Animal Welfare in Multipurpose Cattle Production Systems and Its Implications On Beef Quality
Animal Welfare in Multipurpose Cattle Production Systems and Its Implications On Beef Quality
Animal Welfare in Multipurpose Cattle Production Systems and Its Implications On Beef Quality
Review
Animal welfare and its influence on beef production are major considerations in many developed
countries. In the developing world, where food insecurity and poverty are prevalent, the welfare of
animals receives low priority due to factors such as traditional customs and beliefs, lack of knowledge
in animal handling and sub standard handling facilities. This is worsened by the fact that cattle are used
for several purposes, such as meat production, milk production, draught power and traditional
ceremonies. Research on animal welfare and how it relates to animal production have been done in the
developed world. The objective of this study is to discuss indicators of animal welfare assessment,
their impact on meat quality and applications to improve beef production in developing countries.
Key words: Behavioural assessments, flight zones, meat quality, ritual slaughter.
INTRODUCTION
Animal welfare refers to the physiological or biochemical ment practices should aim at keeping animals free from
state of an animal as it attempts to cope or respond to thirst, hunger and malnutrition, discomfort, pain, injury
internal challenges or ante-mortem conditions at the time and disease, fear and distress, and should also be able
of observation (Gregory, 1998; Broom, 2000; Grandin, to engage in normal pattern of animal behaviour (Bech et
2001). It represents the mental and physical health of an al., 2008). Non-fulfilment of these needs may expose the
animal in relation to its environment (Smith and Pearson, animal to stressors with detrimental effects on meat
2005). Animal welfare also involves the application of production. Prolonged exposure to stressors disrupts
sensible and sensitive animal husbandry practices to the energy mobilization and reactions involved in stress res-
livestock on the farm. Good animal welfare has a positive ponse, thus affecting the normal body functions, for
effect on production. example, immunity, growth, reproduction and expression
Good practices of animal welfare are underpinned by of normal behaviour (West et al., 2003).
the framework provided in the five familiar freedoms that Concerns for animal welfare and its influence on meat
describe an animal’s fundamental needs (Gregory, 1998; quality are major considerations in many developed
Bech et al., 2008; Vessier et al., 2008). Animal manage- countries (Veissier et al., 2008; Muchenje et al., 2009a;
Strappini et al., 2009). In the developing world, where
food insecurity and poverty are prevalent, the welfare of
animals receives low priority due to lack of knowledge in
animal handling, traditional customs and beliefs, sub-
*Corresponding author. E-mail: [email protected]. Tel: +27
40 602 2059, +27 84 422 1487. Fax: +27 86 628 2967.
standard handling facilities and failure of government
legislation (Veissier et al., 2008). Under communal
production systems, for example, cattle tend to have high
Abbreviations: AWOs, Animal Welfare Organizations; DCB,
dry cutting beef; FS, flight speed; FT, flight time; NEFA, non-
levels of parasites (Muchenje et al., 2008a) and face
esterified fatty acid; ACTH, adrenocorticotropic hormone; CK, nutritional deficiencies due to deteriorating rangelands,
creatine kinase; APP, acute phase proteins; Fb, fibrinogen; Hp, particularly in winter (James and Hazel, 2007; Dube,
bovine haptoglobin; SAA, serum amyloid-A. 2008) and during the cropping season where animals are
1050 Afr. J. Biotechnol.
used for draught power (Chimonyo et al., 2002). In addi- ses the methodologies that could be used to assess
tion, health management systems in developing countries animal welfare practices, their impact and application to
are weak or non-existent and draft animals are being improve beef production in developing countries. The
brutally mistreated and totally neglected (Ramaswamy, intention is to open a discussion among practitioners and
1998). researchers on improving handling procedures for beef
In South Africa, ritual slaughtering during traditional animals because most of the available literature is on pig
ceremonies, especially in the Zulu culture and traditions, production, particularly in developed countries. In
sparks a lot of controversy concerning animal rights and addition, the current situation on animal welfare in the
cultural practices (Mnguni, 2006). During marketing of developing countries and possible areas which need
cattle, animals are taken to abattoirs using various trans- research are highlighted.
port means. Animals are improperly loaded in trucks
which are inappropriate, small, poorly ventilated, uncom-
fortable and even open to direct heat and rain. In CATTLE MANAGEMENT ISSUES AFFECTING
addition, overcrowded trucks and on foot driving for long ANIMAL WELFARE IN THE SMALL HOLDER AREAS
distances with untrained stockmen who use sticks and
goads are common, where bruising is not uncommon. Rearing of cattle is an integral part of agricultural produc-
Indigenous African ethnic pastoral groups, such as the tion systems in semi-arid regions of most developing
Maasai people of Kenya, bleed live cattle by opening a countries, such as Zimbabwe and South Africa
vein on the neck or flank with the point of an arrow and (Chawatama et al., 2003; Mapiye et al., 2009b). There
these practices impose pain, thus raising obvious animal are however, little efforts towards raising of animal
welfare concerns (Alana et al., 2008). The reasons welfare issues in most developing countries and animals
behind these practices need to be understood for effect- are reared under very extensive, range-management
tive intervention. conditions, of which their interactions with people are
There is need to raise awareness of these animal wel- infrequent and nearly always aversive (Ali et al., 2006;
fare issues and enforce animal welfare assessment Dwyer, 2009). Under communal production systems that
methodologies in developing countries (Broom, 2000). are common in most developing countries, high parasite
Identification of poor handling procedures, their control loads, feed shortages, provision of draught power, poor
and preventative measures can be implemented effect- housing conditions and cultural practices tend to impact
tively to reduce losses for producers and livestock heavily on the welfare of animals, particularly cattle
keepers and also to improve the quality of the meat pro- (Ramaswamy, 1998; Chimonyo et al., 2002; Muchenje et
duced. Assessment of animal welfare may contribute, not al., 2008a). Cattle in communal production systems are
only to improved beef production but also promotes used for several purposes. These include meat produc-
animal well-being. Ensuring the well-being of the animal tion, milk production, traditional ceremonies, draught
is an ethical requirement for sustainability of biodiversity, power and production of manure that is used for crop
especially when adapted animal breeds are selected and fertilisation (Mapiye et al., 2009a).
reared in production conditions of their ancestral origin.
A number of methods have been developed to assess
animal welfare and authenticity of beef products in the High parasite loads
developed world (Broom, 2000; Tarrant and Grandin,
2000; Campo et al., 2008). These methods evaluate the Cattle in communal areas have been reported to have
behaviors of animals, assess biochemical and physio- high worm and tick loads (Muchenje et al., 2008a; Ndlovu
logical properties and make quantitative observations. et al., 2009a). The impact of these parasites depends on
Novak et al. (2004) used various complementary para- breed, with the indigenous genotypes showing signs of
meters, such as production indicators (performance and tolerance and even resistance to the parasites (Muchenje
yields), physiological indicators (endocrine and cardio- et al., 2008a). As such, imported breeds which have been
vascular responses), pathological indicators (morbidity, bred under relatively benign environments, fail to cope
diseases and mortality) and ethological criteria, such as and their mortalities are high. The high parasite loads are
abnormal behavioural patterns. Besides their use in common among most of the communal farmers, who are
monitoring the welfare of animals on farm, physiological largely resource-poor, failing to afford veterinary drugs
and biochemical measurements can also be used to and the high cost of hiring veterinarians.
monitor the health status of animals on the farm Dipping is mostly infrequent and in some cases, wrong
(Eckersall, 2000) or at slaughter (Campo et al., 2008; dosages of the acaricide are applied (Moyo and Masika,
Muchenje et al., 2009a, b). 2009). Dip tanks are located far away from where cattle
A number of reviews on the impact of animal welfare on are kept and in some cases the dip tanks are non-
meat quality in the developed world are available functional due to the non-availability of water pumps,
(Gregory, 1998; 2007; Broom, 2000; Ferguson and water and acaricides (Moyo and Masika, 2009). As a
Warner, 2008; Strappini et al., 2009). This review discus- result, animals have to be hove driven over long
Ndou et al. 1051
distances to and from dipping areas and during theses Draught power
journeys animals are given water and allowed to rest.
