo
et al., 2005; Nuernberg et al., 2005b; Wachira et al., 2002). The VA
content in different fatty tissues of bulls as measured by Hollo et al.
(2005) was 4.10e7.57% of total fatty acids in subcutaneous haunch
fat, 4.20e11.52% in kidney fat, 3.22e7.91% in visceral fat, and
0.57e1.10% in IM fat. Variations were due to breed and extensive or
intensive production methods. Last but not least, small ruminants
such as lambs, Nubian goats, and crossbred sheep contained higher
amounts of VA (Knight, Knowles, Death, Cummings, & Muir, 2004;
Lee, Kannan, & Kouakou, 2006) in the range of 3.8e5.7% of FAME
(IM fat) (Nuernberg et al., 2005b).
Efforts by the meat and dairy industries to enhance the CLA
content of their food products by adding plant or fish oils into feed,
or even supplementing CLA directly to the animals, proved
successful and this topic has been investigated in many studies that
found several-fold increases in CLA food content using such
measures (Bell, Griinari, & Kennelly, 2006; Boken, Staples,
437
Sollenberger, Jenkins, & Thatcher, 2005; Boles, Kott, Hatfield,
Bergman, & Flynn, 2005; Dhiman et al., 1999, Dhiman et al.
(2005b); Gonthier et al., 2005; Griswold et al., 2003; Poulson
et al., 2001; Ward, Wittenberg, & Przybylski, 2002; Zheng et al.,
2005). Nevertheless, such supplementation of fodder is not dis-
cussed in more detail here as the feeding of grains, seeds, oils and
other additives to ruminants is not considered to constitute
a natural feeding pattern and the objective of this review was to
evaluate the natural content of CLA in our diet that is achievable by
species-adapted feeding regimens and a healthy way of eating.
In summary, the feeding pattern is the most important factor
in CLA content both in milk and meat, though diet does not
necessarily increase CLA and VA in all tissues (Shen et al., 2007).
Animals grazing on pasture had substantially higher CLA in their
milk, muscle and adipose tissue as a proportion of total fatty
acids; albeit at the same time possibly reducing total fat content
(Nuernberg, Dannenberger, et al., 2005). Nonetheless, even if total
fatty acids are reduced, the relative CLA increase is generally so
large that ultimately an increased absolute CLA content is
observed.
5.3. Correlation between CLA and VA in food sources
A close linear relationship between CLA and VA was observed
in milk and meat fat (Chilliard & Ferlay, 2004; Khanal et al., 2008;
Lourenço, Vlaeminck, Bruinenberg, Demeyer, & Fievez, 2005;
Mel'uchová et al., 2008; Nielsen et al., 2006; Noci, Monahan,
French, & Moloney, 2005; Nuernberg et al., 2005a). More
recently, substantial VA contents have been analyzed in cow's milk
(2.9e3.8% of FAME) and cheese (3.1e3.6%) from mountain pasture
(1400e2200 m above sea level) (De Noni & Battelli, 2008). In two
other recent studies, even VA contents of as high as 7.63% and
7.95% of milk fat from cows fed an all-pasture diet were reported
(Dunshea, Walker, Ostrowska, & Doyle, 2008; Khanal et al., 2008).
According to a recent study, a VA:RA ratio of approximately 3:1
was observed in milk, butter, and cheese of cows on a modified
diet to increase CLA in their milk (Tricon et al., 2006). Somewhat
lower VA:RA ratios were found in cheese from yak and cow (2.4:1
and 2.7:1, respectively), with expectedly higher amounts of both
VA (2.27% versus 0.57% of FAME) and RA (6.23% versus 1.35%) in
yak-derived cheese due to altitude and grass diversity (Or-Rashid
et al., 2008). These results correspond to 4.0 and 4.6 times
higher contents of VA and RA, respectively, in yak compared with
cow cheese.
5.4. CLA and VA content in non-domesticated animals
The CLA content of naturally feeding non-domesticated animals
should be indicative of an evolutionarily “planned” normal CLA
content. In this context, Cordain et al. (2002) examined the lipid
composition of different wild North American and African rumi-
nants as well as of cattle fed pasture and grain. The authors
concluded that tissue lipids of wild ruminants were similar to those
pasture-fed, but dissimilar to grain-fed cattle, and that the lipid
composition of wild ruminant tissue may serve as a model for
dietary lipid recommendations. This outcome is in accordance with
a study comparing both range-fed bison and cow meat to wild elk
(Rule, Broughton, Shellito, & Maiorano, 2002). Nevertheless, it has
to be noted that body fat percentage in wild mammals reflects
seasonal fluctuations and may range from as low as 2.5% to more
than 20%, if latitudinal climate conditions require this as a survival
strategy (Cordain et al., 2005). Obviously, this varying fat content
strongly influences the intake of fatty acids.
To illustrate the influence of total fat percentage on a potential
CLA intake, different forage to concentrate ratios (75:25, 50:50,
438 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
25:75) offered to domesticated red deer not only substantially
changed relative CLA content (1.5%, 1.1% and 6.7% of total fatty acids
of dissectible fat, respectively), but also the total amount of dis-
sectible fat (6.2, 7.3 and 10.9 g 100 g 1 of a defined rib section,
respectively). This then led to an absolute CLA content in the low-
forage fed deer that was 8-fold higher (730 mg 100 g 1 of rib
section) than that in the deer fed high forage ratio (Phillip et al.,
2007).
In addition to a higher CLA content, game also has markedly
higher VA content than cattle, as well as higher ALA content
(Cordain et al., 2002; Rule et al., 2002). Remarkable differences can
occur in VA content between muscle of game and cattle that cannot
be accounted for by feeding patterns only, but are distinct to species
(Engelke, Siebert, Gregg, Wright, & Vessby, 2004). The highest
amount of VA hitherto was found in the SF of kangaroo, which is
a non-ruminant. However, the microbial fermentation occurring in
the kangaroo's foregut results in an 12.4% of total fatty acids,
together with a very high 3.8% of RA (Engelke et al., 2004).
Furthermore, the comparison of subcutaneous backfat of barley-fed
beef with that of wild musk ox showed almost double amounts of
total trans C18:1 (4.25 versus 2.67% of FAME) and at the same time
quite the opposite ratio of VA (0.77 versus 1.41%) (Dugan et al.,
2007). As a result, the ratio of total trans C18:1 to VA differed
even more between beef and musk ox (5.5 versus 1.9). Although the
