Sheep: Meat

EN Ponnampalam, Agriculture Research, DEDJTR, Attwood, VIC, Australia

BWB Holman, Centre for Red Meat and Sheep Development, Cowra, NSW, Australia
ND Scollan, Aberystwyth University, Aberystwyth, UK
2016 Elsevier Ltd. All rights reserved.

Introduction
Since antiquity, sheep (Ovis aries) have been domesticated for
the production of meat, albeit often as a secondary product
from wool production. This has resulted in a range of sheep
breeds that vary in terms of production and climatic adaptation
specializations. Therefore, there are inherent differences
between sheep breeds regarding meat productivity and quality.
For example, a purebred Australian Merino is a specialized wool
breed that will produce lighter carcass weight with lean meat
over a longer growth period when compared with a meat specialist breed such as Suffolk or Texel. Using this information,
producers have tailored their production systems to optimize
breed effects, in terms of feed usage and production localities,
to achieve optimum meat yields and quality. Furthermore,
tactical management of feeding systems prior to slaughter that
offers optimum dietary nutrients as protein, energy, and micronutrients can provide more shifts in yield and quality and allow
producers greater responsiveness to changes in market requirements or specification.
Sheep meat production is highly versatile and immediately
employs many livelihoods around the globe, in both small
holdings and large-scale extensive or intensive production systems. This is facilitated by its inclusion within various culinary
diets, based upon sheep meat having a highly nutritious composition and distinctive flavor although ideal flavor is debatable, depending on cultural backgrounds. Consequently, sheep
meat is a key tradable commodity and the focus of much effort to
enhance its nutritional and eating qualities and ensure these are
preserved from production to consumption through processing
and retail. To assist in the clarification of sheep meat, animals are
generally divided into age-dependent categories: lamb, being
meat from animal slaughter prior to 12 months of age; hogget
or yearling, defining the age bracket between lamb and mutton;
and mutton, being meat from animals once the age of 24
months old is surpassed. These definitions highlight the importance of animal age to carcass value, meat composition, and
eating quality characteristics. Herein, the word sheep meat covers
aspects of meat from both lambs (younger age animals lamb)
and sheep (older age animal mutton).
This article aims to detail sheep meat, with reference to
production and processing aspects, in terms of production
types, nutritional composition and health implications, sensory and eating quality traits, and spoilage pathways that
require preservative attention.

Production
In 2010, sheep meat production represented only 3% of the
total global meat production, and this market share emphasizes

750

the huge potential for growth available. This scope for advancement in sheep meat production is encouraging to traditional
sheep meat-producing nations, such as New Zealand and
Australia, which have a strong export focus and access to developing markets in the European Union, the Middle East, North
America, and Asia. However, the global sheep meat industry
faces limitations, which may undermine future growth,
including animal and human welfare considerations, increased
competition for land and feed resources, climatic impacts from
intensified production, and economic restrictions on international trade. Overcoming these hurdles is the responsibility of
all sheep meat stakeholders. Regardless, domestic consumption
of sheep meats is increasing over recent years, and this has
resulted in better prices being offered for sheep meat productions, especially lamb, in comparison with other meats.
Stakeholders across the supply chain from producer,
processor, to retail are vital to the industry, but producers are
particularly important. The production of sheep meat, broadly
speaking, can be categorized into two groups: (1) small holdings, which are more common in developing countries or
hobby farms and generally maintain a small flock (often <20
animals), or (2) large holdings, which can have flock numbers
into the thousands and routinely manage animals on an entire
flock basis. Animals are reared under two main feeding systems:
(1) extensive grazing on traditional pasture feeding and (2)
intensive production systems maintained on concentrate
grain-based diets. However, feeding systems can be combined
depending on availability of resources, quality of feeds, and
the purpose of production. Sheep maintained under extensive
grazing receive diets with high protein (e.g., green pasture)
and grow slower, while animals maintained under intensive
systems receive high energy and grow faster. These in turn can
impact on the muscle composition and eating quality traits
of meat.
Historically, operations with a wool-only production focus
would consider sheep meat as a by-product, but this is no
longer universally true. The advent of synthetic fibers and narrow potential for expansion in traditional or new market share
have both contributed to low wool prices. As a result, many
traditional wool producers have adopted dual-purpose production focus. This entails flocks dominated by dual-purpose sheep
breeds, such as Corriedale and Finnsheep, that equally specialize in both wool and meat production. Another approach to
dual-purpose production uses a base wool specialist flock
joined with a meat specialist breed to achieve progeny with
hybrid vigor and both wool and meat quality characteristics.
Under traditional Australian commercial practices, lambs
under grazing conditions are slaughtered between 4 and 10
months of age, where they reach a live weight to produce
carcasses of 18 kg or more. Sheep production in Australia
uses different genotypes under various production systems.

