Acta Agriculturae Scandinavica, Section A — Animal

Science

ISSN: 0906-4702 (Print) 1651-1972 (Online) Journal homepage: http://www.tandfonline.com/loi/saga20

The flavor of high-quality beef – a review

Margrethe Therkildsen, P. Spleth, E.-M. Lange & P. I. Hedelund

To cite this article: Margrethe Therkildsen, P. Spleth, E.-M. Lange & P. I. Hedelund (2018): The
flavor of high-quality beef – a review, Acta Agriculturae Scandinavica, Section A — Animal Science,
DOI: 10.1080/09064702.2018.1487466

To link to this article: https://doi.org/10.1080/09064702.2018.1487466

Published online: 29 Jun 2018.

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ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE
https://doi.org/10.1080/09064702.2018.1487466

The flavor of high-quality beef – a review

a
Margrethe Therkildsen , P. Splethb, E.-M. Langec and P. I. Hedelundc
a
Department of Food Science, Aarhus University, Tjele, Denmark; bSEGES, Landbrug & Fødevarer F.m.b.A., Aarhus N, Denmark; cTechnological
Institute, Aarhus C, Denmark

ABSTRACT ARTICLE HISTORY

Flavor of beef has been recognized as an equal quality trait to tenderness and lately high Received 1 May 2018
correlations between beef flavor and overall liking of beef have been demonstrated. In this Accepted 4 June 2018
review, we evaluate the effect of breed, sex, age, feeding, growth rate, slaughter method and
KEYWORDS
aging on beef flavor. A common determinator for these factors is their influence on Taste; intramuscular fat;
intramuscular fat (IMF) where more IMF increase the overall flavor liking. Feeding can, however, preferences; pasture-based
affect the flavor of beef beyond that of the amount of IMF. In several studies, pasture or pasture- feeding
based silage lead to a beef flavor characterized as fishy, gamey and grassy in contrast to
concentrate-fed beef cattle. A geographical difference in flavor preferences of high-quality beef
exists, most clearly illustrated among US consumers who prefer beef from concentrate-fed cattle,
whereas European consumers tend to prefer beef from cattle fed a mixed diet on pasture and
concentrate.

Introduction
categories varying in gender and genotype (Gagaoua
In several surveys, tenderness of beef has been rated as et al., 2016), and also Kerth and Miller (2015) demon-
the most important factor affecting palatability, with strated that consumer liking of beef from 20 different
flavor as the second most important trait. However, in cattle categories, cuts and cooking methods was more
the National beef quality audit (NBQA) in 2011 in US, related to consumer flavor liking relative to juiciness
the flavor was identified as more important than tender- and tenderness liking. Altogether, these numbers
ness for market sectors close to the consumer (Igo et al., demonstrate that knowledge of factors that influence
2013). Likewise in the original meat quality model of flavor perception and flavor liking is very important in
Meat Standard Australia (MSA), tenderness was weighted order to produce a beef quality required by the
higher (0.4) than flavor (0.2), but as tenderness improved, consumers.
the model was changed to equal weighting of tender- The beef industry does without doubt contribute to
ness (0.3) and flavor (0.3) (Watson et al., 2008). This illus- the green-house gas emission (GHG); however, there
trates that when tenderness is improved, flavor becomes are large variabilities between production systems
a decisive attribute in eating quality evaluation. Tender- (Nguyen et al., 2010). While beef contributes to a nutri-
ness is dependent on in vivo factors such as breed, sex, tional and healthy diet (Young et al., 2013), the associ-
age and feeding strategy. But whereas the final tender- ation with increased risk of developing certain cancers
ness to a large degree is the result of postmortem pro- (Sødring et al., 2017) and the environmental impact
cesses that can be optimized in order to generate the should not be neglected in the overall quality evaluation
most tender meat, flavor is to a larger extent a function of beef production. Thus, beef productions systems
of an inherent characteristic of the fresh meat that should be developed to deliver high quality, healthy
cannot be changed through postmortem handling, beef with minimal impact on GHG; this also includes
except through curing and cooking. The correlation improved eating experience so waste is minimized.
between overall liking and flavor liking in beef from 10 This review focuses on factors that may have an effect
different quality grades and breeds was found to be on the beef flavor as experienced by either trained
0.96 by Corbin et al. (2015). A positive correlation of sensory panels or consumers. The review (Table 1)
0.81 between beef flavor and overall liking was calcu- includes 1017 references where specific attention has
lated on the basis of data from 265 animals of 8 beef been paid to papers that included flavor profiling and

CONTACT Margrethe Therkildsen [email protected] Department of Food Science, Aarhus University, Blichers Allé 20, Tjele 8830,
Denmark
© 2018 Informa UK Limited, trading as Taylor & Francis Group
2 M. THERKILDSEN ET AL.

