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19 The Nutrition of Companion Animals

The Nutrition of Companion

Animals
A.C. Longland1, M.K. Theodorou1 and I.H. Burger2
1Institute
of Grassland and Environmental Research, Plas Gogerddan,
Aberystwyth, Ceredigion, UK; 2Waltham Centre for Pet Nutrition,
Melton Mowbray, Leicestershire, UK

INTRODUCTION
Companion animals are those species which have a special association with
man and are dependent on him to some degree (Greenhalgh and Corbin,
1998), the most popular in the UK being dogs, cats, fish, birds, horses,
certain rodents and rabbits. Frequently the relationship between man and the
companion animal is sentimental, although for dogs and horses the relation-
ship may be more practical, their being used for racing, hunting, herding, aid-
ing the disabled or for military and police work. The aim of feeding
companion animals, irrespective of their role, is to promote a long and healthy
life and, to achieve this goal, many are fed at or near their maintenance levels
throughout adulthood. Feeding to maintain rather than increase body weight
and to accommodate changes in metabolism during ageing are pertinent
considerations for the companion animal, but are largely irrelevant in farmed
species. Therefore, conceptually, the nutrition of companion animals is more
akin to that of humans than that of the other species described in this book,
which are seldom allowed to achieve their natural life span but are fed to elicit
the most effective production of meat, milk, eggs or wool.
The 1996 statistics for companion animals in the UK suggest that approx-
imately one-quarter and one-fifth of UK households owned dogs and
cats, respectively, representing a total of 7.5 million of both species, and the
number is rising (PFMA, 1997). The size of the companion animal food market
is thus considerable and is estimated to be worth $9.2 billion in Europe alone
(FEDIAF, 1996).
With the exception of horses, which are discussed in Chapter 11, this
chapter will review the nutrition of dogs, cats, birds, fish, rabbits and rats
through their different life stages of growth, maintenance, gestation and
lactation. For dogs, the nutrition of working and senior animals is also
discussed. Clinical nutrition, to combat or aid recovery from specific
CAB International 2000. Feeding Systems and Feed Evaluation Models
(eds M.K. Theodorou and J. France) 435

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436 A.C. Longland et al.

conditions, will only be touched upon where it is normally associated with the
ageing process, but is otherwise outside the scope of this text.

NUTRIENT REQUIREMENTS
The term nutrient requirement refers to the quantity of an essential nutrient
needed by the animal for a given physiological state (Morris and Rogers, 1994).
Although single values for a particular life stage may be presented, these can
only be guidelines, as the nutrient requirements of any given individual within
a population will vary in relation to: (i) its physiological state; (ii) the metabolic
function that is to be sustained or optimized; and (iii) the genetic potential of
the animal. Values quoted by the National Research Council (NRC) usually
reflect minimum requirements and are often determined with highly purified
ingredients; they give no margin of safety for varying bio-availability of
nutrients in relatively unprocessed diets and take no account of variation of
ingredients, interactions between them or indeed interlaboratory variation in
analysis of samples. Therefore, the recommended mimima presented here
should be regarded with caution, particularly when formulating diets from raw
materials. Furthermore, some nutrients are toxic above a certain level and,
in practical terms, the amounts of such nutrients in a diet must lie between
deficiency on the one hand and toxicity on the other. Wherever possible, the
nutrient requirements for companion animals presented here are based on the
results of experimental trials on the target species. Where such information is
not available, recommendations are based on diets that have been shown to
support the target species throughout the different life stages.

ENERGY
Energy is required for growth, maintenance, reproduction and lactation.
Energy is provided by carbohydrate, fat and protein, which typically have
gross energy (GE) values of 17.4, 39.3 and 23.3 kJ g−1, respectively. The differ-
ence between GE intake and that in the faeces is the digestible energy (DE)
of a diet, metabolizable energy (ME) being the DE less the energy in urine
and combustible gases. Due to the complexities involved in measuring the
combustible gases, reported ME values usually reflect DE minus urinary
energy. As ME values are reasonable estimates of the energy available to the
animal, ME values are quoted for preference in this chapter.

Fat
In addition to being a concentrated energy source, fat provides essential
fatty acids (EFAs) and is a carrier for the fat-soluble vitamins. The EFAs are
the (n-6) series, consisting of linoleic, γ-linolenic and arachidonic acids.
Requirements for the (n-6) EFAs are usually expressed in terms of linoleic acid

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The Nutrition of Companion Animals 437

as this can usually be converted into linolenic and arachidonic acids via
desaturation or elongation, respectively (and because linoleate and
arachidonate are normally minor constituents of most fats). The α-linolenate
(n-3) family are also EFAs for some species (Holman et al., 1982; McCartney,
1996). Signs of EFA deficiency include scurfy coats, hair loss, anaemia, fatty
liver and reduced fertility (Wills, 1996a). In addition to the nutritional
characteristics of fat, the organoleptic properties of diets can be enhanced by
its inclusion (NRC, 1986).

Carbohydrates
All animals require glucose to supply energy to the tissues and as a substrate
for the production of glycoproteins, etc. Glucose is provided either directly,
through degradation of carbohydrates or via the metabolism of gluconeogenic
substrates such as glycerol or amino acids. Carbohydrates not digested by the
animals’ enzymes are potentially available for fermentation by the gut
microflora, the resulting volatile fatty acids (VFAs) being used as an energy
source.

Protein and Amino Acids

Plants and microorganisms can synthesize proteins from nitrogenous
compounds, but animals cannot synthesize the amino group and thus require
a supply of dietary amino acids. Some amino acids can, via transamination, be
elaborated from other amino acids, but others, the indispensable amino acids
(IAAs), cannot be produced in this manner in sufficient quantities by the
tissues to support normal function. For most species, there are ten IAAs;
arginine, histidine, isoleucine, leucine, lysine, methionine, phenylalanine,
threonine, tryptophan and valine. Arginine has been shown to be involved in
the release of several metabolic hormones, including insulin, glucagon, growth
hormone and prolactin, and has also been implicated in the immune response
(Barbul et al., 1981). The branched chain amino acids, leucine, isoleucine and
valine, are required for protein synthesis, protein turnover and energy metabo-
lism (Adibi, 1980). Methionine and cysteine are the sulphur amino acids and
are limiting amino acids in many protein sources (Allison et al., 1974). Various
factors can affect the sulphur amino acid requirement, for example in diets
deficient in methyl donors such as choline the requirement for methionine is
higher. Phenylalanine and tyrosine are the aromatic amino acids and may
substitute for one another to some extent.
The IAA profiles required at each of the life stages will differ because the
utilization of amino acids for protein turnover is different from that for net
tissue accretion. Consequently, the amount of protein required depends on the
source: smaller amounts of highly digestible protein with amino acid profiles
which closely complement the animals’ requirement need to be fed than those
which are less digestible and match the animals’ demands less precisely.

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438 A.C. Longland et al.

Minerals
The ash fraction of a diet consists of minerals; the macrominerals that are
required at relatively high concentrations and the microminerals of which only
trace amounts are needed. Minerals are essential for maintaining, at various
levels, acid–base balances, tissue structure and osmotic pressure, in addition to
being key components in many enzyme systems. There are many
inter-relationships between minerals, and feeding excessive levels of some
minerals may prove injurious.

Macrominerals
Calcium and phosphorus have a close metabolic association, and the ratio of
these is of importance for many species. Their utilization is dependent upon
an adequate supply of vitamin D. Calcium deficiency is usually associated with
osteoporosis, detachment of the teeth, bone resorption, rickets in young
animals, tetany and reproductive incompetence. Calcium is also involved in
blood clotting. Phosphorus deficiency can result in rickets, pica, poor growth
and osteomalacia. A high phosphorus:calcium ratio can lead to symptoms of
calcium deficiency. Potassium is required for neural transmission, fluid
balance, muscle metabolism and immune function, and its deficiency results in
poor growth, muscular paralysis and lesions to the kidney and heart. Sodium
and chloride are the major electrolytes of extracellular fluids, deficiencies
leading to fatigue, poor water balance, retarded growth and hair loss. Proper
muscle and nervous tissue function depend on a correct magnesium and
calcium balance, and magnesium ions are essential for enzymes involved with
energy metabolism. Magnesium deficiency can lead to anorexia, poor growth,
ataxia and convulsions, and is enhanced by elevated levels of dietary calcium
and phosphorus (Bunce et al., 1962a,b). Iron is an essential component of
haemoglobin, myglobin and haem enzymes, deficiency being characterized by
anaemia. Iron is more available from animal rather than plant sources, and
soya protein is thought to reduce absorption of iron, zinc and manganese
(Burger, 1995). Iron also interacts with calcium metabolism. Copper is a
constituent of many enzyme systems and is also closely linked to iron metabo-
lism; copper deficiencies can impair haemoglobin synthesis. Excess levels of
both iron and copper can be toxic. Zinc is required for the activity of enzymes
involved in the metabolism of nucleic acids, carbohydrates, proteins and fats.
Zinc is also needed for maintaining healthy keratinized tissues and germinal
epithelia, and deficiencies are characterized by poor growth, anorexia and
testicular atrophy (Burger, 1995). Iodine is needed for thyroid function,
deficiencies leading to hypothyroidism and goitre, compromised utilization of
calcium and phosphorus and reproductive failure. Manganese is important in
bone formation and reproduction. There are strong interactions between
selenium and vitamin E (NRC, 1978a), selenium being a component of
glutathione peroxidase which protects cells against lipid peroxides. Selenium
deficiencies result in muscular weakness, anorexia and, ultimately, coma and
death.

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The Nutrition of Companion Animals 439

Microminerals
Chromium is needed for insulin function; fluoride for tooth and bone develop-
ment and possibly reproductive function; nickel for membrane function;
molybdenum is a cofactor for many enzymes; silica is required for skeletal
development and connective tissue formation; vanadium for reproduction,
growth and fat metabolism.

