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Received: 22 August 2018    Accepted: 25 August 2018

DOI: 10.1111/rda.13335

REVIEW ARTICLE

Ethics in animal breeding

Wenche Farstad

Department of Production Animal

Clinical Sciences, Faculty of Veterinary Contents
Medicine, Norwegian University of Life Ethical breeding involves the use of healthy animals true to their species in behaviour
Sciences, Oslo, Norway
and physical appearance, and when applicable, showing a sustainable performance.
Correspondence The concerns for the species/breed are essential parts of the breeding goals, includ-
Wenche Farstad, Department of Production
Animal Clinical Sciences, Faculty of ing preservation of genetic resources within the species/breed, and the health and
Veterinary Medicine, Norwegian University welfare of the individual animal. Ethical and welfare considerations were often not
of Life Sciences, Oslo, Norway.
Email: [email protected] prioritized in developing new breeds of production or companion animals. As a result,
animal breeding practices are increasingly becoming part of the debate on animal
welfare. In companion animals, breeding for curiosity or “cuteness” may be a goal in
itself, although dogs are also bred for utility. In production animals, breeding focus is
on performance, i.e., quantitative entities and financial income, rather than physical
appearance. For instance, dairy cows are bred to be larger and to have higher milk
yields, sows and ewes to produce more offspring, and horses are designed for riding,
racing, and companionship. Overbreeding in relation to current demand of horses,
cats, and dogs raises welfare issues due to abandonment or killing of horses and mil-
lions of cats and dogs every year. There is variable regulation of health requirements
for breeding animals in different countries of the world. In many countries, consum-
ers are becoming increasingly aware of animal welfare issues such as negative effects
of certain production traits in farm animals, leading to decreased demand for their
meat at a time where increased food production is becoming crucial. Amidst these
dilemmas are the veterinarians. This paper deals with issues connected to traditional
breeding as well as some of the breeding technologies, and includes food safety, eth-
ics, and animal welfare.

KEYWORDS
animal breeding, animal welfare, breeding goals, companion animals, ethics, production
animals

1 | I NTRO D U C TI O N other humans, especially children. Similar core values also exist in
our relationship to animals, with most humans displaying strong
Ethics is a set of moral principles that governs a person’s behaviour empathy with animals, especially with those species, to which
or the conducting of an activity. The moral code of a person or a they are culturally and emotionally attached (Merz-­Perez, Heide,
population is culturally defined, and may depend on religion, pol- & Silverman, 2001). This equates to treating animals we own well,
itics, or nationality (Crony & Millman, 2007). One may argue that so they can serve humans better (utilitarian principles); treating
there is such a phenomenon as core human values or ethics, which them well because we have been given responsibility for them
may be common to all humans irrespective of cultural background, (“parent” principle); or treating animals well because animals have
with most individuals being strongly against hurting or mistreating a value of their own (intrinsic value) and therefore have certain