Table 1. Literature search on WEB of science, spring 2018. Miller, 2015). Maughan et al. (2012) defined 18 attributes,
Search termsa References Without duplicates including the five basic tastes, and here positive attri-
Beef + meat + flavor 1927 butes were associated with brothy, umami, roast beef,
+ breed 272 255
+ sex 86 57
juicy, browned, fatty and salty, whereas negative terms
+ feeding 524 372 were oxidized, bitter, barny, gamey, grassy, livery, met-
+ growth rate 644 232 allic and astringent. Table 2 summarizes the common
+ age 430 76
+ slaughter method 36 1 flavor attributes from these two lexicons and the
+ ageing 446 24 impact on flavor liking. Corbin et al. (2015) concluded
Total 1017
a that a lower score for flavor liking was due to lower
Search terms includes also related terms, for example, beef includes also
cattle, bovine, heifer, bull, steer, cow and veal. score for desirable flavors, rather than higher scores for
undesirable flavors in samples from grain-finished
animals as well as the existence of a positive relationship
also papers most relevant for Scandinavian beef pro-
between fat percentage and positive flavor character-
duction systems.
istics. This may, however, depend on the diet offered

The chemistry and characteristics of beef

flavor Table 2. Flavor attributes associated with beef and their impact
on flavor liking of beef based on the lexicons of Adhikari et al.
Raw beef only possesses characteristics such as salty, (2011) and Maughan et al. (2012).
metallic and/or bloody taste and sweet aroma (Wasser- Impact on
Flavour flavor
man, 1972), whereas a heat treatment is required to gen- characteristic likinga Commentb Reference
erate the characteristic beef flavors (Macleod & Ames, Sour − Basic taste Adhikari et al. (2011)
1986). The characteristic flavors of beef is generated on Maughan et al. (2012)
Salty + Basic taste Adhikari et al. (2011)
the basis of fatty acids (FA), free amino acids and pep- Maughan et al. (2012)
tides, reducing sugars, nucleotides and vitamins (Dash- Sweet + Basic taste Adhikari et al. (2011)
Maughan et al. (2012)
dorj et al., 2015) via complex chemical reactions such Bitter − Basic taste Adhikari et al. (2011)
as Maillard reactions, degradation of FA, oxidation of Maughan et al. (2012)
FA and interaction between the Maillard reaction com- Umami + Basic taste Adhikari et al. (2011)
Maughan et al. (2012)
pounds and FA (Aaslyng & Meinert, 2017). A positive Beefy + Beef flavor identity Adhikari et al. (2011)
relationship has been identified between water-soluble Brown/ + Flavors associated Adhikari et al. (2011)
Roasted with broiled beef Maughan et al. (2012)
carbohydrates and desirable flavor score (Melton et al., with fat
1982a). Likewise, through modeling, it has been demon- Fat-like + Flavors associated Adhikari et al. (2011)
with cooked Maughan et al. (2012)
strated that C18:3 fatty acid generate fishy odor; animal fat
however, in the presence of cysteine and ribose, as Metallic − Flavors associated Adhikari et al. (2011)
with metal Maughan et al. (2012)
examples of a Sulfur amino acid and a sugar, respect- Liver-like − Flavors found in Adhikari et al. (2011)
ively, this odor decreased and the meaty and corned cooked animal Maughan et al. (2012)
beef odor became more apparent (Campo et al., 2003). organs
Bloody/ + Flavors associated Adhikari et al. (2011)
These examples highlight the importance of the maillard serumy with blood Maughan et al. (2012)
reactions in the generation of the characteristic meat on meat
Barnyard − Flavors associated Adhikari et al. (2011)
flavor components. with feces Maughan et al. (2012)
Two recent attempts of making a beef flavor lexicon Rancid − Flavor associated Adhikari et al. (2011)
with oxidized fat
(Adhikari et al., 2011; Maughan et al., 2012) has been (may include fishy)
based on US qualities of beef, high-intramuscular fat Gamey − Flavor associated Maughan et al. (2012)
with wild
beef (Australian wagyu and US wagyu) and grass-fed game meat
beef from NZ (Maughan et al., 2012) combined with Grassy − Flavor associated Adhikari et al. (2011)
impact of different cuts, cooking methods and endpoint with green, Maughan et al. (2012)
plants, grass
temperature (Adhikari et al., 2011). Adhikari et al. (2011) Dairy +/− Flavor associated Adhikari et al. (2011)
defined 26 flavor characteristics, including references of with products made
from cows milk
these flavor attributes. Of these, beefy, brown/roasted, Warmed- − Flavor of reheated Adhikari et al. (2011)
bloody/serumy, fat-like, sweet, salty and umami were over meat Maughan et al. (2012)
a
identified as positive beef flavors, whereas metallic, Impact on flavor liking adapted from Maughan et al. (2012) and Kerth and
Miller (2015) +: positive impact on flavor liking, −: negative impact on
liver-like, sour, barnyard, musty-earthy/humus and flavor liking.
b
bitter was considered negative beef flavors (Kerth & Descriptions modified from Adhikari et al. (2011) and Maughan et al. (2012).
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 3