Vitamins
Vitamins are organic compounds essential to metabolic regulation. They may
be fat soluble (A, D, E and K ) or water soluble (the B vitamins and vitamin C).
The former are capable of being stored in the body, whereas the water-soluble
vitamins are not stored and must be ingested regularly. Vitamin A is necessary
for visual pigments, maintenance of cellular structure and for healthy skin,
mucous membranes, tooth and bone development. Deficiencies cause night
blindness, infertility, seborrhoeic coat conditions and xerophthalmia. Toxicity
results in liver damage and ankylosis of the joints. Vitamin C is required for
protein synthesis and a number of other intracellular processes but, with the
exception of the guinea pig, can be synthesized by most companion animal
species, precluding the need for its dietary provision. Vitamin D requirements
are associated with dietary levels of calcium and phosphorus and it stimulates
calcium absorption from the gut. In some species, vitamin D may be
synthesized in cutaneous lipid after exposure to the sun; deficiencies result in
rickets in growing animals and osteomalacia in adults. Excess vitamin D can
result in hypercalcaemia, calcification of lungs, kidneys and gut, deformation
of teeth and jaw, and death. Vitamin E, together with selenium, protects
cellular membranes against oxidative damage: in diets rich in easily oxidized
polyunsaturated fatty acids (PUFAs) the requirement for this vitamin is
increased. Vitamin E deficiency can cause skeletal muscular dystrophy,
reduced reproductive performance and a compromised immune system.
Vitamin K may be synthesized by the gut bacteria and is involved in blood
clotting; if bacterial synthesis is reduced, haemorrhaging may occur. Thiamin
(B1) is involved in carbohydrate metabolism, symptoms of deficiencies include
anorexia, neurological abnormalities and heart failure. Riboflavin (B2) is
required in oxidative enzyme systems and for cellular growth, deficiencies
leading to lesions of the eye and skin and testicular hypoplasia. Pyridoxine
(B6) is required for protein metabolism; deficiencies are exacerbated with
high-protein diets and cause anorexia, anaemia and kidney damage, especially
in cats. Cyanocobalamin (B12) is involved in fat and carbohydrate metabolism
and is needed for myelinization of nerves. Deficiencies lead to pernicious
anaemia and neurological defects. Pantothenic acid is a constituent of
coenzyme A and is involved in nutrient metabolism; deficiencies are rare but if
they do occur are manifested by reduced growth, fatty liver, gastrointestinal
disorders and convulsions. Biotin is synthesized by gut bacteria and is
involved in lipid and amino acid metabolism. Raw egg contains avidin which
will complex with biotin rendering it unavailable. Biotin deficiencies lead to

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440 A.C. Longland et al.

poorly keratinized structures. Folate can be synthesized by the gut bacteria and
is required for DNA synthesis and maturation of red blood cells in bone
marrow. Symptoms of deficiency include anaemia and leucopenia. Choline is a
constituent of phospholipids and is a precursor of the neurotransmitter,
acetylcholine.

DOGS

Energy
Dogs can live for up to 20 years and exhibit a very large intraspecific variation in
mature body size, ranging from a 1 kg Chihuahua to a 115 kg St Bernard,
necessitating the need to express energy in terms of metabolic body weight (kg).

Maintenance
A number of values for the maintenance requirements for adult dogs have
been reported, reflecting the wide range of breeds (and, therefore, different
body shapes and coat types) kept under a variety of environmental conditions
and the exercise regimes used in the measurements. Values of 423, 490 and
523 kJ of ME W−0.75 day−1 were determined for Labrador–Retrievers, Beagles
and Siberian Huskies, respectively, housed in indoor kennels (maintained at
~ 20°C) with outdoor runs (at −6 to 24°C) (Finke, 1991). Manner (1991),
however, using eight breeds of dog, suggested values of 387 and 450 kJ of ME
W−0.75 day−1, respectively, for resting and active dogs maintained at 20°C.
These values are somewhat lower than the 678 kJW−0.64 day−1 for dogs at rest
and 643 kJW−0.73 day−1 for active dogs (covering 10.5 km day−1) obtained by
Burger and Johnston (1991) using whole-body calorimetry. On the other hand,
Kienzle and Rainbird (1991) observed that apart from Great Danes and
Newfoundlands, which were above and below the average respectively, the
maintenance energy requirement for most breeds was close to 495 kJ of
ME W−0.75 day−1. The above values for the maintenance energy requirement of
dogs are not comprehensive and they are lower than the 552 kJ of ME W−0.75
day−1 given by the NRC (1974). For practical purposes, these values should be
taken as initial guidelines for diet formulation: the amounts fed can then be
adjusted according to the responses of individual animals.

Growth
Post-weaning, puppies grow rapidly and generally attain about half of their
adult weight at 6 months. Thereafter, different breeds of dog will grow at
different relative rates, larger breeds taking longer than smaller breeds to reach
their mature weight. Puppies initially require twice as much energy as an adult
on a body weight basis, but this difference reduces with increased puppy size.
Thus by the time puppies are 40 and 80% of their mature weight, they need
1.6- and 1.2-fold of the adult requirement, respectively (Wills and Morris,
1996). However, research on Great Danes suggests that for this breed at
least, puppies at weaning and at 5 months old (40% of mature weight) require

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The Nutrition of Companion Animals 441

3.6- and 2.8-fold, respectively, of the adult maintenance energy requirement

(Meyer and Zentec, 1991). High levels of energy intake can cause rapid growth
at the expense of proper skeletal development, and several of the large and
giant breeds are disproportionately susceptible to skeletal disorders such as
hypertrophic osteodystrophy and osteochondrosis.

Gestation and lactation

The majority of fetal growth of puppies occurs in the last third of gestation
and, despite early development of mammary and uterine tissues, the energy
requirements of the gravid bitch do not increase markedly before this time
(Wills, 1996b). During the final trimester of pregnancy, the energy require-
ments of the bitch should be satisfied by increasing her ration by 15% per
week so that at parturition she is consuming 60% more than at pre-conception
(Legrand-Defretin and Munday, 1995). At peak lactation, depending on the age
and size of the litter, bitches may need up to four times their maintenance
energy requirement. Thus to enable lactating bitches to ingest sufficient food
to remain in energy balance, they should be offered several small, nutritionally
balanced, nutrient-dense meals per day.

Lipid
The apparent digestibility of various plant and animal fats by dogs has been
shown to vary from 0.8 to 0.95 (James and McCay, 1950; Orr, 1965). The
current NRC (1985) recommendation for dietary fat for dogs at maintenance is
50 g kg−1 (to include 10 g of linoleic acid kg−1); in diets containing 16.7 MJ of
ME kg−1, these values are very similar to the recommendations of Wills (1996b)
of 3.2 g of fat MJ ME−1 (to include 0.66 g of linoleic acid MJ ME−1) for all life
stages. Puppies from 2 to 6 months have been shown to grow well on diets
which supply 13–76% of the energy from fat and 20–25% of the energy from
high-quality protein (Rosmos et al., 1976). This demonstrates that if the IAA
requirements are met, puppies grow well on diets with a wide range of fat
contents. For giant breed puppies, Lepine (1998) has indicated that diets
containing 140 g of fat kg−1 of diet are suitable.
Studies on the appropriate levels of fat for diets for breeding bitches are
few, but it appears that as long as the balance of nutrients to energy are met,
satisfactory reproductive performance will occur. Ontko and Phillips (1958)
fed diets containing 80 and 160 g of fat kg−1 to breeding bitches, resulting in
satisfactory reproductive performance.

Carbohydrates
Carbohydrates are a relatively cheap energy source, and cereal starches,
particularly those which have been dextrinized, are normally well digested by
dogs, coefficients in excess of 0.84 being reported (Rosmos et al., 1981). Dogs

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442 A.C. Longland et al.

do not appear to have a specific requirement for carbohydrate provided that

there are sufficient dietary gluconeogenic amino acids and glycerol (Blaza
et al., 1989); however, adequate levels may be hard to attain at times of high
demand such as gestation and lactation, resulting in bitches becoming
hypoglycaemic and ketotic before parturition, accompanied by high mortality
of pups (Rosmos et al., 1981). As in rats, fetal abnormalities and resorption
may also occur in bitches fed carbohydrate-free diets at conception (Taylor
et al., 1983). It is therefore recommended that bitches receive some available
carbohydrate in their diets, particularly if protein levels are close to the
nutritional requirement.
Although dietary fibre is not normally regarded as essential for dogs, some
inclusion in the diet may be beneficial, though excessive amounts may lead to
a reduction in availability of other nutrients (NRC, 1985). However, there
is recent evidence that feeding fermentable dietary fibre aids in maintaining
glucose homeostasis in diabetic dogs (McBurney et al., 1998) and
fructo-oligosaccharides (FOSs) may have beneficial effects in managing a
number of conditions, including corneal lipidosis (Diez et al., 1998).