Non-dipping of cattle due to these limitations results in The small sizes of arable land and the lack of sophisti-
high tick loads that cause tick damages to parts, such as cation of agricultural operations in most smallholder
udders and scrotums and tick-borne diseases such as farming areas make the use of animal draught power
heart water and red water (Muchenje et al., 2008a). In inevitable. Oxen, bulls, cows and heifers are used for
such cases, it is advisable to use indigenous cattle pulling ploughs, carts, cultivators and sledges (Chimonyo
breeds that are tolerant to ticks and less susceptible to et al., 2002). Although pain is exerted during these
tick-borne diseases. operations, it is crucial to have the users of these animals
trained. During ploughing, for example, whips are used
for the animals to move fast. At the end, draught animals
Feed shortages are left with wounds and scars. In India and Southern
Africa, the use of traditional yokes and harnessing, such
Grazing in most communal production systems are un- as those illustrated in Figure 1, bruise the neck and
controlled and continuous grazing systems are common cause pain to the animal (Ramaswamy, 1998). Other
(James and Hazel, 2007; Dube, 2008). Besides, most agricultural implements attached to draught cattle, such
communities are overstocked, since ownership of grazing as ploughs and carts are of crude design and inefficient,
rangelands is communal; little sense of ownership of the which hurt draught animals.
grazing resources exists (Dube, 2008). The cattle, The working and resting times for draught animals and
therefore, lose body condition, particularly in winter and its impact on beef production has not been extensively
spring (Mapiye et al., 2009b). The poor body condition researched under local conditions. Draught animals are
also exacerbates the impact of the parasites on the also kept in the herd for long periods of time (Mapiye et
welfare of the animals. There is need to investigate the al., 2009b). Draught cattle need to be in appropriate body
welfare status of animals in these deteriorating range- condition (above 3.0) to enhance efficiency, which calls
lands so that appropriate stocking densities can be for providing them with adequate feed resources
formulated. particularly, before the rainy season begins.
1052 Afr. J. Biotechnol.
Poor housing conditions tightly tie all the legs together just below the abdomen
region, such that the knot exert pressure on the abdomen
Kraals used for housing cattle do not have roofs, in many in order to facilitate the rapid release of blood following
instances. During the rainy season, the animals are, slitting without stunning. Alternatively, the Vahera people
therefore, exposed to excessive rain, especially at night may tie three legs only and then the other leg is forced to
and become susceptible to diseases, such as foot rot hook at the back of the head whilst a group of strong men
(Moyo and Masika, 2009). During the rainy season cow pull the three legs from the back of the animal. Then the
dung in the kraals create muddy conditions. This makes it beast is cut on the neck without stunning. In Matabele-
uncomfortable for cattle to lie down, in addition to land South Province of Zimbabwe, the beast is
creating conducive conditions for diseases. Kraaling the restrained, its neck or horns are tied on the tree, and then
cattle for long periods also results in limited time for blood is let out by cutting on the neck in the halal way
grazing. Awareness to these threats to the welfare of following stunning by hitting the cow on the back of the
cattle needs to be raised among the poor farmers. head with a sharp axe. Furthermore, the Maasai people
bleed live cattle to obtain blood for traditional reasons
without any anaesthesia (Fratkin, 2001). In general, the
PRE-SLAUGHTER WELFARE ISSUES IN slaughter practices vary with region and culture. These
SMALLHOLDER AREAS AND THEIR EFFECTS ON practices require investigation and documentation, to
MEAT QUALITY lessen the pain animal suffer during the slaughter pro-
cess.
Pre-slaughter welfare issues are influenced by factors Slaughtering without stunning is associated with stress
including tradition, social customs and beliefs, inefficient of restraint, pain of the cut and undue distress whilst the
policy implementation and government interventions, animal is bleeding out (Gregory, 2005). Cutting the neck
ritual slaughter, lack of education on animal handling, in the unanaethetised state is likely to involve physio-
mixing of animal social groups from different households, logical events which are characterized by undesirable
poor transport condition, inappropriate handling facilities sensations that are likely to be a sense of shock,
and poor slaughterhouse conditions (Mnguni, 2006; comparable to an electric shock (Gregory, 2005). Bleed-
Gregory, 2007; Muchenje et al., 2009a,b). During the ing live animals in the Maasai culture is likely to cause
ante-mortem handling conditions from rearing to slaug- pain which is almost the same as in unanaethetised stun-
hter, cattle are subjected to various internal and external ning (Gregory, 2005).
challenges, which they perceive through specific features In some developing countries in Asia, there are beliefs
or experience. They will subsequently respond or cope that stress benefits some quality traits, such as tender-
with physiological and biochemical changes and products ness resulting in cattle being baited with dogs, chased
that influence the process of transformation of muscle to through streets by mounting stockman equipped with
meat (Broom, 2000; Muchenje et al. 2009a, b). goads (Gregory, 2007). On the other hand, the South
African Zulu slaughtering method, for example, involves
stabbing of the animal on the stomach by using a spear
Tradition, social customs and beliefs and then forcing the animal to move some distance
(Mnguni, 2006). Physical activity for hours before
Failure to implement animal welfare issues can also be slaughter which is common in the Asia, reduces glycogen
ascribed to the weaknesses of traditional customs and concentration and plasma glucose levels below critical
beliefs among some ethnics groups, such as the Zulu, values (Chambers et al., 2001) eventually leading to
Karanga, Fulani and the Maasai in Africa (Mnguni; 2006; reduced lactic acid and increased meat pH above critical
Gregory, 2007; SAPA, 2009). During traditional cere- range of 5.5 to 6.0 (Tarrant and Grandin, 2000; Kannan
monies, human-animal interactions involve constant et al., 2002; Muchenje et al., 2008b) and reduced energy
penetration of animal flight zones, use of sticks and value. Ultimately, pH influences most of the meat eating
goads in driving animals, restraint, feed deprivation, quality traits (Aklilu, 2002; Andersen et al., 2005;
noise, agitation, ululating, shouting and passing angry Muchenje et al., 2008a). On the other hand, Grandin
vocals to the animals when moving them in the required (2000a) established that, when the muscle pH appro-
direction or when expressing some traditional sentiments aches the value of 6.0, the beef tenderness decreases
(Mnguni, 2006; Gregory, 2007). The Fulani culture in and later start to increase as the pH increases from 6.2 to
Africa encourages stockmen to pass threats or willful 6.6, as shown in Figure 2. Although at high pH (6.2-6.6),
behaviors by hitting the animal and failure to do so from the cal pain enzyme system gains more activity and
childhood is regarded as not being courageous (Lott and tenderises meat, the meat becomes dark (Silva et al.,
Hart, 1977). On the other hand, in Asia, cattle awaiting 1999) due to the smaller amount of light reflected where-
slaughter are baited by dogs in the streets to make the as there is high absorption of light radiation.
meat more tender (Gregory, 2005). Napolitano et al. (2002) revealed that, traditional
In Zimbabwe, the Vahera people during slaughter, rearing methods with high levels of animal welfare and
Ndou et al. 1053
Figure 2. Relationship between meat tenderness and ultimate pH. The higher the kgF
value, the less tender the meat (Adapted from Anon, undated).
product specificity may soon assume economic rele- should be nursed. However, it is by logic that animal righ-
vance. Therefore, there is need for research on these ts cannot be reversed but once people appreciate
traditional practices in the developing countries under the civilization in this era of globalization, cultures can be
local conditions and also including the resource users. In adjusted to accommodate welfare of animals. Record
other words, there is need for marriage of stockman, keeping and animal identification is however, still of low
scientists and policy, by incorporating ethnic groups in priority in developing countries but are important to allow
research and formulation of animal welfare policies, for traceability of animal products and conditions that are
this provides an opportunity to capture their perceptions used to produce them in the meat industry.
and aspirations thereby developing shared policies and To enhance traceability during welfare assessment,
improving implementation. According to a report in The smallholder farmers are encouraged to introduce
Citizen (2009), making a judgement between the rights of identification methods such as ear tags from the time of
animals, people’s cultural and religious rights impose a birth up to the time of slaughter (Smith and Pearson,
mammoth task to the judiciary, thus demands careful 2005). The response of animals to handling procedures is
scrutiny. multi-dimensional, thus to fully ascertain cattle welfare
status, all aspects of condition of life, including health and
disease, behaviour, husbandry and management should
Inefficient policy implementation and government be traceable up to the time of consumption of beef
interventions (Fitzpatrick et al., 2006; Smith et al., 2008). Government
and non-profit organizations are invited to formulate legis-
Animal welfare issues are increasingly becoming the sub- lation and raise animal welfare awareness campaigns in
jects of scientific study across the world, receiving major developing countries.
considerations in most developed countries and have led It should also be noted that, most benchmarks on
to government interventions and formation of non-profit animal welfare conditions are based on research which
Animal Welfare Organizations (AWOs) (Ramaswamy, has been done in developed countries under more ad-
1998; Fitzpatrick et al., 2006; Vessier et al., 2008; vanced animal welfare conditions (Grandin, 2007;
Strappini et al., 2009). The impediments to proper welfare Gregory, 2007; Strappini et al., 2009). In developing
in developing countries can also be attributed to poverty, countries, formulation and implementation of appropriate
lack of knowledge on the importance of proper handling, policies that can consider the local conditions and
lack of resources and inefficient government support for practices are of paramount importance to promote the
resource-poor farmers, for example, the budget by the well being of animals and proper use of resources.
