Vet Times

The website for the veterinary profession

https://www.vettimes.co.uk

Palatability: feline food preferences

Author : Tim Watson

Categories : Vets

Date : May 30, 2011

Tim Watson discusses how getting cats to eat can be influenced by factors such as the animal’s
own dietary requirements – and that of its mother

Summary
Palatability describes how readily a food is accepted and measured in terms of its attractiveness
and consumption. It has long been believed that palatability of cat foods is determined by sensory
characteristics – notably smell, taste and mouth-feel – and previous experiences. Cats are highly
specialised carnivores, with high protein requirements, and this is reflected in their sense of taste.
They are attracted to some amino acids, allowing them to identify foods rich in protein, but avoid
bitter stimuli and do not select foods on the basis of their sugar or salt content. Preferences are
also strongly shaped by experiences as kittens, with early exposure influencing food choices in
later life. Cats become averse to long-term feeding of a single food, which may be a mechanism to
avoid potential nutritional imbalances associated with monotonous consumption and reflects an
inherent drive to seek variety. Recent research has shown that cats demonstrate “nutritional
wisdom” and select foods in order to meet specific intake targets of protein, fat and carbohydrate.
This new science may reframe our understanding of palatability and indicates that underlying
nutrition plays a far greater role than previously understood. In fact, tastants or palatability cues
may simply be sensory mechanisms by which cats identify foods with a good nutritional
composition, mediating an initial attraction that is only sustained if the diet proves to be of
appropriate nutritional quality.

Key words

cat, palatability, food preference, macronutrient target

1/8
PALATABILITY, along with nutritional quality and digestibility, is a key attribute of pet foods
and is as important to manufacturers as it is to cats and their owners.

But what does the term palatability actually refer to? In addition, what determines whether a cat will
eat a food and, if so, how much?

Nutrition scientists have worked hard to understand what makes a food attractive to cats. This
knowledge is explored in this article, in terms of the dietary evolution of cats, what we know about
their sensory physiology and how – together with dietary experiences – it shapes domestic cats’
feeding behaviour.

Research has, however, challenged the view that cats simply select foods based on sensory
properties and previous experiences.

It provides compelling evidence that cats, when given freedom to choose their own diet, adapt the
intake of key nutrients to meet specific targets.

This suggests that nutritional content is a key driver of palatability, with tastants acting as a
“navigational aid” to help identify nutritional composition and quality.

What does palatability actually mean?

The purpose of food is to provide energy and essential nutrients, but this can only be achieved if
the animal is willing to eat it. This willingness is not necessarily a measurable property of the food
because it depends on a number of individual factors that influence whether a meal is consumed.

These include the animal’s sensory perception of that food, as well as the shaping of acceptance
by previous experiences and genetic variation.

It is thus an animal’s response to a food that is defined by the term palatability. With regards to pet
foods, this is usually measured by how much is eaten on a meal or daily basis, or whether diet A is
preferred to X, Y or Z when different products are offered simultaneously. A palatable diet could,
therefore, be sufficiently acceptable to cats so that they consume amounts adequate to meet
requirements for energy and essential nutrients – or it might simply be preferred by more cats.

What determines the palatability of a food?

The key determinants of a food’s attraction are its sensory characteristics – notably smell, taste
and mouth-feel. These features, together with learning from previous experiences with the same or

2/8
similar foods and possible genetic factors, have traditionally been believed to shape food choices
made by cats and have been linked to their nutritional evolution.

Cats are highly developed carnivores – technically termed hypercarnivores – that have evolved to
consume almost exclusively vertebrate prey1. This is reflected in their nutritional needs, dental
anatomy and digestive physiology, as well as their taste systems.

Unlike dogs, which hunt in packs, cats are solitary hunters and so usually take live prey, such as
mice, voles, small rabbits and birds, with a body mass less than their own. The habit of consuming
live prey has led to a preference for foods at body temperature and for multiple, small meals
throughout the course of the day.

The feline taste system is similar to other carnivores, but, as they have evolved as obligate meat
eaters (Figure 1), it is further specialised to provide for their high requirements of protein and specific
amino acids, such as methionine and cysteine, and need for taurine and arginine that can only be
met by the consumption of animal tissues2.