4  |  © 2018 Blackwell Verlag GmbH wileyonlinelibrary.com/journal/rda Reprod Dom Anim. 2018;53(Suppl. 3):4–13.
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rights (right based). Hence, ethics of animal husbandry including Breeding is normally performed by natural service in nature, in
animal breeding may be about treating animals so that the animals companion animals and sometimes in livestock. However, assisted
do not suffer, treating them well in order to increase our benefits breeding techniques, such as artificial insemination (AI), in vitro fer-
from keeping them, or because they have a right to have a good tilization, and embryo transfer, are generally accepted by the public
life. Irrespectively, all these approaches claim that we treat ani- and have gained an important role in the last decades in many spe-
mals in such a way that their health and welfare are not compro- cies, especially in cattle, in Europe, the US, and many other coun-
mised during their lifetime. Animal welfare is defined by the World tries. Assisted breeding techniques may also involve manipulation
Organization for Animal Health (OIE) (2013) as “how an animal is of gametes ex vivo. Interventions such as intracytoplasmic sperm
coping with the conditions in which it lives.” injection (ICSI), embryo transfer or cloning by nuclear transfer may
A good definition of breeding per se is not easy to find, although raise ethical concerns.
most people will have a general understanding of the meaning of In the following review paper, some of the ethical issues of
the term animal breeding as the deliberate sexual reproduction of breeding in companion and production animals (including fish) will
animals. Definitions in dictionaries differ but none includes or elab- be presented. Society’s expectations will be discussed, and the role
orates aspects of strategic breeding: i.e., the activity of keeping and of the veterinarian in breeding ethics will be highlighted.
caring for animals in order to produce more animals of a particular
kind or with particular attributes. Strategic breeding involves the
specific selection of parent animals and sometimes includes assisted 2 | B R E E D I N G O F S P O RT S -­A N D
breeding (Cambridge English Dictionary; Merriam Webster and CO M PA N I O N A N I M A L S : D O G S , C AT S , A N D
Oxford Dictionaries). HORSES
From a veterinary perspective, breeding of companion animals
is mainly performed by breeders, who produce a limited number When we want to buy a canine companion, we can choose from
of offspring. In livestock and fish, however, breeding can be con- 350 to 400 different dog breeds. Often, in companion animals,
sidered being part of an extensive industry, aiming to have a high breeding for curiosity or for “cuteness” may be a goal of some
number of offspring output. Task of the veterinary community is to breeders in itself. “Overtyping” is the deliberate breeding of par-
address limitations and animal welfare issues in both types of breed- ticular attributes that usually bear a negative impact on health
ing enterprise. and welfare, and which in the past often led to a significant nar-
When combining the two terms “ethics” and “animal breeding”, rowing of the genetic variation of a population. Positive selec-
the meaning is deliberate production of animal offspring, which in- tion pressure is generally put on those individuals that show the
volves the conscientious consideration of whether it is morally cor- most extreme expression of desired traits, while those individuals
rect or acceptable to produce the intended offspring. Hence, ethical that do not possess these traits are discarded as breeding ani-
breeding needs to involve “the use of healthy animals true to their spe- mals, leading to a reduction in genetic diversity (Fredholm, 2017).
cies in behaviour and looks, and when applicable, showing a sustainable Most modern dog breeds have challenges concerning inherited
performance” (Olsson, Gamborg, & Sandoe, 2006]. defects and diseases, since almost 700 hereditary diseases are
Animal welfare is closely associated with animal breeding, since registered in dogs according to Online Mendelian Inheritance in
in many ways we use animals to fulfil the human demands for com- Animals (OMIA). In this catalogue/compendium of inherited dis-
panionship, beauty, entertainment, research, utility, protection, orders, information on single-­locus traits and genes in 239 animal
food, and financial income. Humans have to make ethical consider- species are collected.
ations when they define breeding goals based on a balance between The Labrador has been the most popular dog in the UK for
our demands and the animals’ requirements. Any domestication af- almost three decades, but according to The Kennel Club (UK) by
fects animal behaviour, which may vary among the different species. the end of 2018 its top place could be challenged by the brachy-
Adaptation to a human environment will occur to a varying degree. cephalic French Bulldog. Kennel Club figures reveal that reg-
Animal breeders often select their breeding subjects based on istrations for this breed rose by 47% from 14,607 in year 2007
specific attributes, aiming to reach individually defined excellence to 21,470 in 2015/2016. The rapid rise of the popularity of the
in production traits, physical appearance, or performance, regard- French Bulldog seems to be based on a combination of celebrity
less whether they breed alpacas, hens, cats, or cows. The reasons impact, commercials, and exposure on social media. As reported
for breeding animals are numerous and differ depending on the by the BBC correspondent Claire Marshall in 2017, “They’ve be-
type of animals. Often financial aspects play an important role. come a sort of fad – but veterinary experts say that the craze for these
Unfortunately, physical, and functional soundness are not always ‘designer’ dogs is helping to create dogs with painful deformities”. The
of high priority when they design new types, for example, in a new major problem is that many brachycephalic dogs have difficulties
cat breed or a high yielding dairy cow. As a result, animal breeding in breathing, and panting to cool down in high ambient tempera-
practices and the outcomes of breeding have become part of the tures is difficult. This is linked to developing upper respiratory
debate dealing with issues of animal welfare and the responsibility tract disorders, the brachycephalic obstructive airway syndrome