to the animals. Meat from grass-fed steers was scored to grassy, gamey, livery, fishy, sour and bitter. The positive
have undesirable flavors, which contributed to a nega- flavors were associated with more MUFA and the nega-
tive overall liking, even when tenderness and juiciness tive flavors were associated with more n-3 PUFA. In con-
were acceptable (Corbin et al., 2015). Rhee et al. (2004) trast, European consumers from Germany, France and UK
studied the palatability of 11 beef muscles from 31 Char- preferred beef from Uruguay raised on pasture plus con-
olais x MARC III steers, and found a strong positive corre- centrate relative to just concentrate (Realini et al., 2009),
lation between beef flavor intensity and lack of off-flavor which otherwise would be expected to give rice to most
in all cases. Across all muscles, a positive correlation IMF and less flavors associated with n-3 PUFA. This pin-
between beef flavor intensity and degree of proteolysis point the issue of past experience, as illustrated by UK
was also demonstrated, underpinning the importance consumers who are used to eating grass-finished beef,
of released peptides and amino acids for the specific with high concentrations of 18:3 n-3 FA producing
beef flavor. strong flavor and who would not be expected to identify
The inherent flavor of beef is dependent on feeding, beef enriched with n-3 FA from, for example, linseed
breed, sex, age at slaughter, slaughter method and (Vatansever et al., 2000).
aging process. Soulat et al. (2016) used rearing factors
to predict meat quality traits of beef based on data
from 688 young bulls and 307 cull cows of 6 pure Factors affecting beef flavor
breeds (Aubrac, Charolais, Blonde d’Aquitaine, Limou-
Breed
sine, Normande and Salers). The prediction of tenderness
and juiciness was low (R 2 < 0.50), whereas flavor intensity Cattle originates from a very large pool of breeds and
could be satisfactorily predicted (R 2 = 0.72) with a model, much beef are also the result of cross breeding. In
including duration of fattening period and concentrate Denmark alone, 42 cattle breeds are registered and
percentage. However, whereas increased concentrate several other breeds can be found in other countries.
percentage had a negative impact on flavor intensity of The use of different breeds and cross-breeds are often
both young bulls and culled cows, the length of the fat- evaluated based on the production and slaughter
tening period had a negative impact on young bulls but quality traits under different environmental conditions
a positive impact on culled cows. This deviation in effect and seldom based on specific meat quality traits. An esti-
of the fattening period, should, however, be interpreted mate of the heritability of flavor intensity and beef flavor
with caution as the period for young bulls spanned from in a population of 1066 cattle representing 12 different
105 to 328 days, whereas for culled cows it was 7 to 114 breeds demonstrated weak to moderate heritability’s of
days. Thus, this may indicate that shorter periods are these traits ranging from 0.06–0.22 to 0.00–0.18, respect-
positive, whereas longer periods are negative irrespec- ively, depending on aging time (Pratt et al., 2013).
tive of the category of cattle. The negative effect of con- Sevane et al. (2014) presented results from the large
centrate is somewhat surprising, as concentrate would European study ‘GeMQual’, which included 436
usually lead to increased degree of fatness, and thus a M. longissimus thoracis samples from 15 European
more intense flavor (Roberts et al., 2009). breeds. Flavor was evaluated by a sensory panel using
To further add to the complexity of beef flavor, the an 8-point scale – the higher the scores the more charac-
flavor preferences may also be dependent on the tra- teristic flavor of beef. The analysis showed significant
dition in a given country or even region. A Spanish positive correlations between beef flavor and fat score
study that included consumers from three north-east (0.12), fat percentage (0.31) and total lipids (0.3), but sig-
Spanish cities (Barcelona, Zaragoza and Pamplona) nificant negative correlations between beef flavor and
demonstrated that the perception of beef flavor is PUFA percentage (−0.25), percentage n-6 PUFA (−0.27),
dependent on the region, which probably reflects the PUFA to SFA ratios (−0.18) and n-6/n-3 ratio (−0.24)
diverging production strategies and cooking method in across all breeds. Thus, across all 15 breeds, the charac-
the specific regions (Perez-Juan et al., 2014). Most consu- teristic beef flavor is associated with increased fatness
mers favor food from their own local region and like and fat content, whereas an increased ratio of PUFA
what they are used to eat. US consumers prefer meat and specifically n-6 FA has a negative impact on the
with increased IMF leading to increased buttery/beef- beef flavor, whereas the percentage of n-3 FA had no
fat flavor and that is associated with increased overall influence on the beef flavor score. This impact of fat on
flavor desirability (O’Quinn et al., 2016). The consumers the beef flavor intensity meant that Angus and Highland
prefer beef with flavors characterized as beefy/brothy, breeds were associated with most flavor intensity in
browned/grilled, buttery/beef fat, nutty/roasted nut opposition to the other breeds. The lack of relationship
and sweet, and dislike flavors as bloody/metallic, between n-3 FA and flavor score is in contrast to other
4 M. THERKILDSEN ET AL.