Protein and Amino Acids

Maintenance levels of protein for adult dogs have been given as 9.6 g of
protein MJ ME−1 (Wills, 1996a), corresponding to 160 g of protein kg−1 in a
diet containing 16.7 MJ of ME kg−1. These values are somewhat higher than
the range of 65–120 g of casein kg−1 quoted by Kade et al. (1948), thus
accommodating the use of protein sources of lower biological value than
casein.
Puppies require protein for tissue synthesis during growth as well as for
maintenance. Protein quality is important, depending upon both the IAA
profile of the protein source and their availability. Work by Nierinck et al.
(1991) illustrates this point for although the amino acid profiles of three
protein sources of animal origin (lung, tripe and mince) and soya meal are
similar, crude protein digestibility coefficients averaged 0.92 for the animal
proteins and 0.76 for the soya meal. The diets in this study were formulated
such that the intakes and digestibility of the four protein sources by Beagles
should have resulted in all of the amino acids being available to them at above
the NRC (1985) recommended minima. However, protein deficiencies may
develop if vegetarian diets are formulated to lower protein levels without
knowledge of protein availability. Indeed, in a field study of diets fed to
vegetarian dogs, Kienzle and Engelhard (1998) found that protein supply to
adults was marginal and some puppies showed signs of protein deficiency.
Minimum protein levels of 110–220 g of protein kg−1 for growing dog diets
have been quoted (Johnson, 1993), and the NRC (1985) concluded that 120 g
of protein kg−1 diet from a high-quality source such as lactalbumin should
meet growth requirements, although 150 g of protein kg−1 may be marginal for
giant breeds. Wills (1996b) regarded 13.2 g of protein MJ ME−1 (equivalent to

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The Nutrition of Companion Animals 443

220 g of protein kg−1 in diets containing 16.7 MJ ME kg−1) as being suitable

for growth of most breeds, although Lepine (1998) suggests that 260 g of
protein kg−1 is more appropriate for giant breeds. As for other species, excess
protein in the diet is either metabolized to produce energy or is stored as fat.
However, when high-fat diets are fed, the protein requirement should be
increased to maintain a balanced protein to energy ratio: a diet containing
200 g of fat kg−1 required protein levels of 250 g kg−1 (Ontko et al., 1957).
There is a paucity of information on the protein requirements of breeding
bitches, but lactating bitches fed a carbohydrate-free diet required 300 g of
protein kg−1 compared with 160 g kg−1 in diets containing carbohydrate
(Kienzle and Meyer, 1989). Wills (1996b) suggested that 13.2 g of protein MJ
ME−1 (equivalent to 220 g of protein kg−1 in a 16.7 MJ ME kg−1 diet) was
sufficient for reproduction. Protein sources fed to lactating bitches should be
of high biological value for maximum utilization, and minimum bulk.
Milner (1979a,b) established that the ten IAAs outlined above were
required for optimum growth and nitrogen balance in growing Beagles.
Current NRC recommendations for IAAs in diets for dogs are summarized in
Table 19.1.

Minerals
As there are limited data on the mineral requirements of dogs, the interactions
between dietary minerals, their bio-availability and the differing requirements
between breeds makes recommendations at best a compromise. The mineral
requirements of dogs presented in Table 19.1 are based on the NRC (1985)
recommendations and those of Burger (1995). The values for calcium and
phosphorus, however, may not meet the needs of some of the large or giant
breeds, and Lepine and Reinhart (1998) recommend that the diet of large or
giant breed dogs should contain 8 g of calcium, 6.7 g of phosphorus, 260 g of
high-quality protein and 140 g of fat kg−1 of diet to allow a moderate growth
rate and enable a managed increase in body mass commensurate with healthy
skeletal development.

Vitamins
The vitamin requirements of dogs at all life stages are given in Table 19.1.
Young, growing dogs require twice as much vitamin E as adults, and Harris
and Embree (1963) have suggested a vitamin E:PUFA ratio (g kg−1) of 0.6:1 as a
minimum to protect against PUFA peroxidation. Dogs cannot synthesize
vitamin D to any significant extent, and it must therefore be supplied in the
diet, although specific supplementation is probably unnecessary as most diets
inherently contain sufficient vitamin D to meet the needs of dogs.

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444 A.C. Longland et al.

Table 19.1. Indispensable amino acid, mineral and vitamin requirements of dogs.
Growtha Maintenanceb Reproductionb

kg−1 diet MJ ME−1 kg−1 diet MJ ME−1 kg−1 diet MJ ME−1

Amino acids
Arginine (g) 5.04 0.33 1.08a 0.07 NS NS
Histidine (g) 1.84 0.12 1.09a 0.07 NS NS
Isoleucine (g) 3.54 0.23 2.31a 0.15 NS NS
Leucine (g) 5.85 0.38 4.16a 0.27 NS NS
Lysine (g) 5.04 0.33 2.47a 0.16 NS NS
Total sulphur 3.85 0.25 1.54a 0.10 NS NS
amino acids (g)
Total aromatic 7.24 0.47 4.31a 0.28 NS NS
amino acids (g)
Threonine (g) 4.62 0.30 2.16a 0.14 NS NS
Tryptophan (g) 1.54 0.10 0.62a 0.04 NS NS
Valine (g) 3.85 0.25 2.92a 0.19 NS NSb
Minerals
Calcium (g) 6.00 0.39b 6.00 0.39 10.16 0.66
Phosphorus (g) 4.4 0.29 4.62 0.30 8.16 0.53
Potassium (g) 4.4 0.29 4.62 0.30 4.62 0.30
Sodium (g) 0.6 0.04 0.62 0.04 0.77 0.05
Chloride (g) 0.9 0.06 0.85 0.055 0.924 0.06
Magnesium (mg) 400 26.0 352.7 22.9 352.70 22.9
Iron (mg) 32 2.10 36.81 2.39 73.92 4.80
Copper (mg) 2.9 0.19 4.62 0.30 6.47 0.42
Manganese (mg) 5.1 0.33 4.62 0.30 4.62 0.30
Zinc (mg) 35.6 2.30 46.05 2.99 46.00 2.99
Iodine (mg) 0.6 0.40 0.62 0.04 1.34 0.09
Selenium (mg) 0.11 0.006 0.11 0.006 0.11 0.006
Vitamins
Vitamin A (IU) 3773b 245b 3773 245 4605 299
Vitamin D (IU) 404 26 400 26 462 30
Vitamin E (mg) 22b 3b 27.72 1.8 46.20 3
Vitamin K (mg) NR NR NR NR NR NR
Thiamin (mg) 1.0 0.06 0.93 0.06 0.92 0.06
Riboflavin (mg) 2.46 0.16 2.31 0.15 2.31 0.15
Pantothenic acid (mg) 10 0.65 10.16 0.66 10.16 0.66
Niacin (mg) 11.0 0.72 11.09 0.72 11.09 0.72
Pyridoxine (mg) 1.1 0.07 1.09 0.07 1.08 0.07
Folic acid (µg) 200 13 200 13 200 13
Vitamin B12 (µg) 25.00 1.62 24.95 1.62 25.00 1.62
Choline (mg) 1247 81 1155 75 1155 75
Biotin NR NR NR NR NR NR
aData from NRC (1985) based on diets containing 15.4 MJ ME kg−1; b data from Burger
(1995).
NS = not stated; NR = no requirement when natural ingredients are fed as sufficient
should be produced by the intestinal microflora.

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The Nutrition of Companion Animals 445

Senior Animals
The population of elderly animals is increasing and, in a survey of the UK
dog population in 1991, 35% were over 7 years old (Davies, 1996). As a
consequence, considerable research is being focused on the nutrition of
elderly dogs with a view to alleviating some of the avoidable effects of ageing.
Defining the elderly dog is best undertaken in terms of breed, as small to
medium sized dogs are known to live longer than the large and giant breeds.
The age at which small and medium breeds are considered ‘geriatric’ averages
at 11 years, whereas for large and giant breeds the onset of old age is some 3–4
years earlier (Goldston, 1995). The maintenance energy requirements of dogs
decrease with age, such that that of Labrador–Retrievers decreased from 593 kJ
of DE W−0.75 for 2-year-olds to 464 kJ of DE W−0.75 for those older than seven
(Kienzle and Rainbird, 1991). This reduction was thought to be due to a
decline in (i) fat free body mass, (ii) basal metabolic rate and (iii) the activity of
elderly dogs. However, the intakes of senior dogs are often reduced and thus it
is not always necessary to lower the energy density of the diet in order to
prevent obesity (Kienzle and Rainbird, 1991). The digestibility of nutrients
does not appear to be altered in ageing dogs (Sheffey et al., 1985) although it
is possible that their protein requirement increases with age (Wannemacher
and McCoy, 1966). However, a balance needs to be struck between feeding
elderly dogs higher levels of protein and inducing renal stress and Mundt
(1991) recommended 4.5 g of highly digestible protein W−0.75. Furthermore,
only highly digestible carbohydrates should be fed to elderly dogs in order to
minimize post-ileal formation of bacterial metabolites (Meyer et al., 1989).
Utilization of fatty acids may change with age, and that of EFAs has been
reviewed by Turek and Hayek (1998). It has been shown that a reduction in
desaturation of fatty acids may occur with ageing (Biagi et al., 1991):
supplementation of fatty acid products of desaturases (e.g. γ-linolenic acid)
may help by-pass such effects, and increased supplementation of diets with
n-3 fatty acids may also be of benefit to the elderly dog (Hayek, 1998).
Comparatively little is known about the mineral and vitamin requirements
of geriatric dogs. However, there may be a requirement for higher levels of
vitamin E (Mundt, 1991), and reduced serum levels of copper and zinc in
ageing dogs may suggest that they have a higher requirement for these (Keen
et al., 1981). Elderly dogs also have a reduced ability to incorporate skeletal
calcium (Weigel and Alexander, 1981), suggesting an extra dietary calcium
requirement. Indeed, Mundt (1991) suggests that the levels of vitamins and
minerals required to support growing dogs are appropriate for senior dogs.

Exercise
The two most extreme forms of exercise for dogs are embodied in racing
Greyhounds which are required to perform intense activity in a sprint for 30 s
or so per race, and sled dogs (Huskies) which may be required to perform in
endurance competitions in which they may run 100 miles per day for

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446 A.C. Longland et al.

