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This paper presents arguments for, and evidence in support of, the important role of pleasure in animals’ lives, and outlines its considerable significance to humankind's relationship to other animals. In the realms of animal sentience, almost all scholarly discussion revolves around its negative aspects: pain, stress, distress, and suffering. By contrast, the positive aspects of sentience – rewards and pleasures – have been rarely broached by scientists. Yet, evolutionary principles predict that animals, like humans, are motivated to seek rewards, and not merely to avoid pain and suffering. Natural selection favours behaviours that enhance survival and procreation. In the conscious, sentient animal, the drives to secure food, shelter, social contact, and mates are motivated by desire (appetitive behaviour) and reinforced by pleasure (consummative behaviour). This is reflected in animals’ behaviour in the realms of play, food, sex, and touch. Despite the heuristic value of interpreting animal behaviour through the proximate (experiential) lens, scholarly study of animals remains entrenched almost exclusively in the ultimate (evolutionary) sphere. Not just science but also ethics suffer for this, for when we see animals as only the products of a competitive struggle for survival, we risk overlooking the positive qualities of their lives. Pleasure has moral import for such practices as factory farming and laboratory research, for it amplifies the moral burden of depriving animals the opportunity to lead fulfilling, enjoyable lives.
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Author's personal copy
Animal pleasure and its moral significance
Jonathan Balcombe *
Physicians Committee for Responsible Medicine, 5100 Wisconsin Avenue, NW, Suite 400, Washington, DC 20016, United States
1. Introduction
At long last, sentience is gaining the importance it
deserves in matters concerning humans’ relationship to
other animals. For much of the 20th Century questions of
animals’ so-called ‘‘private’’ experiences were dismissed as
invalid by leading scientific dogma. Today’s more enligh-
tened science is entertaining questions of animal con-
sciousness, cognition, emotion, and pain. This is significant,
because animals’ capacity to suffer has always been at the
core of animal welfare and animal rights ideologies
(Bentham, 1789; Singer, 1975). This special issue on
animal suffering and welfare is just the latest in a rapidly
growing literature on animal sentience.
Intriguingly lacking from this discourse has been
animals’ capacity for pleasure. With the exception of a
few notable contributions (e.g., Cabanac, 1971; Panksepp,
1998), discussions of animal sentience have been almost
exclusively in the negative realm: pain, stress, distress. At
least nineteen current English language journals (most of
them medical) contain the word ‘‘pain,’’ whereas none deal
with ‘‘pleasure.’’ Granted, pain has more moral urgency
than does pleasure (see Pleasure’s Moral Significance,
below), the role of pleasure in human and non-human
experience nevertheless deserves far more attention than
we have been giving it.
This paper aims to help correct this imbalance by
presenting the argument for animal pleasure, providing
some illustrative examples, and then discussing some of its
moral implications. A more thorough examination of
animal pleasure can be found in my book Pleasurable
Kingdom: Animals and the Nature of Feeling Good (also
available in German translation as Tierisch Vergnu
Applied Animal Behaviour Science 118 (2009) 208–216
Article history:
Available online 12 March 2009
Positive emotions
Animal welfare
Animal ethics
This paper presents argumentsfor, and evidence in support of, the important role of pleasure
in animals’ lives, and outlines its considerable significance to humankind’s relationship to
other animals. In the realms of animal sentience, almost all scholarly discussion revolves
around its negative aspects: pain, stress, distress, and suffering. By contrast, the positive
aspects of sentience – rewards and pleasures – have been rarely broached by scientists. Yet,
evolutionary principles predict that animals, like humans, are motivated to seek rewards,
and not merely to avoid pain and suffering. Natural selection favours behaviours that
enhance survivaland procreation. In the conscious, sentientanimal, the drives to secure food,
shelter, social contact, and mates are motivated by desire (appetitive behaviour) and
reinforced by pleasure (consummative behaviour). This is reflected in animals’ behaviour in
the realms of play, food, sex, and touch. Despite the heuristic value of interpreting animal
behaviour through the proximate (experiential) lens, scholarly study of animals remains
entrenched almost exclusively in the ultimate (evolutionary) sphere. Not just science but
also ethics suffer for this, for when we see animals as only the products of a competitive
struggle for survival, we risk overlooking the positive qualities of their lives. Pleasure has
moral import for such practices as factory farming and laboratory research, for it amplifies
the moral burden of depriving animals the opportunity to lead fulfilling, enjoyable lives.
ß2009 Elsevier B.V. All rights reserved.
This paper is part of a special issue entitled ‘‘Animal Suffering and
Welfare’’, Guest Edited by Hanno Wu
* Tel.: +1 202 686 2210x331; fax: +1 202 686 2216.
E-mail address:
Contents lists available at ScienceDirect
Applied Animal Behaviour Science
journal homepage:
0168-1591/$ – see front matter ß2009 Elsevier B.V. All rights reserved.
Author's personal copy
(Balcombe, 2006a). The current paper includes some
newer ideas and perspectives on animal pleasure and
its moral implications since publication of that earlier
I should briefly allude to the boundaries of what
constitutes an ‘‘animal’’ in the arguments which follow. It
is not the intention of this paper to establish where we
might draw the line as to what species are sentient. It will
be apparent from the examples I give below that I favour a
relatively inclusive definition that embraces all vertebrate
taxa as well as some invertebrates, such as cephalopod
mollusks. For most, the yardstick (if not necessarily the
pinnacle) of sentience is likely to be an animal’s degree of
similarity to humans. What matters for our purposes is
that we can agree to include at least some non-human
animals here. Few would argue, for instance, that a dog or a
giraffe is not sentient.
1.1. Myopic science
Science has and continues to hold too narrow a
perspective in its scholarly interpretation of animal
existence. Specifically, published studies of animal
behaviour are presented almost exclusively in an
ultimate, evolutionary context, without discussion of
the animals’ more proximate, mental and emotional
experiences. This pattern arises from science’s pursuit of
rigour, which is assumed to be found primarily in
the evolutionary context. Schooled on Occam’s Razor,
which holds that the explanation of any phenomenon
should make as few assumptions as possible, scientists
adhere to the law of parsimony like a barnacle to a boat
hull. We do not like making assumptions, and we
certainly do not wish to be anthropomorphic (Kennedy,
For example, when rats show a preference for novel
flavours following three days pre-exposure to a single
flavour, we comfortably propose evolutionary theories to
account for the switch. The rats’ behaviour appears
adaptive for two reasons: (1) it avoids over-dependence
on a potentially short-lived food source, and/or (2) it
reduces the risk of developing a micronutrient deficiency
(Galef and Whiskin, 2003). These are quite sensible
adaptive explanations for the rats’ behaviour. But we also
know with reasonable certainty that rats do not bone up on
Darwinian fitness or ruminate on evolutionary theory.
Thus, we take too narrow a view when we fail to
acknowledge the likely experiential basis for the rats’
behaviour: that they simply got tired of eating the same old
fare and enjoyed something new.