Animal Welfare Board of India could not meet the need of
establishing enough AWOs (Ramaswamy, 1998).
Furthermore, some laws have been enacted in some Ritual slaughter and lack of knowledge in animal
developing countries, such as India and South Africa, but welfare
punishment is nominal and the laws are too cumbersome
to implement and government interventions are lax or Information on ritual slaughters is limited because most
scarce (Ramaswamy, 1998; SAPA, 2009). issues to do with traditional ceremonies tend to be a
In the law enforcement process, animal rights need to closely guarded secret by the elders. According to
be protected, at the same time, the people’s cultures Mnguni (2006), during ritual slaughtering in the Zulu
1054 Afr. J. Biotechnol.
culture, such as at funerals (Umngcwabo), coming of age If cattle are confined in spatially limited areas, such as
(Umemulo) and at weddings (Umshado), a group of crowded waiting pens at high stocking densities or in
people surrounds the animal (usually a bull) to be indoor loose-housing systems common in developing
slaughtered in its visual field and cattle are slaughtered in countries, animals may fail to respect individual distances
the pens without stunning or appropriate handling. The (Calhim et al., 2006; Aschwanden et al., 2008) and sub-
beast is then stabbed using a spear and the animal is not sequently results in animal to animal interactions, such as
expected to be killed instantly to encourage multiple attacks, threats or avoidance behaviours. Fighting and
stabbing (Mnguni, 2006), thus, raising animal welfare bruising is further increased due to the fact that, most of
concerns due to pain caused by frequent stabbing and the cattle slaughtered at small-scale abattoirs come from
not rendering the animal unconscious before slaugh- different herds and they are of different sizes. Bruising
tering. When the animal takes long to die, it would have may also result due to lack of appropriate handling
as many stab wounds as possible and the bridegroom is facilities and the use of whips and sticks in driving cattle
expected to pay more to the in-laws. The bridegroom on hooves. Dark, firm and dry meat and also pale
knows that he is to pay more when there are more stab coloured meat are associated with fighting and bruising of
wounds, this encourages less and less stab wounds cattle depending on the time at which fight occurred prior
which is desirable in terms of animal welfare. to slaughter and these occurrences can cause a loss of
The Ukweshwama ceremony of the Zulu culture of edible beef parts and carcass devaluation (Grandin,
South Africa includes the slaughtering of a bull with bare 2000a).
hands, also without stunning (Mahlangu, 2009; SAPA, Information on the differences in the individual dis-
2009; The Citizen, 2009). During this slaughtering pro- tances of the Tuli, Nguni, Mashona cattle and their
cess, the Zulu warriors strangle a bull to death to crosses is lacking but may be useful in assisting farmers
celebrate the first fruits of the harvest in the first week of to develop welfare standards, such as duration of resting
December (Mahlangu, 2009). This practice led to some in the lairage, stocking densities that conform to the local
animal rights groups in South Africa taking the Zulu king conditions and designing holding pens to reduce social
to court because this was perceived as cruelty to animals conflicts between animals. Scoring of individual distances
(The Citizen, 2009). Those who practise the Zulu culture in conjunction with flight zones will also reduce isolation
argue that, they love animals such that in all ceremonies and crowding at smallholder slaughterhouse.
that are performed animal welfare is seriously taken care
of (The Citizen, 2009). Probably what is required in such
cases is awareness to threats to animal welfare without Poor transport condition
necessarily demeaning one’s culture.
These may not only impart mental well-being of cattle Although vehicle transport on tarred roads are replacing
and cause pain to the animal, but may reduce consumer common methods such as driving on hooves and gravel
acceptability of beef produced from abused animals road in developing countries, Maria et al. (2003) reported
(Gregory, 2007; Bech et al., 2008). Severity welfare that, long transport journeys on tarred roads before
procedures, such as during ritual slaughter in the Zulu slaughter increased the risk of dry cutting beef (DCB).
culture, may influence the ability of meat biochemical Such defaults can be reduced by resting animals in the
components to fall below or above the appropriate lairage for 36 h (Kuzmanvic and Elabjer, 2000). Most of
threshold levels, thereby compromising the beef quality the vehicles used for transporting animals are not roofed
traits of economic importance, such as colour, juiciness, and they expose animals directly to the sun radiation.
tenderness, flavour and shelf life (Lawrie and Ledward, However, there is no information on the response of
2006). It is important for stockpersons to understand the animals or on possible alternatives to ensure animal wel-
behaviour of cattle so that stress can be reduced from fare, for cattle transported by transhumance and gravel
rearing up to the time of slaughter (Grandin, 2006; roads for these methods are still common for transporting
Gregory, 2008). cattle to the markets or abattoirs in developing countries,
especially in the Sub-Saharan African region. Animals
are inevitably transported for long distances from rural
Mixing of different social groups markets to urban smallholder abattoirs, lengthy journeys
place enormous demands for energy metabolism on the
Prior to transportation to the slaughterhouse or at the animal and may be the reason of depleted muscle
slaughterhouse, cattle from different social groups are glycogen pre- and post-mortem thus, less lactic acid and
gathered to a handling point, if there is any or are just consequently high beef pH post-mortem.
driven to the slaughterhouse as groups from different
pens or households (Grandin, 2000c; Lammens et al.,
2007). During this period, different social groups from Inappropriate handling facilities and poor
different pens are mixed thus influencing flight zones and slaughterhouse conditions
individual distances or social distances between and
within animal groups (Aschwanden et al., 2008). In most smallholder farming areas in the Sub-Saharan
Ndou et al. 1055
region, animals in holding pens are stressed due to len- indicators that the animal is having challenges in coping
gthy durations at market places, during auctioning or at with the welfare procedures and in most cases some
the lairage, poor handling facilities and introduction into aspect of the situation is aversive (Broom, 2000). During
different social groups resulting in fights, bullying, bruises the pre-slaughter period, animals perceive specific fea-
and mounting (Grandin, 2000a). Such situations influence tures through previous experience and learning (Gregory,
cattle well-being (Muchenje et al., 2009a) and leads to 2007) and will subsequently react through appropriate
decreases in glycogen levels below the critical threshold behavioral responses that influence meat eating quality
levels, thereby raising meat pH (Warris, 1990) and (Muchenje et al., 2009a). Although evidence exist that
consequently imposes detrimental effects such as man-animal relationships during handling can have major
reduced beef keeping quality and dark cutting. In many impact on both production and welfare of animals
communal areas in Southern Africa, cattle are slaugh- (Munksgaard et al., 2001), information on the behavior of
tered in the kraals and handling facilities are lacking cattle during gravel road transport is scarce, but it is
(Mnguni, 2006). The situation is further exacerbated by important in indicating possible modifications to improve
underdeveloped passageways into the smallholder handling facilities (Tarrant and Grandin, 2000) and how
slaughter-house which allows vision of novelty situations animals acclimatize under extensive management condi-
or unfamiliar stockman and poor operational techniques. tions. The commonly used behavioral measurement
As a result, animals that are easily agitated, such as Bos methods include temperament, flight zones, visual fields
indicus, have greater chances of producing meat with and vocalizing scoring. Compilation of these measure-
defaults (Voisinet et al., 1997a) which may reduce the ments is easy and cheap such that resource-poor
monetary rewards for producers. farmers, if trained properly, can produce accurate on farm
Despite recent reports in Nguni and Bonsmara cattle welfare audits.