CLA content in musk ox was slightly lower (0.50 versus 0.67% of
FAME), it had a higher content of n 3 fatty acids and a lower n 6/
n 3 ratio as would be expected regarding its wild-pasture diet.
5.5. CLA and VA content in meat and dairy foods from animals with
natural feeding practices
Today's agriculture is faced with the inherent problem of
developing production methods that are efficient in terms of
quantity, satisfying from an economical perspective, sustainable,
and at the same time yielding high-quality food for humans. All
these requirements might not be satisfied concomitantly. As an
example, though it seems evident that alpine cheese produced
from the summer milk of cows feeding on fresh grass with a high
clover content may deliver a different fatty acid profile regarding
CLA, n 3, and n 6/n 3 ratio, it is obviously not possible to
produce enough cheese to cover market demands exclusively from
alpine milk, only during summer, and from cows with access to
fresh grass.
Considering only 100% grass-based production methods to
estimate a naturally high CLA intake would, therefore, be far from
realistic. Correspondingly, the present review did not only consider
exclusively grass-based production methods, but allowed up to 50%
grain-based production methods to be included in the “optimal
modern production methods” for the later calculation of a naturally
high CLA intake (see Section 6). Even with this allowance the CLA
content of cow's milk, cheese, beef and lamb meat was estimated to
be about twice than previously estimated (Dhiman et al., 2005a)
(Table 2): 1.25 versus 0.57%, 1.50 versus 0.71%, 1.44 versus 0.69%,
and 0.69 versus 0.38%, FAME for milk, cheese, lamb, and beef,
respectively.
Due to lack of data, no differentiation of feeding strategies could
be made for butter and yoghurt. Likewise, data solely from market
products without feeding strategy differentiation were found for
meat products and non-ruminant meat. Meat products exist in
a vast variety and differ markedly in macronutrient composition.
Consequently, it is not possible to estimate the CLA contribution
from this food group. Nevertheless, as with non-ruminant meat,
CLA seems generally to be rather low in meat products and,
therefore, the lack of data for this food group is not of great rele-
vance for the assessment of CLA contribution in the authors'
opinion. Although in wild animals a feeding strategy does not
apply, husbandry of wild animals and concomitant feeding
methods have also become more relevant. For example, substantial
CLA contents have been found in domesticated deer, possibly the
most common culinary game in Switzerland.
In the present review, VA:CLA ratios similar to those described
in the literature (see Section 5.3) were found in cows' milk, cheese,
beef, and lamb from grass-based production methods of 2.3:1, 2.1:1,
3.4:1, and 4.4:1 respectively (Table 3). A lack of data prevented
calculation of the VA:CLA ratio for other food categories.
6. Achievable CLA availability for humans with regard to
foods from natural animal food production methods
6.1. Bioconversion of VA to CLA
As previously mentioned, monogastric animals and humans
endogenously produce CLA from VA. Nevertheless, until now, VA
was not usually considered in estimates of human CLA intake (see
Section 6.2). As soon as one considers the bioconversion of VA,
however, it seems to be more appropriate to speak about total CLA
availability in contrast to only CLA intake from the diet.
A model to predict exogenous and endogenous contributions to
RA in several tissues was first developed for lambs by Palmquist, St-
Pierre, and McClure (2004). In humans, the bioconversion of VA to
RA was investigated in a study designed with a macronutrient
composition often considered as healthy (i.e., energy percentage
from carbohydrates 55%, from fat 30%, and from protein 15%), and
containing 1.5, 3.0, or 4.5 g d 1 of VA for 9 days. The reported
average bioconversion from VA to RA was 19% (Turpeinen et al.,
2002), which was associated with a linear increase in the propor-
tion of RA with increasing VA supply. On average, the ingested daily
amount of VA was therefore equivalent to 285 mg, 570 mg, and
855 mg of RA, respectively. Based on that work, it was proposed
(Palmquist et al., 2005) that the CLA content of food multiplied by
1.4 would provide an effective estimate of the total available CLA for
humans (thus total available amount being 140% of the analyzed
CLA amount). Turpeinen et al. (2002) similarly suggested that total
CLA availability, including bioconversion of VA, is likely to be on
average 1.5 times the CLA content of ingested ruminant fat.
Recently, Kuhnt et al. (2006) reinforced the results obtained from
this previous study and observed a conversion rate of 25% in serum
and 19% in red blood cell membranes in humans. They also pointed
out that these results do not reflect the gross conversion rate but
the net sum of the surviving end-products. Hence, those biocon-
version rates were estimated only indirectly from either the
increase in serum RA or red blood cell membranes after VA
supplementation and, thus, did not take into account any disap-
pearance of VA-derived RA into organs or tissues. In this context,
the observation that RA is oxidized to a significantly greater extent
than, for example, linoleic acid in rats is also relevant (Kuhnt et al.,
2006). Thus, the real level of RA synthesized from VA might be even
higher. The authors proposed that the VA conversion rate estimated
in human serum is also representative for whole body conversion,
and that approximately one quarter of dietary VA is transformed to
CLA in the human body (Kuhnt et al., 2006). However, how much
the bioconversion rate is influenced by meal-specific differences
in fatty acid profile and the concurrence of the same enzyme
D9-desaturase, given for example that PUFA decreased bioconver-
sion in mice, still needs further examination (Santora, Palmquist, &
Roehrig, 2000). Furthermore, bioconversion efficiency may be
tissue-dependent as observed in rats (Kraft et al., 2006). Though, as
ruminant fat contains at least 2e3 times more VA than RA, even low
rates of conversion contribute significantly to CLA status in the
body.
 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448 439

