8.

Protein Requirements
M.J. Fraga
Departamento de Producción Animal, ETS Ingenieros Agrónomos,
Universidad Politécnica, 28040 Madrid, Spain

Introduction
To synthesize proteins (for example meat, milk and hair proteins) the rabbit
simultaneously requires all constituent amino acids. Some amino acids, those
that the animal does not synthesize, are defined as essential and have to be
supplied by the diet. The requirements of animals therefore are for amino
acids, rather than for protein.
Thus, the dietary protein level necessary to meet the requirements of the
rabbit vary according to: (i) its amino acid profile; (ii) the degree to which the
protein is digested; and (iii) the amount of feed ingested which, in turn, de-
pends on the dietary digestible energy (DE) concentration. Consequently,
information about the digestible essential amino acid levels in relation to the
DE content of diets would be extremely valuable. However, there is only
limited information available on this in rabbits, such that it is not possible to
express requirements in such units.
Currently, information on the dietary digestible protein/digestible energy
(DCP/DE) ratio is valuable because it includes two of the most important
variable factors mentioned above. The utilization of digestible units to
express protein requirements is obviously more appropriate because of the
considerable differences in protein digestibility between feed ingredients, the
average values expressed as coefficients for protein concentrates, cereals,
forages and by-products being 0.79, 0.73, 0.61 and 0.55, respectively
(Villamide and Fraga, 1998).
Information on the optimal dietary DCP/DE ratio should be complement-
ed with the amino acid requirements expressed, if possible, in terms of
digestible amino acids. Although recommended digestible levels of the more
limiting amino acids have recently been published (Taboada et al., 1994,
1996; de Blas et al., 1996), data on amino acid digestibility of different feed-
stuffs are still limited.
This chapter reviews the information that permits the calculation of daily
protein requirements (expressed in g DCP day–1) of rabbits. The practical
© CAB INTERNATIONAL 1998. The Nutrition of the Rabbit
(eds C. de Blas and J. Wiseman) 133
134 M.J. Fraga

recommendations presented in Chapter 13 should be used until more

information about amino acid requirements becomes available.

Maintenance requirements
There are unavoidable (obligatory) losses of amino acids from the body that
require replacement to maintain body conditions constant. These losses are
important in tissues where there is significant sloughing of cells such as in
skin, hair and intestinal mucosae. The requirements to support these activities
are determined as the intercept of the regression equation which relates the
amounts of an amino acid ingested to the quantities retained, and are called
maintenance requirements. There are few data on maintenance requirements
for amino acids in rabbits, but the values for crude protein (CP) are 2.9 and
3.7 g digestible CP kg LW–0.75 day–1 in growing and doe rabbits, respectively
(Box 8.1).

Box 8.1. Protein requirements for rabbitsa.

1. Maintenance
(i) Growing animals: 2.9 g DCP kg LW–0.75 day–1
(see text)
(ii) Pregnant and lactating does: 3.7–3.8 DCP kg LW–0.75 day–1 (refs 1, 2).
2. Milk production
(i) Protein content of milk: 115 g CP kg–1 (360 g kg–1 on a DM basis) (refs 3, 4).
(ii) Efficiency of utilization of dietary digestible protein (DCP) in milk protein synthesis: 0.76–
0.80 (refs 1, 2).
(iii) Efficiency of utilization of body protein for milk protein synthesis: 0.59–0.61 (refs 1, 2).
3. Pregnancy
(i) Efficiency of utilization of dietary DCP for fetal growth: 0.42–0.46 (refs 1, 3).
4. Growth
(i) Protein content in live weight gain: 180 g CP kg–1 (600 g kg–1 DM) (refs 6, 7, 8).
(ii) Efficiency of utilization of dietary DCP for body protein synthesis: 0.56 (see text).
a
If the average live weight of the growing rabbit is known, the maintenance requirements in
DCP day–1 can be calculated (1(i)). If the rate of growth, the protein content in weight gain (4(i))
and the efficiency of utilization of dietary DCP for body protein synthesis (4(ii)) are known, the
growth requirements in DCP day–1 can be calculated. The sum is the total requirement in DCP
day–1 of a growing rabbit. The same procedure may be used for the other physiological
situations.
References: 1, Xiccato et al., 1992; 2, Parigi Bini et al., 1991; 3, Fraga et al., 1989; 4, Xiccato
et al., 1995; 5, Parigi Bini et al., 1992; 6, García et al., 1992; 7, Motta Ferreira et al., 1996; 8,
Fernández and Fraga, 1996b.
 Protein Requirements 135