(Muchenje et al., 2009b), information on the biochemical
changes that take place in relation to stress respon-
siveness and the depletion of glycogen and its effect on Temperament
beef quality of most indigenous cattle breeds, such as
Tuli and Mashona in the developing world, is lacking. Temperament refers to an animal's behavioral expression
Furthermore, there is little or no information on the animal in response to challenging situations such as human
welfare status during transportation and performance of handling or presence (Ferguson and Warner, 2008).
cattle following climatic changes and gradual deteriora- Temperament and stress are strongly related to cattle
ting rangelands of most developing countries. There is behavior and can be used in various aspects to assess
therefore, need to develop animal welfare assessment the previous experience of the animal and predicting
protocols that can be easily adopted in the developing optimum welfare techniques to reduce meat defects.
world. Such animal welfare assessment protocols can be Crush scores can be used to evaluate temperament usi-
developed using relevant, convenient, accurate and/or ng a 1 (calm) to 5 (combative) scales (Campo et al.,
affordable behavioral, quantitative, physiological and bio- 2008). Lower crush scores imply previous optimum
chemical assessment methods that are already available. welfare and consequently chances of obtaining quality
meat which accommodates high consumer concerns.
Although, familiarization of animals to handling
ASSESSMENT OF ANIMAL WELFARE IN procedures makes them easy to handle during the pre-
MULTIPURPOSE CATTLE slaughter period (Grandin, 1993; Gregory, 2008), certain
breeds, such as Bos taurus have calmer temperaments
Due to the multipurpose nature of cattle production sys- than the B. indicus cattle (Fordyce and Goddard, 1984).
tems and limited resources in most developing countries, Studies on the Limousin, Red Bororo and Brahman cattle
the development of animal assessment protocols can be confirm that B. indicus animals are difficult to handle
complex. Despite the possible complications, assessment (Minka and Ayo, 2007) and their meat tends to be
techniques that are developed for typical beef cattle characterized by dark cuttings and toughness. Behrends
production can be adopted to indicate animal welfare et al. (2009) reported that, the response of cattle to novel
status by stockmen and ritual slaughter participants. experience early in life are best predictors of traits
Knowledge of these indicators will go a long way in impacted by temperaments later in life such as beef
raising awareness and minimising pain to cattle pre- tenderness. Docile cattle are associated with greater
slaughter, at slaughter and traditional ceremonies. Cur- average daily gain (ADG) than those agitated during
rent animal welfare assessment methods in use can be routine handling (Voisinet et al., 1997b) and this signifies
categorised into behavioural, quantitative, physiological that, rearing of docile cattle has positive economic
and biochemical classes. implications. Muchenje et al. (2008b) reported that Nguni
cattle raised on natural pastures had acceptable meat pH
Behavioural measurements of animal welfare values which can be used to suggest that they had
calmer temperaments or were properly handled during
Changes in animal behaviors are the most obvious the pre-slaughter period.
1056 Afr. J. Biotechnol.
Figure 3. Diagram of flight zones illustrating the most appropriate positions for moving an animal forward
(Grandin, 2000c).
Animals with aggressive temperament also result in position without moving the eyes or turning the head. The
economic losses to the meat production enterprise due to flight zone and visual fields define the animal’s personal
malicious damaging of the handling facilities, injuries to space such that when a person or any source of danger
the beast or handler, and poor quality meat (Gregory, enters it, the animals will move away, but its size will
2008; Muchenje et al., 2009b). Temperament also corr- slowly diminish with avoidance of the visual fields by
elates well with physiological measures of stress such as stockman, tameness of the animal or when animal
cortisol concentration and catecholamines (Muchenje et previously received frequent and gentle handling
al., 2009a). Animals that are easily stressed are difficult (Grandin, 2000b; Campo et al., 2008).
to handle. Genetic variations in temperament among and Knowledge of flight zones and their association with
within breeds are evident. Heritability estimates as high visual fields of cattle are more useful in preventing stress
as 61% have been reported (Gauly et al., 2001) such than tameness and animal experience especially during
that, selection pressure can be exerted in breeding the pre-slaughter period. Figure 3 illustrates the concept
programmes to improve temperament by selection. There of visual field and flight zones during slaughter of animals
is need to identify and evaluate traits which are correlated (Grandin, 2000c). When driving cattle in their visual field,
with temperament, such as facial hair whorl position in the stockman should stand in the shaded area marked A
beef animals to aid genetic characterization and improve- and B (Figure 3) and approach the animals at the edge of
ment (Lanier et al., 2001). Temperament indicators can the flight zones while staying out of the blind spot
be easily adopted by resource-limited farmers. because, deep penetration of flight zone may cause the
animal to panic (Grandin, 2000c) and induce undesirable
physiological and biochemical changes which may
Flight zones and visual fields compromise with beef meat quality traits. Application of
cattle flight zones and visual fields is more useful in
The flight zone is the area inside the distance which replacing traditional methods employed in developing
when a person or any source of danger enters, the countries, such as using goads, sticks, dogs and making
animal will maintain it or move away (Albright, 2000; aggressive threats or willful behaviors to move the
Grandin, 2007). The visual field is the area of the external animals in a desired direction as well as ululating to the
environment that is visible to the animal at any given animals during traditional ceremonies (Mnguni, 2006;
Ndou et al. 1057
Warris et al. (1994) reported that, vocalizations are noti- Mortality rates
ceably correlated with physiological measures of stress in
pigs. In cattle, vocalization scorings of mooing and Mortality rates are useful tools in assessing poor welfare
bellowing are one of the most obvious if not potential associated with disease and lack of care in farm animals
1058 Afr. J. Biotechnol.
Table 1. Mortality rates that might be tolerated in industrialized welfare of animals is suboptimal. Gregory (2007) pro-
nations before concerns are raised about the welfare of the posed mortality rates which when exceeded, will be
animals or profitability of the enterprise.
cause for concern amongst stock owners (Table 1).
When incidences of deaths increase beyond the accep-
Class of stock or situation Mortality rate (%)
table levels, their causes which are likely to indicate poor
Calves raised of milk replacer 10 welfare to the animals should be investigated.
Feedlot cattle 4
Fat cattle transported for slaughter 0.005
Breeding cattle-ranching system 4 Bruises or body damages
(Gregory, 2007).
Occurrence of bruises, broken limbs or footpad lesions
are sources of pain and have negative impact on animal
welfare as well as on the quality of beef (Gregory, 2007;
(Gregory, 2007), especially during transportation and in Strappini et al., 2009). The thickness of bovine skin
calves during their first 6 months of life (Ortiz-Pelaez et makes it difficult for bruises to be seen during the ante-
al., 2007). In most cases, mortalities during handling and mortem period but their presence, position and age can
transport are usually preceded by a period of poor be detected successfully in the carcasses (Strappini et
welfare due to failure of the physiological activities to al., 2009). Whilst the presence of bruises indicates aver-
cope or withstand external or internal challenges. Poor siveness of the welfare procedure, age and position of
and prolonged cattle transportation using vehicles on the bruises provide information on when and where the
gravel roads or on hoof, especially when it is hot, could welfare is suboptimal.
cause high mortalities before slaughter in most under- Bruising and foot pad lesions common in cattle in
developed countries. In the case of vehicle transporta- communal production systems results in lameness,
tion, mortalities can be caused by the accretion of wounds, reluctance to eat and drink, thereby increasing
exhausts fumes and road dust in respiratory tract with the risk of acquiring infectious diseases (Gregory, 2007)
subsequent poisoning. This poisoning combined with and subsequently reducing the available meat due to
inadequate escape of gaseous exchange, especially on high mortality rates and trimming off of affected meat
stationary vehicles, can then be large enough to cause portions. In cattle, bruises result due to lack handling
the animal to faint or even die from suffocation before facilities, use of whips and sticks as well as poor res-
even getting to the slaughterhouse (Berg, 2001). training of draught animals or during routine husbandry
Mortalities impose negative implications on the (Ramaswamy, 1998; Gregory, 2007). In addition, bruising
economy of the farm because they result in delayed reduces the economic value of the meat (Grandin,
breeding, reduction in number of offspring, losses when 2000b). Reducing bruising improves animal welfare, so
carcasses are disposed at arrival at the abattoir (Broom livestock handlers need to use and understand principles
and Fraser, 1997) and they also reduce selection of cattle handling, such as visual fields, flight zones and
intensities. Analysis of mortality records are not important their point of balance, flock instincts and loading densi-
in investigating the severity of the welfare methods only ties. Most bruises can be prevented by careful, quiet
but if combined in different regions, they can provide handling, dehorning of calves, fixing broken equipment
useful information for national mortality statistics, eco- and shunning traditional customs of handling animals,
nomic impact and disease surveillance and control (Ortiz- such as animal abuse (aggression, assertion and intimi-
Pelaez et al., 2007). Unfortunately, records or estimates dation) (Lott and Hart, 1977), inappropriate restraint and
of livestock mortality rates in developing countries are confinement and routine husbandry procedures (Mnguni,
lacking but a study of traditional cattle herds in Ethiopia 2006; Gregory, 2007).