of breeders and veterinarians. (BOAS), because of their short skulls, extreme flat short noses,
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and narrow lanynges, (Koch, Arnold, Hubler, & Montavon, 2003; are significant (Chastant-­Maillard, Guillemot, Feugier, Mariani, &
O’Neill et al., 2015). The same problem occurs in brachycephalic Mila, 2017; Fournier et al., 2017). High fertility and high demand
cat breeds, e.g., Persian cats (Schlueter et al., 2009). make dog-­ and sometimes cat-­breeding very profitable in terms of
high puppy prices for popular breeds. However, overproduction of
undesired animals may lead to crowded animal shelters and eutha-
2.1 | Dogs
nasia or killing of millions of cats and dogs every year due to loss
of interest or for economic reasons. Consequently, regulations on
2.1.1 | Conventional breeding
overbreeding and selling by third parties via so called “Puppy Mills”
Conventional breeding or natural breeding is for many dog breeders and illegal import have now been introduced in some countries, such
and the public the most preferable way to produce offspring, since as the United Kingdom, the USA, and Australia, although the busi-
it allows normal sexual behaviour and interaction between males ness of puppy mills is yet not under control. The rapid growth of
and females. Sexual behaviour in such a context may elicit positive the internet and social media offering direct contact with the buyers
emotions, which is an important aspect of animal welfare (Mellor, may have worsened the situation. The lack of regulation of the trade
2015). However, if breeders force mating or use a male much larger of pedigree dogs, including specific requirements for health certif-
or smaller than the female, this may not be in the best interest of icates, is of increasing concern. Therefore, international directives
the bitch, it may impede good socialization of the animals, and may should be introduced to both to legitimate responsible breeders and
interfere with natural behaviours which consecutively may hamper to address the concerns of the public.
a successful natural mating. This may lead to the wish for assisted
breeding, i.e., artificial insemination.
2.1.3 | Artificial and assisted breeding
The welfare aspect of a bitch needing to experience pregnancy,
which is often argued by owners of family dogs, is not scientifically Artificial insemination (AI) is the procedure of instrumentally in-
proven. However, in the discussion of the significant effect of posi- serting semen collected from a male into the female’s reproductive
tive emotions on animals welfare, the effect of experiencing mater- tract. In dogs, the use of AI has increased steadily and the prepara-
nal care cannot be entirely disregarded (Mellor, 2015). tion of fresh, fresh chilled, and frozen-­thawed semen increasingly
Some breeds have developed a predisposition for dystocia or supports breeding and sharing genes between countries and conti-
other periparturient diseases. Typically, this concerns the brachy- nents (Payan-­C arreira, Miranda, & Niżański, 2011) The use of AI to
cephalic breeds and breeds, such as the Scottish Terrier or Collie, overcome physical or mental defects or lack of libido in dogs that
where foetal heads are relatively wide (brachycephalic breeds) or preclude natural sexual behaviour or mating may not be acceptable
long (Scottish Terrier, Collie), and the pelvis of the dam has a small if the defect is considered hereditary. However, for most of the con-
(brachycephalic) or triangular diameter (Scottish terrier). These ditions that prevent natural mating scientific data on the heritability
breeds have an increased risk of dystocia and consequently assisted is scarce.
delivery or Caesarean section (Bergström, Nødtvedt, Lagerstedt, &
Egenvall, 2006; O’Neill et al., 2017). The study by O’Neill et al. (2017)
2.1.4 | Cloning (somatic cell nuclear transfer)
confirmed the welfare concerns related to BOAS as well as other
health problems described earlier to be common in these breeds The cloning of dogs was first reported in 2005 by Lee et al. (2005)
(Koch et al., 2003; Pratschke, 2015), and highlighted the need for and is reviewed in the current issue of this journal entitled “Dog clon-
breeders to address these issues when selecting dogs for breeding. ing – no longer science fiction” by Lee et al. (2018). Although the
pregnancy rate and number of live born puppies is below that oc-
curring after natural service and AI, a 50% full term pregnancy rate
2.1.2 | Fertility and overbreeding
and 5%–6% survival rate to term of cloned embryos was reported
Except for working dogs, i.e., sniffer dogs, police and military guard by Jeong et al. (2016). The health issues of cloned offspring at birth
dogs, hunting dogs, and dogs bred for human assistance (guide dogs), described in some of the larger domestic animal species have yet
the vast majority of domestic dogs are kept as companion animals. not been reported in dogs so far. However, more research by more
In working dogs, where high performance in working tasks is essen- research groups is warranted. Cloning of dogs was initially devel-
tial, assortative breeding is often used, a selection strategy based oped to create individuals that could be used as medical research
on animal performance or an expectation of performance (Bourdon, models, since dogs and humans have many genes involved in disease
2000). development in common, and for special tasks, such as rescue dogs
The market of breeding dogs is regulated by supply and demand, and sniffer dogs. The ethical aspect of dog cloning for these pur-
and is influenced by popularity as well as the wish and request for poses has so far not been questioned very much by the scientific
companion or working dogs. Two large data surveys conducted on community or the public. In addition, cloning of individual pets has
feline and canine fertility were recently published, showing that fer- so far not become a much sought for service due to the high costs.
tility in cats and dogs is high in terms of pregnancy rate, birth rate, In the future, more debates concerning the ethics of animal cloning
litter sizes and low abortion, and stillbirth rates, but breed variations may arise.
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TA B L E   1   The Modified ethical matrix is using dog as an example, but can be applied to all animals