Requirements for growth

Body composition

The amino acid composition of whole body protein of growing rabbits is

presented in Table 8.1. There is a notable similarity with the amino acid
pattern of other species such as pigs or rats; the main difference is related to
the high cystine content (a high proportion of hair to rabbit whole body, and a
high level of cystine in hair protein). On the other hand, the methionine
content of the body of rabbits is low. There is a close agreement between the
sum of the individual amino acid levels and the CP content.
When comparing rabbits of different breeds and strains at the same live
weight, the differences are very small for body CP content. However, a higher
content of body fat and energy and a lower content of water is observed in
fast-growing rabbits.
In the same way, the influence of body weight and sex on the body CP
content is small. The males had a slightly higher body CP content than females
(1–2%; Fraga et al., 1983; Fernández, 1993), but the differences for amino
acids due to sex are small and generally not significant (Moughan et al., 1988).

Table 8.1. Amino acid composition (mg g–1 N) of the whole growing bodya and of the milk of doeb
rabbits.

Whole body Milk

Absolute Relative Absolute Relative
Amino acids value to lysine value to lysine
Lysine 383 100 451 100
Alanine 365 74 228 50
Arginine 415 108 328 73
Aspartic acid 467 121 451 100
Histidine 193 50 159 35
Isoleucine 194 51 304 67
Leucine 429 112 567 125
Methionine 77 20 150 33
Cystine 158 41 175 39
Glutamic acid 788 205 1220 270
Glycine 466 121 106 23
Phenylalanine 249 65 281 62
Serine 283 74 228 50
Threonine 245 64 305 67
Tyrosine 192 50 332 73
Valine 239 62 382 85
a
53-day-old New Zealand rabbits (Moughan et al., 1988).
b
New Zealand × Californian doe rabbits (M.J. Fraga, unpublished data).
136 M.J. Fraga

The diet does not affect the body CP content, except when the rate of
growth is modified. In any case, a variation of 10 g day–1 in the rate of growth
(i.e. from 35 to 45 g day–1) results in a variation in the body CP content of
rabbits of only 2% (Fraga et al., 1983).

Retained protein and efficiency of growth

As a consequence of the lack of variation in body CP content at slaughter, the

CP content of live weight gain is generally constant (Box 8.1) and, therefore,
daily protein retention (and daily requirements) depend on the rate of growth
of the rabbit. Following from this, the efficiency of utilization of total
digestible CP ingested increases with the rate of growth because the relative
contribution of maintenance requirements of protein decreases.
Protein retention decreases linearly with an increase in slaughter weight
(Table 8.2). In the later stages of the growing period the rate of growth is
lower. As a consequence, the efficiency of DP utilization decreases regularly
as slaughter weight increases. Males retain more protein, grow faster (8.7%,
Partridge et al., 1989; 4%, Fernández, 1993) and are slightly more efficient at
DP utilization (Table 8.2) than females.
From the data obtained by Partridge et al. (1989), Motta Ferreira et al.
(1996) and Fernández and Fraga (1996b), the following regression equation
was derived relating retained protein and total digestible protein ingested:
DCPI = 2.88 + 1.78 CPR, P < 0.0001, R2 = 0.85, n = 17,
where DCPI and PR are digestible crude protein ingested and protein
retained, respectively, as g kg LW–0.75 day–1. From this equation, the intercept
(2.9 g kg LW–0.75 day–1) corresponds to the maintenance requirements of
growing rabbits, and 0.56 (1/1.78) is the amount of dietary digestible CP used
for growth that is retained in the body of rabbits (Box 8.1).

Table 8.2. Influence of slaughter weight and sex on retained protein and on overall efficiency of
digestible crude protein utilization for growing (New Zealand × Californian) rabbits.

Slaughter weight (kg) Sex

2.0 2.5 Male Female
Daily protein retention
(g kg LW–0.75 day–1)a 5.4 4.7 5.2 4.8
Overall DCP efficiencya 0.39 0.35 0.38 0.37
Daily protein retention
(g kg LW–0.75 day–1)b 4.6 3.9
Overall DCP efficiencyb 0.37 0.32
a
Fernández and Fraga (1996b).
b
García et al. (1992).
 Protein Requirements 137