Encyclopedia of Food and Health

http://dx.doi.org/10.1016/B978-0-12-384947-2.00620-6

Sheep: Meat

Table 1
Comparative percentage muscle (lean (%)) and fat (fat (%))
contents of different sheep genotypes that were slaughtered when
having a hot carcass weight of 20, 30, or 40 kg
Weight (kg)

Genotype

Age (months)

Lean (%)

Fat (%)

20

M
BL
M
PD
M
PD
M
PD
(BL
M)
M
M
BL
M
PD
M
PD
(BL
M)
M
BL
M
BL
M
PD
M
PD
M
PD
(BL
M)
PD
(BL
M)

8
8
4
8
4
14
22
14
14
8
22
14
22
14
22
14
22

76.4
74.9
77.5
77.1
76.8
71.5
67.2
68.3
71.3
70.8
65.0
64.0
60.0
67.8
63.9
65.9
61.9

20.4
22.2
19.5
20.1
20.6
24.3
28.5
28.1
25.2
26.0
30.2
32.1
36.0
28.4
32.3
30.7
34.6

30

40

The genotypes investigated were Australian Merino (M), Border Leicester (BL), and Poll
Dorset (PD).
Adapted from: Ponnampalam, E. N., Butller, K. L., Hopkins, D. L., Kerr, M. G., Dunshea,
F. R. and Warner, R. D. (2008). Genotype and age effects on sheep production. 5. Lean
meat and fat content in the carcasses of Australian sheep genotypes at 20-, 30- and
40-kg carcass weights. Australian Journal of Experimental Agriculture 48, 893897.

The major types of animals produced for sheep meat are

about 40% from second cross (Poll Dorset
(Border
Leicester
Merino)), 25% from first cross (Poll Dorset
Merino and/or Border Leicester
Merino), and 20%
from pure Merino genotypes. This latter approach is popular
among traditional wool producers as it allows their original
wool flock genetics to be preserved, it avoids a full production
system overhaul, and it allows better flexibility to match market
flux (Table 1).
Sheep meat-only production focus routinely uses meat specialist breeds. This may include wooled breeds such as Dorset or
Cheviot that produce coarse wool that can be harvested as a byproduct of production. Otherwise, haired breeds that shed their
coats may be used, for example, Dorpers and Barbados these
are gaining popularity as labor prices increase and wool prices
remain low. As a sidenote, dairy sheep (e.g., East Friesian) are
another source of sheep meat as they are typically slaughtered at
the end of their primary usefulness and may be considered a byproduct of the production focus just as described with a woolonly production focus. Selection of a production focus ultimately is regulated by market demands and preferences.

Composition
Sheep meat has a complex structure and consists of water
(73%) and protein (21%) fractions, with lipid (5%),
carbohydrate, and many vitamins and minerals also present
but in smaller amounts. The major water, protein, and fat of
meat (or muscle) composition can be affected by sheep genetics, feeding background, and age of sheep. The fat content of
meat ranges from 2% to 10% and can substantially influence

751

the meat quality aspects such as flavor/aroma, juiciness, and

texture. It was observed that when sheep are reared under the
same paddocks, within a given age and genotype category,
sheep with smaller carcasses deliver less fat and more lean
than sheep with larger carcass counterparts. This implies that
within a genotype at a particular age, there is variation in fat
and muscle deposition, with faster growing animals diverting
more energy into fat deposition than muscle growth compared
with smaller growing animals.
The protein-rich muscle fibers (myofibrils) of sheep meat
mostly contribute to its distinctive texture (tenderness of meat).
Myofibrils are arranged into bundles or sheets and are individually surrounded by an endomysium and sarcolemma
(a plasma membrane); in turn, these are encapsulated in groups
by a perimysium and as a whole muscle by an epimysium. A
collective term for endomysium, perimysium, and epimysium
is connective tissues. There are three myofibril types, and the
prevalence of these has been found to vary depending on muscle function and location; these are (1) type 1/slow twitch,
using aerobic metabolism and found in postural muscles; (2)
type 2A/fast twitch, which are similar to type 1 but found
mostly in limb muscles; and (3) type 2B/fast twitch, using
anaerobic metabolism and found in locomotive muscles. The
density of myofibril types can affect the sensory characteristics
and nutritional quality of meat. This protein fraction is highly
digestible, more so than plant protein, and provides consumers
with the essential amino acids, lysine, threonine, methionine,
phenylalanine, tryptophan, leucine, isoleucine, and valine.
As a red meat, sheep meat is a rich source of dietary iron
(Fe) within a heme (Fe) complex containing comparatively
higher levels of Fe than beef. Heme (Fe) is readily absorbed
when compared with nonheme (Fe), partly due to sheep meat
being rich in glutathione in which its activity promotes iron
absorption. Humans require iron to avoid anemia and the
inhibition of several cellular processes including oxygen
transportation throughout the cardiovascular system. The consumption of heme (Fe) has, however, been associated with an
increased incidence of colorectal cancer and congenital heart
diseases. But evidence suggests that provided sheep meat such
as trim lamb cuts or lean meat is consumed as part of a wellbalanced diet, the risk from these diseases can be ameliorated.
Sheep meat is also a valuable source of selenium and can
provide approximately 20% of the daily recommended intake
for an adult per 100 g serving. It also provides zinc that can
be more readily absorbed than plant-sourced zinc because of
the absence of many phytates and fiber that hinder its uptake.
Sheep meat is also an important source of calcium, magnesium, copper, phosphorus, and potassium, all being essential
to human health. Interestingly, the presence of these minerals
does not only solely contribute to nutritional health benefits
but also play a role in many sensory characteristics and keeping
quality of meat.
Sheep meat, as with other red meats, has an unmerited
notoriety for its high content of unhealthy saturated fatty
acids (SFA); these are unfavorable because of their association
with cardiovascular disease and antagonism of several cancers.
However, recent efforts are making progress to reverse this negative reputation by developing sheep meats with an improved
healthy fat content through genetics and nutritional improvement. It is also worthy to state that lean meat cuts from sheep are