Wellbeing (W) Autonomy (A) Fairness (F)

Modified ethical matrix: Translation of the ethical principle s for the corresponding interest group (England & Millar, 2008)
Dog (D): stud, dam and Dog welfare (DW) Behavioural freedom dog(DA) Intrinisic value of dog (DF)
pups
Breeder (B) Satisfactory income and working conditions (BW) Mangerial freedom and Fair regulation and trade
creativity (BA) (BF)
Owner(O) Safety and quality of one’s life (OW) Freedom of choice (OA) Affordability of animal
products (OF)
Veterinarans (V) Satisfactory income and working conditions (VW) Professional freedom (VA) Equitable standards of
practice (VF)
Society (S) Safety and social harmony (SW) Democratic choice (SA) Fair resource allocation
(SF)

From the breeders point of view, the decision to breed an individual animal is purpose, why one wants to breed an animal. Then breeding decision must
be taken based on risk assessment of compromising the dam’s welfare, the welfare of the offspring and the quality of life of the offspring. Secondly,
how to breed, natural service or assisted breeding techniques, economic cost. Thirdly, the consequences for the parties involved: first of all, the indi-
vidual animal, then the breeder, the new owner, the veterinarians and society. Affordability of animals products, and economic cost and fair resource
allocation are important factors for society when considering breeding ethics in production animals. (Reproduced with written persmisssion from the
authors England and Millar (2008)).