studies (Warren et al., 2008), but might be explained with doubled-muscle genotype, a meat purpose breed from
the non-grass-based diet used in the ‘GeMQual’ study in Spain, a dual-purpose breed (Brown Swiss) and a rustic
contrast to studies which have presented an n-3 impact breed from Spain. There was no effect of breed on the
on flavor scores. The importance of the fat content for overall flavor intensity of beef aged from 1 to 21 days,
the flavor in beef is further demonstrated by the fact but the doubled-muscle bulls had more acidic flavor at
that 16% of the variation in flavor was explained by the some aging times compared with the other genotypes,
amount of IMF using data from an experiment with whereas the rustic breed had more liver flavor at some
large variation in IMF from 0.3% to 15% (Thompson, aging times compared with the other genotypes. Like-
2004), whereas only 3% of the variation was explained wise Nuernberg et al. (2005) compared the flavor of
using a dataset with minor variation in IMF (on average German Holstein and Simmental bulls slaughtered at
1.5%) (Hocquette et al., 2011). The latter study evaluated the same weight (620 kg) fed either a concentrated
correlations between IMF and flavor scores from the BIF- based or a grass-based diet. Among 13 flavor character-
BEEF database and showed that among young bulls istics, only the metallic flavor was more intense in the
there is a positive correlation between flavor and IMF Holstein compared with the Simmental bulls, with no
in the interval from 1% to 2.5% of IMF, whereas below effect on the overall liking. Furthermore, differences in
and above that, the relationship does not exist. Likewise, overall flavor intensity or in specific flavor characteristics
when data from heifers and steers, characterized with among breeds may depend on feeding, for example, a
more IMF, were included, no relationship could be ident- difference in beef flavor was found between Charolais
ified (Hocquette et al., 2011), illustrating that other and Limousine steers when they were fed barley,
factors such as sex, age and feeding also have an whereas there were no differences when they were fed
impact on flavor score. a corn diet (Mandell et al., 1997a). Likewise the bloody
There are several studies comparing different breeds flavor intensity was higher in Holstein compared with
under the same conditions like feeding and age, where Simmental bulls when they were fed a concentrate-
a difference in flavor liking is measured, and where the based diet, whereas if they were on a grass-based diet,
difference can be attributed to difference in IMF, for the Holstein had less bloody flavor compared with the
example, Hanwoo vs Angus cattle (Van Ba et al., 2013) , Simmental bulls (Nuernberg et al., 2005).
Hereford vs Simmental bulls (Mandell et al., 1997b), Also, age may be a factor that interacts with breed on
and crossbred steers Angus x Holstein vs Belgian Blue x the effect of specific flavor characteristics. In a study
Holstein (Keady et al., 2017). In the latter study, the comparing Angus and Holstein steers, a fishy flavor
higher flavor liking of Angus x Holstein meat coincidence was more intense in 14-months-old Holstein steers com-
with higher scores for greasy, sweet and dairy flavor and pared with Angus, but this difference disappeared in the
lower scores for bitter, acidic, cardboard and vegetable older steers (19 and 24 months), whereas among the 19-
flavor. If IMF was included in the model as a covariate, months-old steers, Holstein was characterized with more
the two genotypes only differed in the intensity of abnormal and less beef flavor, and in the 24-months-old
acidic and dairy flavor and this meant that the difference steers, Holstein was characterized with less acidic but
in flavor liking was only 0.29 units on an 8-point scale more rancid flavor (Warren et al., 2008).
(Keady et al., 2017). In addition, when meat from Here- With the low heritability of flavor in mind and the
ford and Simmental steers were compared, no flavor results from comparisons between breeds, it is clear
difference was found in agreement with lack of differ- that specifically the fatness and the composition of fat
ence in IMF (Mandell et al., 1997b). Overall, small flavor plays a major role in the differences in flavor between
differences might be identified in muscles rich in IMF, breeds and genotypes, and as such is affected by sex,
whereas this difference will not be identified in leaner feeding and age of the animals.
muscles from the same animals (Sinclair et al., 2001).
The impact of IMF on flavor of meat may explain differ-
Sex
ences in flavor intensity between some breeds, for
example, cull cows of Holstein and Salers breeds (Jurie As illustrated above, the flavor of the meat is highly
et al., 2007), Holstein and Holstein x Jersey bulls (Nian dependent on the amount of fat in the meat, and
et al., 2018) and not among others, for example, Holstein sex is one way to manipulate with the fat content,
and Welsh Black steers (Vatansever et al., 2000). for example, steers vs bulls. Thus, in a comparison of
Other studies have found differences in specific flavor the flavor of LD from heifers and bulls of low-fat car-
characteristics among breeds but not in the overall flavor casses of the Pirenaica breed (1.72% and 1.61% fat in
intensity; for example, Campo et al. (1999) compared the LD, respectively) and the Friesian breed (2.87% and
flavor of M. longissimus dorsi (LD) from bulls from a 1.09% fat in LD, respectively), Gorraiz et al. (2002)
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 5