752

Sheep: Meat

low in fat and can contribute to the intake of key nutrients in a

diet. Omega-3 polyunsaturated fatty acids (n3) are one such
healthy fat, as these have been linked with improved resilience
against coronary disease and diabetes, better mental status and
infant development, and the maintenance of neural and optical
tissues. Many types of n3 exist, but those associated with these
benefits are long-chain (>C13) n3, which include eicosapentaenoic acid (EPA: 20:5n3) and docosahexaenoic acid (DHA:
22:6n3). In general, people fail to consume the recommended
EPA DHA levels by the World Health Organization (0.52% of
total caloric intake), the United States (0.252%), or the United
Kingdom and Australia (12%). Lamb has been identified as a
good source of these n3s where a good source refers to n3
content above 60 mg/100g of food according to Food Standards
Australia New Zealand. Furthermore, n3 content can be
improved by supplementing lambs with feeds rich in n3 (e.g.,
some algae and flax) as part of a flock dietary management
strategy. It should be noted that meat cut selection will influence
the available n3 as EPA and DHA are primarily located in
membrane phospholipids and therefore are found in greater
quantity in muscles rich in slow-twitching fibers (type 1 myofibrils). Another healthy fat found in sheep meat is conjugated
linoleic acid (CLA), which has been found to reduce the risk of
some cancers, diabetes, and arthritis. The amount of CLA in
sheep meat is highly correlated with fat content and is influenced
by animal diet, but generally, 0.22% of total tissue fat is CLA.
Animal genetics and rumen flora also play major roles in determining the fat content and composition of sheep meat (Table 2).

Table 2
This information was sourced from Food Standards
Australia New Zealand (2010) and describes the nutrient content of raw
meat (per 100 g fresh weight) sampled from 11 different cuts that were
from various regions of the sheep carcass (lamb and mutton)

Nutrient
Energy (kJ)
Moisture (g)
Protein (g)
Total fat (g)
Saturated fat (g)
Polyunsaturated fat (g)
Monounsaturated fat (g)
Cholesterol (mg)
Total long-chain omega-3 (mg)
Minerals
Sodium (mg)
Iron (mg)
Selenium (mg)
Zinc (mg)
Phosphorus (mg)
Calcium (mg)
Vitamins
Retinol (vitamin A) (mg)
Thiamine (B1) (mg)
Riboflavin (B2) (mg)
Niacin (B3) (mg)
Pantothenic acid (B5) (mg)
Pyridoxine (B6) (mg)
Cobalamin (B12) (mg)
Vitamin E (mg)

Lamb

Mutton

2315
31
10.8
57.6
25.7
2.7
20.8
62
140.7

2522
28.8
8.2
64.4
28.6
3.7
24.1
72
163.0

22
0.41

22
56
5

33
0.77
10
0.99
88
10

34

0.07
2
0.43
0.1
2.9
1.1

64

0.07
5
0.46
0.1
2.9
0.7

Sheep meat is a valuable source of several key vitamins. A

100 g serving can provide nearly two-thirds of an adults daily
recommended intake of vitamin B12 and one-quarter of their
daily vitamin B2, B3, B5, and B6 requirements. Sheep meat is
also a useful source of many lipid-soluble vitamins, including
vitamin A, but efforts to produce leaner sheep meat have had a
negative influence on its content. Leaner sheep meat also has
implications on some sensory characteristics and is described
in the following section.