When conducting an ethical analysis on each of the artifi- cat breeds, physical mutations that were previously allowed to perish,
cial breeding techniques in connection with the welfare of dogs, are now being developed merely for the sake of difference. Not all are
England and Millar (2008) introduced a modified ethical matrix. This harmful, but some are achieved at considerable cost to the cat.” One
may be used to assess the concept of wellbeing, autonomy, and fair- example of a controversial cat type are twisty cats, also known as
ness of a particular action such as breeding, AI, or other Assisted squittens or kangaroo cats (Leary, 2016). These cats are born with
Reproduction Techniques (ARTs) or treatments to the correspond- short forelegs that are the result of conditions such as radial hypo-
ing interest group. This matrix is useful to assess the ethical aspects plasia, radial aplasia, radial agenesis, or foreleg micromelia. Due to
of a particular intervention and is applicable irrespective of species their short forelegs, they often sit in an upright posture similar to a
(Table 1). kangaroo or squirrel (Hartwell, 2012). Another curiosity is the nude
cat, which has no body hair except whiskers. These cats need special
care due to their lack of fur in cold climates and protection against
2.2 | Cats
the sun in warmer climates. Persian cats are in many ways similar to
Cats have traditionally been a natural companion animal in rural brachycephalic dogs with their large round eyes, flat faces, and short
areas where they are used to control rodent numbers. Compared noses, making them susceptible to traumatic keratitis, dyspnea, and
to dogs, breeding cats for specific traits is less common although dystocia. Hence, many of the designed cat breeds will need special
cats are also used as a model animal in biomedical research. The care and medical attention.
Researchers at the University of Texas produced the first feline
clone named “Copy Cat” (Shin et al., 2002); although in common with
2.3 | Horses
other species, the efficiency was as low. There is no documentation
in the scientific literature or public media that the welfare of cloned In Western countries, the focus in horse breeding is usually not on
cats (and dogs), neither the donors nor recipients, is compromised in meat quality, but rather on sports-­or working performance, e.g., rac-
a way it is in livestock, such as cattle and pigs. However, the principle ing, pulling, riding, or herding. As other companion animals, horses
of cloning and the unforeseen future consequences of breeding by are subjected to selective breeding for traits that humans consider
cloning in any species led to a legal ban on somatic cell nuclear trans- desirable, such as extreme sizes. In some regards, the breeding for
fer (SCNT) cloning in 1997 in Norway, whereas in other countries originality or curiosity also plays a role in horses, as it does in smaller
such as The USA, The UK, and South Korea, it is legal in animals, but companion animals.
not in humans.
Nowadays, cats are valued pet animals in both urban and rural
2.3.1 | Breeding exceeds demand
areas, and they have a high fertility rate. A variety of domestic cat
breeds and domestic and wild hybrids have been bred, of which not Overbreeding in relation to demand creates welfare issues of horse
all are well adapted to life with humans. Roger Tabor (UK), biolo- abandonment and/or killing for example in the UK. Unwanted horses
gist and author of “The Rise of the Cats” (Tabor, 1995) stated: “In are often rehomed, but frequently also left to themselves, or are
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euthanized, because of the high cost of keeping, feeding, and car- embryonic cloning may possibly be painful to donor mares as sug-
ing for them, sometimes due to injuries or old age, or because of a gested by Campbell and Sandoe (2015) and Campbell (2018). This
change in the life status of the owner. Horsemeat is considered ac- points to possible welfare concerns and ethical considerations in the
ceptable for human consumption in some countries and, therefore, equine embryo technology industry.
horses are slaughtered in abattoirs. In countries, such as the UK
and the USA, where horses are considered not suitable for human
consumption, a system of slaughtering and processing for other 3 | B R E E D I N G O F PRO D U C TI O N A N I M A L S
use, e.g., in dog food, may exist (Montpoli,2013). Abandonment and
neglect, as a result of overbreeding, has been identified by various In production animals, the breeding focus is on production perfor-
equine charities as a major welfare issue (e.g., World Horse Welfare, mance, i.e., quantitative entities, yield, (meat or milk, eggs or fleece)
2013). At the end of 2007, a number of 11,476 horses were in the rather than physical appearance. Some cattle breeds are bred to
care of member charities of the National Equine Welfare Council in reach very high production levels (Lucy, 2016). As an example, one
the United Kingdom. Similar to dogs and cats, high demand makes Wisconsin Holstein cow set The United States of America national
horse breeding profitable, but overproduction leads to the slaughter milk annual production record of 35,000 kg of milk, whereas the
of thousands of horses every year. actual production average for all the USA Holstein herds in 2014
was 11,318 kg of milk. A yearly increase of 12% in annual milk yield
rate per cow has been recorded the last 10 years (2008–2017) ac-
2.3.2 | Breeding practices
cording to The USA Department of Agriculture National Agricultural
According to Campbell and Sandoe (2015) welfare problems re- Statistics Service (2018). Consumers are becoming increasingly
lated to breeding, for example by natural mating after long road aware of the welfare issues of modern breeding and keeping prac-
or air transport, or by the application of artificial reproduction tices. These concerns have led to a decrease in demand for conven-
techniques (ARTs), have been given little attention in the academic tional farm products and an increase in demand for organic food
literature. Potential welfare issues are associated with breeding, products in some countries.
regardless whether the technique used is “natural” or “artificial”.
This relates to reasons, such as selecting inappropriate breeding
3.1 | Cattle
animals, excessive frequency of breeding, and health issues re-
lated to parturition.
3.1.1 | Dairy cattle
High yielding cows have been bred because of their large genetic
2.3.3 | Artificial insemination and embryo transfer
potential for increased milk production and current herd manage-
Transport of breeding animals for long distances is common in horse ment strategies focus on “high income over feed costs” (IOFC). This
breeding. Artificial or assisted reproductive techniques, such as AI development led to higher requirements in terms of food quality
and oocyte retrieval, may help to overcome some of the disadvan- and fertility management. For example, impaired ovulation and si-