10–14 days (Hinchcliff et al., 1998). Other working dogs such as foxhounds
and herding dogs perform somewhere in between these two extremes.
Although the exhaustion of glycogen reserves is the chief limiting factor for
any continued effort, other factors such as providing a suitable electrolyte/
water regime to reduce dehydration and to buffer acid accumulation must also
be catered for when formulating diets for racing animals. Greyhounds may
require 10–20% more energy above their maintenance requirements, and
Grandjean (1996) has suggested that Greyhound diets should contain
400–450 g of carbohydrate, 200–250 g of fat and 350 g of protein kg−1 of diet,
with short and or medium chain fatty acids contributing up to 25% of the total
dietary lipid. Unlike human athletes, carbohydrate loading in dogs can lead to
exertional rhabdomyolysis, and therefore should not be employed to enhance
the performance of racing dogs (Kronfeld, 1973). However, high-fat diets
combined with appropriate training appear to result in glycogen sparing,
reduced lactic acid accumulation and thus decreased muscle fatigue (Kronfeld
et al., 1994). Hultman et al. (1971) and Mutch and Bannister (1983)
demonstrated that overall carbohydrate oxidation accounts for 10–15% of the
energy supply to muscles in Huskies, whereas free plasma fatty acids account
for 70–90%. Therefore, as a general rule, as the length of a race increases and
the intensity of performance decreases, the content of dietary lipid should be
elevated at the expense of carbohydrate (Grandjean, 1996). Recommendations
for racing huskies are 300 g of highly digestible carbohydrate and 350 g of
fat kg−1 of diet which, as for Greyhounds, should contain 25% short and
medium chain fatty acids; the EFAs should be supplied at twice the
maintenance value, the ratio between n-6 and n-3 fatty acids being maintained
at 5.5:1 (Grandjean, 1996). For endurance races, protein should be supplied at
about 21.5 g of MJ ME−1. For other working dogs, dietary fat contents should
rise from 140 to 220 g kg−1 as work becomes more prolonged, while protein
should rise from 300 to 400 g kg−1 of diet to provide a high protein:energy ratio
(Grandjean, 1996). Calcium, phosphorus and trace element levels should be
twice, and those of magnesium three times the maintenance levels in diets
for prolonged endurance race dogs, with intermediate levels for racing Grey-
hounds or sprint sled dogs (Grandjean, 1996). Although dogs do not sweat in
the same way as humans and horses, sled dogs produce large volumes of
urine, resulting in substantial sodium loss, which, if not supplied in the diet,
may cause hyponatraemia (Hinchcliff et al., 1998). In order to ensure adequate
amounts of vitamins for racing dogs, dietary levels of vitamins A, D and K can
be doubled and those of vitamins B, C and E tripled (Grandjean, 1996).
The energy costs of thermoregulation in dogs are high, and thus a drop in
ambient temperature from 15 to 8°C results in a 25% increase in the ME
requirement (Blaza, 1982). Dogs working in hot, humid conditions may
require 50–100% more energy than similar dogs in less stressful circumstances
(Costill, 1981). Furthermore, there is an increased need for water in working
dogs. Compared with a 20 kg house dog which loses about 1400 ml of water
per day via urine, faeces and respiration, 20 kg sprint and distance sled dogs
will lose about two- and fourfold this amount, respectively (Reynolds et al.,
1998), and thus suitable hydration is essential for dogs undergoing strenuous

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The Nutrition of Companion Animals 447

work for many hours. There is evidence to suggest that inducing a mild state of
hyperhydration via dilute glycerol solutions may result in more enduring
rehydration than more traditional glucose–electrolyte solutions (Reynolds
et al., 1998).

CATS
Domestic cats (Felis domesticus), unlike dogs, are obligate carnivores in that
although they can obtain substantial amounts of nutrients from foods of plant
origin, they require certain fatty acids and proteins that can only be obtained
from animal sources.

Energy
The mature body weight of the domestic cat ranges from 2 to 6 kg and,
because there is little variation in body size compared with dogs, many authors
express energy requirements in terms of body weight as opposed to metabolic
body weight. Kendall et al. (1983) found no benefit in terms of precision of
using mass exponents of body weight (in kg) of W0.75 or W0.67 compared with
unity.
The maintenance energy requirements of large (Earle and Smith, 1991) or
inactive (Kendall et al., 1983) cats range from 209 to 293 kJ of ME W−1 day−1,
whereas slightly higher values of 335–380 kJ of ME W−1 day−1 have been
reported for active cats with access to runs (Miller and Allison, 1958).
Neutering cats reduces their energy requirements, such that the resting
metabolic rates of gonadectomized male and female cats were 28 and 33%
less, respectively, than their sexually intact counterparts (Root et al., 1996).
Thus energy intakes of neutered cats should be regulated to prevent their
becoming obese.
Kittens grow rapidly in the first 6 months, during which time they attain
three-quarters of their mature size, weight gains after this time being associated
with developmental rather than skeletal changes (Legrand-Defretin and
Munday, 1995). From 6 to 8 weeks of age, males weigh more than their female
littermates, and this pattern continues throughout life. Kittens normally start to
eat solid food at about 4 weeks old, and intakes initially are in the region of
10–40 kJ W−1 day−1. By 6 weeks, they may consume between 250 and 350 kJ
W−1 day−1, and by 8–10 weeks they may ingest in excess of 800 kJ W−1 day−1
(Munday and Earle, 1991). Thereafter, ME intakes decline per unit of body
weight, although they remain fairly high during the rapid growth phase in the
first 6 months, ME requirements at 25–30 weeks are approximately 420 kJ of
ME W−1 day−1, but by the time cats are 1 year old, their requirements stabilize
to around 250 kJ of ME W−1 day−1 for confined cats and 355 kJ of ME W−1 day−1
for those which are more active (Miller and Allison, 1958).
During gestation and lactation, nutrients must be supplied to meet the
needs of the mother, to support fetal growth and allow for effective milk

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448 A.C. Longland et al.

production. It is therefore recommended that the diet during this period

is nutrient dense, highly palatable and digestible. The normal gestation
period of the cat is 64 days and, unlike most mammals, as soon as a queen
has conceived she will increase her food intake, such that her body weight
increases steadily, virtually from the first day of gestation. In a comprehensive
review of the subject, Loveridge and Rivers (1989) showed that over the
entire pregnancy, the average weight gain was 39% of the pre-conception
weight of the queen. Following parturition, only 40% of this extra weight
was lost, the remainder being lost during lactation. These authors concluded
that this unusual weight gain profile during gestation in cats was due to
net uterine tissue accretion during early pregnancy, which was presumed
to act as an energy store to be mobilized during lactation. To accommodate
the steady increase in maternal body weight during gestation, queens require
a rising level of nutrition, and Loveridge, (1986) showed that the
voluntary intakes of 64 queens rose steadily from 1.13 to 2 MJ of ME per cat
per day (equivalent to 389 to 456 MJ of ME W−1 day−1). Despite being offered
ad libitum access to nutrient-rich, palatable diets during lactation, queens
often lose body weight during this time. Loveridge (1986) reported that the
mean weight of queens decreased from 4.2 kg post-partum to 3.4 kg after 6
weeks lactation. The energy requirements of lactating queens depends on the
size and age of the litter, as kittens may gain more than 100 g per week. Thus,
a queen with one kitten in the first 2 weeks of lactation required 48% of the
energy of her counterpart with six kittens, corresponding values being 45, 41,
33 and 28% in weeks 3, 4, 5 and 6 of lactation, bearing in mind that from week
4 some 5% of the energy from solid food would have been ingested by the
kittens, rising to 30% during weeks 6 and 7 (Legrand-Defretin and Munday,
1995; NRC, 1986).

Lipid
Cats have a high tolerance to fat, diets containing 640 g of fat kg−1were fed
successfully to cats without causing ketonuria, increased fat excretion or
cardiovascular changes (Humphreys and Scott, 1962). Diets containing
250–300 g of fat kg−1 are often fed, and high-fat diets tend to be palatable
(Kendall, 1984). Kane et al. (1981) found that cats preferred diets containing
250 g of fat kg−1 compared with those containing 100 or 500 g of fat kg−1. The
fat contents of commercially available cat foods range from 75 to 300 g of fat
kg−1 of diet (Kelly, 1996).
Cats are deficient in certain desaturase enzymes and are unable to convert
linoleic acid to arachidonate, which must therefore be supplied in the diet;
arachidonate is only found in animal sources (MacDonald et al., 1983,
1984a,b,c). Without a source of arachidonate, queens cannot produce viable
young (MacDonald et al., 1984c). The NRC (1986) regard 5 and 0.2 g of linoleic
and arachidonic acids per kg of diet, respectively, as the minimum levels of
inclusion for cats. However, Wills (1996a) suggests that 1 and 0.02 g of linoleic
and arachidonic acids, respectively, MJ ME−1 (equivalent to 20.9 and 0.42 g in a

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The Nutrition of Companion Animals 449

diet containing 20.9 MJ of ME kg−1) should be supplied in diets for cats at all
life stages.

Carbohydrates
Cats do not necessarily require carbohydrates, if sufficient fat and protein
are provided from which the metabolic requirement for glucose can be met.
However, commercial cat diets frequently contain more than 400 g of carbo-
hydrate kg−1 which they may utilize well: digestibility coefficients for glucose
and starch being 0.99 and 0.96, respectively (Trudell and Morris, 1975).
However, consistent with their carnivorous nature, cats are unable to tolerate
very high carbohydrate diets (Wills, 1996a). Dietary fibre is fermented to vary-
ing degrees by cats, and some dietary inclusion of fibre may be useful to
encourage peristalsis, improve faecal consistency and to lower the energy
density of diets for inactive cats.