Quality
The quality of sheep meat is fundamentally determined by its
appeal to consumers and their tangible experience following
mastication. To better understand the appeal of sheep meat,
consumer responses are routinely defined as major sensory
characteristics: (1) tenderness, also known as mouthfeel or
texture; (2) flavor, which is the intrinsic combination of both
smell and taste senses; (3) appearance or color when sheep
meat is retailed or on display; and (4) juiciness or moistness.
These four sensory characteristics provide insight into the overall consumer acceptability of sheep meat.
Tenderness refers to the ease by which sheep meat can be
severed and chewed during mastication. This is widely considered to have the most authority over perceived quality when
compared with other sensory characteristics and hence acts as a
key discriminating factor for consumers selecting sheep meat. In
Australia, for example, lamb maintains a significantly larger
portion of the sheep meat market than mutton, largely related
to improved tenderness. This highlights the influence animal age
has on tenderness, but other factors such as breed, genetics,
muscle type, and even location within a muscle can affect tenderness. The pathway by which these shape meat tenderness
revolves around their influence on myofibril type, distribution,
and amount and intramuscular fat (IMF) content. Evidence
suggests that tender sheep meat results from a greater density of
thinner myofibrils with improved collagen solubility and when
IMF levels exceed 3% which is the threshold for consumer
acceptability. Therefore, much investigation toward understanding genes controlling tenderness and their inheritability has been
undertaken for inclusion into breeding programs. However,
genetic contribution to tenderness must be balanced with management practices to achieve optimum tenderness.
The management of animal stress prior and during slaughter, particularly during lairage and on the kill floor, is important
to improve meat tenderness. It is also common following
slaughter and during the processing stage to apply other
methods to further enhance tenderness. These methods include
(1) mechanical stretching, wherein a carcass is positioned (e.g.,
pelvis hanging, or tenderstretch) or physically manipulated
(e.g., carcass pull); (2) temperature manipulation, which
avoids toughening in response to cold shortening or heat shortening in combination with various hygienic and spoilage considerations; (3) pH modification, to prompt a better and
hygienic pH and temperature decline rate and prevent cold
shortening; and (4) aging; increasing the period between
slaughter and consumption of meat kept refrigerated allows
myofibrils and muscle structures to tenderize due to changes

Sheep: Meat

in enzymatic activity and provide less resistance to mastication

although this approach may allow contamination and faster
spoilage. With the exception of aging, these methods aim to
prevent myofibril shortening during which their density and
therefore toughness increase. Research has found that a combination of these approaches results in most significant increases
in meat tenderness, especially when electrical stimulation of a
carcass is used to manage the pH of carcass post-slaughter. This
proves an effective and cheap way to rapidly change pH via the
stimulation of glycogen metabolism.
Flavor refers to the combination of consumer smell/aroma
and taste sensations immediately prior to and during consumption. Smell is stimulated by the numerous volatile compounds
released from sheep meat, mostly released following cooking,
perceived by the olfactory receptors within consumer nasal cavities. These volatile compounds often result from protein and
lipid lyses during cooking, but not much information exists
linking specific compounds with sheep meat aroma with
exception to 2-ethyl-3,6-dimethylpyrazine and 2-pentylpyridine
(heterocyclic compounds) and 3-methylindole acid and
4-methylnonanoic acid (MNA) and 4-methylphenol (skatole
compounds), which have been associated with a mutton
aroma. Nevertheless, it can be construed that their prevalence
relies upon the base compounds of sheep meat, which may be
influenced by animal diet. Taste refers to the stimulation of taste
bud receptors within the mouth and throat to sheep meat compounds, especially branched-chain FA, which are identified as a
precursor to the unique sheep taste. Therefore, taste relies on
many of the same compounds as smell and is also manageable
with dietary controls. Hence, smell and taste are often referred to
as flavor and are influenced by animal diet. For example,
4-methylphenol, methyloctanoic acid, and MNA content are
intensified when animals are fed with pasture-based diets and
result in a pastoral sheep flavor. Also, it is general perception
that when sheep are fed with canola and other oil-rich feeds,
their meat can develop a foreign or atypical lamb/mutton flavor.
Nonvolatile compounds from sheep meat can also influence
flavor, including antioxidants, carotenoids, and lipids. The contribution from these compounds can result from their opposing
influence on lipid peroxidation from which many off-odor and
sour or off-tasting compounds are a by-product. As a rule, however, it is difficult defining an ideal sheep meat flavor because
consumer culinary preferences, cooking methods, and cultural
heritage can all influence its acceptability. This can affect the
objective measurement of flavor, and therefore, it is typically
measured using a trained sensory panel and reported within
the context of the sample represented. This is necessary as, for
example, British and Australian sheep meat consumers prefer
stronger sheep flavors than Asian consumers, and if both these
sampled the same sheep meat, they could provide different
results on consumer perception and acceptability.
Many consumers first encounter with sheep meat is in a
retail environment, wherein an impression of freshness,
quality, and value is often formulated based on appearance
and color. Research has demonstrated that sheep meat must
have a bright cherry red color and bright white fat content to
match consumer ideal or preference. Achieving this appearance
is essentially a function of myoglobin (Mb) oxidation with
Mb being a complex protein with a central iron atom and a role
in oxygen transportation that is plentiful in sheep meat. This Mb