tages of live animal transport, and also to prevent the spread of lent oestrus led to the increased use of hormonal treatments. One
equine contagious diseases transmitted by direct contact. Campbell of the suggested regimens for programmed ovulation is the Double
and Sandoe (2015), nevertheless, point to research indicating that Ovsynch protocol, which includes as many as four GnRH and two
oocyte retrieval by ovum pickup may cause discomfort to the mare, PGF2a injections per female to induce ovulation that can be used
highlighting possible welfare concerns of ART in horses. for timed AI (Souz, Ayres, Ferreira, & Wiltbank, 2008). From an
animal welfare point of view this may not be acceptable, and there
is a risk of reducing the ability to ovulate naturally since ability of
2.3.4 | Cloning
spontaneous ovulation is not part of a deliberate genetic selection
In equine reproduction, production of embryos using somatic cells as anymore (Lucy, 2016). The concern is that timed AI programmes
the source of nuclear transfer is nowadays both a research tool and that are commonly used in the United States and other parts of
a clinical service (Hinrichs, 2012). Although the number of cloned the world may mask infertility by enabling otherwise infertile cows
foals was small in 2012 (estimated at 100–200 worldwide; Hinrichs, (non-­cyclers, etc.) to become pregnant. These infertile cows may
2012), Johnson, Clark-­Price, Choi, Hartman, and Hinrichs (2010) re- have a high daughter pregnancy rate (DPR) because they received a
ported that only 26% of cloned embryos resulted in the birth of a timed AI and become pregnant early post-­partum, sometimes even
live foal. with sexed semen. In consequence, the reproductive system of the
Possibly because there is not the same public concern about the offspring may not function independently of external intervention
possible negative effects on human health when consuming food (Lucy, 2016). Although pregnancy rates have increased in Holsteins
products derived from cloned animals, such as cattle and pigs (EFSA, (Norman, Wright, Hubbard, Miller, & Hutchison, 2009), they are sig-
2012), data about the health and welfare of equine clones is scarce. nificantly lower than in cattle breeds that produce less milk. For ex-
The hormonal treatment for superovulation and ovum pickup in ample, the dual purpose cattle Norwegian Red averages 8,000 kg/
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year with a Non-­Return Rate (NRR) on day 56 of 72% and a calving respiratory failure, and limb malformations. In cattle and other
rate of 65% (Holstein cows that produce an average of 11,300 kg ruminants, these abnormal phenotypes are referred to as the
(23,000 pounds) of milk annually, (Holstein, 2018, USA; The USA large offspring syndrome (LOS) (Young, Sinclair, & Wilmut, 1998).
Department of Agriculture National Agricultural Statistics Service Improper development of the placenta may play a major role in
(2018) show NRR percentages as low as 35%–40% (Fouz, Gandoy, foetal abnormalities. It has been suggested that the pathological
Sanjuán, Yus, & Diéguez, 2011). The consequence of strict selection phenotypes in the placental and foetal development of clones are
for increased milk production in Holstein dairy cows has been a very associated with abnormal reprogramming by the host ooplasm of
noticeable reduction in reproductive performance (Berry, Friggens, the donor cell used for nuclear transfer (Humphreys et al., 2001;
Lucy, & Roche, 2016), and for this reason selection of reproductive Rideout, Eggan, & Jaenisch, 2001; Suzuki et al., 2009). Abnormal
efficiency and daughter fertility has been emphasized in Holstein offspring in some domestic animal clones entails a cost of cloning
breeding programmes since 2010. A consideration of a combination to the animal, the species (increased risk of reduced genetic di-
of milk yield and health parameters in the context of breeding is nec- versity), the breeder, as well as the veterinarian and society that
essary to prevent such unwanted effects on cow health and welfare are concerned with the health and welfare of the clones and their
(Berry, 2017). offspring (EFSA,2012).
Several terms describe genetically engineered animals: ge-
netically modified, genetically altered, genetically manipulated,
3.1.2 | Beef cattle
genome edited, transgenic, and biotechnology-­d erived, among
Many beef cattle breeds are bred to produce a maximum of daily others. In the early stages of genetic engineering, the primary
weight gain. Some breeds have special attributes that make them at- technology used was transgenesis, literally meaning the trans-
tractive on the beef market, but some traits are also associated with fer of genetic material from one organism to another (Waigmann
health problems. The probably most referred to breed in the context et al., 2012). However, with advances in the field, new technolo-
of animal welfare issue, is the Belgian Blue. Its double muscling is gies have emerged which do not necessarily require transgenesis:
due to a mutation of the myostatin gene, which codes for the protein recent applications allow for the creation of genetically engi-
myostatin that inhibits muscle development (Kambadur, Sharma, neered animals via the gene editing (CRISPR) insertion or deletion
Smith, & Bass, 1997). This mutation also interferes with fat deposi- of genes, or the manipulation of genes already present (Ormandy,
tion, resulting in very lean meat and high protein value. The meat, Dale, & Griffin, 2011; Sander & Joung, 2013). Clustered regularly
therefore, has an extraordinary tenderness and taste, which makes interspaced short palindromic repeats (CRISPR) was used to pro-
it attractive on the beef market. However, myostatin mutations can duce two Holstein calves without horns in Minnesota USA in 2016
affect a number of different body systems, and have a negative im- (Carlson et al., 2016). Gene direction can be used to increase dis-
pact on the welfare of cattle. The most widely known symptom in ease resistance and other attributes that may be of advantage.
Belgian Blues is dystocia and the extraordinarily high rate of cae- However, there is a potential for altering genes that may compro-
sarean sections due to the large muscle mass on the back end of mise the intrinsic value of cows, such as insertion of genes from
the calf (West, 1997). Furthermore, problems with deformations of other species, or creating traits that are more beneficial to produc-
the jaw and over-­enlarged tongues, which affect the calf’s ability to tion profit than to the cows themselves (Eriksson, Jonas, Rydhmer,
suckle and hence its survival, as well as respiratory, heart, and re- & Röcklinsberg, 2018). In these cases, insertion of genes may be
productive problems, have all been documented in the Belgian Blue considered unethical or controversial, opening up issues relating
breed. The welfare issues related to the mutations have prompted to animal integrity and/or dignity (Ormandy et al., 2011).
veterinarians to discourage crossing these breeds with other double
muscled breeds.
3.2 | Swine