Table 2
Conjugated linoleic acids (CLA) content in food in respect of feeding strategies.

Food source Production method CLA contenta Reference

Range (median)
Milk and dairy products
Cows' milk Grass-based feeding 0.47e3.52 (1.25) Collomb et al. (2002); Couvreur et al. (2006); Dhiman et al. (1999);
Dhiman et al. (2005a) Kelly et al. (1998); Khanal and Olson (2004);
Leiber et al. (2004); Lourenço et al. (2005); Prandini, Sigolo, Tansini,
Brogna, and Piva (2007); Rego et al. (2004); Staszak (2005); Stockdale
et al. (2003); Talpur, Bhanger, Khooharo, and Memon (2008); Troegeler-
Meynadier and Enjalbert (2005); van Dorland et al. (2007); Vanhatalo,
Kuoppala, Toivonen, and Shingfield (2007); White et al. (2001)
Fluid milk products Market products 0.34e0.80 (0.57) Dhiman et al. (2005a)
Various cheese types Grass-based feeding 1.04e2.27 (1.50) Addis et al. (2005); Buccioni et al. (2007); De Noni and Battelli (2008);
(cow and sheep milk) Khanal and Olson (2004); Lucas et al. (2006); Or-Rashid et al. (2008);
Tyagi et al. (2007)
Various cheese types Market products 0.34e1.07 (0.71) Dhiman et al. (2005a)
Yoghurt No differentiation 0.38e1.05 (0.47) Fritsche and Steinhart (1998); Khanal and Olson (2004); Martins et al.
of feeding strategies (2007); Prandini et al. (2007)
Fermented dairy foods Market products 0.36e0.94 (0.65) Dhiman et al. (2005a)
(including butter)
Butter No differentiation 0.47e2.09 (0.94) Bertschi et al. (2005); Fritsche and Steinhart (1998); Khanal and Olson
of feeding strategies (2004); Martins et al. (2007); Seçkin, Gursoy, Kinik, Akbulut (2005);
Troegeler-Meynadier and Enjalbert (2005)
Butter Market products 0.47e0.94 (0.71) Dhiman et al. (2005a)
Ruminant meat
Beef, with 10% Grass-based feeding n.a. (0.69) Aldai et al. (2007); French et al. (2000); Giuffrida et al. (2005); Hollo
dissectible fatb et al. (2005); Latimori et al. (2008); Leheska et al. (2008); Lorenzen et al.
(2007); Martins et al. (2007); Noci et al. (2005a); Noci, O'Kiely,
Monahan, Stanton, and Moloney (2005); Nuernberg et al., 2005a;
Poulson et al. (2001, 2004); Realini et al. (2004); Rule et al. (2002);
Shantha et al. (1994); Steen and Porter (2003)
Beef Market products 0.12e0.64 (0.38) Dhiman et al. (2005b)
Lamb, intramuscular Grass-based feeding 0.56e1.90 (1.44) Garcia et al. (2008a); Knight et al. (2004); Martins et al. (2007);
Nuernberg et al. (2005b); Serra, Mele, La Comba, Conte, Buccioni, and
Secchiari (2009); Troegeler-Meynadier and Enjalbert (2005)
Lamb Market products 0.18e1.20 (0.69) Dhiman et al. (2005a)
Deer, 9.1% dissectible fat Domesticated, n.a. (3.90) Phillip et al. (2007)
grass-based feeding
Non-ruminant meat
Turkey, pork, chicken, rabbit Market products 0.06e0.25 (0.16) Dhiman et al. (2005a)
Meat products
Smoked turkey, Market products 0.17e0.24 (0.21) Dhiman et al. (2005a)
smoked bacon
Salami Market products n.a. (0.42) Fritsche and Steinhart (1998)
Salami (Milano-type) Market products n.a. (0.13) Herranz, Ordóñez, de la Hoz, Hierro, Soto, and Cambero (2008)
Sausage (Wiener) Market products n.a. (0.36) Fritsche and Steinhart (1998)
Smoked ham Market products n.a. (0.29) Fritsche and Steinhart (1998)
Overall meat products Median n.a. (0.29) Dhiman et al. (2005a); Fritsche and Steinhart (1998); Herranz et al.
(2008)
a
Values are given as g 100 g 1 fatty acid methyl esters; n.a. ¼ not applicable.
b
10% Total fat (Gerber et al., 2006) with a contribution ratio of inter-, intramuscular, subcutaneous fat of 14%, 68%, and 18% respectively (mean of 3 genotypes, 14% total fat)
(Aldai et al., 2007).