reported mortality rates from diseases of 24 months of
age to be 19% (Donaldson, 1996). Calf mortalities in
communal production systems, though not well docu- Loading densities
mented and quantified, are unacceptably high mainly due
to the inevitable competition between the household and Among other regulations for beef cattle transportation,
the calf for milk and colostrums (Roderick et al., 1999; the amount of space provided for animals is one of the
Gregory, 2007). The situation is exacerbated in small most important aspects influencing animal welfare
herds which are characteristic of most smallholder far- (Broom and Fraser, 2007). High or low loading densities,
2
mers than in large herds especially when cows are also such as 600 and 200 kg/m , increases the bruising
used for draught work during the ploughing season scores (Tarrant et al., 1988). In communal production
(Chimonyo et al., 2002). systems in the developing world, high loading densities
Farmers in developing countries are encouraged to are likely to result, due to shortage of vehicles. Carcass
keep records of deaths during handling procedures or in bruise score are elevated above basal levels with
cases of disease outbreaks. Record keeping, eventually increase in loading densities, as illustrated in Table 2.
opens a way for diagnosis and correction of where the Animals with reduced capacity to withstand the rigors of
Ndou et al. 1059
Table 2. The effect of stocking density during 24 h road journeys on plasma constituents and
carcass bruises in Friesian steers.
Stocking density
Plasma constituents
Low Medium High
-1
Plasma cortisol (ng ml ) 0.1 0.5 1.1
-1
Plasma glucose (mmol l ) 0.81 0.93 1.12
-1
Plasma creatinine kinase (units l ) 132 234 367
Carcass bruise score 3.7 5.0 8.5
Values for plasma cortisol, glucose and CK are the difference between the pre- and post-transport values
(Tarrant et al., 1992).
Table 3. Consumable feed production and sustainable stocking areas (Ezanno et al., 2003; Gregory, 2007), selection of
rates in different agro-ecological zones of Africa. breeds with sound walking ability, small-body framed
animals with competing meat quality such as the Nguni
Zone Consumable feed Estimated sustainable cattle breeds (Muchenje et al., 2008a, b, c; 2009c),
(t dry matter per stocking rates
2
Mashona, Tuli and Maasai, raising awareness program-
ha) (TLU per km ) mes through extension officers and implementing long
Arid 0.19 8 term grazing management schemes. The Nguni, Masho-
Semi-arid 0.51 22 na and the Tuli in Southern Africa and their crosses graze
Sub humid 0.72 31 during hot day periods, whilst some of the large-framed
Highland 0.76 33 breeds would spend some of their grazing time in
(Gregory, 2007) shades. Maasai zebu cattle in Kenya walk for up to 16 km
per day during grazing under semi-arid conditions
(Homewood et al., 1987). Large body-framed breeds will
not withstand such grazing distance and the welfare
transport, such as aggressive breeds extensively reared problems can be worsened further in large herds (>100)
and horned animals should be provided with segregated (Ezanno et al., 2003), especially those which are kraaled
space sufficient for the animal to lie down in a normal at night.
position.
bility are associated with the activation of sympathetic- hot weathers should be avoided at all costs.
adrenomedullary system (Tarrant and Grandin, 2000)
and are other forms of autonomic response that are
initiated by acute stressors such as human contact or Biochemical measurements of animal welfare
disturbing situations that need rapid response in farm
animals (Broom, 2000). The normal heart rate for resting Biochemical properties have normal basal levels which
animals is 76.5 beats per minute. Elridge et al. (1988) fluctuate with differences in the severity of the welfare
showed that, the heart rates of cattle during transpor- procedure such that any deviations from normal basal
tation were 15% above those recorded, while the animals levels indicate that some aspect of the situation is
were grazing at pastures. Confinement of animals into aversive. To optimize the welfare of animals, it is neces-
the vehicle therefore, had direct effects on the physio- sary to determine the physiological response in relation to
logical status of the cattle, so there is need for resting the the biochemical changes and products that affect the
animals after transportation so that they return to their meat eating quality especially with extensively kept ani-
basal levels during the pre-slaughter period. In cattle, mals (Muchenje et al., 2009a; Ndlovu et al., 2009b).
heart rate can be used for the assessment of progress or Measurement of metabolites, such as acute phase
well being of animals especially during situations like proteins, hormonal concentrations, blood glucose levels,
intra-specific grooming of young cattle with constant non-esterified fatty acids (NEFA), urea, meat pH and
invasion of its flight zones or during transportation. They glycogen concentrations can be used to monitor beef
can also be used to assess animal welfare in animals that cattle health and welfare status both on the farm and at
are used for draught power (Dube et al., 2000). slaughter (Eckersall, 2000; Chimonyo et al., 2002;
Campo et al., 2008). Although, biochemical assessments
are more expensive and difficult to apply in resource-
Temperature and respiratory rate limited conditions, they are more accurate indicators of
animal welfare than behavioral assessments.
Temperature and respiratory rate can be recorded during
handling or transportation directly through measurements
of rectal temperature and by direct observation, respect- Hormonal levels
tively. Martinez et al. (2006) introduced ingestible pills,
which can be used to monitor the welfare and health Hormones such as cortisol, adrenaline, creatine kinase,
status of cattle under research through records of core dehydrogenase, prolactin, beta-endorphin and gluco-
temperature. Meat quality is associated with the rate of corticoid are good indicators of acute stress experienced
glycolysis and temperature both in the ante and post by animals. These hormones increase substantially when
mortem conditions. For example, Mounier et al. (2006) cattle are exposed to various welfare procedures, such
revealed that, body temperature above normal for bulls as being handled, castrated, feeding, regrouped, trans-
on arrival of the truck at the slaughterhouse was asso- ported and receiving veterinary attention (Corkum et al.,
ciated with greater pH of the longissimus muscle. Since 1994; Boe and Faerevik, 2003; Muchenje et al., 2009b).
climatic conditions of most developing countries are Fluctuations of hormonal levels are important indicators
characterized by hot sub-tropics, temperature of the of the activity of the parasympathetic and sympathetic
animals has to be monitored as a welfare measure, nervous systems, because they result from neuronal
roofed vehicles should be used and transhumance during washout of tissues as the animal tries to cope or respond
Ndou et al. 1061
Breed
Catecholamine
Nguni Bonsmara Angus
Nor-epinephrine (nmol/mmol) 4.3 9.7 6.5
Epinephrine (nmol/mmol) 5.1 10.8 6.7
Dopamine (nmol/mmol) 4.0 14.8 7.2
(Muchenje et al., 2009b).
to welfare procedures. Relationships between changes in mortem lactic acid concentration in the blood system. The
hormonal concentrations and meat quality traits are quite glycogen depletion process may start from the first few
noticeable (Muchenje et al., 2009b) and these can be minutes of confinement, especially to cattle from unso-
used to predict optimum welfare methods suitable for phisticated management such as those of small-scale
attainment of particular levels that are associated with farmers with different social status and continues during
beef quality which meet consumer demands. handling and transportation thus causing high post-
The release of adrenaline and nor-adrenaline due to mortem beef pH (Kannan et al., 2002).
pre-slaughter stress results in depletion of muscle glycol-
gen causing increased meat pH, dark meat (Muchenje et
al., 2008; 2009b) and mobilization of energy which is Glycogen and glucose levels
further amplified with the increase in glucocorticoids
secretions. On the other hand, evidence exist that, dopa- Blood or muscle glycogen concentrations in cattle ranges
mine regulates the cortisol secretion (Ahmadzadeh et al., between 75 and 120 mmol/kg but fluctuate with changes
2006) and glycogen metabolism thereby causing in psychological and physical response (Immonen et al.,
quantifiable effects on tenderness, ageing potential, color 2000; Ndlovu et al., 2009b). Muscle glycogen concen-
and water-holding capacity (Gregory, 2007). Increased trations are the most important factors affecting beef
heart rates may arise from increased movement and quality at slaughter (Immonen et al., 2000). For example,
exercise but rough journeys are more stressful than animals with muscle concentrations that are not within
smooth ones, as demonstrated by elevated plasma corti- the critical threshold of 45 to 57 mmol/kg will not attain
sol (Ruiz-de-la-Torre et al., 2001). the desired ultimate pH (pHu) of 5.5 to 5.6 in meat
Elevation is the concentrations of catecholamines in (Tarrant et al., 1988; Mounier et al., 2006). It can be
urine (Muchenje et al., 2009b, Table 5), saliva and blood suggested that, animals within the normal threshold
(Grandin, 2000b), that can be used as physiological levels should be slaughtered if desirable meat pH is the
indicators of welfare or stress. Secretion of cortisol is target of the producer and this can be achieved by
time-dependent taking approximately 15 to 20 min for it to optimization of welfare procedures.