found more intense livery and bloody flavor and less out of total fat and not on total amount of specific FA
beef characteristic flavor in bulls compared with in meat.
heifers. On the other hand, no flavor differences in The use of glycerin in bovine diets is increasing, as it is
either M. semimembranosus (SM) or LD was found a byproduct from the biodiesel production from veg-
between Holstein x Limousine bulls and heifers slaugh- etable and animal fat. The glycerin in bovine diets is
tered at the age of 16.9 months (Therkildsen & Vester- either fermented to propionate in the rumen by the
gaard, 2014). Steers are usually more fat than bulls microorganisms (Chung et al., 2007) or absorbed over
(Mandell et al., 1997b), and this led to a better score the rumen wall and directed to the liver to glucose syn-
for beef flavor for Simmental steers compared to thesis (Remond et al., 1993). Prado et al. (2016) tested the
bulls, whereas among Hereford steers and bulls this effect of substitution of 16% dry matter of corn with gly-
difference was not seen. The comparison between cerin on the meat quality. There were no effects on mar-
meat from heifer and steers was done by Jeremiah bling and color and minor differences in chemical
et al. (1997), and even though some differences were composition of the M. longissimus. A consumer panel
seen in flavor attributes with more samples with evaluated the eating quality of the M. longissimus, and
bloody flavor and less with off-flavor and browned scored the meat from the glycerin diets to have more
flavor among the heifers compared with the steers, meat flavor. The explanation for this could be the
this was not effects that were recognized among increased content of two FA myristic (C14:0) and marga-
consumers. ric (C17:0) acid in meat from the glycerin diets, although
As conclusion, the effect of sex on flavor is very much Melton et al. (1982a) did not find any significant corre-
affected by the degree of fatness of the animals, often lation between these FA and flavor score in ground
heifers and steers score better compared with bulls beef. Also, van Cleef et al. (2017) found increased flavor
because of more IMF. intensity with the increased exchange of corn and
soybeen hulls with sunflower meal and glycerin to
Nellore bulls, and in this case it was associated with an
Feeding
increased juiciness and greasy intensity, but no differ-
Feeding of beef producing cattle varies around the ence in C14 and C17.
world, including pure concentrate diets high in energy Likewise, with the aim to change the fatty acid com-
and a minimum amount of roughage, introduction of position in beef to a more unsaturated and healthy com-
by-products from different food or energy production position, inclusion of specific feed components in the
sites, roughage based on corn, grass or legumes, diet of beef cattle has been tested. For example,
pasture on arable land or natural grass land low in inclusion of 10% linseed in a concentrate diet to young
energy. Despite this diversity in feeding, it is only Holstein bulls from 200 to 325 days of age changed
reflected in the flavor of meat when the more extreme the content of n-3 FA from 12 mg/100 g muscle to
diets are compared. In a high-energy diet, the grain 30 mg/100 g muscle, but did not have an effect on the
source (corn vs barley) did not affect any attributes flavor perception among consumers (Perez-Juan et al.,
(cooked beef, bloody, grainy, cardboardy, painty, fishy, 2014). The diet also included Vitamin E, which will
livery, soured, corn, barley or metallic flavor) of beef protect the unsaturated FA from oxidation. Similar
steaks (Miller et al., 1996) or only increased livery flavor results were seen by Alberti et al. (2017) and Vatansever
(among 13 flavor characteristics) in barley diets com- et al. (2000). In a study which included a comparison of a
pared to corn (O’Quinn et al., 2016). concentrate diet (barley, sugar beet pulp and soya) and a
On the other hand, there are explanations for flavor grass silage diet, the content of vitamin E as a protector
differences caused by difference in diets, for example, of fatty acid oxidation may also have played a major role
more reducing sugars are found in meat from concen- in the perception of the flavor (Warren et al., 2008).
trate-fed beef cattle in comparison with beef cattle fed Silage-fed steers had more vitamin E in the meat and
grass silage which had higher levels of free amino were scored with more livery flavor at the age of 14
acids (Koutsidis et al., 2008). Likewise, the fatty acid com- months and 24 months, less abnormal and more beef
position in beef can differ depending on diet and treat- flavor at the age of 19 months, and at all age groups
ments, and a significant correlation with beef flavor the silage-fed steers were scored with better or no differ-
attributes have been shown by O’Quinn et al. (2016), ence in overall liking. Some effects of silage feeding were
whereas an attempt to correlate off-flavor with fatty found by Nuernberg et al. (2005). An increased intensity
acid composition or ratios in LD did not show any of fishy flavor was the result when bulls were fed a grass-
pattern (Stelzleni & Johnson, 2010); however, the based diet with pasture in summer followed by wilted
attempt was done on the percentage distribution of FA silage, barley and cracked linseed in the winter period
6 M. THERKILDSEN ET AL.