Protein and Amino Acids

Although minimal requirements of the ten essential amino acids have
been determined in growing cats, there are few corresponding values for
maintenance and reproduction. Compared with less strict carnivores, cats have
a limited ability to regulate transaminase and urea cycle enzyme activity to
accommodate changes in dietary protein levels, resulting in comparatively
high nitrogen losses (Rogers et al., 1977). As a consequence, both kittens and
adult cats have higher protein requirements than most other companion
mammals (Rogers and Morris, 1982).
The protein content of diets for cats at maintenance recommended by
Kelly and Wills (1996) and Burger (1995) is 15 g of protein MJ ME−1, which
equates to 250 g of protein kg−1 in diets containing 16.7 MJ of ME kg−1. The
recommendations for the protein contents of diets of both growing kittens and
breeding females are the same; the NRC (1986) suggests 240 g of protein kg−1
diet, whereas Kelly and Wills (1996) and Burger (1995) recommend 17 g of
protein MJ ME−1 (equivalent to 284 g of protein kg−1 in diets containing 16.7 MJ
of ME kg−1). The IAA requirements of growing cats are summarized in Table
19.2. It is noteworthy that arginine is an IAA for cats evidenced by the onset of
hyperammonaemia within 3 h of feeding a single arginine-free meal (Morris
and Rogers, 1978a,b).

Taurine
Taurine is an amino sulphonic acid which occurs in the free form in all animal
tissues. Most mammals can synthesize adequate levels of taurine from
methionine and cysteine. Additionally, the bile salts of most mammals contain
a mixture of taurine and glycine conjugates and, when in a state of taurine
deficiency, preferentially synthesize glycine conjugates. However, in the cat,
bile salts are conjugated exclusively with taurine (Rabin et al., 1976), and thus

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450 A.C. Longland et al.

Table 19.2. Indispensable amino acid, mineral and vitamin requirements of cats.
Growtha Maintenancec Reproductionc

kg−1 diet MJ ME−1 kg−1 diet MJ ME−1 kg diet−1 MJ ME−1

Amino acids
Arginine (g) 10.03 0.48 10b 0.48b NS NS
Histidine (g) 2.93 0.14 NS NS NS NS
Isoleucine (g) 5.02 0.24 NS NS NS NS
Leucine (g) 11.91 0.57 NS NS NS NS
Lysine (g) 7.94 0.38 3.3b 0.16b NS NS
Total sulphur 7.52 0.36 3.2b 0.16b NS NS
amino acids (g)
Total aromatic 8.57 0.41 NS NS NS NS
amino acid (g)
Threonine (g) 7.11 0.34 NS NS NS NS
Tryptophan (g) 1.46 0.07 NS NS NS NS
Valine (g) 6.06 0.29 NS NS NS NS
Minerals
Calcium (g) 8.00 0.38 8.40d 0.40d 12.50 0.60
Phosphorus (g) 6.06 0.29 6.30 0.30 10.00 0.48
Potassium (g) 3.97 0.19 6.30 0.30 6.30 0.30
Sodium (g) 0.48 0.023 0.50 0.023 1.0d, 2.5c 0.05d, 0.12c
Chloride (g) 19.01 0.91 NS NS NS NS
Magnesium (mg) 397 19 376 18 627 30
Iron (mg) 79.42 3.80 84c, 125d 4c, 6d 125 6
Copper (mg) 5.02 0.24 5c, 6.3d 0.24c, 0.3d 6.30 0.30
Manganese (mg) 5.02 0.24 5c, 6.3d 0.24c, 0.6d 12.50 0.60
Zinc (mg) 50.16 2.40 50.00 2.40 50.16 2.40
Iodine (mg) 0.355 0.017 0.4c, 1.25d 0.02c,0.06d 1.25 0.06
Selenium (mg) 0.10 0.005 0.10 0.006 0.10 0.006
Vitamins
Vitamin A (IU) 3333 1600 3333 160 6000 329
Vitamin D (IU) 500 24 500d, 1000c 12d, 24c 1254 60
Vitamin E (mg)e 30 1.43 17 0.8 30d, 100c 3d, 5c
Vitamin K (µg) 100 5.00 125 6 125 6
Thiamin (mg) 5 0.24 5 0.24 6 0.3
Riboflavin (mg) 4 0.20 4 0.20 6 0.3
Pantothenic acid (mg) 5 0.24 5 0.24 12.5 0.6
Niacin (mg) 40 2.00 40 2.0 56 2.7
Pyridoxine (mg) 4 0.20 4 0.20 5 0.24
Folic acid (µg) 800 38.00 800 38 1250 60
Vitamin B12 (µg) 20 1.00 21 1.0 25 1.2
Choline (mg) 2400 114.00 2508 120 2508 120
Biotin (µg) 70c, 125d 3.34c, 6d 90c, 125d 4.2c, 6d 90c, 125d 4.2c, 6d
Taurine (mg) (canned) 3114c 150c 3114 150 3114 150
Taurine (mg) (dry food) 1254c 60c 1254 60 1254 60
Taurine mg (all diets) 400 19 4001 191 4001 191
Data from NRC (1986) based on diets containing 20.9 MJ of ME kg−1 ; bData from Burger and Smith
a

c d e
(1990); Data from Burger (1995); Data from Kelly and Wills (1996); For diets containing high levels
of polyunsaturated fatty acids, vitamin E levels should be quadrupled for all life stages.

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The Nutrition of Companion Animals 451

production of bile salts represents a net loss of taurine to the cat (Sturman
et al., 1978). Cats also exhibit poor hepatic synthesis of taurine from
methionine and this, together with losses via bile salts, accounts for the
susceptibility of felines to taurine-deficient diets which is manifested in feline
central retinal degeneration where affected cats can become blind. Dietary
levels of taurine recommended by NRC (1986) are 400 mg kg−1 diet for growth
and maintenance and 500 mg kg−1 diet for reproduction. However, the taurine
requirements reported in Burger (1995) are much higher at 60 and 149 mg MJ
ME−1 for commercial dry and canned foods respectively (equivalent to 1000
and 2500 mg kg−1 diet containing 16.67 MJ of ME kg−1diet).

Minerals
Calcium and phosphorus deficiencies result in hyperparathyroidism in the cat
(Bennet, 1976) with a generalized decrease in bone density due to resorption
of bone calcium to restore serum levels. Dietary magnesium levels of 400 mg
kg−1 diet are recommended for growth (NRC, 1986) but excessive magnesium
intakes are associated with the formation of urinary calculi (Kallfellz et al.,
1980). The mineral requirements of cats are given in Table 19.2.

Vitamins
Cats require pre-formed vitamin A for, unlike many other mammals, they are
unable to convert β-carotene into vitamin A (Ahmad, 1931), deficiency leading
to conjunctivitis and photophobia (Scott et al., 1964). Cats cannot synthesize
sufficient vitamin D on exposure to sunlight and again require a dietary
supply: both vitamins A and D are present in animal tissues and, as cats are
obligate carnivores, they obtain sufficient amounts of these from their prey.
Growing kittens require greater dietary levels of niacin than many other
mammals, as felines are unable to synthesize adequate levels of niacin from
tryptophan. Excessive levels of vitamins, especially A and D, can be toxic;
feeding cats liver or certain canned fish diets for long periods can lead to A
and D hypervitaminosis, respectively. Many cats are fed fish-based diets which
contain high levels of PUFAs which can auto-oxidize to produce potentially
harmful peroxides and, therefore, appropriate intakes of vitamin E are
essential for cats fed such diets. Of the water-soluble vitamins, thiamin is most
likely to be lacking, and certain species of fish contain thiaminase (Smith and
Prout, 1944) and thus should not be fed raw. Furthermore, as thiamin is very
sensitive to heating (up to 90% may be lost during canning), it is important that
sufficient thiamin is added to processed diets to ensure adequate availability.
Thiamin deficiency leads to neurological defects and death (Everet, 1944). The
vitamin requirements of cats are given in Table 19.2.

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452 A.C. Longland et al.

Senior cats
Cats are generally considered to be geriatric from 12 years old (Goldston,
1995). The incidence of disease is greater with elderly cats, chronic renal
failure and hyperthyroidism being fairly common disorders (Peterson and
Graves, 1992) leading to low intakes and weight loss. Feeding highly digest-
ible, palatable diets is recommended as senior cats have a reduced digestive
capacity. Neoplasia is also prevalent in the geriatric cat population, and
feeding vitamin A and E may help to inhibit this and certain other degenerative
diseases (Legrand-Defretin and Munday, 1995).

BIRDS
The most popular species of companion bird in Northern Europe are the
budgerigar (Melopsittacus undulatus) and the canary (Serinus canarius), the
former making up approximately 65% and the latter 15–20% of the total
population, the balance largely consisting of finches, parrots, parakeets,
pigeons and quails (Nott and Taylor, 1995). In the wild, these birds eat seeds
from a wide variety of plant species (Nott, 1992). South American and African
parrots eat fruit, shoots (Nott and Taylor, 1995) and insects (Long, 1984) as
well as seeds, whilst the lories and lorikeets are nectar feeders (Nott and
Taylor, 1995). The naturally varied diets of such birds in the wild ensures that
their nutrient requirements are met but, if young captive birds are offered diets
of limited diversity, they may become loath to accept new foods, leading to
nutritional deficiencies (Lawton, 1988). The information below largely pertains
to the psttiacines and canaries and is not comprehensive. As there is a paucity
of data on the nutrient requirements of companion birds, other authors
frequently present data extrapolated from the NRC (1984) to which the reader
is referred for additional information.
The avian metabolic rate is higher than that of mammals, with body
temperatures averaging 41–42°C (Nott and Taylor, 1995), and birds excrete
surplus nitrogen in the form of uric acid, a process requiring 3.25 times more
energy than the excretion of an equivalent amount of urea. Consequently, as
birds require more energy pro-rata than mammals, they must process, digest and
absorb food with greater efficiency. The nutrient requirements of birds are
affected by the physiological state of the animal, and factors such as age, breed-
ing and moulting may result in increased nutrient demands.