753

will adopt a deoxymyoglobin (DMb) structure when sheep

meat is stored in an anaerobic environment but will change to
oxymyoglobin (OMb) when exposed to oxygen in a reversible
reaction. This causes a color shift from purplish to cherry red.
Prolonged exposure to oxygen, however, will result in OMb
assuming a metmyoglobin (MMb) structure and will consequently instill a brownish color to sheep meat. The concentration of MMb is directly associated with this discoloration; hence,
work has been undertaken to find a threshold value indicative of
perceived quality and methods to prevent this threshold from
being breached. For example, improving sheep meat antioxidant
(e.g., vitamin E) and other micronutrient (e.g., beta-carotenoids)
contents with dietary management (Figure 1) and using innovative meat packaging technologies have provided means to
manage its appearance. This proves true for both meat and fat
appearances.
When juiciness is used within the context of a sensory quality,
it describes the amount of fluid released from sheep meat upon
maceration in combination with that contributed from the consumer via salivation. Salivation is prompted by the IMF content,
and a threshold level of 3% of IMF has been identified as necessary for juicy meat. The fluid discharged from sheep meat when it
is chewed is a function of its water-holding capacity (WHC),
which is the potential of meat to withhold liquid during (1)
storage periods, otherwise lost as drip and purge; (2) heating,
referred to as cooking loss although the cooking method can
have significant implications on this factor; and (3) chewing and
mastication, with fluid released promoting perceived juiciness.
Ideally, sheep meat should be juicy and this requires a high
WHC. The WHC can be influenced by animal carcass energy
reserves, which are measureable as pH with high pH equating
to high energy reserves and therefore heightened WHC. This may
be improved using electrical stimulation technology. However,
the WHC does contribute to carcass yield, and when the WHC is
low, meat yield as well as subsequent profits are less. Not only
the amount of water but also the distribution of water in muscle
tissue has an effect on meat color, firmness, and drip loss. Under
postmortem conditions with the onset of rigor, water moves
from the myofibrillar compartment into the intermyofibrillar
space and then in turn into the extracellular space. The degree
of myofibril shrinkage will impact on the reflection of light from
the muscle surface, and the ultimate pH will impact on this by
affecting the amount of water loss.
Ultimately, an accurate indication of overall liking or
achievement of a consumer-acceptable product can be provided by combining these four major sensory characteristics.
However, this relies upon accurate assessment, which is often
difficult due to the sometimes subjective nature of sensory
panel protocols, experience of panel members conducting the
evaluation, and expense needed to facilitate a statistically sound
insight. As a result, efforts have been distinctly focused toward
developing and validating objective measures as a proxy for
consumer responses.

Spoilage and Preservation

All meat is susceptible to spoilage, and this can have detrimental implications upon perceived value, consumer safety, and
wastage. Spoilage refers to the degradation or downgrade of

754

Sheep: Meat

P (slope) = 0.14, P (diet) = 0.0021

P (slope) = 0.42, P (diet) = 0.00047

16
a*-value day 3 and day 4

a*-value day 3 and day 4

16
15
14
13

Pasture

12
11

Grain

15
Pasture
14
13
Grain
12
11

0.0

0.2

0.4

0.6

0.8

0.8

1.0

Heme iron (mg/100 meat)

(a)

P (slope) = 0.07, P (diet) = 0.0006

a*-value day 3 and day 4

a*-value day 3 and day 4

Pasture

Grain

12

80

70

90

100

15

Pasture

14
Grain
13
12

100

110

Total n-3 (mg/100 g meat)

(c)

140

160

180

P (slope) = 0.097, P (diet) = 0.029

16

16
a*-value day 3 and day 4

a*-value day 3 and day 4

120

Total n-6 (mg/100 g meat)

(d)

P (slope) = 0.05, P (diet) = 0.014

Pasture

14
Grain
13
12
11

160
(e)

1.6

11

11

15

1.4

16

14
13

1.2

P (slope) = 0.097, P (diet) = 0.029

16
15

1.0

Non heme iron (mg/100 g meat)

(b)

15
14
13 Pasture
12

Grain

11

180

200

220

240

260

Total PUFA (mg/100 g meat)

280
(f)

Vitamin E (mg kg1 meat)

Figure 1 Scatter plots of redness (a*) of sheep meat after 3 and 4 days of retail display with (a) heme iron, (b) nonheme iron, (c) total omega-3 (n3),
(d) total omega-6 (n6), (e) total polyunsaturated fatty acid (PUFA), and (f) vitamin E, measured in the M. longissimus lumborum. Parallel
regression lines are fitted for lambs grazing pasture only (dashed) or offered with hay supplemented with grain (solid). Adapted from Ponnampalam, E.
N., Butler, K. L., McDonagh, M. B., Jacobs, J. L. and Hopkins, D. L. (2012). Relationship between muscle antioxidant status, forms of iron,
polyunsaturated fatty acids and functionality (retail color) of meat in lambs. Meat Science 90, 297303.