Similarly, while there may be good adaptive bases for
the broad and diverse human cultural practice of adding
spices to our foods (Billings and Sherman, 1998), I am not
aware that anyone reaches for the oregano or the curry
powder with the conscious intent of warding off intestinal
microbes. We spice our food because it enhances the taste.
I propose that Occam’s Razor has been blunted by
overuse. As evidence of animals’ cognitive and emotional
capacities snowballs (now that we have ended the
embargo on addressing these concepts), it becomes
increasingly clear that animals are conscious, experiencing
individuals. And it follows that to deny them such
experiences – as science tacitly does by neglecting to
discuss them – we become increasingly outmoded and
anthropocentric, and science loses rigour (Burghardt,
2007). Whenever we address questions about how an
animal might be feeling, we inevitably make assumptions
because we cannot know absolutely what the animal is
feeling. And we inescapably anthropomorphize because
we are anthropoid apes. These are not sins so long as we
make reasonable assumptions backed by good science, and
so long as we anthropomorphize critically, from a
foundation of knowledge of the animals we were describ-
ing (Burghardt, 1991). In the face of current knowledge, it
is a bigger assumption that animals are unconscious,
unfeeling things than that they are sentient, emotional and
aware. Furthermore, ascribing conscious experience to
animals, and the sentience that ensues, need not be seen to
violate the law of parsimony, for cognitive explanations
can be simpler than cumbersome stimulus–response
explanations (de Waal, 1991).
2. Arguments for a pleasurable kingdom
Because an appreciation of animal pleasure and its
ethological study – we might call it Hedonic Ethology – is a
nascent and as yet under-represented field, it helps to
present some arguments for its presence and significance.
In addition to the issues of parsimony and anthropomorph-
ism mentioned above, there are still influential scholars
who deny animals a conscious experience of life (e.g.,
Budiansky, 1998; Macphail, 1998; Wynne, 2004). Thus, the
argument still needs making.
Before I proceed with the arguments I find to be most
compelling for animals’ experience of pleasure, let me
point out the hypocrisy of any argument for the
unprovability of animal conscious experience that never-
theless accepts its existence in humans. In the purest
sense, human sentience is no more scientifically provable
than is non-human sentience. While our sophisticated
language likely permits us to share our pain/pleasure
experiences more discretely on the psychological plane
than for other taxa, the physical privacy of individual
experience is as unbridgeable between two humans as it is
between a human and a non-human animal. Quite literally,
one person cannot actually feel another’s pain or pleasure.
Do we thereby accept the non-existence of human
sentience? No. We accept it, and with good reasons even
though the solipsist’s claim that ‘‘my mind is the only thing
that I know exists’’ is logically incontrovertible. Thus, to
accept the existence of sentience in humans while denying
it in certain animals (say, birds) is scientifically and
ethically inconsistent.
2.1. Pleasure is adaptive
Pleasure is a product of evolution. The evolution of
motility was a key step towards the eventual evolution of
complex sensory systems that could perceive environ-
ments and detect noxious and rewarding stimuli. Just as
the capacity for pain is adaptive for a mobile organism that
can move away from aversive things, so too is pleasure
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
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beneficial by rewarding the individual for performing
behaviours that promote survival and procreation (Caba-
nac, 1971). Because animals can actively avoid and
approach things, they benefit from being able to have a
qualitative experience of those things. Pain’s unpleasant-
ness helps steer the animal away from ‘‘bad’’ behaviours
that risk the greater evolutionary disaster of death
(Dawkins, 1998). Similarly, pleasure encourages animals
to behave in ‘‘good’’ ways, such as feeding, mating, and –
depending on climatic conditions – staying warm or cool.
There are of course energetic costs to evolving and growing
complex neurological organs and sensory systems, but
clearly the benefits have outweighed the costs, at least for
members of Phylum Chordata.
2.2. Humans feel pleasure
Arguing the case for animal pleasure would be more
difficult were it not for the fact that humans experience
pleasure. That we know and accept the existence of this
sensory phenomenon in one species provides a foundation
for its presumptive existence in others. That human
languages contain rich vocabularies for describing good
feelings attests to the diversity of both physical and
emotional pleasures that can be felt by humans. English
includes: happiness, delight, surprise, anticipation, pride,
satisfaction, joy, elation, ecstasy, thrill, euphoria, exulta-
tion, jubilation, excitement, rapture, fulfillment, gratifica-
tion, and comfort, among others. Though the writer is
unilingual, he is aware of at least one German word –
funktionslust: the pleasure in what one does best – that
captures a sort of pleasure for which English has not a word
(Masson and McCarthy, 1995). It seems likely that other
languages contain terms for distinctive pleasures.
Having evolved in diverse environments, where differ-
ent niches present different adaptive challenges and
opportunities, different animals have different sensory
and perceptual skill sets. It follows that some animals
might be able to experience realms of pleasure unfamiliar
to humans. The echolocation abilities of bats and
cetaceans, electric communication in fishes (Hopkins,
1974), and the ability of pinnipeds to detect fish turbulence
trails with their vibrissae (Dehnhardt et al., 2001; Schulte-
Pelkum et al., 2007) are representative examples. Some
animals can tune into the earth’s magnetic field to help
navigate, and there is now evidence that birds perceive this
visually (Heyers et al., 2007). These examples do not
explicitly involve pleasure, but they illustrate the potential
for pleasures unknown to us. Two leading neuroscientists,
and an ethologist, have recently suggested that other
animals may experience some feelings more intensely than
we do (Burgdorf and Panksepp, 2007; Bekoff, 2007).
2.3. Animals feel pain and distress
A heuristic argument for animals’ experience of
pleasure is that they feel pain. Pain in animals is well
studied, and there is solid empirical evidence for its
existence in vertebrate groups (Braithwaite, 2007). The
detection of a nociceptive stimulus is highly adaptive for
motile organisms that can move away from the source of
the stimulus. We can imagine an ancient, primitive animal
flinching and fleeing from the bite of another. By escaping
more readily than would a non-perceiving conspecific, her
genes are more likely to show up in future generations. For
animals with highly developed nervous and sensory
systems, being able to learn from an earlier painful
experience is a further refinement of the stimulus–
response dynamic. Cognitive creatures can recognize
and remember the source of an earlier hurt, and make
adjustments to their behaviour to reduce the chances of a
repeat experience. The conscious pain perceiver is once
again more likely to survive future encounters and to be
favoured by natural selection.
Indirect evidence for animal pain is widespread in
nature. Plants, for example, have exploited animals’
capacity for pain and discomfort with the evolution of
thorns, and bitter tasting chemical compounds in their
tissues. Similarly, many animals have well-developed
spines, stingers, horns, and tusks which, like thorns, not
only inflict pain, but also signal ‘‘don’t touch.’’
More direct evidence for animal pain resides in animals’
behavioural responses to noxious stimuli. Their vocaliza-
tions and their movements – retreating from and avoiding
the source of pain, flinching, limping, protecting the
injured part – are all consistent with what we may expect
of an individual who experiences pain.