reach peak value (Lay et al., 1998) and 1 h to return to its Glycogen concentrations and metabolizable energy
baseline level (Veissier and Le Neindre, 1988), during intake are correlated. It is therefore suggested that, im-
this time, dopaminergic mechanisms results in the re- provement of nutrition through supplementing with
lease of -endorphin hormone above basal levels (5 concentrates for 17 days before slaughter as a welfare
ng/ml) (Ladewig and Smidt, 1989). Records on measure- measure can help prevent depletion of glycogen levels
ments of blood -endorphin levels are useful as a backup and reduce the effects of stress in raising pH (Immonen
for adrenocorticotropic hormone (ACTH) or cortisol et al., 2000; Gregory, 2008) and eventually, preventing
measurement in assessing welfare. A rise in ACTH is dark cutting and poor keeping quality. Glycogen concen-
often accompanied with a rise in -endorphin. Moreover, trations influence post-mortem glycolytic rate in ovine
-endorphin is involved in the regulation of various muscle (Ferguson and Warner, 2008) and possibilities of
reproductive hormones. Gregory (2007) suggested that, the same effects are expected in bovine muscles.
records of -endorphin are vital tools to explain the
reduction in reproductive performance during poor wel-
fare conditions. Increase in the number of injuries, such Acute phase proteins
as bone breakage, can contribute to metabolic exhaust-
tion and consequently stimulates the release of creatine Acute phase proteins (APP), is a group of proteins that
kinase (CK) (Elrom, 2001) into the blood system. The CK change in concentration in animals subjected to external
will catalyse the conversion of creatine to phos- and internal challenges, such as infection, trauma, inflam-
phocreatine for energy reservoir in tissues as a response mation or stress (Campo et al., 2008), they act as
to physical stress. This process causes diminished post- inhibitors or mediators of inflammatory processes. Their
1062 Afr. J. Biotechnol.
concentrations can be used to assess and monitor the Africa Union/Inter-Africa Bureau for Animal Resources. Available
online:
livestock welfare and health status (Eckersall, 2000). For
http://www.unsudanig.org/darfurjam/trackII/data/cluster/development/
example, measurement of plasma APP has been used as An%20Audit. Accessed on 6 July 2009.
stress indicators during weaning and transportation of Alana A, Corrie G, Laura L, Natalie S, Taunya V (2008). Maasai.
calves (Arthington et al., 2005), physically stressed cattle Available on:
www.nipissingu.ca/faculty/arohap/aphome/NURS3036/Resources/Ma
(Alsemgeest et al., 1993) and in nutritional trials (Campo
asaiPresentation.ppt
et al., 2008). Once the APP have been assayed, plasma Accessed on the 3rd December 2009.
concentrations of acute phase proteins such as cerulo- Albright JL (2000). Dairy cattle. In: Grandin T (Ed). Livestock Handling
plasmin, fibrinogen (Fb), bovine haptoglobin (Hp) and and Transport Livestock, Second edition. CABI Publishing, New York,
USA.
serum amyloid-A (SAA) in beef animals fluctuate with Ali BH, Al-Qarawi AA, Mousa HM (2006). Stress associated with road
various welfare procedures such that their identifications transportation in desert sheep and goats, and the effect of
are important during meat inspection. pretreatment with xylazine or sodium betaine. Res. Vet. Sci. 80: 343-
The proposed respective acceptable values of bovine 348.
Alsemgeest SP, Taverne MA, Boosman R, Van Der Weyden BC, Gruys
haptoglobin, serum amyloid A and fibrinogen in monitor-
E (1993). Peripartum acute-phase protein serum amyloid-A
ring the calf health status are Hp: 0.13 g/l, SAA: 25.6 mg/l concentration in plasma of cows and fetuses. Am. J. Vet. Res. 54:
and Fb: 6.45 g/l (Ganhaim et al., 2003). Although, there is 164-167.
evidence that APP concentrations act as non-specific Andersen H, Oksbjerg N, Young J, Therkildsen M (2005). Feeding and
meat quality-A future approach. Meat Sci. 70: 543-554.
biosensors of any source of internal or external challenge
Arthington JD, Spears JW, Miller DC (2005). The effect of early weaning
(Gregory, 2007), there is need to link APP concentration on feedlot performance and measures of stress in beef calves. J.
to predict future performance of calves so that un- Anim. Sci. 83: 933-939.
precedented levels may be monitored through handling Aschwanden J, Gygax L, Wechsler B, Keil NM (2008). Social distances
of goats at the feeding rack: Influence of the quality of social bonds,
and welfare of animals. According to Campo et al. rank differences, grouping age and presence of horns. Appl. Anim.
(2008), animals with high APP values also have high pH Behav. Sci. 114: 116-131.
values (48), indicating that stress plays an important role Bech JM, Bennett A, Bouchard R, Condron R, Dabirian S, Dornom H,
in the transformation of muscle to meat. Erlacher-Vindel E, Jorn H, Laura K, Brian L, Cheryl M, Nitya NP, Jorg
S, Girish KS, Girish KMS, Leopold SJ, Gwyneth V (2008). Guide to
good animal welfare in dairy production. International Dairy
Federation, Brussels, Belgium.
CONCLUSIONS Behrends SM, Miller RK, Rouquette Jr. FM, Randel RD, Warrington BG,
Forbes TDA, Welsh TH, Lippke H, Behrends JM, Carstens GE
Holloway JW (2009). Relationship of temperaments, growth, carcass
Animal welfare procedures have aversive result in poor characteristics and tenderness in beef steers. Meat Sci. 81: 433-438.
meat quality that may impose negative economic Berg EP (2001). Running Head: Effect of stress on meat yield and
implications on the beef industry due to failure to meet quality. Influence of stress on composition and quality of meat,
consumer expectations. Creating an environment which poultry, and meat products. Available online:
http://www.fass.org/fass01/pdfs/Berg.pdf. Columbia: Animal Science
minimizes livestock discomfort and enhance their produc- Research Centre. Accessed June 21, 2008.
tivity is important. Furthermore, effective animal welfare Boe KE, Faerevik G (2003). Grouping and social preferences in calves,
assessment techniques will help in developing animal heifers and cows. Appl. Anim. Behav. Sci. 80: p. 175.
husbandry practices that will promote quality animal Broom DM (2000). Welfare assessment and welfare problem areas
during handling and transport. In: Grandin T (Ed). Livestock Handling
products from ethically acceptable production systems. and Transport Livestock, Second edition. CABI Publishing, New York,
Research activities that promote best practices from pp. 43-61.
behavioral, biochemical and physiological assessment Broom DM, Fraser AF (2007). Domestic Animal Behaviour and Welfare,
techniques are therefore encouraged in the developing CAB International, Wallingford, pp. 180-207.
Calhim S, Shi J, Dunbar RIM (2006). Sexual aggression among feral
world. There is need for research that focuses on addres- goats: testing between alternative hypotheses. Anim. Behav. 72: 31-
sing issues surrounding animal welfare during production 41.
of multipurpose cattle, including ritual slaughter. For Campo M, Soares de Lima JM, Brito G, Manteca X, Hernandez P,
successful implementation of the animal welfare aware- Sanudo C, Montossi F (2008). Effect of different feeding strategies on
animal welfare and meat quality in Uruguayan steers. In: Proceedings
ness programmes, governments in developing countries of the 54th International Congress of Meat Science and Technology
have to implement schemes to promote well-being during held at Cape Town, South Africa.
management of livestock. Chambers PG, Grandin T, Heinz G, Srisuvan T (2001). Guidelines for
human handling, transport and slaughter of livestock. Food and
Agriculture Organisation (FAO) Publishers, Thailand.
Chawatama S, Ndlovu LR, Richardson FD, Mhlanga F, Dzama KA
REFERENCES (2003). A simulation model of draught animal power in smallholder
farming systems. Part I: Context and structural overview. Agric. Syst.