3.1.3 | Artificial and assisted breeding 3.2.1 | Sustainable breeding goals

Cloning and genetic engineering Most pigs are bred for meat production, a smaller number are bred
While cloning and genetic engineering are only briefly dealt with in either for research purposes or for companion animals (minipig).
this review, it is important to note that these new technologies are These three categories of production have their own challenges and
becoming increasingly prominent in modern assisted breeding pro- have different breeding perspectives and ethical issues.
cedures in livestock (Hasler, 2014; Perry, 2017).
The procedure of cloning by SCNT in most mammals results
3.2.2 | Ethical issues in industrial pig breeding
in pregnancy rates much lower than those obtained in vivo after
insemination and from transfer of embryos derived in vitro (Kruip Litter size in pigs varies between breeds, is regulated by ovulation
& den Daas, 1997). Furthermore, in some species a significant rate, embryo survival, and uterine capacity, and is an important eco-
proportion of cloned foetuses that survive to term have disorders nomic factor in swine production. While selection for increased litter
such as oversized organs, increased or decreased overall growth, size yields more born piglets, large litters are also characterized by a
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higher risk of intrapartum and postpartum death (Rootwelt, Reksen, fish do not show vocalization or physical symptoms of pain, and
Farstad, & Framstad, 2013) as reduced placental area per piglet leads rarely show behavioural reactions that humans can easily interpret.
to lower birthweights. These results may indicate that there is an upper Most importantly, until recently, the general view was that fish could
uterine limitation of litter size and that placental area and placental not experience pain in the same way as higher vertebrates. The ex-
weight influence postpartum survival (Rootwelt et al., 2013). Selection istence of pain receptors (nociceptors) in fish was only confirmed in
for increasing number of teats in sows may be a way to increase teat 2002 (Sneddon, Braithwaite, & Gentle, 2003) leading to the recogni-
availability per piglet, although this may also affect vertebral develop- tion of fish as sentient animals. Also, large scale industrialized aqua-
ment (Duijvesteijn, Veltmaat, Knol, & Harlizius, 2014). Hence, selection culture is a relatively recent farming method, and the animal welfare
for increased little size in maternal pig lines may be limited. Crossing and health challenges of fish farming have only recently come to the
this limit may not be ethical because the number of piglets weaned is attention of the public (Huntingford et al., 2006). Even though the
compromised due to sibling competition, piglet birthweight differences welfare of farmed fish has improved significantly in recent years in
and uneven growth, which hampers piglet welfare and leads to in- many countries with new legislation, a number of welfare issues still
creased mortality and increased financial losses for the owner (Ocepek, exist, including skeletal deformities, cataracts, and heart disease,
Newberry, & Andersen, 2017). Selection for larger litters should remain hormone treatments – induced spawning (GnRH), triploidity and
within reasonable borders and be accompanied by better surveillance transgenic technologies.
of parturition and low weight piglets, although these measures are One area of public concern has been the development of trans-
costly in high intensity large swine production systems. genic fish. While transgenic fish are currently banned in Europe,
the first genetically modified salmon (AquAdvantage-­s almon de-
veloped by AquaBounty Technologies) was put on the food mar-
3.3 | Chickens and hens
ket in Canada in 2015. This genetically engineered Atlantic salmon
grows faster due to a growth hormone (GH) gene insert, which
3.3.1 | Featherless chickens and killing of males
functions to increase endogenous GH production. While produc-
While featherless broiler chickens are believed to be a more tion of sterile transgenic salmon in land-­b ased tanks in Panama has
resource-­efficient and, according to the breeders, suffer less wel- now been approved by the USDA, major concern still persists over
fare problems (Olsson et al., 2006), many people have major con- the potential environmental impact of escaped transgenic fish.
cerns because the presence of feathers is considered an essential Due to their larger size and increased aggression, escaped trans-
characteristic of birds. While genetic selection is not generally un- genic fish may have a serious impact on wild conspecifics (Upton
acceptable, brought to extremes, it may be objectionable because & Cowan, 2015).
questions arise about the conditions in and about the purpose for
which animals are bred. This is also an example that the animals’
3.4.2 | Triploid salmon
original characteristics and dignity may be compromised.
In egg laying stock production nowadays usually all males are There is growing interest in the use of triploid fish in aquaculture.
sorted out and killed – similar to male dairy goat kids, as well as sur- There is concern regarding the large size of the farmed stocks rela-
plus dairy breed bull calves – due to the costs-­outcome imbalance of tive to wild fish, particularly over potential adverse impacts of es-
raising the male animals for meat production (Turner, 2010). In Norway, caped farmed salmon through potential interbreeding with wild fish
which has no tradition for consuming goat kid meat, nearly 31,000 sur- (Green et al., 2012).The production of triploids, which are function-
plus newborn male goat kids are killed annually (Roed, 2017). To many ally sterile, would result in two major benefits. Firstly, the use of
people this gender sorting and killing practice is ethically unaccept- triploid fish would negate the problem associated with early sexual
able because it implies that these animals are surplus by-­products and maturation in aquaculture, and secondly triploids, in being sterile,
therefore this practice violates the intrinsic value of animals. would mitigate the main environmental concerns of potentially es-
caped fish (Benfey, 2015; Iversen, Myhr, & Wargelius, 2016).