Sheep: Meat

sheep meat quality, both sensory and nutritional, to an extent

after which a meat product fails to meet safety and quality
standards dictating consumption. The spoilage of sheep meat
can be assessed as discoloration, a loss of integrity and texture,
the development of sour and off-flavors and odors, loss of
moisture, and/or visible bacterial and fungal infection. Bacterial and fungal infection could indicate the contamination of
meat with pathogenic bacteria or toxic by-products from
fungal and bacterial growth. As a general observation, meat
spoilage results in a compromise of core compound structures,
such as the lipid and protein fractions, from oxidation and/or
microbial infection.
Sheep meat is a rich source of SFA and UFA, the latter being
especially prone to lipid peroxidation. Lipid peroxidation is
the oxidation of a fatty acid molecule and occurs as three
sequential phases: (1) initiation, when a UFA reacts with a
free radical species and donates a hydrogen ion to become a
fatty acid radical; (2) propagation, which occurs during a fatty
acid radical reaction and acceptance of an oxygen atom (or
oxidizes) to become a peroxyl-fatty acid radical this can then
be used in the former phase and create a cyclic reaction; and
(3) termination, when a fatty acid radical reacts with a nonradical (e.g., antioxidant), which interrupts the initiation and
propagation reciprocal interactions. In instances where the
termination phase fails to occur, the previous phases will continue until the original UFA is completely degraded into its
many potential secondary oxidative products, which include
reactive aldehydes, ketones, alcohols, hydrocarbons, epoxy
compounds, and volatile organic acids. Lipid peroxidation of
meat does compromise not only nutritional and healthpromoting advantages from UFA but also meat structural integrity during prolonged storage and cooking and can affect
perceived juiciness and tenderness qualities. Furthermore, the
secondary products from lipid peroxidation can have a negative influence over sheep meat flavor and cause atypical and
undesirable tastes and aromas. These can also be employed to
initiate protein oxidation, and in this respect, lipid peroxidation acts as a precursor reaction.
Protein oxidation, or proteolysis, is catalyzed when protein
peptides are exposed to oxygen and reactive radical species.
Hence, this commonly follows lipid peroxidation because of
the high reactivity of peroxyl-fatty acid radicals and other secondary oxidative products with protein peptides. This reaction
causes the protein structure to change as peptide bonds are
cleaved and amino acid side chains are modified this latter
change being the most common and can result in the synthesis
of protein carbonyl groups. As an upshot from protein oxidation, many covalent intermolecular cross-linked protein secondary species are created. These include disulfide, dityrosine, and
protein hydroperoxide species groups. An outcome from protein
oxidation is a change in protein structure and a resultant inactivation of several enzymatic functions, although often the
proteins become more prone to the effects of proteolytic
enzymes potentially sourced from microbial infections. It is
interesting to note that some amino acids are more susceptible to
oxidation than others, for example, cysteine, tyrosine, arginine,
lysine, and methionine, and therefore, proteins rich in these
amino acids will be preferentially oxidized. The presence of
transition metals can also influence protein oxidation, and this
has implications on heme proteins such as Mb. Mb is oxidized

755

when, as OMb, it autoxidizes before MMb enzymatic reduction,

when it is catalyzed by the degradation of lipid peroxyl-fatty acid
radicals, or when Mb acts as a pseudoperoxidase. However, it is
the central Fe molecule within Mb structures that significantly
contributes to Mb oxidation, as this is readily oxidized from
MbFe(II) to MBFe(III) as OMb becomes MMb. Nonheme Fe
and other transition metal ions can contribute to protein oxidation, provided hydrogen peroxide is present resulting in a
Fenton reaction.
The end result from protein oxidation on sheep meat quality is most apparent on its sensory qualities. For instance, when
meat is aged, its structural proteins within myofibrils and
connective tissues will degrade resulting in increased tenderness. It should be noted that this response is mediated by the
calpain system wherein calpain enzymes, specifically m-calpain
and m-calpain, and several cathepsins sever myofibrils. Protein
oxidation can interfere with the calpain system, with research
showing that oxidation can inhibit m-calpain activity and therefore causes the tenderness response to aging to be lessened.
Protein oxidation also affects juiciness by prompting changes
to myofibril diameters and protein cross-linkage connections,
which can alter sheep meat WHC. Consequently, drip loss is
found to increase during protein oxidation, and less fluid is
retained within sheep meat, which can be released upon chewing; hence, consumer-perceived juiciness is reduced. The effect
of protein oxidation on appearance is centered upon its effects
on Mb as discussed.
Practicality dictates that sheep meat cannot be processed
without experiencing some degree of bacterial or fungal (microbial) contamination. However, not all microbial contaminations contribute to spoilage, and therefore, efforts have been
made to identify causal agents. Examples of spoilage bacteria
that have been identified as hazardous to sheep meat preservation against spoilage include lactic acid bacteria (LAB),
Enterobacteriaceae, Brochothrix thermosphacta sp., and Clostridium spp., as these often continue to proliferate during cold
storage. As a rule, these contribute to spoilage as a result of
their metabolism and subsequent breakdown of sheep meat
protein and lipid fractions and glycogen reserves, which can
influence integrity and texture. Furthermore, a by-product of
metabolism by some microbes is the synthesis and deposition
of often rancid-smelling or rancid-tasting compounds, which
makes sheep meat unacceptable these are commonly sulfurrich metabolism products. In other instances, the by-products
or the microbes themselves may be toxic to consumers and pose
health risks. For instance, E. coli, Salmonella, and Campylobacter
jejuni are three examples of pathogenic bacteria that if found
above threshold levels on sheep meat will cause it to be inedible. Fortunately, microbial contamination can be controlled by
direct application of health organization-approved antimicrobial agents to a sheep meat carcass or, more commonly, the
manipulation of storage conditions to match those that inhibit
microbial proliferation. This latter control focuses on temperature and oxygen availability factors primarily during the storage
stage in the journey of sheep meat from processor to consumer
through retailing sector.
With these effects of oxidation and microbial infection on
sheep meat understood, it becomes apparent that spoilage is
unavoidable. Instead, delaying spoilage onset and restricting
its extent provide the best measure of combating spoilage until