The dietary intake of VA from bovine milk fat was assessed in 15
European countries and ranged from a low 0.3 g (Spain) to a high
0.9 g (Finland) per person per day (Wolff & Precht, 2002). Notably,
these intakes did not include VA from ewe and goat's milk which
are non-negligible dietary sources in some countries, for example
Greece (Wolff, 1995). In the TRANSFAIR study, total VA intake, i.e.,
from animal fat and industrially processed plant oil sources in 14
Western Europe countries was estimated to range from 0.82 g d 1
in Greece to 4.23 g d 1 in Iceland, with most other countries being
between 1.1 and 3.5 g d 1 (Hulshof et al., 1999). The contribution of
natural animal-derived TFA intake ranged from the lowest at 28% in
Norway to the highest at 79% in Germany, translating into an
absolute intake of C18:1 trans fatty acids between 0.98 g d 1
(Norway) and 1.07 g d 1 (Germany). These quantities do not only
reconfirm, but even exceeded the estimated intakes of the first
mentioned European study (Wolff & Precht, 2002) and indicate that
VA intake is not negligible for calculating total CLA availability as is
discussed in the following.
The importance of the contribution of VA to total CLA intake is
further reinforced by a French study (Weill et al., 2002), in which
a huge 233% increase of VA intake (0.39 versus 1.30 g d 1) was
shown, along with 117% increase of RA (0.17 versus 0.37 g d 1),
which was caused by adding extruded linseeds into the animals'
fodder without changing the human diet. Notably, 35% of daily
dietary energy stemmed from fat (mean of 2.9 MJ) and a 24% from
animal fat. There was no difference in serum and plasma VA despite
its increase in the daily regimen, while serum RA content increased
by 50% and 57% in the two assay groups versus control respectively.
Therefore, efficient bioconversion to RA in humans was confirmed.
At a presumed bioconversion rate of 20% the ingested 1.3 g of VA
provided about 260 mg additional, endogenously formed CLA, and
the total available CLA was up to 630 mg d 1.
440 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
Table 3
Vaccenic acid (VA) content and VA to conjugated linoleic acids (CLA) ratio in food from grass-based feeding strategies.
Food source VA contenta
Range (median)
Cows' milk 0.95e7.95 (2.88)
Various cheese types 2.21e6.23 (3.15)
(cow and sheep milk)
Beef, with 10% dissectible fatb n.a. (2.34)
c
Lamb racks, with 8.3% dissectible fat n.a. (5.48 )
a
Values are given as g 100 g 1 fatty acid methylester; n.a. ¼ not applicable.
b
10% Total fat (Gerber et al., 2006) with a contribution ratio of inter-, intramuscular, subcutaneous fat of 14%, 68%, and 18% respectively (mean of 3 genotypes, 14% total fat)
(Aldai et al., 2007).
c
Median of intramuscular and subcutaneous fat, no value for intermuscular fat available.
6.2. CLA intake studies
Twenty-three studies on recorded CLA intake in humans were
found during the literature search (Table 4). In 16 of them, food
frequency questionnaires (FFQ) and/or diet records were used. Five
were related to national nutrition surveys, and two stemmed from
experimental diets. In one further study, the CLA intake was esti-
mated based on recommendations for a balanced, normal daily
diet. In all except two studies, present day food products and their
generally low CLA content were considered. VA and its contribution
to total CLA availability by endogenous synthesis were not
considered in any of the 24 studies.
The pooled median CLA intake of all 24 studies was 278 mg d 1,
if gender was not taken into account and dietary record methods
were neglected (Table 4). However, the intakes diverged strongly
from less than 200 mg d 1 (Jiang, Wolk, & Vessby, 1999; Larsson,
Bergkvist, & Wolk, 2005; Ritzenthaler et al., 2001) to more than
1000 mg d 1 (Desroches et al., 2005; Fremann, Linseisen, &
Wolfram, 2002). The reason for this fivefold difference in CLA
intake is likely to be not only due to varying diets and CLA contents
in food sources, but also because of different study designs and
differences in the quality of the food composition data used to
evaluate the diet. Fritsche et al. (1999) estimated CLA intake based
on analysis of 139 foods from the German market and data from
a German national consumption survey. However, as the authors
acknowledged, the analyzed CLA values were not necessarily
representative. Obviously, this would also not reflect fatty acid
profiles as would be expected by a natural animal feeding regimen.
Yet, the daily CLA intake was 360 mg for women and 440 mg for
men. An estimation of the European Food Safety Authority indi-
cated a possible CLA consumption as high as 1.5 g d 1 in Australia
(European Food Safety Authority, 2004) and a diet applied by
Desroches et al. (2005) reached the highest intake of 2590 mg d 1
using a modified macronutrient profile (dietary energy: carbohy-
drates 45%, fat 40%, protein 15%). Notably, this latter study con-
cerned food production methods that enhance CLA content.
A general problem when estimating CLA intake in dietary intake
studies is that the resulting CLA intake seems to strongly depend on
the method used. The experimental analysis of food duplicates
revealed 20e45% higher CLA intakes compared with an estimation
by dietary record or FFQ (Ritzenthaler et al., 2001). The authors
suggested that the latter methods are not reliable estimators of
individual CLA intakes and may underestimate the total CLA intake
of population groups.
It is also striking that CLA intake studies mostly relied on the
same older analytical food database containing CLA values that
were taken from analytical data from 1992 (Chin, Liu, Storkson, Ha,
& Pariza, 1992) and 1997 (Fritsche & Steinhart, 1998). Due to
Ratio VA:CLA Reference
2.3 De Noni and Battelli (2008); Dunshea et al. (2008); Ferlay et al. (2008);
Khanal et al. (2008); Rego et al. (2004, 2008); Talpur et al. (2008);
Vanhatalo et al. (2007)
2.1 Addis et al. (2005); Buccioni et al. (2007); De Noni and Battelli (2008);
Lucas et al. (2006); Or-Rashid et al. (2008)
3.4 Hollo et al. (2005); Latimori et al. (2008); Leheska et al. (2008); Noci,
et al. (2005a)
4.4 Nuernberg et al. (2005b); Serra et al. (2009)
improved analytical methods, it is reasonable to speculate that if
those “older” foods were reanalyzed today, slightly higher total CLA
values would be obtained. However, it is still not a simple task to
analytically resolve the different CLA isomers and possible losses of
c9,t11 CLA by isomerisation during an analysis such as that
described by Lee and Tweed (2008) could still lead to a slight
underestimation of CLA. The growing analytical data also indicate
substantial differences in the meat CLA content depending on the
examined muscle. IM fat in leg muscles (semitendinosus, semi-
membranosus, rectus femoris, gluteus, and tensor fascia latea)
seem to contain 30e70% more CLA (1.19e1.54% versus 0.9% of total
fatty acids) than the longissimus dorsi muscle of lambs (Garcia
et al., 2008a). According to the authors such muscle differences in
CLA concentration may indicate potential tissue differences in
expression or activity of D9-desaturase. Overall, the CLA intake
reported in many studies does not necessarily correspond to the
real CLA intake and is very likely to be substantially lower than the
possible total CLA availability including bioconversion from VA as
discussed before.
In the present review, the results of the different models chosen
to evaluate CLA supply or intake are summarized in Fig. 1. In the
pooled CLA intake studies, the median CLA intake increased from
278 mg d 1 to a total available CLA intake of 650 mg d 1, resulting
from multiplication of the CLA intakes of each single study by
a factor of two for foods naturally high in CLA (see Section 5.5), plus
a corresponding multiplication of CLA intake with factor 1.5 for VA
bioconversion (see Section 6.1). As mentioned above, the multi-
plication of exogenous CLA intake with a factor 1.4 or 1.5 was
formerly suggested to correct for bioconversion from VA (Palmquist
et al., 2005; Turpeinen et al., 2002). Interestingly, in reinforcement
of such an outcome, 32 out of 77 subjects consuming “normal”
amounts of dairy foods (300e400 g week 1 of cheese) had RA
blood plasma levels equal to or higher than the lower limit of
subjects receiving and additional 1.4 g of supplementary CLA
(Zlatanos, Laskaridis, & Sagredos, 2008). It should be noted,
however, that the production methods of those foods were not
assessed.
6.3. CLA supply from fat and food disappearance data
Total CLA availability from fat and food disappearance data using
the example of Switzerland, including VA bioconversion, was
707e1107 mg d 1 (Fig. 1). These values seem rather low, in partic-
ular if one considers that food or nutrient disappearance data
usually grossly overestimates intake obtained from more direct
dietary assessment methods (FFQ, 7-d protocols, etc.). For example,
in a Danish nutrition report examining both food disappearance
and intake data from 1985e2001 (Fagt et al., 2004), it was noted
Table 4
Estimated conjugated linoleic acids (CLA) intake of different populations.
1
Population CLA intake (mg d ) Method Remarks Reference