Energy
Various estimates have been made of the maintenance energy requirements
for different types of companion birds. Aschoff and Pohl (1970) quoted a value
of 535 kJ W−0.715 day−1 for finches, and Earle and Clarke (1991) reported the
maintenance energy requirement of budgerigars to be 1.59 MJ W−1 day−1, a
value some threefold that of a domestic chicken (NRC, 1984). Young birds

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The Nutrition of Companion Animals 453

grow rapidly, budgerigar chicks increasing their birth weight more than 12-fold
in the first 10 days of life. In order to achieve this growth rate and simulta-
neously developing plumage, young birds devote a large proportion of energy
intake to growth (Brooke and Birkhead, 1991). Breeding substantiality raises
energy demands; over an average 78-day breeding period from pairing to
fledging of 28 budgerigar families (hens average body weight 50 g, cocks aver-
age body weight 56 g and three chicks), mean intakes were 242 kJ per family,
with a peak of daily energy demand at day 60 of approximately 500 kJ per
family (Earle and Clarke, 1991).

Protein
Protein has an important role in birds in terms of beak, feather and claw
production (Earle and Clarke, 1991). The protein requirement of the adult
budgerigar has been estimated as 100 g kg−1 of the diet (Drepper et al., 1988),
and this may increase two- or threefold during moulting or breeding, especially
when crop milk is produced. There are scant data on the amino acid require-
ments of companion birds but, in a study on budgerigars, values of 20, 35 and
35 g kg−1 of crude protein were given for lysine, arginine and methionine
plus cysteine (the sulphur amino acids being required for feather formation),
respectively (Drepper et al., 1988). Seeds commonly fed to companion birds,
such as red and white millet, groats and canary grass seed, typically contain
from 110 to 160 g of protein kg−1, with apparent protein metabolizability in
budgerigars of 0.72–0.90 (Earle and Clarke, 1991). It has been estimated that
200 g of protein and a minimum of 8 g of lysine kg−1 of diet is required for
optimal growth of cockateils (Nymphicus hollandicus) (Roudybush and Grau,
1991), although Ullrey et al. (1991) recommend 240 g of protein kg−1 for all life
stages of psittacines. Thirteen grams of protein MJ ME−1 with 0.62 and 0.82 mg of
lysine and sulphur amino acids, respectively, enabled rapid growth rates in
canaries (Kamphues and Meyer, 1991). In addition to the ten IAAs required by
adult birds, glycine is also regarded as indispensable for growing chicks as its
hepatic synthesis is limited in these birds, yet it is essential for collagen, feather
and uric acid production; 10 g of glycine kg−1 diet is recommended
(Featherstone, 1976). Ullrey et al. (1991) reported that growth and reproduction
of psittacines was supported by an extruded cereal-based diet, the amino acid
content of which is shown in Table 19.3.

Lipid
Fat is a useful source of energy for birds, and Ullrey et al. (1991) reported that
20 g of fat kg−1 supported satisfactory performance in psittacines; it has been
suggested that linoleic acid should constitute 10 g kg−1 of the diet (NRC, 1984).
Companion birds are often prone to obesity, and restricted fat intakes are
therefore recomended. The seeds usually fed to budgerigars contain 35–70 g
fat and 17–24 g of linoleic acid kg−1 of diet (Earle and Clarke, 1991), which
suggests that such birds are unlikely to suffer EFA deficiency.

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454 A.C. Longland et al.

Table 19.3. Indispensable amino acid, mineral and vitamin requirements of

−1
companion birds, ornamental fish, rabbits and rats (kg diet).
Rabbitsc Ratsd

Birdsa Fishb Growth Reproduction Growth Reproduction

Amino acids
Arginine (g) 13 16 6.0 NS 4.3 4.3
Histidine (g) NS 8 3.0 NS 2.8 2.8
Isoleucine (g) 11 9 6.0 NS 6.2 6.2
Leucine (g) NS 13 11.1 NS 10.7 10.7
Lysine (g) 12 22 6.5 NS 9.2 9.2
Total sulphur amino acids (g) 9 12 6.0 NS 10.0 10.0
Total aromatic amino acids (g) NS 25 11.1 NS 10.2 10.2
Threonine (g) 9.5 15 6.0 NS 6.2 6.2
Tryptophan (g) 2.4 3 2.0 NS 2.0 2.0
Glycine (g) 10 NS NS NS NS NS
Valine (g) NS 14 7.0 NS 7.4 7.4
Minerals
Calcium (g) 11 3–7 4.0 4.5 (7.5) 5.0 6.3
Phosphorus (g) 8 4–6 2.2 3.7 (5) 3.0 3.4
Potassium (g) 7 6–12 6.0 6.0 3.6 3.6
Sodium (g) 2 12–30e 2.0 2.0 0.5 0.5
Chloride (g) 2 NR 3.0 3.0 0.5 0.5
Magnesium (mg) 1500 400–700 350 350 500 600
Iron (mg) 150 200 50e 50e 35 75
Copper (mg) 20 3.0 3.0 3.0 5.0 8.0
Manganese (mg) 65 13 8.5 2.5 10.0 10
Zinc (mg) 120 80–200 50e 50e 12 25
Iodine (mg) 1 NS 0.2 0.2 0.15 0.15
Selenium (mg) 0.3 0.5–1.0 0.1 0.1 0.15 0.40
Vitamin
Vitamin A (IU) 8000 10,000* 580$ >1160 2333 2333
Vitamin D (IU) 500 2400 1000e 1000e 1000 1000
Vitamin E (IU) 250 100 40 40 18 18
Vitamin K (mg) 4000 10,000 0.2 0.2 1000 1000
Thiamin (mg) 6 10 10e 10e 4.0 4.0
Riboflavin (mg) 6 10 10e 10e 10e 4.0
Pantothenic acid (mg) 20 50 20e 20e 10 10
Niacin (mg) 55 50 180 NS 15 15
Pyridoxine (mg) 6 10 39 NS 6.0 6.0
Folic acid (µg) 900 10 1000e 1000e 1000 1000
Vitamin B12 (µg) 25 20 10e 10e 50 50
Choline (mg) 1700 2000 1200 1500e 750 750
Biotin (µg) 300 1000* 200e 200e 200 200
aData from Ullrey et al. (1991) of the nutrient composition of an extruded diet fed to birds
(psittacines) which supported good performance at all life stages, as data on exact requirements
for each nutrient are unavailable. For information on nutrient requirements of poultry, readers are
advised to consult the NRC (1984).
bData for indispensable amino acids are for carp reported by Nose (1979). Data for mineral

requirements are those quoted by Burger (1995), the values being ranges depending on species
and life stage, and are not maxima and minima. Data for vitamin requirements of ornamental fish
are estimated minima quoted by McCartney (1996), except where marked with an asterisk where
data is from Burger (1995).
cData are from NRC (1977), values for vitamins in parentheses denote requirements for lactation.
$Rabbits at all life stages also have a requirement for 30 mg carotene kg−1 of diet.
dData are from NRC (1993); edata from Kelly and Wills (1996).

NS = not stated; NR = not required.

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The Nutrition of Companion Animals 455

Minerals
The calcium requirement for birds has been calculated as 6–10 g kg−1 of diet
depending on factors such as egg laying, etc. Most birdseeds are deficient in
calcium, hence the need to provide appropriate supplements, but excess
calcium can reduce iodine absorption (Taylor, 1954). Phosphorus in seed is
often in the form of phytate and therefore of limited availability, values of
0.1–1 g kg−1 of diet as available phosphorus being quoted by Earle and Clarke
(1991). Kamphues and Meyer (1991) suggested a phosphorus requirement of
0.32 and 0.57 g MJ ME−1 for slow and rapidly growing canaries, respectively.
The mineral content of the extruded diet of Ullrey et al. (1991) which
supported good performance of psittacines is given in Table 19.3.

Vitamins
Seeds lack β-carotene and do not contain vitamin A and, therefore, vitamin A
deficiencies are relatively common in seed-fed birds (Pitts, 1983). Supple-
menting budgerigars with an extra 12 µg of vitamin A per day per bird was
regarded as sufficient (Baker, 1990), with 750 µg being the maximum
tolerated, as oversupplementation can prove lethal. Vitamin D can be
synthesized from sterols in the cutaneous layer of budgerigars on exposure to
UV light or sunlight, and a daily requirement of 25 mg per bird has been
suggested (Baker, 1990). Vitamin D deficiency can lead to poor feathering,
reduced egg production, thin shells and rickets (Garlisch and Wyatt, 1971).
The vitamin content of the diet fed to psittacines by Ullrey et al. (1991) is
shown in Table 19.3.

ORNAMENTAL FISH
The nutrition of ornamental fish is one of the least researched areas of
companion animal nutrition; much of the information given in various articles
on the subject is derived from data obtained for food fish (McCartney, 1996).
As food fish are dealt with in Chapter 18, only some general points will
be addressed here. The interest in ornamental fish keeping has increased
dramatically worldwide over the past 20 years, such that it is estimated that
there are more ornamental fish kept than all the other companion animals
added together (Pannevis, 1995).
A number of fish species at different life stages with different nutritional
requirements and feeding habits are often kept together in a community tank,
yet they frequently are fed a single diet. Moreover, the diet must be able to
satisfy the needs of fish which are surface, middle or bottom feeders and
which may have different diurnal feeding patterns. The safest option is to feed
diets with specifications that will satisfy the needs of the most nutritionally
demanding fish and yet will not result in environmental pollution, either
directly in the form of uneaten food or indirectly through excretion.

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456 A.C. Longland et al.