Scientific studies of animal pain reinforce what
observations suggest. Laboratory studies have shown
repeatedly that injured rats will favour the bitter taste
of water that contains a pain-relieving drug over una-
dulterated water (Persinger, 2003; Colpaert et al., 1980,
1982). Chickens also self-administer pain-killers. These
birds often experience leg problems associated with
breeding, growth and husbandry, and can develop highly
abnormal gaits, or become completely unable to walk.
Lame birds ingest more drugged food than non-lame birds,
and as the severity of the lameness increases, lame birds
consume a greater proportion of the drugged food (Dan-
bury et al., 2000). Fishes can learn to avoid noxious stimuli
such as electric shocks (Ehrensing et al., 1982) and anglers’
hooks (Beukema, 1970a,b), and the evidence for fishes’
experience of pain is mounting (Braithwaite, 2007).
Empirical evidence is now emerging that suggests that
some invertebrates experience pain (Elwood, 2007).
2.4. Animals behave as if they feel pleasure
2.4.1. Play
Play behaviour is widespread in mammals, and has also
been described in about half of all avian families (Ortega
and Bekoff, 1987). Behaviour suggesting play has been also
observed in other taxa, including reptiles, fishes, and
cephalopods (see Burghardt, 2005). Because play tends to
occur spontaneously and unpredictably, it is difficult to
measure, and most published studies of animal play are
anecdotal. While the accumulation of well-documented
anecdotes has scientific value, there is also a growing
database of empirical studies of play. For example, a three-
year study of aerial drop-catching behaviour by Herring
Gulls in Virginia concluded that it was play. These birds
will drop clams onto hard surfaces to smash them and
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
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access the soft parts; they also will swoop to catch clams
and other objects they have dropped before they hit the
ground. This latter behaviour appears to be playful (as
opposed to kleptoparasitic, for example) because: drop-
catches were performed more by younger birds, drop-
catches were not necessarily made over a hard substrate,
the behaviour was sometimes performed with non-food
objects, and it occurred more often during warm, windy
weather (Gamble and Cristol, 2002).
There are good adaptive reasons for the existence of
play, and scholarly discussions of play are usually in the
context of its evolutionary benefit (Bekoff and Byers, 1998;
Burghardt, 2005). Playing games of chase is no doubt
beneficial for healthy, well-fed lambs, as it is for fox cubs,
even though as adults these species may deploy these skills
in different contexts (escape and pursuit, respectively).
Yet, animals (including humans) do not consciously play
for ultimate reasons; they play because it is fun to do so.
Enhanced survival can be seen as a positive reward in the
evolutionary sense, but not in the sense of experiencing a
pleasurable sensation.
Thus, what is largely lacking in discussions of animal
play is its affective element. Inasmuch as the context and
expression of animal play often resemble that of human
play, we may expect that it is pleasurable for them, too.
Having fun seems to many of us the main characteristic of
animal play (Burghardt, 2005). Burgdorf and Panksepp
(2001) found that rats would run to the hand, on average,
four times as quickly to receive tickles to their bellies
(which mimic rats’ rough-and-tumble play behaviour)
than would rats trained to expect dorsal strokes. Several
species are known to calibrate the boisterousness of their
play, apparently to sustain the activity, which also suggests
that they are enjoying it. Adult red-necked wallabies self-
handicap against smaller and weaker playmates. For
example, rarely does the larger partner rear up or kick;
littler playmates are also much more vigorous in their
play-fighting (Watson and Croft, 1996). Younger chimps
are more likely to engage older chimps who are soliciting
play if the latter shows self-handicapping behaviour
(Mendoza-Granados and Sommer, 1995). Young chimps
(and other species) tested in the laboratory will choose
play over food unless they are very hungry (Goodall and
Bekoff, 2002), and there are anecdotal accounts of captive
animals working for the chance to play (Linden, 1999).
We may say that the adaptive benefits of play are what
sustains its expression in a gene pool, but if we are dealing
with conscious, emotional animals, we can and should
acknowledge the proximate role of pleasure (another
product of evolution) in the interaction. This more
inclusive interpretation is more thorough because pleasure
acts as the proximate mechanism that causes animals
(including humans) to behave adaptively—in this case, by
2.4.2. Food
Because food is so indispensable to an animal’s survival,
we may expect that conscious animals are highly
motivated to obtain it, that they anticipate it, and that
its consumption brings rewards. We ‘‘know’’ (though the
solipsist could challenge us) that humans have such a
relationship to food. But the experience of food pleasure in
animals is almost wholly unexamined.
There are innumerable clues that animals favour the
flavour in their food. They show food preferences (e.g.,
Johnston and Fenton, 2001), tastes that change through
time (Galef and Whiskin, 2003), and the anticipation of
food (Tinklepaugh, 1928). Some, such as language-trained
apes and parrots, can tell us their reactions to food (Barber,
1993; Linden, 2003). It has also been shown that animals
produce pleasurable compounds known as opioids during
both the search for food (the appetitive phase) and its
consumption (the consummatory phase) (Berridge, 1996).
Facial responses to tastes are similar between rodents,
primates and humans, suggesting shared evolutionary
origins (Steiner et al., 2001; Cabanac, 2005). Enjoyably
sweet flavours elicit characteristic licking responses, while
aversive bitter tastes elicit gaping and head shaking. These
behavioural responses are accompanied by activity in
brain ‘hedonic hotspots,’ such as the nucleus accumbens
and ventral pallidum, where neurotransmitters coordinate
the ‘‘liking’’ of tastes (Pecin
˜a et al., 2006; Norgren et al.,
2006). This linking of brain activity with positive patterns
of behaviour (Berridge, 1996) suggests the conscious
experience of pleasure.
A study of juvenile green iguanas showed that these
animals would trade off the palatability of a bait (lettuce)
with the disadvantage of having to venture into a very cold
area to retrieve it. As the ambient temperature at the bait
decreased, the lizards visited the bait less frequently and for
shorter periods, choosing instead to stay under the heat-
lamp where nutritionally complete reptile chow was freely
available. Moreover, time interval between sessions with
bait (ranging from 1 to 8 days)had no effecton the duration of
stay on the bait, suggesting that the lettuce was more of a
luxury rather than an indispensable nutritional food source
´and Cabanac, 1998). Earlier experiments found that
rats invariably shunned laboratory chow and ran into a cold
environment to consume highlypalatable foods,and that the
animals’ individual preferences (which varied considerably)
were reflected by the amount eaten, the number of
excursions, and the time spent feeding in the cold (Cabanac
and Johnson, 1983).