Cats have highly sensitive taste receptors to acidic and bitter stimuli, which allow them to identify
foods rich in selected amino acids and avoid bitter or sour stimuli associated with potentially toxic
ingredients3,4. It is, therefore, not surprising that in palatability tests, cats prefer foods that contain
substances associated with meat and fishbased diets5.

Cats lack sweet taste receptors and do not select foods on the basis of their sugar content6. This
insensitivity to sugars is unlikely to be a problem for cats, as they derive little nutritional benefit from
sugar-containing foods, such as fruits. Their preybased diet contains less than five per cent
carbohydrate on a calorie basis7. It has been suggested that the inability to taste sugars enables
better appreciation of essential amino acid balance within foods by removing any masking effect
from sugars8.

Although the precise contribution of odour to palatability is less clear, cats are believed to have an
acute sense of smell that works in tandem with taste to inform about a food’s suitability and safety.
Form and texture are important, with the particle size of dry foods and viscosity or stickiness of wet
foods being prominent factors (Figure 2)9.

Effects of experience and genetics

It is clear that cats do modify their food preferences based on experience. In the wild, this ensures
they avoid repeating disadvantageous feeding experiences associated with nutritionally incomplete
or potentially toxic prey. Domesticated cats also express this behaviour, although it is most
commonly manifested as a growing aversion towards foods that form a large part of their diet10.

This so-called “monotony effect” reduces the perceived palatability of habitual foods in favour of a

3/8
novel food with contrasting sensory characteristics. This could represent a mechanism by which
cats avoid long-term consequences associated with consuming foods that are unbalanced in one
or more nutrients. Alternatively, it may reflect a hard-wired drive to seek variety – a survival
strategy that avoids dependence on a single food source or, indeed, minimises the risk of depleting
a single source of prey.

Powerful monotony effects are observed in kittens as well as adult cats, and appear to be stronger
in free-ranging cats than those living in households and exclusively fed nutritionally complete
diets11.

This supports the belief that the monotony strategy works on the assumption that two foods, with
markedly different flavours, are not both likely to be nutritionally incomplete.

It is during early life that food selection is most plastic and easily modified, with kittens strongly
influenced by the preferences of their mothers (Figure 3)12. This effect stems from the mother bringing
food to the kitten, as well as her simply being present when a new food is introduced13.

It is also possible that maternal influence starts before weaning, with the transfer of flavours via
amniotic fluid and milk. This is supported by the observation that kittens born to mothers trained to
eat unusual foods, such as potato and banana, will select their mother’s diet in preference
to meat14. A more recent study confirmed this, showing that kittens exposed to cheese flavours
during uterine and early postnatal life via their mother’s diet were more likely to select cheese-
flavoured foods15.

The bias that cats show towards their mother’s diet may be a manifestation of neophobia, the
magnitude of which is related to the contrast between the new foods’ sensory attributes and those
of the foods they, or their mother, were accustomed to. This effect can be reduced by weaning
kittens to varied diets, which results in them having broad food preferences throughout life8. The
triggering of neophobia appears to be reduced in these instances, because any new food is likely
to have some similarity to those already experienced. Genetics appear to play only a small role in
determining food preferences, with differences between cat populations more readily explained by
their feeding experiences, rather than genetic diversity16.

Importance of nutrition in determining food palatability

Compared with other species, relatively little information is available about the role of specific
nutrients in food selections made by cats.

It is known that they do not select foods on the basis of their sweetness3 or salt content17, and
some evidence suggests cats can develop learned aversions or preferences when foods are
lacking in certain nutrients.

4/8
Because of the cat’s high protein requirement and nutritional peculiarities related to amino acid
requirements, it is the response to dietary amino acids that has been most frequently studied. Cats
exhibit a severe depression in food intake, together with a learned taste aversion, when fed a diet
deficient in arginine18.

They also show initial preference for diets containing adequate methionine over methionine-free
diets, although this preference is not maintained beyond two days of feeding19 and cats will tend to
avoid diets lacking isoleucine20.

By contrast, kittens willingly consume diets with excess leucine21 and adult cats continue to
consume diets deficient in essential taurine, despite their taste system having the ability to detect
this amino acid22.