Ahmadzadeh A, Barnes MA, Gwazdauskas FC, Akers RM (2006). 76: 415-440.
Dopamine antagonist alters serum cortisol and prolactin secretion in Chimonyo M, Kusina NT, Hamudikuwanda H, Ncube I (2002). Changes
lactating Holstein cows. J. Dairy Sci. 89: 2051-2055. in stress-related plasma metabolite concentrations in working
Aklilu Y (2002). An audit of the livestock marketing status in Kenya, Mashona cows on dietary supplementation. Livest. Prod. Sci. 73:
Ethiopia and Sudan. Vol. 1. Nairobi, Kenya: Community-Based 165-173.
Animal Health and Participatory Epidemiology Unit, Pan African Corkum MJ, Bate LA, Tennessen T, Lirette A (1994). Consequences of
Programme for the Control of Epizootics and the Organization of reduction of number of individual feeders on feeding behaviour and
Ndou et al. 1063
stress level of feedlot steers. Appl. Anim. Behav. Sci. 41: 27-35. Gregory NG (1998). Animal welfare and meat science. CAB
Dean WRJ, Macdonald IAW (1994). Historical changes in stocking rates International, Wallingford, Oxfordshire, UK, p. 298.
of domestic livestock as a measure of semi-arid and arid rangeland Gregory NG (2005). Recent concerns about stunning and slaughter.
degradation in the Cape Province, South Africa. J. Arid Environ. 26: Meat Sci. 70: 481-491.
281-298. Gregory NG (2007). Animal Welfare and Meat Production, second ed.
De Harn C (1995). Development support and livestock services. In CAB International, Wallingford, UK.
Wilson RT, Ehui S and Mack S (eds) Livestock development Gregory NG (2008). Animal welfare at markets and during transport and
strategies for low income countries.FAO, Rome, Italy, pp. 23-27. slaughter. Meat Sci. 80: 2-11.
Donaldson KN (1986). Pastoralism and drought. A case study of the Homewood K, Rodgers WA, Arhem K (1987). Ecology of pastoralism in
Borana of southern Ethiopia. MPhil thesis, University of reading, UK. Ngorongoro Conservation area, Tanzania. J. Agric. Sci. 108: 47-72.
Dube S (2008). Communal rangelands of Eastern Cape Province: Is Immonen K, Ruusunen M, Hissa K, Puolanne E (2000). Bovine muscle
there interaction between resource users, science and policy? A case glycogen concentration in relation to finishing diet, slaughter and
study in South Africa. In: Emergent practice of Adaptive collaborative ultimate pH. Meat Sci. 55: 25-31.
Management in natural resources management in Southern Africa: James B, Hazel B (2007). Rangeland as a common property resource:
Eight case studies. Wageningen International, Wageningen Contrasting insights from communal areas of Eastern Cape Province,
University and Research Centre, Netherlands. South Africa. Hum. Ecol. 35: 97-112.
Dube Z, Ndlovu LR, Muchenje V (2000). The Potential of single-hitched Kannan CB, Chawan CB, Kouakou B, Gelaye B (2002). Influence of
donkeys (Equuas asinus) in cultivation tasks in Zimbabwe. J. Appl. packaging method and storage time on shear value and mechanical
Sci. S. Afr. 6: 45-54. strength of intramuscular connective tissue of chevon. J. Anim. Sci.
Dwyer CM (2009). Welfare of sheep: Providing for welfare in an 80: 2383-2389.
extensive environment. Small Rumin. Res. 86: 14-21. Knowles TG, Warriss PD (2000). Stress physiology of animals during
Eckersall PD (2000). Recent advances and future prospects for the use transport. In: Grandin T, ed. Livestock Handling and Transport. 2nd
of acute phase proteins as markers of disease in animals. Rev. Med. ed. Cambridge, MA: CABI Publishing. pp. 385-407.
Vet. 151: 577-584. Kuzmanvic Z, Elabjer I (2000). Influence of preslaughter resting time on
Elridge GA, Winfield CG, Cahill DJ (1988). Responses to cattle to bulls on meat temperature, colour and pH value. Czech J. Anim. Sci.
different space allowances, pen sizes and road conditions during 45: 511-518.
transportation. Aust. J. Exp. Agric. 28: 155-159. Ladewig J, Smidt D (1989). Behaviour, episodic secretion of cortisol and
Elrom K (2001). Review: Handling and Transportation of Broilers adrenocortical reactivity in bulls subjected to tethering. Horm. Behav.
Welfare, Stress, Fear and Meat Quality. Part VI: The consequences 23: 344-360.
of handling and transportation of chickens (Gallus gallus domesticus). Lammens V, Peeters E, De Maere H, De Mey E, Paelinck H, Leyten J,
Isr. J. Vet. Med. p. 56. Geers R (2007). A survey of pork quality in relation to pre-slaughter
Ezanno P, Ickowicz A, Bocquier F (2003). Factors affecting the body conditions, slaughterhouse facilities, and quality assurance. Meat Sci.
condition score of N’Dama cows under extensive range management 74: 381-387.
in Southern Senegal. Anim. Res. 52:37-48. Lanier JL, Grandin T, Green R, Avery D, McGee K (2001). A note on
Ferguson DM, Warner RD (2008). Have we underestimated the impact hair whorl position and cattle temperament in the auction ring. Appl.
of pre-slaughter stress on meat quality in ruminants? Meat Sci. Anim. Behav. Sci. 73: 93-101.
80:12-19. Lay Jr. DC, Friend TH, Randel RD, Bowers CL, Grissom KK,
Fitzpatrick J, Scott M, Nolan A (2006). Assessment of pain and welfare Nevendorff DA, Jenkins OC (1998). Effects of restrictive nursing on
in sheep. Small Rumin. Res. 62: 55-61. physiological and behavioural reactions of Brahman calves to
Fordyce G, Goddard ME (1984). Maternal influence on the subsequent restraint and nursing. Appl. Anim. Behav. Sci. 56: 109-
temperament of Bos indicus cross cows. In: Proc. Aust. Soc. Anim. 119.
Prod. pp. 345-348. Lawrie RA, Ledward DA (2006). Lawrie’s Meat Science. 7th ed.
Fratkin E (2001). East African pastoralism in transition: Maasai, Boran, Woodhead Publishing, Cambridge, England.
and Rendille cases. Afr. Stud. Rev. 44: 1-25. Lott DF, Hart BL (1977). Aggressive domination of cattle by Fulani
Gauly M, Mathjak H, Hoffman K, Kraus M, Erhardt G (2001). Estimating herdsmen and its relation to aggression in Fulani culture and
genetic variability in tempermental traits in German Angus and personality. Ethos, 5: 174-186.
Simmental cattle. Appl. Anim. Behav. Sci. 74: 109-119. Mahlangu D (2009). ANC joins row over bull-killing ritual. TimeLIVE.
Grandin T (1993). Behavioural acclimation during handling of cattle is Available online http://www.timeslive.co.za/news/article218055.ece.
persistent over time. Appl. Anim. Behav. Sci. 36: 1-9. Accessed on 6 December 2009.
Grandin T (1998). The feasibility of vocalization scoring as an indicator Mapiye C, Chimonyo M, Dzama K (2009a). Seasonal dyanamics,
of poor welfare during slaughter. Appl. Anim. Behav. Sci. 56: 121- production potential and efficiency of cattle in the sweet and sour
128. communal rangelands in South Africa. J. Arid Environ. 73: 529-536.
Grandin T (2000a). Handling facilities and restraint of range cattle. In: Mapiye C, Chimonyo M, Dzama K, Raats JG, Mapekula M (2009b).
Grandin T (Ed). Livestock Handling and Transport Livestock, Second Opportunities for improving Nguni cattle production in the smallholder
edition. CABI Publishing, New York, pp. 103-125. farming systems of South Africa. Livest. Sci. 124:196-204.
Grandin T (2000b). Handling and Welfare of livestock in slaughter Maria GA, Villarroel M, Sanudo C, Olleta JL, Gebresenbet G (2003).
plants. In: Grandin T (Ed). Livestock Handling and Transport, CABI Effects of transport time and ageing on aspects of beef quality. Meat
Publishing, New York, pp. 409-439. Sci. 65: 1335-1340.
Grandin T (2000c). Behavioural principles of handling cattle and other Martinez A, Schoenig S, Andresen D, Warren S (2006). Ingestible pill
grazing animals under extensive conditions. In: Grandin T (Ed). for heart rate and core temperature measurement in cattle. In: Proc.