Requirements of does
Milk production

Amino acids are necessary to synthesize milk protein in the mammary gland.
Some amino acids may also be used for gluconeogenesis by does fed diets
with low starch levels. In these diets the uptake of glucose from the gut may
be insufficient to meet the requirements for milk lactose synthesis. De Blas et
al. (1995) obtained a decrease of feed efficiency in doe rabbits fed diets with
low levels of starch (the use of amino acids for gluconeogenesis implies a low
efficiency in digestible energy utilization); the optimal value being obtained
with a level of 190–210 g dietary starch kg–1. During lactation, mobilization
of body protein may occur to support synthesis of milk protein, mainly when
lactation and pregnancy are simultaneous.
Milk production is high in hybrid does, the main factors contributing to
its variation being the litter size and the remating interval. Total milk
production can be predicted from easily measurable traits such as litter live
weight at 21 days, which is when the pups began to consume solid feed. Using
the data of Méndez et al. (1986), Fraga et al. (1989) and de Blas et al. (1996),
the following regression equation was obtained (to a litter weight at 21 days
from 2 to 3 kg):
MP = 1.77 + 1.39 LW21, P < 0.001, R2 = 0.88, n =13,
where MP is total milk production (kg) and LW21 is the litter weight (kg) at
21 days of lactation.
Daily milk production continuously increases from parturition to 17–21 days.
The relatively low incremental requirements for milk yield in early lactation
allows milk yield and live weight gain to occur simultaneously.
From 21 to 30 days, milk yield decreases but at different rates in pregnant
and in non-pregnant does. Does re-mated 1 day after parturition showed a
lower total milk yield than those re-mated 8 days later (approximately 0.5 kg,
with 0.95 of the decrease occurring during the last week of lactation). Does
that are not pregnant during lactation yield about 1 kg more than those re-
mated at parturition (0.76 of the difference occurring during the last week).
These amounts should be used to correct the values obtained by applying the
equation.
The CP content of milk of does approaches 115 g kg–1 (Box 8.1).
However, a high CP content (about 140 g kg–1; Partridge and Allan, 1982;
Pascual et al., 1996) was observed during the last week of lactation in
lactating pregnant does. The efficiencies of utilization of dietary digestible
protein and body protein for milk protein synthesis are shown in Table 8.1.
The type of diet will not substantially alter the CP content and the amino
acid composition of milk. The more noticeable differences in the amino acid
patterns of milk of does (Table 8.1), cows and pigs are the low contents of
138 M.J. Fraga

proline and serine in doe milk. The methionine/methionine + cystine ratio is

also lower and can be related to the high requirements for the postnatal
growth of pups, which is characterized by the rapid growth of skin and hair.
With respect to lysine content, the threonine and sulphur amino acid levels of
doe milk are higher than in other species.
During lactation, in non-pregnant does, there is a mobilization of fat
deposits, and the empty body shows an increase in water. However, the body
protein content remains virtually unchanged, or shows a positive protein
balance (Box 8.2).

Pregnancy

During pregnancy, amino acids are retained in the conceptus, in the mammary
glands and in the maternal body. The importance of protein changes in the
body of the does during pregnancy is higher whenever the does are
concurrently lactating.

First gestation
During the initial two thirds of pregnancy, body protein retention of does is
higher than that of the conceptus (Box 8.2). However, to support the high
growth of this in the last third of pregnancy, a portion of body protein is

Box 8.2. Protein balance in does.

(a) Partition of weight gain and retained protein during the first pregnancy of does (Parigi Bini et
al., 1990).
0–21 days 21–30 days
Doe’s empty Doe’s empty
body Conceptus body Conceptus
Empty body gain (g) 180.2 192.3 –90.5 453.4
CP content (g kg–1 DM basis) 580 620 630 610
Protein retained (g) 60.3 18.3 –21.9 54.3
(b) Protein balance during the first lactation of does (Xiccato et al., 1995).

Physiological state
Lactating and pregnant Lactating non-pregnant
Empty body gain (g) –131 184
Retained protein (g) –38 75
Protein balance (%) –6 11
(c) Composition of litters (Xiccato et al., 1995).
Water: 808 g kg–1, CP: 123 g kg–1, 615 g kg–1 DM, fat: 50 g kg–1, 250 g kg–1 DM, energy: 4.56 MJ
kg–1 (22.8 MJ kg–1 DM).
 Protein Requirements 139

mobilized. From data on the body composition of pups at birth (Box 8.2),
Partridge and Allan (1982) have determined that between 0.52 and 0.84 of
retained protein is used to support fetal growth, with the remainder being for
intrauterine deposition. The rapid turnover in fetal protein may explain why
the efficiency of utilization of dietary digestible protein for pregnancy is
lower than that for growth (Box 8.1).