3 months before slaughter compared with a concentrate 70% of corn, soybean meal and corn silage) compared
diet (semi ad libitum access to maize silage and barley with a pure pasture/hay diet (Bjorklund et al., 2014).
rich concentrate). This difference was the only difference These results were further supported by a study where
among 13 flavor characteristics (beef, abnormal, greasy, steers fed grass for 4 months were finished with a
bloody, livery, metallic, bitter, sweet, rancid, acidic, whole shelled corn diet for 0, 28, 56, 84, 112 or 140
grassy and dairy); however, the fishy flavor probably con- days (Melton et al., 1982b). Here, an immediate effect
tributed together with more toughness to the negative of the change to the corn diet on the intensity of
impact on the overall liking of meat from the silage-fed milky-oily, beef-fat, sour, liver and fishy flavor was illus-
bulls. The origin of the silage may affect the flavor as trated; thus the milky-oily, sour and fishy flavor
Lee et al. (2009) compared the flavor of meat from Hol- decreased already after 28 days of corn diet, whereas
stein-Friesian dairy cull cows finished for 12 weeks with beef-fat and liver flavor increased after 28 days of corn
ad libitum access to grass silage or red-clover silage. diet. Most changes occurred the first 56 days, and after
The two diets had an effect on the fatty acid composition 84 days there were little changes in the flavor intensity
in LD, with more PUFA and especially more n-3 FA in scores. There were no changes over time on the metallic
meat from red-clover fed cows, and this might explain and off-flavors whereas days on corn increased the con-
the difference in flavor characteristic with more fishy centration of glucose in meat and in desirable beef
flavor intensity and less acidic flavor in the red-clover flavor. A positive correlation was found between
beef. Fishy flavor is often associated with increased specific FA (C15, C18 and C18:3) and milky-oily flavor
levels of long chain PUFA, which is susceptible for oxi- and fishy flavor and negative correlation with cooked
dation, and in the study by Lee et al. (2009) a lover con- beef-fat flavor.
centration of vitamin E was also measured. In some feed-lot systems, a period with pasture is
included before finishing with diets rich in grain. This
Fresh pasture will have an overall effect on the age of the animal at
In early comparisons of grass-based diets with grain- slaughter, and the overall growth rate. However, in the
based diets, it was demonstrated that the amount of study by Durunna et al. (2014a, 2014b), the eating
C18:3 correlated negatively with desirable beef flavor, quality of steers born either early spring or summer
when evaluated by a trained flavor profile panel from and exposed to a period with pasture (approximately
North Carolina State University and by an un-trained 100 days) or not, and all finished on high concentrates
panel from Tennessee (Melton et al., 1982a). In the diets in feed-lots to a target of 8 mm back fat for 105–
study by Melton et al. (1982a), grass-finished steers had 195 days, there were no effect on the beef flavor or
less carbohydrates and more C18:3 FA in the meat com- off-flavor of the LD. Likewise the other way around, a pre-
pared with grain-finished steers. Meat patties prepared vious difference in feeding, for example, forage with or
from ground beef from SM from grass-finished steers without concentrate supplement, followed by finished
were characterized with more intense dairy flavor and on a similar diet (pasture and barley for 163 days) did
off-flavors compared to the grain-finished steers not lead to any differences in beef flavor intensity, liver
(Melton et al., 1982a). Another study from Utah (Tansa- flavor intensity or abnormal flavor intensity (Panea
wat et al., 2013) found similar results, that pasture et al., 2012). In other words, it is the feeding prior to
feeding was associated with the flavor descriptors of slaughter that has an effect on the flavor of beef. An
barny, gamey and grassy compared with grain feeding example of this is if culled beef cows were scored as
which was associated with umami and juicy flavors. either being fed or non-fed prior to slaughter, then this
When three different US groups (trained sensory panel, was reflected in the intensity of off-flavor. More off-
consumers and informed consumers) evaluated the flavor was identified among the non-fed beef cows,
liking of flavor of beef from pasture-fed versus concen- which was expected to be slaughtered from pasture;
trated fed steers, the trained panel and the uninformed however, this was not the case in the comparison
consumers preferred the flavor of beef from the concen- between fed and non-fed culled dairy cows, which was
trate-fed steers, whereas among the informed consu- expected to be slaughtered from a concentrate diet no
mers there were no difference in liking (Medeiros et al., matter what condition at time of slaughter (Stelzleni &
1987). This pattern was also seen among another Johnson, 2010). This suggests a dilution effect of concen-
group of US consumers, who gave higher scores for trate feed on the possible off-flavor components related
flavor liking and less off-flavor in beef raised either con- to pasture as supported by the results of Larick et al.
ventional (67% increasing to 80% concentrate and the (1987) and Mezgebo et al. (2017), where Larick et al.
rest being roughage of corn silage and grass hay) or in (1987) saw decreased scores for grassy flavor following
an organic system (pasture supplemented with up to 56, 84 and 112 days in feedlot where the animals were
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 7