A compelling and ecologically important example of the
hedonic aspects of food to animals is the evolution of fruit as
a seed-dispersal mechanism. Fruit, to paraphrase botanist
and science writer Michael Pollan, evolved as the product of
a grand co-evolutionary bargain struck between plants and
animals (Pollan, 2001). Desire, according to Pollan, is built
into the very nature and purpose of fruit. He is correct, but
you wouldhave to search hard to find anyscientists sayingas
much in a peer-reviewed journal. When I was researching
this paper, I encountered a brief essay titled ‘‘Seed dispersal
by animals: Behaviour matters,’’ by a then-doctoral student
of botany at Washington State University. While lucid on
the evolutionary facets of plant-animal coevolution, the
writer nevertheless makes this characteristically agnostic
‘‘Fruit appearance provides the cue for fruit choice, but
nutrients provide the positive feedback that maintains
an animal’s preference for the fruit.’’(Yang, nd)
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
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Surely, if our own experience is any guide, what
provides the animal with (immediate) positive feedback is
not nutrients, but the taste of the fruit. Nutrients are the
mediator; taste is the sentience pay-off. The more proximal
experience of taste is an evolutionary product of what is
nutritionally beneficial to the forager. And its flavour, not
nutrient content directly, is what an animal experiences
consciously when she selects and eats a fruit.
2.4.3. Sex
It is hard to overestimate the importance of reproduc-
tion to an organism. Without it, species would cease to
exist. Because reproduction is so important, natural
selection should strongly favour behaviours that promote
mate-seeking, mating and, where necessary, the raising of
young. In conscious, feeling creatures, we may expect that
what motivates them to behave in these ways is a
combination of instinct on the one hand, and a powerful
desire to attain reward on the other.
When it comes to sex, science is once again fixated on
the ultimate, evolutionary context to explain how animals
behave, and not the proximate, experiential context.
Sexual activity in animals is conventionally portrayed as
all business and no pleasure. The Encyclopedia of Animal
Behaviour, for example (Bekoff, 2004), contains several
entries about animal sexual behaviour, all of which
interpret that behaviour solely in evolutionary contexts.
From journal articles to textbooks to television documen-
taries, the idea that animals may be enjoying themselves is
not explicitly rejected; it is not mentioned at all. The result
is that there is precious little consideration of how animals
might be feeling during sexual encounters, and how this, in
turn, might affect their behaviour.
What, then, might be said of the role of pleasure in
animals’ sex lives? We do know that pleasure is a powerful
reinforcer of sex drive in humans. Furthermore, the human
quest for sexual gratification (and profit) manifests itself in
an astonishing diversity (and, one might add, perversity) of
sexual behaviour—witness a thriving pornography indus-
try that caters to a wide range of individual sexual leanings
and fetishes.
Against this human backdrop, sexual behaviour in non-
human animals may appear both perfunctory and passion-
less, when it is observed at all. Yet, a good deal of animal
sexual behaviour appears not devoted directly to the
reproductive fitness (Bagemihl, 1999). In Biological Exu-
berance: Animal Homosexuality and Natural Diversity,
Bagemihl surveyed of homosexuality and other forms of
non-procreative sexual behaviour among mammals and
birds. He found a lot of it.
And there is a lot of it. Many animals routinely copulate
or engage in other sexual activities outside of the breeding
season, including during pregnancy, menstruation (in
mammals), and egg incubation. Such non-procreative
activity may even constitute a large proportion of the
animals’ sexual behaviour, as it does, for instance, for
common murres, proboscis monkeys, addax antelopes,
rhesus macaques, wildebeest, golden lion tamarins, and
mountain goats (see Bagemihl, 1999 for citations). Another
variation on the theme of wasteful sex is group sexual
activity wherein few if any participants are passing along
genes. These sorts of orgies have been observed in spinner
dolphins, gray and bowhead whales, swallows, and herons
(ibid). Variations on non-copulatory mounting, include:
mounts without erection, mounts with erection (but with
no penetration), reverse mounting in which a female
mounts a male, mounting from the side or in positions
from which penetration is impossible, which has been
reported in mammals such as Japanese macaques, water-
buck, mountain sheep, takhi (Przewalski’s horse), collared
peccaries, warthogs and koalas, and birds including ruffs,
hamerkops and chaffinches. Animals also engage in
various forms of oral sex, stimulation of partner’s genitals
using the hands, paws, or flippers, and various forms of
anal stimulation. There are also observations of inter-
species sexual coupling in the wild (Bagemihl, 1999).
Skeptics might claim that these misguided antics are
the fumblings of young, confused or otherwise inexper-
ienced individuals, though cases of oral or manual
stimulation and cross-species interactions strain such
claims. Behaviours that probably most suggest that sexual
stimulation is pleasurable are various forms of auto-erotic
behaviour. Owing to its utter futility for procreation,
masturbation suggests that the performer merely seeks
pleasure. Masturbation is widespread in mammals, and
practiced about equally by both females and males
(Bagemihl, 1999; Judson, 2003). It is known from at least
seven mammalian orders, including primates, carnivores,
bats, walruses, ungulates, cetaceans, and rodents. Mas-
turbation appears less common in birds, but is not absent
(e.g., Burger, 2001; Gaston and Kampp, 1994; Winterbot-
tom et al., 2001).
Most biologists today recognize same-sex sexual
interactions as being part of the normal, routine beha-
vioural repertoire of the animals who engage in it.
Currently, at least 300 species of vertebrates are known
to practice homosexuality (Bagemihl, 1999). In most cases,
participating individuals will also engage in heterosexual
behaviour, and their sexual life history can most accurately
be categorized as bisexual. Homosexual behaviour is
clearly maladaptive from a strictly procreational stand-
point, and while practitioners may benefit by gaining
practice, or by releasing sexual tension, pleasurable
rewards seem a likely motivation for engaging in it.
There are other bases for sexual pleasure in animals,
including the prevalence of a clitoris in female mammals,
and evidence for female orgasm in various primate groups,
including apes, monkeys, lemurs and marmosets (Dixson,
1998; Bagemihl, 1999). In one study of Japanese macaques,
female orgasmic responses were observed in 80 of 240
copulations (33%) (Troisi and Carosi, 1998).
2.4.4. Touch
Unlike food or sex, touch is neither indispensable to
survival nor to reproductive success. However, touch
sensitivity is very useful. It allows animals to react
adaptively to their environments. For example, being able
to detect water movements helps fishes orient themselves
in murky streams and on migration routes, and to detect
the movements of other nearby animals, including
potential predators. But for social species especially, touch
often plays a key role in interactions with other
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
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individuals. Chimpanzees and other primates are well
known for their proclivity to groom one another, which
some species spend on average about 20% of their waking
time doing (Uhlenbroek, 2002). Given its importance in
these social contexts, we may expect that touch is a source
of pleasure, as we know it to be from our own experience.
Very few investigators have addressed the pleasure of
touch. In some cases, animals’ liking of tactile contact may
reveal itself by accident. For example, in a study in which
dolphins could request rewards (pleasures) by pressing
plastic symbols on a keyboard with the tips of their beaks,
some animals favoured getting a rub to getting a fish
(Linden, 2003). In some locations, whale-watchers have
gained the trust of gray whales, who ride up against the
sides of boats to have their bodies stroked and patted
(Nollman, 1986). These examples do not prove that
cetaceans enjoy the feeling of touch, but they are
consistent with it.
There have been a few attempts to measure positive
physiological responses in groomed animals. In primates,
the release of pain-relieving endorphins has been demon-
strated when they groom each other (Keverne, 1992).
Evidence from horses supports a pleasant, calming effect.