Simultaneous pregnancy and lactation

The overlapping of pregnancy and lactation increases the protein
requirements in response to the concurrent high demands for protein for fetal
growth and milk yield. As a consequence, a negative balance of body protein
was observed (Box 8.2) in pregnant lactating rabbits.

Digestible protein/digestible energy ratio of rabbit diets

The protein requirements, estimated as g DCP day–1 using the data in Box 8.1,
can be converted to dietary concentrations for the purposes of feed
formulation if the actual daily feed intake of rabbits is known. As the quantity
of energy ingested daily in terms of DE tends to be constant, the feed intake of
rabbits can be predicted from the energy concentration of the diet. As a
consequence, it is advisable to express the protein requirements as a DCP/DE
ratio.
According to de Blas et al. (1985), the requirements in terms of DCP/DE
ratio for maintenance are close to 6.8 g MJ–1, implying that maintenance
represents a high energy cost in relation to protein; for average growth the
requirement for protein is much higher. The young rabbit multiplies its birth
weight sixfold during the first 3 weeks of life, in which milk (with a ratio of
13–14 g CP MJ–1 GE) is its only feed. From 21 days to weaning a progressive
change is made from milk to doe feed. Later, the rate of growth in relation to
live weight decreases (30–45 g day–1 up to 8 weeks). The best results in
growing rabbits measured in productivity terms were obtained by de Blas et
al. (1985) using Spanish Giant rabbits with diets containing about 10 g DCP
MJ–1 DE (see Chapter 13). In countries where rabbits are raised up to 2.5–2.8 kg
live weight, and using two or three different growing feeds, a decrease in
dietary protein levels should be considered in the latter stages (see earlier,
‘Retained protein and efficiency of growth’).
However, the recommended value should be revised when using current
fast-growing commercial strains. It is possible that the daily protein
requirements do not vary, but the higher fat content in the live weight gain of
fast-growing rabbits can decrease the recommended value of the DCP/DE
ratio. However, more information is necessary on the effect of selection for
growth on body composition.
Daily protein requirements for lactating does are relatively higher than
140 M.J. Fraga

for growing rabbits. Because of this, and taking into account the difficulty of
obtaining a high feed intake during the lactation of highly productive does,
the DCP/DE ratio recommended is higher than for growing rabbits, and varies
from 11.0 to 12.5 g MJ–1 according to performance (see Chapter 13).

Amino acid requirements

The actual amino acid intake of rabbits is the sum of amino acids provided by
the diet and from reingestion of soft faeces. In rabbits fed conventional diets
the contribution of soft faeces to total intake of CP is around 0.17 (see Chapter
3) but the amino acid intake and its relation to dietary composition remains to
be established, because of the limited information concerning the variation in
amino acid composition of soft faeces. As a consequence, the amino acid
requirements of growing and doe rabbits that have been determined to date,
mainly by dose–response trials (see Chapter 13 for practical recommenda-
tions), should be used with caution.
The data obtained by Moughan et al. (1988) allow estimation of the
amino acid requirements of growing rabbits according to their respective
proportions of body protein. To apply this method it is necessary to know the
optimal level, measured in productivity terms, for a single amino acid (for
example lysine). From the data on amino acid composition of doe milk that
are presented in Table 8.1, the amino acid requirements of lactating females
may also be estimated according to their respective ratios in milk protein.
In fact, the values obtained for threonine and sulphur amino acids with
respect to lysine content in milk (see Table 8.1) compare well with the
optimal relative requirements of these amino acids (0.68 and 0.76 for
threonine and sulphur amino acids, respectively) expressed in digestible units
obtained by Taboada et al. (1966) and de Blas et al. (1996), respectively. In
the same way, the optimal values of amino acids for growth obtained by the
same authors were consistent with the data on body amino acid composition
(Table 8.1).
However, the use of the pattern of amino acids in doe milk and in the
whole body of growing rabbits to define their amino acid requirements
assumes that the maintenance requirements are only a small proportion of the
total amino acid requirements during lactation and growth. This proportion is
low in pigs (approximately 0.05 in lactating sows; Pettigrew, 1995) but there
are no data for rabbits. However, using the data in Box 8.1, the protein
maintenance requirements of growing and doe rabbits are, respectively, about
0.30 and 0.25 of total protein requirements. The contribution of doe hair loss
(approaching 300–500 mg day–1 in the 5 days before parturition; Sawin et al.,
1960) and its growth during the next lactation to maintenance requirements
may partially explain these differences. In the same way, there is a
mobilization of body protein stores in pregnant and lactating doe rabbits to
 Protein Requirements 141