offered a diet of corn-silage and whole shelled corn. the SM (Hansen et al., 2006). Part of this difference
Likewise, when mature beef cows raised on pasture could most probably be ascribed to the amount of IMF,
were offered a high-energy diet for 0, 28, 56 or 84 with less IMF in the meat from the compensatory
days, this increased the flavor intensity dependent of growth strategy. Keady et al. (2017) studied the effect
time on the high-energy diet and decreased the off- of a compensatory growth strategy on, among others,
flavor intensity after only 28 days on the high-energy the flavor of the M. longissimus from steers, with no sig-
diet (Boleman et al., 1996). In support of the importance nificant effect on 12 flavor characteristics evaluated, but
of fat in relation to flavor intensity, there was also an also no difference in daily gain the last 100 days prior to
increase in fat thickness on the carcass and in the mar- slaughter and no difference in percent fat in the
bling score with time on the high-energy diet (Boleman M. longissimus.
et al., 1996). Thus unless the growth rate or path leads to a differ-
In contrast to these results, French et al. (2000, 2001) ence in IMF, it does not seem to have an impact on the
demonstrated that when steers are fed to the same flavor of the meat, whereas with more IMF a more
growth rate, irrespective of the diet (grass, grass + con- intense beef flavor can be expected.
centrate, grass silage + concentrate or concentrate)
there were no difference in the flavor liking among a
Age
trained Irish sensory panel. Likewise, even though
flavor became more intense and the desirable sensory Flavor scores increase with age of the animal (Dran-
attributes became more consistent with more IMF, sfield et al., 2003), but this can be linked to fatness
when adjusted to same IMF only very few differences and maturity (Zembayashi, 1994). As such, no difference
between grass- and grain-fed Angus steers were found was found in meat flavor among culled dairy cows from
in flavor evaluated by an Australian sensory panel first parity compared with older cows (Vestergaard
(Frank et al., 2016). However, low-fat, grass-fed Angus et al., 2007), nor in taste or off-flavor among bulls of
were characterized by liver flavor, barnyard odor, 258 days and 326 days old (Vestergaard et al., 2000),
acidity, astringency and metallic after taste (Frank et al., or in beef flavor among steers of 14, 19 and 24
2016). months (Warren et al., 2008) or among bulls of 16.2
In conclusion, the flavor is dependent on the diet prior or 19.2 months old (Mezgebo et al., 2017). Interestingly
to slaughter, and characteristic flavors from pasture though, Nian et al. (2017) found that meat from older
feeding like fishy, gamey and grassy are recognized. bulls (22 months) vs young bulls (15 months) resulted
However, these characteristics will be diluted within a in more residual roasted beef flavor after swallowing
month of feeding with a more concentrate-rich diet. the sample. This is a trait which is not always evaluated
by trained panels.
Growth rate
Transport and slaughter procedure
Growth rate and growth path in itself does not seem to
have an effect on the flavor of beef (French et al., 2001; Only few studies address the effect of transportation
Sinclair et al., 2001; Moloney et al., 2008), unless it conditions on the flavor of meat (Schwartzkopf-Gen-
courses differences in the amount of fat. This was the swein et al., 2012). No effect on flavor has been found
case in a study with dairy cows that were dried off and in studies with transportation of bulls (Villarroel et al.,
finished for 9 weeks compared with dairy cows that 2003) or steers (Alende et al., 2014) for either 30 min,
were slaughtered while lactating (Therkildsen et al., 3 or 6 h or 3 h vs 8 h, respectively, and likewise no
2011). The meat from the dried off cows had more effect on flavor of lairage time at the slaughterhouse
beef flavor and fresh flavor and less off-flavor and liver (2 vs 14 h) (Alende et al., 2014). In those studies, the
flavor, with the changes being more intense in LD rela- transport and lairage time did not affect the final pH
tive to the changes seen in SM. This could be linked to of the meat. Any effect of transportation and slaughter
the changes in IMF which doubled in LD and did not procedure on the flavor of meat is expected to be
change significantly in SM. Likewise, Vestergaard et al. related to the level of stress and as such an effect on
(2007) found an increase in meat flavor in finishing-fed the pH decrease post mortem and the final pH. A
culled dry dairy cows, compared with dairy cows dried clear effect of increased pH from 5.66 to 6.89 result in
off for one week. Also a compensatory growth strategy increased rancid, musty/earth/hummus, fat-like, metallic
for bulls resulted in less beef meat flavor, and more and overall sweetness flavor and decreased brown/
steer taint flavor in M. supraspinatus compared with ad roasted, sour and salty flavor (Yancey et al., 2005;
libitum feeding, less effect in the LD and no effect in Grayson et al., 2016), with all characteristics deemed
8 M. THERKILDSEN ET AL.