Like many other mammal species that form social groups,
Camargue horses regularly engage in mutual grooming.
When human researchers experimentally groomed them,
the animals’ heart rates dropped significantly, but only
when the touch was directed at those areas of the neck that
are the preferred grooming sites in this species (Feh and de
`res, 1993).
As part of the study of tickling in rats mentioned earlier,
tickled rats were trained for nine days to press a bar to
receive tickling. A second, inactive bar elicited no tickling
when pressed. On the test days that followed, the rats
pressed the active bar repeatedly to receive tickling; they
almost never pressed the inactive bar, and their bar-
pressing increased during this period. For seven days of
extinction testing, during which time bar-pressing elicited
no tickling, use of the active bar declined significantly and
by the fifth day was equivalent to presses on the inactive
bar (Burgdorf and Panksepp, 2001).
Some interesting evidence for positive affect associated
with touch has been documented in interspecies interac-
tions in aquatic environments. At a freshwater spring in
Kenya, fishes of various species gravitate to wallowing
hippopotamuses, nibbling and plucking at their skin.
Observations by biologists/photographers Mark Deeble
and Victoria Stone indicate that the hippos are far from
passive participants in these cleaning services. They
deliberately splay their toes, spread their legs and hold
their mouths open to provide easy access for the fishes, and
they appear to solicit cleanings by visiting ‘cleaning
stations’ where fish congregate (Deeble and Stone,
2001). The hippos were so relaxed during these ‘‘spa
treatments’’ that they would sometimes fall asleep.
The widespread phenomenon of fish cleaning stations
not only gives us a fascinating glimpse into the experiences
of animals still assumed by many to have no feelings (Rose,
2007), they offer compelling evidence for the experience of
tactile pleasure in fish. Cleaner fish of a variety of species
nibble loose skin, fungal growths, and fish lice from other
fish ‘‘clients.’’ Wounds may also be plucked at, which may
relieve infection and speed healing. It is a mutualism:
cleaners benefit by getting food – delivered buffet-style by
clients who line-up patiently to await their turn – and
clients get a body-cleansing service. Different species of
customers (or ‘‘hosts’’) are cleaned in a highly specific
manner by cleaner fish (and shrimps), who advertise their
services with brightly colored uniforms, and perform
bobbing/fussing movements to signal their willingness to
attend to clients. Clients also may signal their readiness, for
instance by orienting themselves vertically in the water,
and opening their mouths and gills at appropriate times to
allow access to the cleaner fishes (Potts, 1973).
There are features of the cleaner–client relationship
that suggest positive feelings are involved. Invitation
postures indicate that cleaners may be anticipating the
attentions of clients, and clients the services of their hosts
(Potts, 1973). Recent study of one of the principal cleaner
species, the cleaner wrasse, supports the notion that tactile
stimulation is an important motivator for the interaction.
Based on 112 h of observation of 12 different cleaners,
cleaners appear able to alter client decisions over how long
to stay for an inspection, and to stop clients from fleeing or
responding aggressively to a cleaner bite that made them
jolt (Bshary and Wuerth, 2001). There is also evidence that
cleaners and clients recognize each other (Tebbich et al.,
2002), and that they return to their favoured business
partner (Bshary and Shaeffer, 2002), much as we return to
a favourite barber or hairdresser. Finally, cleaners some-
time cheat by nipping off some of the client’s own skin;
established clients behave as if taking serious offense at
this, chasing the cleaner around, and/or shunning the
cleaner’s future solicitations (Bshary and Grutter, 2005).
This punishment helps to stabilize the relationship
between cleaners and their clients.
2.4.5. Other pleasures
Animals’ lives afford them the opportunity to experi-
ence a wealth of other pleasures beyond the realms of food,
sex and touch. Animals seek comfort, pursuing rewarding
sensations that help maintain homeostasis (Cabanac,
1971), such as basking in the sun or seeking shade.
Animals may also appreciate esthetic beauty; for example,
female preference for showy male displays has driven
sexual selection for elaborate beauty. And animals may
experience a broad spectrum of emotions (Masson and
McCarthy, 1995; Bekoff, 2007). Demonstrating these
phenomena is challenging in species which cannot report
their experiences in the way that humans can. As interest
in the sensory experiences of animals rises, we may expect
more ingenious methods to probe them, and more data to
shed light on these topics.
3. Pleasure’s moral significance
What, if any, are the moral implications of animal
pleasure? Does being not merely a pain-avoider but also a
pleasure-seeker add to whatever moral implications may
follow from the capacity for pain and distress? Beginning
with 18th Century philosopher Jeremy Bentham’s (1796)
now famous appeal to animals’ capacity to suffer (‘‘The
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
Author's personal copy
question is not: Can they talk, or Can they reason, but Can
they suffer’’), most of the recent scientific and philoso-
phical discourse about animal welfare and rights has
focused around pain and suffering. This is intriguing, for
the consideration of pleasure has made some major
contributions to moral philosophy. Utilitarianism, first
expounded as a formal philosophy by Bentham, favours
actions that optimize pleasurable outcomes while mini-
mizing negative ones. Notably, Bentham regarded animals
as serious objects of moral concern, based on their capacity
for both pain and pleasure. That Bentham’s utilitarianism
has also been called Hedonistic Utilitarianism is a further
acknowledgment of the importance of pleasures to
considerations of right and wrong.
Peter Singer, the most influential contemporary utili-
tarian philosopher, has argued persuasively that sentient
animals have interests, and that those interests involve not
just avoiding physical pain and/or psychological suffering
but also the experience of pleasure (Singer, 1975). But
Singer himself admits to a general preoccupation with
negative experiences: ‘‘ focus has always been on
animals’ capacity to suffer, from the time I started thinking
seriously about the ethics of how we treat them’’ (Singer,
Some imbalance here is to be expected, given the
greater urgency and gravity attached to pain and suffering
when compared to pleasure. The avoidance of pain relates
to matters of immediate survival; pain evolved as a means
to avoid the greater penalty of death. Pain teaches
individuals to avoid dangerous or harmful behaviours
and encourages prudence in the decisions they make, for
instance, in the pursuit of survival and social interactions.
Pleasure, by comparison, does not normally hinge on the
grave matter of survival, at least not in the immediate
sense; obviously, if an animal were to ignore the
motivations provided by the anticipation of food and the
pleasure of its consumption, then that animal would soon
But the utter absence of any scholarly journal dedicated
to positive feelings is surprising given pleasure’s important
– arguably central – role in sentient experience. Readers
might ask themselves what drew them to their career
choice, and what motivates their preferences on a
restaurant menu, their choice of clothing, or their weekend
The capacity for good feelings has important corollaries
for ethics. Lives that contain pleasure are lives with
intrinsic value (Regan, 1983). That is to say, an individual
who can experience good feelings has a life that is of value
to that individual, independent of any value that indivi-
dual’s life might have to another, for example as a source of
entertainment, or revenue (ibid). Put another way, an
animal who can experience pleasure has the capacity for a
quality of life (McMillan, 2005). It is a life worth living, one
in which there are better and worse days, and moments
that are more or less pleasurable than others.