All Male Female

France 213 178 Food composition data Individual and national Laloux, du Chaffaut, Razanamahefa,
combined with food survey food survey (INCA1) and Lafay (2008)
127 Assuming a typical daily Dhiman et al. (2005a)
diet with low-CLA products
441 Assuming a typical daily

R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
diet with high-CLA products
Great Britain 370 Food frequency questionnaire (FFQ) Meat-eaters Rosell et al. (2005)
Canada 290 3-d record Low dairy Desroches et al. (2005)
240 Experimental diet according to food Low-CLA dairy fat
2590 recommendations High-CLA dairy fat, CLA
(carbohydrates 45% of energy adjusted by feed regimen
intake, protein 15%, fat 40%)
Sweden 51 Prospective cohort study, FFQ Low dairy Larsson et al. (2005)
177 High dairy
Europe 300 e Average intake estimate European Food Safety Authority (2004)
Netherland 70 Prospective cohort study, FFQ 1st quintile, median 200 mg Voorrips et al. (2002)
290 5th quintile, median 200 mg
15 EU countries 140e380 Only milk consumption considered Correlating with 300e900 mg VA Wolff and Precht (2002)
(per capita and year)
France 370 Guided diet, linseed supplemented Associated with 1300 mg VA Weill et al. (2002)
feeding strategy
Germany 246 FFQ Mean c9,t11; range 81e481 mg Fremann et al. (2002)
323 7-d record Mean c9,t11; range 135e496 mg
1
Canada 95 2  7-d record Only c9,t11 isomer, range 15e174 mg d Ens et al. (2001)
USA 212 151 3-d record, FFQ, food duplicate analysis Ritzenthaler et al. (2001)
Germany 452 512 399 7-d record, national food Mean c9,t11; range 92e1338 mg Fremann (2000)
consumption survey 1985e1988
Germany 430 350 National food consumption Ratio meat:meat products 40:60, Fritsche and Steinhart (1998)
survey 1985e1988 pork:beef 80:20, other species disregarded
Sweden 160 7-d record, 24-h recall Jiang et al. (1999)
USA 291 3-d record Guided high-dairy diet Park et al. (1999)
49e659 FFQ Chronic intake estimate
Finland 310 Diet record (fat 32% of Salminen et al. (1998)
energy intake), food duplicate
analysis
USA 139 3-d record Herbel et al. (1998)
USA 221 265 172 Continuing Survey of Food Beef-eating subjects only Agrawal, Heimbach, Amann, Mohammedshah,
Intake of Individuals (CSFII) and Douglas (1998) (cited in National
Cattlemen's Beef Association, 1999)
Australia 1000 e Values 500e1500 mg Parodi (1994) (cited in European Food
Safety Authority, 2004)
Overall median 278 Pooled data of men, women, and
both sexes; no correction for
methods used

441
442 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448

1200 450
Availability of CLA (mg d -1)

1107
1000
923 400
800

Total available CLA (mg)

774
650
707 350
600
300 e n - CL A
400 e x- CLA
615 m g 344 m g
200 250
(64%) (36%)
0 200
i es