These studies indicate that the metabolic feedback from specific amino acids is complicated and
raises the question of whether they select proteins, rather than essential amino acids. A prev ous
attempt to answer this led researchers to conclude that cats do not select for or against protein,
even if this results in the consumption of protein-deficient diets, with the consequent detriment to
growth23.

Evidence has emerged that minks, which are also hypercarnivores, are able to modify feeding
behaviour to balance protein and fat intake24.

This prompted re-evaluation of the effects of protein, as well as fat and carbohydrate, on cats’ food
selection25.

This showed that cats modify choices and consumption of foods to regulate their intake of protein,
fat and carbohydrate and achieve a target balance. Cats consistently choose to eat diets with
specific macronutrient profiles equivalent to a daily intake of around 420kJ protein, 280kJ fat and
100kJ carbohydrate (Figure 4). These values equate to a diet containing 52 per cent of energy as
protein, 36 per cent as fat and 12 per cent as carbohydrate.

The research also revealed how cats prioritise nutrient intakes when fed diets that do not enable
them to meet targets. When given foods high in carbohydrate and lower in protein, they did not eat
more to achieve the protein intake target because of a ceiling on carbohydrate intake of 300kJ/d
(20.5g/d). There was more flexibility in consumption of fat and protein, with cats overshooting their
target intakes to meet protein or energy requirements.

What does research tell us about nutrition and palatability?

These latest findings indicate that the palatability of cat foods is intimately linked to their nutritional
profile, with cats choosing foods to meet an “intake target” of protein, fat and carbohydrate. The
benefit to the cat of achieving the intake target is not clear, although in other species it has been

5/8
suggested as a target to support optimal performance26.

Given how cats have evolved, it comes as no surprise that the targets expressed are similar to the
composition of their natural vertebrate prey, particularly in relation to protein and fat, and queens’
milk (Table 1). They do, however, differ with regard to carbohydrate.

While domestic cats selected diets with 12 per cent energy from carbohydrates (8g/day to
10g/day), a prey-based diet of six to 12 small rodents would contain – at most – 3g of carbohydrate
that is equivalent to just four per cent of energy. The reasons for this discrepancy remain unclear.

When the macronutrient targets are compared with the composition of commercially available cat
foods, meat-based wet foods have protein content similar to the target, but they contain slightly
more fat (Figure 5). Dry diets have less protein, but a similar fat content. The most obvious difference
is in relation to carbohydrate – wet foods contain minimal amounts of carbohydrate (less than 15
per cent energy) compared with up to 40 per cent in dry foods.

This raises the question of whether wet cat foods are inherently palatable because their nutritional
profile is more closely aligned to intake targets.

If this is the case, palatability may reflect a response to nutritional adequacy, rather than just
sensory properties. Indeed, the inherent nutritional wisdom shown by cats suggests that no amount
of tastants or attractants will overcome the food’s basic nutritional quality.

While the majority of cats readily consume foods in either wet or dry formats, it is interesting to
speculate whether macronutrient targets are linked to the monotony effect seen with long-term
feeding of a single food. Could this be a mechanism by which owners are forced to offer alternative
foods, thereby satisfying the cat’s need for variety while allowing them to select foods that enable
them to meet nutritional targets?

References
1. Holliday J A and Steppan S J (2004). Evolution of hypercarnivory: the effect of
specialization on morphological and taxonomic diversity, Paleobiology 30: 108-128.
2. Zoran D L (2002). The carnivore connection to nutrition in cats, J Am Vet Med Assoc 221:
1,559-1,567.
3. Beauchamp G K, Maller O and Rogers J G (1977). Flavor preferences in cats (Felis catus
and Panthera species), J Comp Physiol Psych 91: 1,118-1,127.
4. Carpenter J A (1956). Species differences in taste preference, J Comp Physiol Psych 49:
139-144.
5. Mugford R A (1977). External influences on the feeding of carnivores. In Kane M R and
Maller O (eds), The Chemical Senses and Nutrition, Academic Press, New York: 25-50.
6. Li X, Li W, Wang H, Bayley D L, Cao J, Reed D R, Bachmanov A A, Huang L, Legrand-