Humankind’s relationship to animals has been and
continues to be one based mostly on a ‘‘might-makes-
right’’ or ‘‘bright-makes-right’’ ethic. With our large brains
and manipulative hands, our sophisticated language and
technologies, we have evolved the capacity to control other
animals completely. And we wield that control to the
fullest. According to the Food and Agriculture Organization
of the United Nations (FAO), over 50 billion land animals
worldwide were killed for food in 2005 (Steinfeld et al.,
2006). The numbers of individual fishes killed by humans
are probably higher based on FAO statistics; global
consumption in 2004 was 141.6 million metric tons
(FAO, 2007). Close to 100 million animals are consumed
yearly in laboratory experimentation in the United States
(Carbone, 2004). The Humane Society of the United States
estimates that each year over 50 million animals are killed
for their fur (HSUS, 2007). Paradoxically, the number of
animals we kill continues to rise, despite our greater
appreciation for and ethical concern towards animals than
ever before.
While there have been some scientific advancestowards
better understanding the animal’s experience of slaughter
(Grandin, 1994; Gregory, 2004), modern methods of killing
still involve intense levels of stress, pain and suffering
(Warrick, 2001; Patterson, 2002; Scully, 2004). We may
naturallyconclude that harm is done to them. To the degree
the victims can also feel pleasure, it may notfollow obviously
that the harmof a painful, stressful end is amplified. Surely, it
is suffering, not pleasure, that weighs on death.
Pleasure’s moral significance resides in two aspects of
our relationship to other animals. Each relates to the denial
of pleasures. First, when we keep animals in impoverished
conditions, as we do in factory farms, laboratory cages
(Mason, 1991), and zoos (Jensen and Tweedy-Holmes,
2007), we deny them the opportunity to express natural
behaviours. At least in rats and mice, studies show that
animals bred for many generations in captive confinement
retain behaviours of their wild ancestors (Estep et al.,
1975; Boice, 1977), and that they are highly motivated to
engage in behaviours that are a routine part of their day-
to-day lives in nature, such as foraging, exploring, and
social behaviour (Dawkins, 1988; Balcombe, 2006b).
One objection to this denial-of-pleasure argument is
that captivity may offer the animal safe shelter from the
vicissitudes of life in the wild (Tannenbaum, 2001). Wild
existence can be hazardous, and some species may live
longer in captivity (though others, such as cetaceans, do
not). The problem with this objection is that a safer life is
by no means a better life. We would be safer if we stayed
home and never ate food prepared by others, for instance,
but few of us would choose this mundane sort of existence.
Another problem is that animals – like humans – prefer not
to be confined; it is fairly axiomatic that animals prefer
freedom to being in a cage.
The second form of denial of pleasure relates to death.
In killing animals – especially young, healthy animals – we
cause harm by denying them the opportunity to experi-
ence rewards that life would otherwise offer them. It may
be claimed that a dead animal misses nothing. But the
main reason that our criminal system treats murder so
seriously is not that the victim may suffer—though that
certainly compounds the crime. Murder is wrong because
life, specifically that portion of life yet to be experienced,
has value.Thus, killing is the greatest harm that can be
done to conscious, autonomous beings (Balluch, 2006), and
pleasure is firmly rooted in the harm committed.
J. Balcombe / Applied Animal Behaviour Science 118 (2009) 208–216
Author's personal copy
4. Conclusion
With the removal of former behaviourist taboos against
subjective experiences in animals and the subsequentrise of
cognitive ethology, there is an accelerating accumulation of
information supporting animal sentience and awareness.
Mice show empathy(Langford et al., 2006), sheep respondto
the emotional states of other sheep (Da Costa et al., 2004),
monkeys show a sense of justice (Brosnan and de Waal,
2003), starlings can become optimistic or pessimistic
(Bateson and Matheson, 2007; Matheson et al., 2007), and
rats show mirthful responses to tickling (Burgdorf and
Panksepp, 2001). Science shows us that humans are not the
only autonomous, sentient beings, or the only hedonists.
Many other animalsalso have lives worth living. They do not
speak as we do, but the scientific case that they have
experiences is far stronger than the case that they do not.
The inevitable moral question arises: How can we
reconcile our current treatment of animals with this
knowledge? The answer, I believe, is that we decisively
cannot. The reason for this lies in the principles that inform
human law and policy. As autonomous, sentient beings,
humans have intrinsic value. It follows that we have
certain basic, inviolable rights (provided we behave within
the confines of the law), such as the right to freedom and
the right to life. What grants us these rights is our intrinsic
worth. Our reasoning and linguistic abilities are not the
foundation of such rights, else we would deny them to the
many humans who lack these capacities (Singer, 1975).
Despite this, our treatment of animals continues to
languish in a Cartesian framework, one that permits the
sacrificing of animals’ most precious possessions (their
freedom and their lives) for such relatively trivial human
ends as gustatory pleasure, and recreation (Regan, 1983).
Each hen in a battery house of 30,000 individuals has the
capacity and potential to perform a range of highly
motivated, rewarding behaviours, such as scratching,
dustbathing and uttering and responding to a broad
repertoire of social calls (Hughes and Channing, 1998;
Widowski and Duncan, 2000; Evans and Evans, 1999). That
she is unable to do so constitutes a denial of pleasures by
human interference. Her suffering is compounded by the
frustration of rewards. If we view animals’ interests solely
in terms of avoiding pain and suffering, then the case for
their moral protection appears sound. When we include
their capacity for pleasure, the case is made stronger.
I thank Lori Marino, Hanno Wu
¨rbel, and two anon-
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... Sex for reproductive concerns is at the heart of life not only for human but also for other species (Launer, 2014). In contrast, nonreproductive sex for sexual pleasure is also dominant particularly among humans and the closest extant human relatives such as bonobos or Japanese macaques (see Manson and Parish, 1997;Hull et al., 2002;King, 2004, Sommer andVasey, 2006;Balcombe, 2009 for primate sexual behaviors). According to psychologist Garza-Mercer (2007), sufficient frequency in sexual pleasure drives human behavior. ...
... In another study, Annicchiarico et al. (2020) found out that homosexual ventro-ventral, or genito-genital rubbing between female bonobos facilitates conflict resolution, anxiety reduction and social bonding. As observed among female Japanese macaques (Sommer and Vasey, 2006), same-sex sexual behavior, or sex for pleasure, demonstrates us that pleasure is dominant in ventro-ventral sexual behavior (see also Hull et al., 2002;Balcombe, 2009). Miller (2001, p. 241) describes penis "as a metaphor for the mind's sexually selected entertainment abilities", and "clitoris as a metaphor for the mind's judgment and discrimination abilities". ...
... While the role of positive emotional states and pleasure has been emphasized in animal welfare [163][164][165], wellbeing is elusive and multi-faceted and difficult to define precisely. It is especially hard in species that do not frequently display positive emotional states, social and play behaviour. ...