Livestock Handling and Transport, CABI Publishing, New York, pp. Annu. Int. Conf. of the IEEE Eng. Med. Biol. Soc. pp. 3190-3193
63-85. Minka NS, Ayo JO (2007). Effects of loading behaviour and road
Grandin T (2001). Cattle vocalizations are associated with handling and transport stress on traumatic injuries in cattle transported by road
equipment problems at beef slaughter plants. Appl. Anim. Behav. Sci. during the hot-dry season. Livest. Sci. 107: 91-95.
71: 191-201. Mnguni ME (2006). An investigation into the commercial and the Zulu
Grandin T (2006). Progress and challenges in animal handling and traditional modes of slaughtering, butchering, culinary properties and
slaughter in the US. Appl. Anim. Behav. Sci. 100: 129-139. service with special reference to socioculturasl ritual behaviors in
Grandin T (2007). Introduction: effect of customer requirements, KwaZuluNatal. Available on http://ir.dut.ac.za/bitstream/handle/
international standards and marketing system structure on the 10321/77/Mnguni_2006.pdf?sequence=40.
handling and transport of livestock and poultry. In: Grandin T (ed). Accessed on the 6 December 2009.
Livestock Handling and Transport, 3rd edn. CAB International, Mounier L, Dubroeucq H, Andanson S, Veissier I (2006). Variations in
Wallingford, UK. meat pH of beef bulls in relation to conditions of transfer to slaughter
1064 Afr. J. Biotechnol.
and previous history of the animals. J. Anim. Sci. 84: 1567-1576. Ruiz-de-la-Torre JL, Velarde A, Diestre A, Gispert M, Hall SJG, Broom
Moyo B, Masika PJ (2009). Tick control methods used by resource- DM, Manteca X (2001). Effects of vehicle movements during
limited farmers and the effect of ticks on cattle in rural areas of the transport on the stress responses and meat quality of sheep. Vet.
Eastern Cape Province, South Africa. Trop. Anim. Health Prod. 41: Res. 148: 227-229.
517-523. SAPA (2009). Judgment on Zulu bull killing expected Friday. News.
Muchenje V, Dzama K, Chimonyo M, Raats JG, Strydom PE (2008a). South Africa. In Time LIVE. Available on
Tick susceptibility and its effects on growth performance and carcass http://www.timeslive.co.za/news/local/article216261.ece. Accessed
characteristics of Nguni, Bonsmara and Angus steers raised on on 3 December 2009.
natural pasture. Animal, 2: 298-304. Silva JA, Patarata L, Martins C (1999). Influence of ultimate pH on
Muchenje V, Dzama K, Chimonyo M, Raats JG, Strydom PE (2008b). bovine meat tenderness during ageing. Meat Sci. 52: 453-459.
Meat quality of Nguni, Bonsmara and Angus steers raised on natural Smith DG, Pearson RA (2005). A review of the factors affecting survival
pasture in the Eastern Cape, South Africa. Meat Sci. 79: 20-28. of donkeys in Semi-arid regions of Sub-Saharan Africa. Trop. Anim.
Muchenje V, Dzama K, Chimonyo M, Strydom PE, Hugo A, Raats JG Health Prod. 37: 1-19.
(2008c). Sensory evaluation and its relationship with physical meat Smith GC, Pendell D L, Tatum JD, Belk KE, Sofos JN (2008). Post-
characteristics of beef from Nguni and Bonsmara steers raised on slaughter traceability-review. Meat Sci. 80: 66-74.
natural pasture. Animal, 2: 1700-1706. Strappini AC, Metz JHM, Gallo CB, Kemp B (2009). Origin and
Muchenje V, Dzama K, Chimonyo M, Strydom PE, Hugo A, Raats JG assessment of bruises in beef cattle at slaughter. Animal, 3: 728-736.
(2009a). Some biochemical aspects pertaining to beef eating quality Tarrant V, Grandin T (2000). Cattle transport. In: Grandin T (ed).
and consumer health: a review. Food Chem. 112: 279-289. Livestock Handling and Transport Livestock, Second edition, CABI
Muchenje V, Dzama K, Chimonyo M, Strydom PE, Raats JG (2009b). Publishing, New York, USA.
Relationship between pre-slaughter stress responsiveness and beef Tarrant PV, Kenny FJ, Harrington D (1988). The effect of stocking
quality in three cattle breeds. Meat Sci. 81: 653-657. density during 4h transport to slaughter on behaviour, blood
Muchenje V, Hugo A, Dzama K, Chimonyo M, Raats JG, Strydom PE constituents and carcass bruising in Friesian steers. Meat Sci. 24:
(2009c). Cholesterol levels and fatty acid profiles of beef from three 209-222.
cattle breeds raised on natural pasture. J. Food Comp. Anal. 22: 354- Tarrant PV, Kenny FJ, Harrington D, Murphy M (1992). Long distance
358. transportation of steers to slaughter: effect of stocking density on
Munksgaard L, DePassille AM, Rushen J, Herskin MS, Kristensen AM physiology, behaviour and carcass quality. Livest. Prod. Sci. 30: 223-
(2001). Dairy cows’fear of people:social learning, milk yield and 238.
behaviour at milking. Appl. Anim. Behav. Sci. 73: 15-26. The Citizen 2009. Bull killing saga won't end after Friday judgment:
Napolitano F, Cifuni GF, Pacelli C, Riviezzi AM, Girolami A (2002). expert. News. South Africa. Available on
Effect of artificial rearing on lamb welfare and meat quality. Meat Sci. http://www.thecitizen.co.za/index/home.aspx?pDesc=2,1,27.
60: 307-315. Accessed on 4 December 2009.
Ndlovu T, Chimonyo M, Muchenje V (2009a). Monthly changes in body Veissier I, Le Neindre P (1988). Cortisol responses to physical and
condition scores and internal parasite prevalence in Nguni, Bonsmara pharmacological stimuli in heifers. Reprod. Nutri. Dev. 28: 553-562.
and Angus steers raised on sweetveld. Trop. Anim. Health Prod. 41: Vessier I, Butherworth A, Bock B, Roe E (2008). European approaches
1169-1177. to ensure good animal welfare. Appl. Anim. Behav. Sci. 113: 279-
Ndlovu T, Chimonyo M, Okor AI, Muchenje V, Dzama K, Dube S, Raats 297.
JG (2009b). A comparison of nutritionally-related blood metabolites Voisinet BD, Grandin T, Tatum TD, O’Connor SF, Struthers JJ (1997a).
among Nguni, Bonsmara and Angus steers raised on sweetveld. Vet. Feedlot cattle with calm temperaments have higher average daily
J. 179: 273-281. gains than cattle with excitable temperaments. J. Anim. Sci. 75: 892-
Novak P, Vokralova J, Kunc P (2004). Welfare conditions of dairy cows 896.
from the point of performance efficiency. Research Institution of Voisinet BD, Grandin T, Tatum TD, O’Connor SF, Tatum JD, Deesing
Animal Production Prague, Czech Republic. MJ (1997b). Bos indicus-cross feedlot cattle with excitable
Ortiz-Pelaez A, Pritchard DG, Pfeiffer DU, Jones E, Honeyman P, temperaments have tougher meat and a higher incidence of
Mawdsley JJ (2007). Calf mortality as a welfare indicator on British borderline dark cutters. Meat Sci. 46: 367-377.
cattle farms. Vet. J. 176:177-181. Warris PD (1990). The handling of cattle pre-slaughter and its effects on
Petherick JC, Holroyd RG, Doogan VJ, Venus BK (2002). Productivity, carcass and meat quality. Appl. Anim. Behav. Sci. 28: 171-186.
carcass and meat quality of lot-fed Bos indicus cross steers grouped Warris PD, Brown SN, Adams SJM (1994). Relationship between
according to temperament. Aust. J. Exp. Agric. 42: 389-398. subjective and objective assessment of stress at slaughter and meat
Ramaswamy NS (1998). Draught animal welfare. Appl. Anim. Behav. quality in pigs. Meat Sci. 38: 329-135.
Sci. 59: 73-84. West JW, Mullinix BG, Bernard JK (2003). Effects of hot, humid weather
Roderick S, Steveson P, Ndungu J (1999). The production parameters on milk temperature dry matter intake, and milk yields of lactating
influencing the composition and structure of pastoral cattle herds in a dairy cows. J. Dairy Sci. 86: 232-242.
semi-arid area of Kenya. Anim. Sci. 68: 201-209.