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Sheep: Meat

consumption. This is routinely achieved by using specialized

storage containments with inherent preservation and quality
enhancement properties. For example, keeping sheep meat in
chilled or frozen storage is a time-tested approach to restrict
spoilage, as these low temperatures limit oxidation and microbial growth and therefore slow their contribution to spoilage.
Recent efforts have used vacuum packaging or modified atmospheric packaging to shift the atmospheric gas surrounds of
sheep meat to lower its exposure to oxygen a necessity for
propagation and growth of many spoilage bacteria and oxidation. Increasingly, intelligent sheep meat packaging is emerging as a means to restrict spoilage by better informing
stakeholders along the supply chain to the extent of spoilage
using embedded sensory equipment. These packages may also
have antioxidants or antimicrobial agents incorporated into
packaging films to directly reduce spoilage. However,

a*- value day 3 and day 4

18

Adjusted for heme iron

15
12
9
6
3
0
0

Muscle vitamin E (mg kg1 meat)

Figure 2 Relationship (P 8.9
107) between redness (a*) of sheep
meat after days 3 and 4 of retail display and M. longissimus lumborum
vitamin E, after a* was adjusted for muscle heme iron for lambs
offered with pasture only () or offered with hay supplemented with
grain (D). Adapted from Ponnampalam, E. N., Butler, K. L., McDonagh,
M. B., Jacobs, J. L. and Hopkins, D. L. (2012). Relationship between
muscle antioxidant status, forms of iron, polyunsaturated fatty acids and
functionality (retail color) of meat in lambs. Meat Science 90, 297303.

a*-value day 3 and day 4

18

Adjusted for vitamin E

15
12
9
6
3

preserving against spoilage is not solely a role for processors

as producers make significant contributions using feeding
management. Research has demonstrated that sheep fed with
diets rich in antioxidants and vitamin E (a-tocopherol) will
have increased antioxidant levels in their meat and hence
better resistance to oxidation when compared with control
animals (Figures 2 and 3). This has shown that when vitamin
E levels exceed 2.95 mg kg1 of muscle, sheep meat will be
better preserved against lipid peroxidation. All these preservative methods share the common priority to interrupt or delay
oxidation and microbial growth, and, in doing so, prolong
sheep meat longevity.

Conclusion
The production and consumption of sheep meat support many
persons around the globe. This support may be economic, with
a section of most communities basing their livelihoods on
sheep meat retailing, processing, and/or producing, or support
may be nutritional, as sheep meat proves to be a highly nutritious source of many vitamins, minerals, protein, and healthy
fats such as omega-3s. Consequently, it is meritorious for all
stakeholders to explore production strategies, animal management, welfare of animals from farm to slaughter, and technological methods at processing and retail to improve sheep meat
quality and achieve a shared priority to deliver a product that a
consumer finds attractive and acceptable. This has resulted in
much research into what describes consumer preferences for
sheep meat and the underlying principles of preservation,
spoilage, or the degradation of these quality traits. Understanding spoilage has allowed steps toward preserving or maintaining sheep meat quality until consumption to be successfully
achieved, as fundamentally, it is consumer preference and
satisfaction with a sheep meat product upon which the entire
industry is based.