Energy
As fish are poikilotherms, their energy requirements change with environ-
mental temperature, although it is generally accepted that the maintenance
energy requirements of fish are usually 5–10% of that of birds and mammals
(Smith, 1989). Other contributory factors to explain such comparatively low
energy requirements are that fish excrete ammonia rather than urea and their
movement through water is more energetically efficient than that of land
animals (Pannevis, 1995). The maintenance energy requirement of goldfish
increased approximately threefold when water temperature was raised form 20
to 24°C, and that of pond-kept Koi carp more than doubled when the water
temperature increased from 10 to 20°C but, when water temperatures dropped
below, 5°C these fish barely fed at all, living off body reserves for many
months (McCartney, 1996). However, for a 5 g goldfish at 20°C, McCartney
(1996) suggests that the maintenance energy requirement is 40 kJ of DE W−1.
Calculating from results reported by Pannevis and Earle (1994a) for the
maintenance energy requirements of four popular ornamental fish at 26°C, the
following values (J of DE g body weight−1 day−1) were obtained: Neon Tetras
(374), Leopard Danio (422), Kribensis (178) and Moonlight Gourami (272). To
put this into context for the fish owner, in the case of average sized adult Neon
Tetras and Moonlight Gouramis, these values equate to 0.6 and 4.9 flakes of a
commercial fish food per day. Although for reasons of economy, carbo-
hydrates are fed as energy sources to food fish, carbohydrates are relatively
poorly utilized by many fish. Therefore, the majority of the dietary energy for
ornamental fish is provided in the form of lipid and protein. Optimum ratios
for digestible protein to digestible energy are believed to be in the region of
19–28 mg of protein kJ DE−1 (NRC, 1993a).

Protein
The requirement for protein by fish ranges from 250 to 500 g kg−1 and should
be of high quality to prevent pollution; indeed many of the health problems
encountered with ornamental fish are related to nitrate build up due to over-
feeding and/or insufficient filtering (McCartney, 1996). The quantitative
requirement by fish for IAAs is high (NRC, 1993a), and feeding a protein
source that closely matches the needs of the species in question will result in
maximal growth and minimal excretion. For these reasons, fish meal is
regarded as the most appropriate protein source in ornamental fish diets.

Lipid
Fat is required as a major energy source by ornamental fish and it is
recommended that diets contain between 100 and 200 g of fat kg−1 diet
(Cowey and Sergeant, 1979). It is of note that fats with a low melting point
should be avoided when feeding fish in cold environments, as solidification in

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The Nutrition of Companion Animals 457

the digestive tract could prove fatal. The EFAs required depend on both the
species and the feeding habits of the fish. Freshwater species are generally
able to elongate fatty acids or desaturate the shorter chain EFAs and therefore
require either linolenic (n-3) or linoleic (n-6) acids, or both. Marine herbivores
require a source of n-3 fatty acids, and linolenic acid is usually sufficient. The
piscivores, however, require an EFA profile that reflects the composition of
their prey, and need the longer, more unsaturated eicosapentanoic acid (EPA)
(20:5) (n-3) or docosahexaenoic acid (DHA) (22:6) (n-3). McCartney (1996)
recommends the inclusion of 10 g each of linoleic and linolenic acids per kg
in freshwater fish diets and 20 g of linolenic acid per kg of diet for marine
herbivores, this being replaced with DHA or a mixture of EPA and DHA for
marine carnivores.

Minerals
Unlike terrestrial animals, fish can absorb some minerals from the environment
via fins, gills or oral epithelia, or by drinking, thus reducing their dietary
requirement for minerals. Calcium deficiency has seldom been detected in
ornamental fish due to their being fed high levels of calcium-rich fish meal.
Deficiencies of sodium, potassium and chlorine are also rare, as they are
readily absorbed from the environment. Requirements of other minerals by
ornamental fish are shown in Table 19.3. A problem encountered with present-
ing food in water is leaching of water-soluble nutrients into the environment
prior to their ingestion by the fish. For example, Pannevis and Earle (1994b)
showed that 90% of vitamin B12, 66% of vitamin C, 47% of pantothenic acid
and up to 27% of choline, folate and pyridoxine was lost from the diet after
only 30 s exposure. Furthermore, freshwater fish cannot absorb leached
vitamins, and the dilution effect of a large body of water would render drink-
ing by marine species ineffective in terms of leached vitamin recapture. As the
average time for flaked feeds to reach the bottom of a 30 cm deep tank is 90 s
(Pannevis, 1995), bottom feeders are potentially most at risk from
hypovitaminosis when flaked diets are fed, due to both longer dietary expo-
sure to water and competition from surface and middle water feeders. It is
recommended that both flaked diets which float and fast sinking pelleted diets
are fed in community tanks to allow fish of all feeding habits maximum benefit
from their diets. When formulating diets for ornamental fish estimated leaching
characteristics should be considered.

RABBITS
The domestic rabbit (Oryctolagus cuniculus) varies in mature size from 1 to
more than 6 kg. Rabbits are herbivores, and practise coprophagy from the time
they start to eat solid food, whereby, in addition to hard faecal pellets, they
produce soft, dark pellets which are ingested by the rabbit on their emergence
from the anus. Coprophagy, together with hind-gut fermentation, is thought to

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458 A.C. Longland et al.

provide most of the B vitamin requirements of the rabbit and some bacterial
protein. Coprophagy also allows further digestion of nutrients by virtue of their
being passed through the digestive tract on successive occasions.

Energy
Rabbits derive their dietary energy largely from fresh and conserved forages
and cereal grains. Rabbits are less efficient at utilizing dietary fibre compared
with other herbivores, the average crude fibre digestibility coefficient in the
rabbit being 0.14 compared with 0.22 for pigs, 0.33 for the guinea pig and
0.41 for the horse (Maynard and Loosli, 1969). The gut microflora of the rabbit
differs from that of other herbivorous species in that it consists almost entirely
of Bacteriodes species (Fuller and Moore, 1971), whereas Escherichia coli and
Lactobacillus sp. are either absent, or present in very low numbers (Smith,
1965). However, it has been calculated from caecal VFA concentrations
that 10–12% of the daily maintenance energy requirement of rabbits may be
met from fermentation of fibre (Lebas, 1975a), and some fibre in the diet is
necessary for maintaining normal digestive function, a minimum of 120 g of
crude fibre kg−1 of diet being recommended (Lebas, 1975b). Rabbits digest
both starch and sugars efficiently (NRC, 1977), and like other species will
utilize protein for energy when other sources are scarce. The energy require-
ments of rabbits have been reported as: 418.4 kJ W−0.75 day−1 for maintenance,
798–882 kJ W−0.75 day−1 for growth, 567 and 840 kJ W−0.75 day−1 for early
and late gestation respectively, and 1260 kJ W−0.75 day−1 for lactation (Tobin,
1996).

Lipid
Rabbits showed a preference for diets containing up to 100 g of oil kg−1 of diet
over those containing 0, 50 or 200 g of oil kg−1 (Cheeke, 1974), and young
rabbits grew faster on diets containing 100–250 g of oil kg−1 than those on diets
containing 50 g of oil kg−1 (Thacker, 1956). However, the tendency for rabbits
to become obese has led to a recommendation of 20–50 of oil kg−1 in diets for
non-lactating animals (Tobin, 1996). Recent work by Pascual et al. (1999)
showed that lactating does had higher milk yields with greater energy content,
enabling higher litter weight gains and reduced mortality of nursing young,
when fed diets containing 99 or 117 g of lipid kg−1 of diet compared to those
fed a diet containing 26 g of lipid kg−1.

Protein and Amino Acids

Rabbits require essential amino acids in the diet, as the bacterial protein
obtained via coprophagy is inadequate in this regard. The NRC (1977)
recommendations for amino acids for rabbits as a percentage of the diet are:

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The Nutrition of Companion Animals 459

arginine, 0.6; lysine, 0.65; and methionine plus cysteine, 0.6, and these
levels will both support rapid growth and provide a safety margin for those
at maintenance. In contrast to the situation with fibre, rabbits are able to
utilize forage protein more efficiently than a number of other monogastrics.
Thus protein digestibility coefficients for alfalfa meal were 0.75 and 0.50
for rabbits and pigs, respectively (Slade and Hintz, 1969). Although
coprophagy does not appear to provide many essential amino acids, it
may help to maintain the nitrogen balance in mature animals fed low-quality
protein sources. For example, adult rabbits fed gelatin which were allowed
to engage in coprophagy were able to maintain a positive nitrogen balance
but were unable to do so when coprophagy was prevented (Kennedy and
Hirschberger, 1974). Although exact recommendations as to appropriate levels
of dietary protein in diets for rabbits at different life stages depend on protein
quality, the generally accepted levels of protein are 120, 160, 150 and 170 g of
crude protein kg−1 of diet for maintenance, growth, pregnancy and lactation
(NRC, 1977).

Minerals
Calcium and phosphorus levels of 4.5 and 3.7 g kg−1, respectively, were
regarded as adequate for growth and gestation (Chapin and Smith, 1967), the
corresponding values for lactation being 7.5 and 5.0 (Lebas et al., 1971).
Although these values are slightly lower than those recommended by Tobin
(1996), these amounts should not be exceeded greatly, for rabbits excrete
excess dietary calcium in the urine and thus high-calcium diets can lead to the
formation of uroliths. The mineral requirements of rabbits are summarized in
Table 19.3.