ce

diet
mid
tic
ran
tud

tatis 150

y ra

tive
pe a
s

dp
ly s
led

ec
100
sap

foo

Sel
Poo

upp
di

i ss
ds

50
Fat

Sw
Foo

0
r t) t) i lk t
Fig. 1. Availability of conjugated linoleic acid (CLA) from standard and 50% grass- tte fa fa m ur se
bu L gh ee
based food production methods with consideration of vaccenic acid (VA) bioconver- g . 3% 10
% m yo ch
sion: (,), exogenous CLA from standard feeding strategy, values taken from Dhiman 15 (8 f( 0 g g
s e 40 0 40
et al. (2005a); Fritsche and Steinhart (1998); ( ), exogenous CLA from 50% grass- ck be 15
based feeding strategy; (-), total available CLA (sum of exogenous CLA intake plus
ra g
b
m 75
endogenously formed CLA from VA using a bioconversion factor of 1.5). For the pooled la
studies, the median of pooled CLA intake studies was used for the exogenous CLA data
g
75
(see Table 4), while the median CLA availability was based on pooled studies and
considering enhanced CLA intake from grass-based production methods as well as
Fig. 2. Total available conjugated linoleic acids (CLA) under consideration of exoge-
bioconversion from VA (each single study value was multiplied with a factor of 2 and
nous, food-derived CLA (-) and CLA endogenously formed from vaccenic acid ( ) in
1.5 prior to median formation). For fat disappearance, fat supply data were from
respect of (histogram bars) a selective diet with natural-high CLA food sources and (pie
Switzerland from animal-derived fat (Jacob, 2005; Schweizerischer Bauernverband,
chart) from grass-based production methods. For the pie chart, ex-CLA refers to
2005) and a 50:50 beef meat to meat products ratio was assumed. Data for food
exogenous, food-derived CLA (medians taken from Table 2), and en-CLA refers to
supply statistics were from FAOSTAT (FAOSTAT, 2005) for all food groups, except for
endogenously bioconverted CLA from vaccenic acid at a 20% conversion rate. Median
cheese from 2001/02 which was from Gremaud et al. (2005). For the Swiss Food
vaccenic acid contents were taken from Table 3: the value from milk was also used for
Pyramid (Walter et al., 2007), servings corresponding to the daily recommendations
butter and yoghurt. Total fat intake was 45 g with fat contents of foods derived from
were: 10 g butter, 120 g beef, 200 mL partly skimmed milk, 180 g yoghurt and 60 g
the Swiss Food Composition Database (Bundesamt für Gesundheit and ETH Zürich,
cheese. The fat content of these foods, derived from the Swiss Food Composition
2003) for butter, 86%, milk, 2.8%, and yoghurt, 1.5%; from Gerber et al. (2006) for
Database (Bundesamt für Gesundheit and ETH Zürich, 2003), were: 86%, 10%, 2.8%,
lamb racks, 8.3%; and from Dhiman et al. (2005a) for cheese, 32%.
1.5%, and 30%, respectively. For the selective diet, 45 g d 1 of animal fat, corresponding
to 17% of energy of a 10 MJ diet, the data were taken from Fig. 2.