3.4 | Fish
4 | D I S CU S S I O N
3.4.1 | Environmental concerns
4.1 | Veterinary involvement
Compared to land farmed animals the welfare of fish and breeding
ethics have not traditionally been an important issue for consum- The primary task of veterinarians is to protect and monitor the health
ers, producers, and legislators. Basically, until recently, fish were not and the welfare of animals. As such, veterinarians give professional
much taken into consideration regarding welfare issues (Huntingford advice to owners, the society and governmental institutions regard-
et al., 2006). A number of factors may explain the reasons for the ing possible positive and negative effects and welfare issues of treat-
lack of public concern regarding fish welfare, some of them based on ments, breeding practices and production systems. In that regard,
ethical views. Fish do not elicit compassion in humans in the same veterinarians have to communicate evidence-­based information
way as other warm-­blooded animals do. In addition, unlike mammals, on potential risks to owners (breeders), media, the animal breeding
FARSTAD |
      11

industry, the Food Safety Authority and other stakeholders such as independence of the wealthier countries allows the inhabitants to
the general public, animal welfare activists and politicians. share large volumes of food with companion animals that are not
In recent years welfare issues related to animal breeding are in- producing food, but rather consuming it and competing for the
creasingly discussed. In 2017, 1,500 Norwegian veterinarians signed same food source with people from poor countries. The extent to
a petition against breeding brachycephalic breeds (Strand, 2017). which people will be able to keep family pets as now for a fore-
Furthermore, during the 2017 FECAVA congress a note was posted seeable future, is not known if food resources relying on animal
that aims at encouraging veterinarians to speak out against unethical protein is becoming scarce. However, breeding of peculiarities
breeding practices related to brachycephalic dog breeds (FECAVA, (designer dogs and cats) and allowing advanced medical treatment
2017). At the EVSSAR congress 2017 ethics of animal breeding was of pet animals with poor health most certainly and hopefully, will
addressed, and this topic will be continued also in 2018. be challenged. The resulting health problems must be addressed
In everyday contact with animal owners and breeders, ad- appropriately and effective improvement of the situation will not
dressing animal welfare and ethical issues may be a difficult task. occur until all stakeholders acknowledge their own responsibility
Some potential dilemmas that veterinarians are facing with regard in this process.
to breeding are highlighted in Table 1. For instance, regarding the
statement Satisfactory income and working conditions (VW) vets are
AC K N OW L E D G E M E N T S
involved in many aspects of the breeding and production systems
and may partially rely financially on the economic success of these The author is grateful to professor Ian Mayer, Professor of Fish re-
systems. However, of course regarding the statement Professional production at the Norwegian University of Life Sciences, Faculty
freedom (VA) they have the possibility to decline required interven- of Veterinary Medicine, Department of Production Animal Clinical
tions and should be intolerant towards systems that compromise an- Sciences for contribution and discussion on ethical aspects of fish
imal welfare exceedingly. In that regard also the statement Equitable breeding.
standards of Practice (VF) is important because it means fair and im-
partial attitudes towards all parties.
C O N FL I C T O F I N T E R E S T
The dilemmas listed here are applicable to any species, since
they also exist when dealing with all production animals (including None of the author have any conflict of interest to declare.
fish), and animals used in research. All unsound breeding practices
should be under scrutiny irrespective of species. As genetically en-
AU T H O R C O N T R I B U T I O N
gineered animals are released on the food market, such as Atlantic
salmon, special care, biosecurity and management may be required. The paper is authored by Wenche k Farstad only, acknolwledgment
Veterinarians not only are responsible for health care of these ani- is made to professor Ian Mayer’s discussion and knowledge con-
mals, but also contribute to policy discussions related to their health veyed to the author on the topic and considerations on ethical as-
and welfare, including breeding practices. In some instances, veter- pects on breeding and welfare of fish.
inarians may have to refrain from profit when owners and breeders
need telling about the welfare implications of poor health due to
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