Vitamins
Although adequate supplies of the vitamin B complex should be supplied via
coprophagy, it is prudent to supply some dietary B vitamins, especially for
animals housed on wire-mesh floors. Although it might also be expected
that coprophagy would supply the vitamin K requirement, there have been
reports of deficiencies affecting reproduction, and therefore there is a case for
providing dietary vitamin K. Rabbits are highly sensitive to vitamin E
deficiency resulting in muscular dystrophy and cardiac problems, therefore
diets should always be supplemented with this vitamin, taking into account its
relatively rapid decline in feeds (5–20% per month) during storage (Tobin,
1966). Rabbits also require β-carotene which, in addition to being a precursor
of vitamin A, also appears to enhance reproductive performance, and levels of
at least 30 mg of β-carotene kg−1diet have been recommended (NRC, 1977). It
is of note that although β-carotene is present in fresh green feed, up to 80%
may be lost during haymaking. The vitamin requirements of rabbits are given
in Table 19.3.

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460 A.C. Longland et al.

RATS
Domestic rats (Rattus norvegicus) are omnivorous monogastrics which in their
natural habitat eat grains, seeds, small invertebrates and vertebrates. Most
commercial diets for rats, however, are largely vegetarian, being based on milk
products, vegetable oils, cereal grains and legumes such as peas, beans and
alfalfa.

Energy
The maintenance energy requirement for rats quoted by NRC (1993b) is 470 kJ
W−0.75 day−1, whereas that for growth is reported to be 606 kJ W−0.75 day−1
(Tobin, 1996). Although protein appears to be more important for successful
reproduction than energy (Menaker and Navia, 1973), the daily ME require-
ment for early gestation is approximately 600 kJ W−0.75 day−1, but may increase
to 1110 kJ W−0.75 day−1 during later stages of pregnancy (NRC, 1993b). The
energy requirement of lactating females depends on the number of pups
suckled but is usually in the order of two- to fourfold that of dry females
(Nelson and Evans, 1961). In addition to large increases in feed intake, during
lactation, female rats mobilize large stores of body fat that they accrue during
gestation. Naismith et al. (1983) reported that such body fat supplied much of
the energy required during lactation. Nevertheless, it has been estimated that
the daily ME requirement of rats at peak lactation ranges from 1.3 MJ W−0.75
day−1 (NRC, 1993b) to 1.8 MJ W−0.75 day−1 (Tobin, 1996).

Lipid
Digestibility of lipids by rats varies with source, processing and level of
dietary inclusion. Thus digestibility coefficients for lipid from diets containing
either 50 g kg−1 of cocoa butter or corn oil by adult rats were 0.92 and
0.59, respectively, though in the latter case when the inclusion level was
increased to 200 g kg−1 the corresponding coefficient was 0.71 (Apgar et al.,
1987). The digestibility coefficient of lipid from bland lard was 0.94, but
only 0.63 for hydrogenated lard in adult female rats (Crockett and Deuel,
1947). Linolenic acid (n-6) is the major EFA required by rats, and work
by Bourre et al. (1989, 1990) indicated that 1.2 and 0.2 g of linoleic and
linolenic acids, respectively, per kg of diet were the minimum requirements of
EFA by rats, but these levels are substantially lower than the 6.0 g of linoleic
acid kg −1 recommended by the NRC (1978a). Dietary levels of 50 g of lipid
kg −1 appear to be sufficient for all classes of rat, and many sources of lipid
provide adequate amounts of the EFA at this level of inclusion (NRC, 1993b).
A number of studies have shown that high levels (up to 500 g kg−1) of lipid
inclusion in the diet can have deleterious effects on health (Kollmorgen et al.,
1983), reproduction (Richardson et al., 1964) and growth of nursing pups
(Rolls and Rowe, 1982).

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The Nutrition of Companion Animals 461

Carbohydrates
Rats require dietary carbohydrates and can utilize glucose, fructose, starch,
maltose and a number of sources of dietary fibre. However, poor performance
was observed in galactose-fed rats (Day and Pigman, 1957), and xylose was
found to be toxic to adult rats (Booth et al., 1953). Carbohydrate-free diets
cannot support reproduction (Taylor et al., 1983) or lactation in rats (Koski
et al., 1990).

Protein and Amino Acids

Protein contents of 50 and 70 g kg−1 of diet for high-quality and natural
ingredient protein sources, respectively, have been reported as being suitable
for maintenance of rats (Bricker and Mitchell, 1947; Dibak et al., 1986). Early
work suggested that the least amount of protein which had a balanced amino
acid profile required to support maximum weight gain in young rats was
100–150 g kg−1 of a low fibre diet containing 17 MJ of ME kg−1 (Mitchell and
Beadles, 1952). This is consistent with the NRC (1978a) recommendation of
120 g of protein kg−1 for growing rats when a highly digestible protein source
with a balanced amino acid profile was used. However, when non-purified
dietary ingredients are used, 180–250 g of crude protein kg−1 is required.
Studies by Nelson and Evans (1958) on lactating females indicated that
although diets containing 180 g of casein kg−1 supported maximum growth in
suckling young, 240 g was required for the dam to maintain or gain body
weight.
The current (NRC, 1993b) recommendations for IAAs to support growth
and reproduction in rats are largely the same, with lower values for
maintenance (Table 19.3.) In addition to these requirements, there appears to
be a need for dietary asparagine during pregnancy as asparagine-free diets
were associated with impaired neurological development of the young
(Newburg and Fillios, 1979).

Minerals
Dietary concentrations of calcium and phosphorus of 5 and 3 g kg−1 of
diet, respectively, are recommended by the NRC (1993b) for growth and
maintenance of rats, a calcium:phosphorus molar ratio of at least 1.3:1 being
required. The lactating rat has an increased demand for these minerals, and
Brommage (1989) indicated that a lactating female could pass 200 mg of
calcium and 140 mg of phosphorus into the milk in 1 day, and that this
increased need was met via higher feed intakes and enhanced absorption of
these minerals: NRC (1993b) suggests 6.3 g of calcium and 3.7 g of phosphorus
kg−1 of diet as being suitable. Magnesium-deficient rats may have reduced litter
sizes or malformed young (Hurley et al., 1976a,b). Rats have a specific require-
ment for iodine, their utilization of this mineral being high and being largely

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462 A.C. Longland et al.

used for thyroid hormone function (Gross, 1962). Although Tobin (1996)
suggests that 75 mg of manganese kg−1 of diet is appropriate for all classes of
rat, the current NRC recommendation is 10 mg kg−1, due to reports by Davis
et al. (1971) that excess manganese has a deleterious affect on iron
metabolism. Table 19.3 shows the mineral requirements of rats.

Vitamins
High levels of vitamin A and E appear to interfere with vitamin K metabolism
such that the onset of vitamin K deficiency is more rapid in diets rich in
vitamins A and E (Ralli and Dumm, 1953). Biotin-deficient rats have a
depressed immune response, develop alopecia and may develop an abnormal
gait (Rabin, 1983). Pantothenic acid deficiencies in rats result in death after 4–6
weeks exposure to the deficient diet, largely due to heamorrhagic necrosis of
the adrenals, reduced antibody production (Ralli and Dumm, 1953) and
thiamin deficiency. The vitamin requirements of rats are given in Table 19.3.

CONCLUSIONS
The diversity of species addressed in this chapter dictated that each was dealt
with in less depth than in those chapters dedicated to a single species. The
overriding feature to emerge from this chapter was the variability in nutrient
requirements both within classes of companion animals and also within a
species, particularly dogs. The data reported here are valid for the target
animal within a prescribed set of experimental conditions, but clearly it would
be a gargantuan and probably impossible task to determine the nutritive
requirements of every breed at each life stage under every conceivable
exercise regime and set of housing conditions. Therefore, the values presented
here should be regarded as guidelines. The unusual metabolism of the cat,
rendering it dependent on at least some animal-derived nutrients, is of interest,
and the specific needs of the increasing population of geriatric dogs and cats
are areas for further research. The requirement of companion birds from the
time they are hatched for a variety of seed types suitably supplemented to
meet their nutritional demands must be considered and, as more avian species
become popular as companion birds, further investigations as to their specific
needs will be necessary. The challenges of feeding communities of many
species of ornamental fish with diametrically opposed needs continue to
require detailed research into effective feeding strategies as well as nutrient
requirements. Although much of the data on rabbits was obtained from
animals kept under laboratory conditions, many companion rabbits live in
outdoor hutches, and therefore allowances must be made for extra nutrient
demands under variable conditions. However, several points have emerged
which are common for all companion animals in that highly digestible diets
with ‘ideal protein’ in the form of amino acids which closely conform to the
animals requirements are desirable to reduce pollution and to prevent

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The Nutrition of Companion Animals 463

imbalances with associated wastage of poorly utilized nutrients. From the

foregoing chapters, it is clear that companion animal nutrition is a relatively
recent area of research in comparison with that of farmed livestock for which
models to predict the responses of animals to different feeding systems
abound. It is to be hoped that future research will yield similar models for
companion animals.

REFERENCES
Adibi, S.A. (1980) Roles of branched chain amino acids in metabolic regulation. Journal
of Laboratory Clinical Medicine 95, 475–477.
Ahmad, B. (1931) The fate of carotene after absorption in the animal. Biochemical
Journal 25, 1195–1204.
Allison, J.B., Anderson, J.A. and Seely, R.D. (1947) Some effects of methionine on the
utilisation of nitrogen in the adult dog. Journal of Nutrition 33, 361–370.
Apgar, J.L., Shively, C.A. and Tarka, S.M. Jr (1987) Digestibility of cocoa butter and corn
oil and their influence on fatty acid distribution in rats. Journal of Nutrition 117,
660–665.
Aschoff, J. and Pohl, H. (1970) Rhythmic variation in energy metabolism. Federation
Proceedings 29, 1541–1552.
Baker, J. (1990) Dangers in vitamin overdose. Cage Aviary Birds 31, 5–6.
Barbul, A., Sisto, D., Wasserkrug, H.L. and Efron, G. (1981) Arginine stimulates
lymphocytic immune response in healthy human beings. Surgery 90, 244–257.
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