additionally available CLA (20% bioconversion rate). This would add

that cheese intake was only about half of cheese disappearance. up to a total of z168 mg available CLA. This quantity corresponds to
This outcome has been seen twice, in 1995 and 2000/01. Similar or even exceeds the daily intake estimated by some studies that
losses have to be expected with meat, where asymmetric losses of only looked at exogenous CLA (Ens, Ma, Cole, Field, & Clandinin,
fat and protein probably result from leftovers. 2001; Herbel, McGuire, McGuire, & Shultz, 1998; Larsson et al.,
In consequence, a CLA intake estimated from the retrieved 2005; Ritzenthaler et al., 2001). In contrast to lamb, beef seems to
disappearance data in the present review would be perhaps up to provide only z85 mg 75 g 1 (10% dissectible fat content) of the
50% lower than the supply data themselves, yielding values of total available CLA as shown in Fig. 2. Another example illustrating
about 350e550 mg d 1. the necessity of considering VA is given by the analysis of a mean
VA content of z1000 mg 100 g 1 of cheese (32% fat) from grass-
6.4. CLA intake from dietary guidelines and a selective diet aiming based milk production methods, providing >80 mg CLA per 40 g
to maximize CLA availability serving of cheese (Lucas et al., 2006). Converting these examples
from grass-based production methods in a selective diet aimed at
Using the Swiss Food Pyramid (Walter et al., 2007) as an a maximal CLA availability by choosing food products naturally
example of a food based dietary guideline, it is evident that a CLA high in CLA and VA (and using a 20% VA to CLA bioconversion rate)
availability of >770 mg d 1 (Fig. 1) would be possible if foods as shown in Fig. 2, one obtains 959 mg of total available CLA, along
derived from animals on at least partly grass-based feeding strat- with 45 g of animal fat representing 17% of energy (%E) of
egies were consumed. Assuming standard feeding strategies, a 10 MJ d 1 diet.
however, also the CLA intake estimated from the Pyramid model In this example, the two different approaches to estimating total
resulted in a similar CLA value of between 250 and 350 mg d 1 as available CLA by means of either multiplying exogenous CLA with
with the three other models for estimating total CLA availability a factor of 1.4e1.5 as proposed elsewhere (Palmquist et al., 2005;
(Fig. 1). However, only a slight dietary modification of the Swiss Turpeinen et al., 2002), or by adding analyzed VA at a conversion
Food Pyramid dietary guidelines, as presented in Fig. 2 (i.e., þ5 g rate of 19e25% to CLA (Kuhnt et al., 2006), do match very closely
butter (þ75 g lamb racks e 45 g beef) ¼ þ30 g “meat”, þ200 mL (861e923 mg versus 942e1045 mg). Importantly, this range of total
milk, 30 g yoghurt, 20 g cheese) and the selection of CLA-rich CLA gives a strongly diverging picture from existing intake studies
food sources would yield an intake of more than 900 mg CLA per such as those summarized in Table 4. The non-consideration of VA
day (Fig. 1). Importantly, such a dietary modification would only limits the value of those studies and also substantially underesti-
slightly increase the animal-derived fat contribution to total energy mates total CLA availability. The example presented in Fig. 2
intake. For example, the consumption of a rather small serving of suggests an important 35% contribution of total CLA originating
75 g lamb racks, assuming a total fat content of 8.3% (Gerber, from endogenous VA bioconversion.
Scheeder, & Wenk, 2006), could deliver 100 mg of CLA and also Applying a macronutrient profile diverging from present dietary
z340 mg of VA (Nuernberg et al., 2005b), resulting in z68 mg of guidelines (45%E, 15%E, and 40%E from respectively carbohydrates,
 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
protein, and fat; thereof 24%E of the test butter fat) resulted in
a daily CLA intake of 2590 mg (Desroches et al., 2005). This high
intake of animal-derived fat did not show any deleterious effects on
blood lipid profile. The additional contribution of VA to total
available CLA from that study, using the multiplication factors of
1.4e1.5, would result into a total of 3626e3885 mg d 1.
Actually, the estimates of total available CLA in the present work
have to be considered as fairly conservative considering the
weaknesses described in sections 3 and 4 and strategies discussed
herein would increase total CLA supply and intake, rather than the
opposite. Indeed, while considering partly grass-based feeding
strategies instead of standard methods we could easily more than
double the total available CLA (factor 1.9e2.1, data not shown).
Thus, the sole consideration of all-pasture feeding would obviously
lead to three- to four-fold higher CLA availability.
6.5. Influence of nutritional behaviour on total available CLA
Whereas ancestral hunteregatherers could not rely on dairy
foods, one can assume that they appreciated fatty tissues like liver,
bone marrow, and brain and not only ate the lean meat parts as is
mostly the case nowadays (Eaton, 2006). This, obviously, would
have influenced absolute intake of specific fatty acids including
CLA. For example, high-fat bone marrow from wild animals
contains up to 1.4% RA (Cordain et al., 2002). Before the emergence
of the bovine spongiform encephalopathy crisis during the late
1980s, marrow of bovine origin was appreciated in diverse cultures
as a side dish with soups or meat, and when taking reference values
from wild ruminants (Cordain et al., 2002) it could provide
z150 mg CLA per 20 g portion.
In modern times, the changes in cattle feeding strategies from
pasture to forage and concentrates indeed can influence CLA
availability to humans eating a typical Western diet. This is
reflected in a Dutch study reporting increased RA and VA contents
of about 20e35% in the milk of lactating women, when following
a diet with dairy and meat products of >90% organic versus
conventional origin (Rist et al., 2007). Yet, not only food production
methods but also nutritional behavior influences CLA intake in
humans. Ruminant milk fat and adipose tissues are the most
abundant source of CLA. Dairy products are estimated to contribute
to z60% or more of total CLA intake (Fremann et al., 2002;
Ritzenthaler et al., 2001). Thus, it is not surprising that milk and
dairy food consumption positively correlates with CLA status in
human adipose tissue (Jiang et al., 1999), in blood plasma (Zlatanos
et al., 2008), as well as in human milk (Bertschi et al., 2005; Park
et al., 1999). However, eating patterns shifting from animal- to
plant-derived fat and from high- to low-fat dairy foods constitute
the dietary trends of our society, and often serve as a way to cut
energy intake and (tentatively) to improve dietary quality.
Milk consumption amongst American female teenagers
decreased by 36% from the late 1970s to the mid 1990s, and the
same percentage of 19 year old girls did not drink milk at all
between 1994 and 1996 (Bowman, 2002). The same picture applies
for Switzerland where in 2002 nearly two thirds of the population
were not consuming milk or dairy products on a daily basis, along
with 36% and 40% of men and women not drinking any milk at all,
respectively (Eichholzer, Bernasconi, Jordan, & Gutzwiller, 2005).
Another important factor influencing CLA intake in humans is
cultural animal meat preference. Nowadays, a much higher relative
amount of non-ruminant meat (pig and poultry) is consumed in
Europe than in traditional beef-eating countries like Australia or
Argentina. Shifting culinary habits likely contribute to a decreased
CLA intake, since meat-containing meals rich in SF and IT fat lost
popularity not only because they were considered unhealthy, but
mainly due to changes in the taste preferences of younger
443
generations and the higher consumption of fast or easily prepared
convenience foods. Similarly, entrails high in CLA/VA such as
marrow (Cordain et al., 2002), liver (Dannenberger et al., 2005;
Khanal & Olson, 2004; Mir et al., 2004), and kidney (Hollo et al.,
2005), are likely to disappear from our table.
A clear trend to consume less meat in general, and the substi-
tution of ruminant meat in favour of poultry was shown in the
Swiss annual meat market statistics from Proviande (2004) and in
the latest edition of the Swiss Nutrition Report (Eichholzer et al.,
2005). This substitution along with a general and small decrease
in daily consumption of animal fat can also be observed in FAOSTAT