6/8
 Defretin V, Beauchamp G K and Brand J G (2006). Cats lack a sweet taste receptor, J Nutr
136: 1,932S-1,934S.
7. Eisert R (2011). Hypercarnivory and the brain: protein requirements of cats reconsidered, J
Comp Physiol B 181: 1-17.
8. Bradshaw J W S (2006). The evolutionary basis for the feeding behaviour of domestic dogs
(Canis familaris) and cats (Felis catus), J Nutr 136: 1,927S-1,931S.
9. Kane E (1989). Feeding behaviour of the cat. In Burger I H and Rivers J P W (eds),
Nutrition of the Dog and Cat, Cambridge University Press, Cambridge: 147-158.
10. Thorne C J (1982). Feeding behaviour in the cat – recent advances, J Small Anim Pract 23:
555-562.
11. Church S C, Allen J A and Bradshaw J W S (1996). Frequencydependent food selection by
domestic cats: a comparative study, Ethology 102: 495-509.
12. Stasiak M (2002). The development of food preferences in cats: the new direction, Nutr
Neurosci 5: 221-228.
13. Wyrwicka W and Long A M (1980). Observations on the initiation of eating of new food by
weanling kittens, Pavlov J Biol Sci 15: 115-122.
14. Wyrwicka W (1978). Imitation of mother’s inappropriate food preference in weanling
kittens, Pavlov J Biol Sci 13: 55-72.
15. Becques A, Larose C, Gouat P and Serra J (2010). Effects of pre- and postnatal
olfactogustatory experience on early preferences at birth and dietary selection at weaning in
kittens, Chem Senses 35: 41-45.
16. Bradshaw J W, Healey L M, Thorne C J, Macdonald D W and Arden-Clark C (2000).
Differences in food preferences between individuals and populations of domestic cats (Felis
silvestris catus), Appl Anim Behav Sci 68: 257-268.
17. Cook N E, Roger Q R and Morris J G (1996). Acid-base balance affects dietary choice in
cats, Appetite 26: 175-192.
18. Morris J G and Rogers Q R (1978). Arginine: an essential amino acid for the cat, J Nutr
116: 655-667.
19. Rogers Q R, Wigle A R, Laufer A, Castellanos V H, Morris J G (2004). Cats select for
adequate methionine but not threonine, J Nutr 134: 2,046S-2,049S.
20. Hargrove D M, Rogers Q R, Calvert C C and Morris J G (1988). Effects of dietary excesses
of the branched-chain amino acids on growth, food intake and plasma amino acid
concentrations of kittens, J Nutr 118: 311-320.
21. Hargrove D M, Rogers Q R, and Morris J G (1994). Kittens choose a high leucine diet even
when isoleucine and valine are the limiting amino acids, J Nutr 124: 689-693.
22. Sturman J A, Garano A D, Messing J M and Imaki H (1986). Feline maternal taurine
deficiency: effect on mother and offspring, J Nutr 116: 655-667.
23. Cook N E, Kane E, Rogers Q R and Morris J G (1985). Self-selection of dietary casein and
soy-protein by the cat, Physiol Behav 34: 583-594.
24. Mayntz D, Nielsen V H, Sørensen A, Tøft S, Raubenheimer D, Hejlesen C and Simpson S J
(2009). Balancing of protein and lipid intake by a mammalian carnivore, the mink, Mustela
vison, Anim Behav 77: 349-355.

7/8
 25. Hewson-Hughes A K, Hewson-Hughes V L, Miller A T, Hall S R, Simpson S J and
Raubenheimer D (2011). Geometic analysis of macronutrient selection in the adult
domestic cat, Felis catus, J Exp Biol 214: 1,039-1,051.
26. Simpson S J, Sibly R M, Lee K P, Behmer S T and Raubenheimer D (2004). Optimal
foraging when regulating intake of multiple nutrients, Anim Behav 68: 1,299-1,311.
27. Keen C, Lonnerdal B, Clegg M S, Morris J G, Rogers Q R and Rucker R B (1982).
Developmental changes in the composition of cats’ milk: trace elements, minerals, protein,
carbohydrate and fat, J Nutr 112: 1,763-1,769.

//

8/8
Powered by TCPDF (www.tcpdf.org)