To understand animal wellbeing, we need to consider subjective phenomena and sentience. This is challenging, since these properties are private and cannot be observed directly. Certain motivations, emotions and related internal states can be inferred in animals through experiments that involve choice, learning, generalization and decision-making. Yet, even though there is significant progress in elucidating the neurobiology of human consciousness, animal consciousness is still a mystery. We propose that computational animal welfare science emerges at the intersection of animal behaviour, welfare and computational cognition. By using ideas from cognitive science, we develop a functional and generic definition of subjective phenomena as any process or state of the organism that exists from the first-person perspective and cannot be isolated from the animal subject. We then outline a general cognitive architecture to model simple forms of subjective processes and sentience. This includes evolutionary adaptation which contains top-down attention modulation, predictive processing and subjective simulation by re-entrant (recursive) computations. Thereafter, we show how this approach uses major characteristics of the subjective experience: elementary self-awareness, global workspace and qualia with unity and continuity. This provides a formal framework for process-based modelling of animal needs, subjective states, sentience and wellbeing.
... No harm has been done before death, and from the moment of death, there is no living animal anymore to have an interest in welfare, and therefore no welfare of an animal is taken away (Broom 2011;Webster 1994). This view is opposed by those who claim that cutting an animal's life short is taking away its future welfare, and therefore is wrong (Balcombe 2009;Bovenkerk and Braithwaite 2016;Deckers 2016;Kagan 2016;Kasperbauer and Sandøe 2016;Singer 1981Singer /2011Višak 2015;Yeates 2010). For preference-utilitarians, fulfilment of preferences is the central value that should be maximized (Singer 1993(Singer /1999. ...
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The aim of this paper is to take normative aspects of animal welfare in corporate practice from a blind spot into the spotlight, and thus connect the fields of business ethics and animal ethics. Using insights from business ethics and animal ethics, it argues that companies have a strong responsibility towards animals. Its rationale is that animals have a moral status, that moral actors have the moral obligation to take the interests of animals into account and thus, that as moral actors, companies should take the interests of animals into account, more specifically their current and future welfare. Based on this corporate responsibility, categories of corporate impact on animals in terms of welfare and longevity are offered, including normative implications for each of them. The article concludes with managerial implications for several business sectors, including the most animal-consuming and animal-welfare-threatening industry: the food sector. Welfare issues are discussed, including the issue of killing for food production.
... Finally, most scholarly work focuses on suffering, pain and stress in animals as negative aspects. Balcombe (2009) argues that positive aspects of sentience such as the seeking of rewards and pleasure through play and touch require a greater degree of investigation in order to amplify "the moral burden of depriving animals the opportunity to lead fulfilling, enjoyable lives" (p. 208). ...
Research on animal ethics continues to escalate in tourism but does so in the absence of a comprehensive statement on animal suffering. This paper aims to rectify this deficiency through two broad aims. The first is to define suffering and related terms and provide a brief statement on the science of suffering; highlight issues related to the ethics of animal suffering; and provide a brief discussion on policy and suffering. The second aim is to develop a roadmap to eliminate animal suffering in tourism as a priority. The theoretical framework adopted in this paper rests within care ethics and empathy.
... Hence, there is a possible limitation of the interpretation of physiological parameters as being unspecific proxies of the emotional intensity. However, positive emotions have received little attention so far (de Vere & Kuczaj, 2016), leading to less available information about physiological reactions to positive than to negative stimuli, but recent reviews should further stimulate such research (Ahloy-Dallaire et al., 2018;Balcombe, 2009;Berenbaum et al, 2016;Boissy et al., 2007). Moreover, biomarkers of emotions in humans, which could help to distinguish between positive and negative emotions, are currently investigated for their usefulness in animal research (e.g., immunoglobulin A: Staley et al., 2018). ...
Les mécanismes qui sous-tendent la personnalité animale (c.-à-d., les différences individuelles de comportement stables à travers le temps et les contextes) sont encore mal compris. Il a été suggéré que la personnalité pourrait émerger à partir de différences individuelles dans les réactions émotionnelles. Cette thèse a pour objectif d’étudier comment la tendance à l’exploration, l’un des traits de personnalité les plus étudiés, est liée aux différences individuelles d’émotions, à différentes classes d’âge chez deux rongeurs d’origine sauvage. Chaque chapitre aborde un composant d’une réaction émotionnelle (comportement, cognition et physiologie), afin d’évaluer la valence ou l’intensité de l’expérience émotionnelle. Tout d’abord, nous avons montré que le taux d’appels d’isolement pouvait être utilisé pour caractériser les profils émotionnels de jeunes souris domestiques, celui-ci étant stable durant trois jours et dans trois situations stressantes. Deuxièmement, nos résultats ont suggéré qu’une tendance plus forte à l’exploration pourrait être liée à une plus grande tendance à exprimer des états affectifs négatifs (c.-à-d., un biais de jugement plus négatif).Troisièmement, nous avons constaté que les souris glaneuses plus exploratrices étaient caractérisées par une réactivité plus forte du système sympathique, exprimée par des températures périphériques de la queue plus basses, peu de temps après une procédure de manipulation brève. Dans l'ensemble, les résultats de ce projet de recherche contribuent à la compréhension de la base émotionnelle des traits de personnalité et soulignent l'importance de prendre en compte l'individualité lors de l'évaluation des émotions.
... Exsitu conservation is a complementary technique to in situ conservation which consists of conserving the biological diversity outside natural habitat, targeting all levels of biodiversity such as genetic, species, and ecosystems [159,160] . The NCL has undergone this kind of conservation, and referring to the studbook of the species by Columbus Riverbed Zoo of South ...