(FAOSTAT, 2005) both within the European Union and Switzerland
(data from 1990e2005). Total meat consumption per capita in
Switzerland (as sold to consumers, not corrected for losses of water,
bones, fat, leftovers, etc.) decreased z14% between 1991 and 2004,
but in the same time period there was an increase of roughly 20% in
poultry meat consumption (Proviande, 2004). This shift is another
contributing factor to a loss of total CLA in our diet, as poultry is not
a valuable source of CLA.
Regarding CLA content in non-bovine meat and products as well
as CLA intake originating from these food groups, multiple
assumptions had to be made when using food disappearance data
to calculate estimates. Nevertheless in the authors' opinion this is of
only minor importance, because from other data it is estimated that
only about 8% of CLA derives from food sources other than dairy
(60%) or beef (32%) (Ritzenthaler et al., 2001). Moreover, when
using food supply data from FAOSTAT (FAOSTAT, 2005) a much
lower contribution of meat to CLA intake was found: in
Switzerland, only 4.7% of total available CLA originated from meat,
of which only about one third was beef-derived as shown in Fig. 3.
Similar data were found for the European Union (data not shown).
This discrepancy may be due to the assessment method (3-d
record: Ritzenthaler et al., 2001), overestimated beef-derived fat
content from food databases, or a combination of both.
For all these reasons, a lower CLA intake than is naturally
achievable may be a rather general phenomenon nowadays where
energy-restricted dieting is normal, and dietary fat balance is
possibly impaired by reduced animal/dairy fat and the concomitant
high intake of industrial plant oils from snack foods. From a nutri-
tional point of view, it may be interesting to determine if CLA may
be looked at as a marginal missing nutrient in present food sources.
This raises the question of whether an enrichment of foods with
CLA, in doses naturally found in food from animal-adapted feeding
strategies, could be used to correct the presently low content in
food caused by inadequate feeding of animals, contemporary
human lifestyle, and changes in our diets. However, before CLA
Cheese
33.9%
Bovine meat
1.7%
Pig meat
1.7%
Meat
Sheep & goat meat
4.7%
0.7%
Chicken & turkey
meat 0.6%
Game meat
0.1%
Milk, whole, fresh
61.4%
Fig. 3. Contribution of different food sources from grass-based production methods to
conjugate linoleic acids (CLA) supply in Switzerland. All food supply data were from
FAOSTAT (2005), except for cheese, which was from Gremaud et al. (2005) for 2001/02.
444 R.P. Jutzeler van Wijlen, P.C. Colombani / International Dairy Journal 20 (2010) 433e448
enrichment of foods could be envisaged, the question about which
isomer to use for this enrichment would have to be solved.
To properly estimate the potentially missing CLA quantities in
our diet, it seems important to elucidate the naturally accessible
CLA contents in our food. This necessitates improved production
methods with natural, species-adapted feeding and no supple-
mentation or enrichment of the animals' feed with plant oils, seeds
or directly with CLA. Simultaneous analysis of different CLA isomers
and VA would be of great importance, given that the evidence on
VA content of any source is far more scarce, and because of its
consequence for endogenous synthesis of CLA. Once natural CLA
including VA quantities in food are determined, a preferable dietary
CLA intake in humans in consideration of dietary recommendations
may be established. Subsequently, it may be a practical and
meaningful approach to substitute the CLA precursors lost in feed
sources such as concentrate compared with fresh species-rich
grass, thereby increasing CLA content in ruminant-derived foods
reflecting the values of wild animals. In the same context, feeding
CLA at a content as low as 1%E to non-ruminants such as swine or
poultry allows a several-fold increase at far higher rates than in
grazing ruminants (Dugan, Aalhus, & Kramer, 2004; Khanal &
Olson, 2004). Further studies examining CLA and especially VA in
swine and poultry fed a natural diet, oriented on the feeding
patterns of comparable wild animals, will be helpful in answering
the question of what those contents are expected to be.
Furthermore, a slightly elevated dietary fat intake of z35e40%E
may be recommended. Provided that the fatty acid profile is
adequate, such an approach takes into account newer evidence on
health issues related to fat (Appel et al., 2005; Kopp, 2006;
Mozaffarian, 2005; Westman et al., 2007), reflecting most prob-
ably a diet more closely adapted to our genes than presently pre-
dominating dietary recommendations. This suggestion is
supported when considering the most plausible percentages of fat-
derived dietary energy ranging from 38 to 49% in worldwide
hunteregatherers, where z50% is estimated to be animal fat, and
a dependence on animal foods of 50% of total energy (Cordain et al.,
2000).
Finally, increasing CLA intake in the human diet may not only be
desirable because of its potential health-promoting effects, but also
in respect to the health aspects linked to the consumption of its
main food carrier, the dairy products. High intake of dairy foods and
consequently high milk calcium is thought to be correlated with
lower prevalence of obesity, higher bone density, and (potentially
related to the higher CLA content in full-fat dairy foods) decreased
risk for colorectal cancer (Larsson et al., 2005). Therefore, simply
giving as much pasture access to ruminants as possible would be
a meaningful and simple approach to enhance not only the CLA and
VA content of meat and dairy products, but also to produce a more
desirable profile of MUFA, PUFA, SFA, and n 3 fatty acids (Alfaia
et al., 2007a, 2007b; Garcia et al., 2008b).
7. Conclusions
Feeding strategy is the main factor influencing the CLA content
both in milk and meat and several-fold increases likewise in CLA
and VA are achievable if partly or all-pasture based production
methods are employed. Consequently, nowadays modern grain-
based production methods cause a lower than presumably normal,
and naturally achievable, CLA food intake.
A more than doubled exogenous CLA intake is achievable from
grass-based production methods for dairy food and meat. Inclusion
of VA in estimates of CLA supply and intake is recommended
because of its high contribution to total available CLA via endoge-
nous bioconversion. Total available CLA calculation by means of
integrating VA-derived quantities discussed in this review (20%
synthesis rate) closely matched with the postulated multiplication
of exogenous CLA with factors of 1.4 or 1.5. Considering grass-based
production methods and VA bioconversion, the estimated total
available CLA supply found in this review was 707e1107 mg d 1
from food and fat disappearance data in Switzerland, and
774e923 mg d 1 from dietary recommendations and a selective
diet. Such quantities are substantially higher than estimated in
most previous intake studies. It is hypothesized that a more
selective, all-pasture derived food choice providing about 35e40%E
from fat, may even have the potential to supply more than
2e3 g d 1 of total available CLA. Future research will need to
elucidate if CLA should be considered to be a marginally missing,
semi-essential nutrient in our present diet (because of lower than
normally achievable intakes). However, CLA and VA may be only
a few of many substances whose intake is “artificially” reduced by
current dietary recommendations and animal feeding practices.
These data may add to a reconsideration of the actual dietary fat e
especially of animal origin e and meat intake recommendations.
Assuming a dietary zero energy balance, an adequate daily
consumption of full-fat dairy foods and meat from at least partly
grass-based production methods is encouraged. Apart from being
the major CLA providing foods, tasty and high-fat dairy products
have well-known health benefits and are available in abundance
and great variety. In contrast, meat products are of minor impor-
tance with respect to CLA supply in our present diet, except for the
choice of VA and/or CLA-rich sources such as lamb or game with
a substantial fat percentage. However, derived from grazing
animals and apart from its importance as a protein and vitamin
source, meat can provide further benefits with regard to n 3,
n 6/n 3 ratio, and MUFA.
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