See also: Beef; Fats: Classification and Analysis; Fats: Production and
Uses of Animal Fats; Fatty Acids: Determination and Requirements;
Fatty Acids: Essential Fatty Acids; Fatty Acids: Fatty Acids; Fatty Acids:
Trans Fatty Acids; Iron: Biosynthesis and Significance of Heme; Iron:
Physiology of Iron; Iron: Properties and Determination; Meat:
Conversion of Muscle into Meat; Meat: Eating Quality and Preservation;
Meat: Role in the Diet; Meat: Structure; Protein: Digestion, Absorption
and Metabolism; Protein: Food Sources; Protein Quality and Amino
Acids in Maternal and Child Nutrition and Health; Protein:
Requirements; Proteins: Chemistry, Characterization, and Quality;
Sensory Evaluation: Sensory Rating and Scoring Methods.

0
0

0.2

0.4

0.6

0.8

1.2

Heme iron (mg/ 100 g meat)

Figure 3 Relationship (P 0.0031) between redness (a*) of sheep
meat after days 3 and 4 of retail display and M. longissimus lumborum
heme iron, after a* was adjusted for muscle vitamin E for lambs offered
with pasture only () or offered with hay supplemented with grain (D).
Adapted from Ponnampalam, E. N., Butler, K. L., McDonagh, M. B.,
Jacobs, J. L. and Hopkins, D. L. (2012). Relationship between muscle
antioxidant status, forms of iron, polyunsaturated fatty acids and
functionality (retail color) of meat in lambs. Meat Science 90, 297303.

Further Reading
Daley CA, Abbott A, Doyle PS, Nader GA, and Larson S (2010) A review of fatty acid
profiles and antioxidant content in grass-fed and grain-fed beef. Nutrition Journal
9: 10.
Dixon RM and Stockdale CR (1999) Associative effects between forages and grains:
consequences for feed utilization. Australian Journal of Agricultural Research
50: 757774.
Faustman C, Sun Q, Mancini R, and Suman SP (2010) Myoglobin and lipid oxidation
interactions: mechanistic bases and control. Meat Science 86: 8694.

Sheep: Meat

Hopkins DL, Stanley DF, Martin LC, Ponnampalam EN, and van de Ven R (2007) Sire
and growth path effects on sheep meat production. 1. Growth and carcass
characteristics. Animal Production Science 47: 12081218.
Howe P, Buckley J, and Meyer BJ (2007) Long-chain omega-3 fatty acids in red meat.
Nutrition and Dietetics 64: 135139.
Howes NL, Ahmed AED, Burritt DJ, and Campbell AW (2015) Opportunities and
implications of pasture-based lamb fattening to enhance the long-chain fatty acid
composition in meat. Comprehensive Reviews in Food Science and Food Safety
14: 2236.
Lobley GE (1998) Nutritional and hormonal control of muscle and peripheral tissue
metabolism in farm species. Livestock Production Science 56: 91114.
McAfee AJ, McSorley EM, Cuskelly GJ, Moss BW, Wallace JMW, Bonham MP, and
Fearon AM (2010) Red meat consumption: an overview of the risks and benefits.
Meat Science 84: 113.
Montossi F, Font-i-Furnols M, Campo M, San Julian R, Brito B, and Sanudo C (2013)
Sustainable sheep production and consumer preference trends: compatibilities,
contradictions, and unresolved dilemmas. Meat Science 95: 772789.
Ponnampalam EN, Butler KL, Hopkins DL, Kerr MG, Dunshea FR, and Warner RD
(2008) Genotype and age effects on sheep meat production. 5. Lean meat and fat
content in the carcasses of Australian sheep genotypes at 20, 30 and 40 kg carcass
weights. Australian Journal of Experimental Agriculture 48: 893897.
Ponnampalam EN, Butler KL, McDonagh MB, Jacobs JL, and Hopkins DL (2012)
Relationship between muscle antioxidant status, forms of iron, polyunsaturated

757

fatty acids and functionality (retail color) of meat in lambs. Meat Science
90: 297303.
Ponnampalam EN, Norng S, Burnett VF, Dunshea FR, Jacobs JL, and Hopkins DL
(2014) The synergism of biochemical components controlling lipid oxidation in
lamb muscles. Lipids 49: 757766.
Scollan ND, Hocquette JF, Nuernberg K, Dannenberger D, Richardson I, and Moloney A
(2006) Innovations in beef production systems that enhance the nutritional and
health value of beef lipids and their relationship with meat quality. Meat Science
74: 1733.
Watkins PJ, Frank D, Singh TK, Young OA, and Warner RD (2013) Sheepmeat flavor
and the effect of different feeding systems: a review. Journal of Agricultural Food
Chemistry 61: 35613579.
Wood JD, Enser M, Fisher AV, et al. (2008) Fat deposition, fatty acid composition and
meat quality: a review. Meat Science 78: 343358.

Relevant Websites
http://ec.europa.eu European Commission.
www.fao.org Food and Agriculture Organization of the United Nations.
www.foodstandards.gov.au Food Standards Australia New Zealand.
www.mla.com.au Meat and Livestock Australia.
www.usda.gov U.S. Department of Agriculture.