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Carnivores have always undergone interspecific and intraspecific threats that seem tough to detect since carnivore studies often rely on passive sampling when investigating spatiotemporal threats or interactions with human activities. Studies on carnivores’ niche have been an important ecological topic for a long time as carnivore species are crucial in the functioning of ecosystems. This study focused on analysing the coexistence patterns of the North China leopard (Panthera pardus japonensis), the leopard cat (Prionailurus bengalensis) and the red fox (Vulpes vulpes) in a human-dominated landscape, the Tieqiaoshan Natural Reserve (TNR), and to provide insights for implications of carnivores conservation. The objectives of the study were: 1) to make insights on the North China leopard in distribution, threats, conservation and population status; 2) to characterize the spatiotemporal coexistence of the North China leopard, the leopard cat and the red fox in one season of data collection and depict effects of environmental factors on species’ site occupation; 3) to make a multi-year assessment of occupancy, detection and coexistence across three years and evaluate anthropogenic disturbances on carnivores estimates; 4) to analyse the threats of invasive species, including humans, livestock, and domestic dogs, on native carnivore in spatiotemporal patterns. We used the documentary method to bring out results related to the first objective. For the three remaining objectives, in the spatial patterns, we performed the occupancy models, the single-season single-species and single-season two species (where 589 independent photographs from 81 camera traps were analysed), multi-season single-species and multi-season two species (where 81, 62 and 62 camera traps were respectively used in season one, two and three, with 589, 496 and 472 independent photographs respectively) from 2017 to 2019. We estimated three carnivores' site occupation, the environmental factors’ and human disturbances effect on species’ occupancy and detectability. In the last objective, we also estimated the site occupation of invasive species (humans, livestock, and domestic dogs). On the other hand, we calculated the temporal overlap between species using the Kernel Density Estimate through the overlap package in the temporal patterns. The mainly results of this study are followings: 1) We suggested that the North China leopard's current distribution has drastically changed and only 2 % of its historical distribution remains occupied. Extant patches are in continual danger as the proximity index of patches was small which implying lack of connectivity. Habitat fragmentation, retaliation, and decline in prey species are the main threats. However, there is hope in conservation and long term existence in the area for this leopard sub-species for its survival because new management policies are being undertaken and will eradicate or reduce threats. 2) Our study revealed extensive and simultaneous presence, implying high overlapping for space and activities during a broad time period (dawn-morning, and crepuscular) between fox and leopard. The North China leopard and the leopard cat avoided each other. The leopard cat and the red fox independently co-occurred with an overlap in nocturnal time. There was true coexistence between the North China leopard and the red fox. The vegetation continuous cover degree was found to be the most important factor in candidate models for site occupation. 3) In a multi-year pattern, the North China leopard occupancy probability did not markedly change with time as the occupancy equilibrium was constant or slightly enhanced. The occupancy of the leopard cat decreased with time. The occupancy equilibrium of the red fox alternately increased and decreased. However, all species presented a slight level of occupancy stability due to their small values of rate of change in occupancy. Environmental factors and anthropogenic disturbances slightly influenced the occupancy of all species across the years. The colonisation and local extinction for all species were relatively more strongly affected by the distances to villages and roads. Moreover, elevation increased the colonisation and decreased the extinction of the leopard cat. Species interaction factors increased with time for all species. 4) In the invasive and carnivore species’ encroachement, the invasive species did not show substantial changes in the occupancy rate and were well detected. Still, invasive species depicted higher values of occupancy equilibrium than carnivores in both interseasons. Domestic dogs directly co-occur with native carnivores (SIF > 1) while humans and livestock presence have direct (SIF > 1) and indirect (SIF not very high than 1) co-occurrence respectively with the North China leopard and leopard cat and red fox. The leopard cat was the least spatially affected carnivore by the invasive species interactions. In temporal patterns, the North China leopard depicted real temporal activities overlap (high Δ4) with all invasive species compared with the leopard cat and red fox (low Δ4). 5) This research confirmed that environmental factors and human perturbations are vital factors to wild carnivores' coexistence. It also exposes the negative impact of free-ranging invasive species across this protected area on native wild carnivores. An evaluation of how a carnivore species is studied and its coexistence with sympatric and invasive species across diverse protected areas management regimes is crucial to develop robust landscape-scale conservation strategies.
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This paper seeks to expand traditional aesthetic dimensions of design beyond the limits of human capability in order to encompass other species’ sensory modalities. To accomplish this, the idea of inclusivity is extended beyond human cultural and personal identities and needs, to embrace multi-species experiences of places, events and interactions in the world. This involves drawing together academic perspectives from ecology, neuroscience, anthropology, philosophy and interaction design, as well as exploring artistic perspectives and demonstrating how these different frames of reference can inspire and complement each other. This begins with a rationale for the existence of non-human aesthetics, followed by an overview of existing research into non-human aesthetic dimensions. Novel aesthetic categories are proposed and the challenge of how to include non-human aesthetic sensibility in design is discussed.
There is public interest for the welfare of dogs that spend at least part of their lives housed in kennel facilities, such as working, shelter and sporting dogs. The impacts of living in environments that limit social, physical, and behavioral opportunities are generally well understood in other animals, such as livestock and zoo animals. Research exploring the effects of the kennel environment and its enrichment on the behavior and physiology of dogs is emerging. However, human perceptions concerning what is important to the welfare of kenneled dogs have been overlooked. What people believe is important will influence their behavior, with direct relation to care provided to animals and the underlying social license of related industries to operate. This study evaluated the perceived importance of specific kennel management practices relating to canine health, kennel facility design and routine, social interactions, and environmental enrichment. Over 2,000 self-selected adults completed a voluntary, internet-based questionnaire. Differences in beliefs and attitudes were identified based on kennel facility experience, employment role, age, and gender, highlighting potential areas of discordance that may contribute to occupational stress and staff turnover. The results also suggest that research findings published in the scientific literature may not be successfully translating into evidence-based changes in industry practice. Future models to assess animal welfare should include the critical dimension of human-animal interaction. The beliefs, attitudes, and consequent behaviors of people interacting with dogs housed in kennels will determine how living in captivity impacts upon the experiences and welfare of the resident dogs.
An in-depth exploration of animals' capacity for pleasure.
The past few decades have seen a virtual explosion of scientific research in the area of cognition, emotions, suffering, and mental states in animals. Studies in the field, laboratory, and clinical medical practice have amassed an overwhelming body of evidence demonstrating that mental well-being is of paramount importance in all aspects of animal care. There is no longer any reasonable doubt among researchers that mental health is of equal importance as physical health and animal well-being. Recent research convincingly shows that physical health is strongly influenced by mental states, thereby making it clear that effective health care requires attention to the emotional well-being as well as physical. Yet, for its vast importance, mental health in veterinary medicine has to date not been compiled and structured into an organized field or body of knowledge. This information, so critical to the formal establishment of the field of mental health and well-being in animals, remains scattered throughout a wide array of scientific journals. This book represents the first authoritative reference text bringing together the most up-to-date information in the variety of subjects comprising the field of mental health and well-being in animals. Bringing together a host of distinguished experts internationally noted in the fields of animal emotion research, animal behavior, cognitive science, and neuroscience, the book represents the first authoritative reference compiling the diverse information on the animal mind and combining the revolutionary advances in the cognitive sciences with the knowledge in veterinary medicine and clinical animal behavior. This book takes a descriptive and proscriptive approach to mental health, mixing the scientific research with practical information with clinical applications for veterinary health professionals to use in practice.
Biological Exuberance by Bruce Bagemihl. textlessPtextgreaterA Publishers Weekly Best BookOne of the New York Public... Bonus Publisher Materials: Excerpt, Praise, Author Biography, Awards
Suffering is a state of mind that is difficult to measure and analyse in human beings and considerably more so in animals. It is related to the environment in which we live and our physical and mental states. Understanding the physiology of suffering in animals is crucial in assessing animal welfare. Written by an expert in applied welfare aspects of physiology, this book is the first to address the physiological aspects of suffering in animals. It explores the different causes of suffering - physical discomfort, thirst and hunger, the responses in the body that lead to suffering and it offers insight into how suffering can be managed. The second book in a major new animal welfare series Draws together information that is scattered across the literature Written for the specialist and non-specialist alike Includes colour pictures This book is part of the UFAW/Wiley-Blackwell Animal Welfare Book Series. This major series of books produced in collaboration between UFAW (The Universities Federation for Animal Welfare), and Wiley-Blackwell provides an authoritative source of information on worldwide developments, current thinking and best practice in the field of animal welfare science and technology. For details of all of the titles in the series see