important for overall liking according to a beef flavor no changes in flavor from 2, 14, 21, 42 to 63 days of
lexicon (Adhikari et al., 2011). If the meat has a higher aging in either GM or M. longissimus lumborum. In con-
content of carbohydrate, the potential to form the Mail- trast, studies related to the MSA (Hughes et al., 2015)
lard reaction is assumed to increase and thus the gener- showed that meat flavor was improved from 2 to 12
ation of the favorable meat flavor. Meat with pH below weeks of aging as evaluated by the consumer panel.
5.55 obtained a higher score for flavor quality than meat The results from Yancey et al. (2005) may explain
with pH higher than 5.55 (Villarroel et al., 2003), which these different results as they found different response
could be explained by depletion of the glycogen among muscles and depending on the pH of the
stores prior to slaughter by stressful handling, which muscle. At high pH (> 6.0), IS and M. psoas major
might lead to higher pH meat and less flavor quality (PM) decrease in beef flavor intensity with aging (7,
due to the potential of carbohydrates to form Malliard 14, 21 and 35 days), whereas at normal pH (<5.7),
reaction compounds. these muscles and normal and high pH GM did not
change, except for normal PM after 35 days of aging
which became more beef flavor intensive.
Aging
Thus, there seem to be a general pattern of increased
Flavor compounds are related to late postmortem times beef flavor intensity with increased aging, although this
(Campo et al., 1999). Thus, an aging period is required to may depend on the pH and muscle, and there might
develop the flavor characteristics of meat (Gorraiz et al., be critical points for optimal flavor, which occur earlier
2002). This is in accordance with a study from Belgium in the aging period relative to, for example, optimal
(Raes et al., 2003), where meat from Double muscled aging for tenderness.
cattle and limousine cattle were compared with beef
from Argentine and Ireland. The beef from Argentine
and Ireland were scored to have more flavor intensity, Conclusion
supported by increased concentrations of very potent
What is the flavor of high-quality beef? This review has
aroma compounds such as pyrazines, saturated and
demonstrated that it depends very much on the
unsaturated aldehydes. The beef from Belgium were
degree of fatness, which again are determined based
aged for 14 days compared to 29 and 22 days for the
on the breed, the sex and the feeding, and in the end
Argentine and Irish beef; however, other than a differ-
on the consumer preferences, which reflects previous
ence in aging time, there might also have been differ-
experience. Thus, the literature demonstrates that a com-
ences in sex and feeding. In another study, which
parison between grain-fed animals and grass-fed animals
included beef from four breeds and 6 aging times, 3
will most probably elucidate differences in flavor charac-
flavor intensities (beef, liver and bitter) were evaluated
teristics, and dependent on the previous experience the
by a trained panel, but only bitter flavor seemed to
consumer will favor the flavor of one or the other. There
increase with aging time from 1 to 35 days, with the
is a tendency that studies performed in the US demon-
difference being recognized between 21 and 35 days
strate preferences for meat based on concentrate
of aging (Monsón et al., 2005). A consumer test of the
feeding, whereas European consumers are more accus-
same beef suggested that a critical point between 14
tomed to meat from beef cattle fed a combination of
and 21 days of aging exist for flavor acceptability, fol-
pasture and concentrate.
lowed by a decreased acceptability probably due to
generation of off-flavors. Consumer ratings of flavor
liking was also increased from 7 to 21 days of aging
Acknowledgements
of meat from young Holstein bulls (Perez-Juan et al.,
2014), whereas no difference in flavor was recognized Associate professor Jette F. Young, Aarhus University is
acknowledged for comments to the manuscript.
by a trained panel evaluating meat from Holstein
young bulls aged for either 7 or 14 days (Hansen
et al., 2006). Likewise, Adcock et al. (2015) evaluated
Disclosure statement
the eating quality of meat aged 14, 28 and 42 days
from Angus beef cattle and found that there were no No potential conflict of interest was reported by the authors.
change in flavor in M. infraspinatus (IS) and LD with
aging time, whereas for M. longissimus thoracis and
M. gluteus medius (GM) the flavor preference score Funding
was higher for the meat aged only 14 days vs 28 and This work was supported by Interreg - Öresund-Kattegat-Skagerak
42 days. Colle et al. (2015), on the other hand, found European Regional Development Fund [20200994].
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 9

ORCID Dransfield, E., Martin, J.F., Bauchart, D., Abouelkaram, S., Lepetit,
J., Culioli, J., Jurie, C. & Picard, B. (2003). Meat quality and
Margrethe Therkildsen http://orcid.org/0000-0002-2427-0131 composition of three muscles from French cull cows and
young bulls. Animal Science 76, 387–399.
Durunna, O.N., Block, H.C., Iwaasa, A.D., Thompson, L.C., Scott,
S.L., Robins, C., Khakbazan, M. & Lardner, H.A. (2014a).
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