support protein synthesis. However, in does fed diets with high CP contents,
slightly positive balances can be obtained. More information is necessary to
establish a method, similar to the one proposed by Pettigrew (1995) for pigs,
where the amino acid requirements determined for different functions
(replenishment of obligatory losses, mobilization of body protein) could be
integrated with data on the amino acid composition of different products. Any
improvement in the knowledge of amino acid requirements would allow a
more appropriate use of synthetic amino acids with consequent reductions in
the nitrogen content of diets and excreta, in the energy losses and in the risks
of digestive disturbances.

References
de Blas, J.C., Fraga, M.J. and Rodríguez, J.M. (1985) Units for feed evaluation and
requirements for commercially grown rabbits. Journal of Animal Science 60,
1021–1028.
de Blas, J.C., Taboada, E., Mateos, G., Nicodemus, N. and Méndez, J. (1995) Effect
of substitution of starch for fiber and fat in isoenergetic diets on nutrient
digestibility and reproductive performance of rabbits. Journal of Animal Science
73, 1131–1137.
de Blas, J.C., Taboada, E., Nicodemus, N., Campos, R., Piquer, J. and Méndez, J.
(1996) The response of highly productive rabbits to dietary threonine content for
reproduction and growth. In: Lebas, F. (ed.) Proceedings of the 6th World Rabbit
Congress, Toulouse, Vol. 1. Association Française de Cuniculture, Lempdes, pp.
139–143.
Fernández, C. (1993) Efecto de la incorporación de grasa en piensos fibrosos sobre la
utilización digestiva de la dieta, crecimiento y calidad de la canal de conejos en
cebo. PhD Thesis, Universidad Politécnica de Madrid, Spain.
Fernández, C. and Fraga, M.J. (1996a) The effect of fat inclusion on growth
performance, carcass characteristics, and chemical composition of rabbits.
Journal of Animal Science 74, 2088–2094.
Fernández, C. and Fraga, M.J. (1996b) Effect of fat inclusion in diets for rabbits on
the efficiency of digestible energy and protein utilization. World Rabbit Science
4, 19–23.
Fraga, M.J., de Blas, J.C, Pérez, E., Rodríguez, J.M., Pérez, C.J. and Gálvez, J.F.
(1983) Effect of diet on chemical composition of rabbits slaughtered at fixed
body weights. Journal of Animal Science 56, 1097–1104.
Fraga, M.J., Lorente, M., Carabaño, R.M. and de Blas, J.C. (1989) Effect of diet and
remating interval on milk production and milk composition of the doe rabbit.
Animal Production 48, 459–466.
García, G., Gálvez, J.F. and de Blas, J.C. (1992) Substitution of barley grain by
sugar-beet pulp in diets for finishing rabbits. 1. Effect on energy and nitrogen
balance. Journal of Applied Rabbit Research 15, 1008–1016.
García, G., Gálvez, J.F. and de Blas, J.C. (1993) Effect of substitution of sugarbeet
pulp for barley in diets for finishing rabbits on growth perfomance and on energy
142 M.J. Fraga

and nitrogen efficiency. Journal of Animal Science 71, 1823–1830.

Méndez, J., de Blas, J.C. and Fraga, M.J. (1986) The effects of diet and remating
interval after parturition on the reproductive perfomance of the commercial doe
rabbit. Journal of Animal Science 62, 1624–1634.
Motta Ferreira, W., Fraga, M.J. and Carabaño, R. (1996) Inclusion of grape pomace,
in substitution for lucerne hay, in diets for growing rabbits. Animal Science 63,
167–174.
Moughan, P.J., Schultze, W.H. and Smith, W.C. (1988) Amino acid requirements of
the growing meat rabbit. 1. The amino acid composition of rabbit whole-body
tissue – a theoretical estimate of ideal amino acid balance. Animal Production
47, 297–301.
Parigi Bini, R., Xiccato, G. and Cinetto, M. (1990) Energy qnd protein retention and
partition in rabbit does during the first pregnancy. Cuni-Science 6, 12–29.
Parigi Bini, R., Xiccato, G. and Cinetto, M. (1991) Utilizzazione e ripartizione
dell’energia e della proteina digeribile in coniglie non gravide durante la prima
lattazione. Zootecnica e Nutrizione Animale 17, 107–120.
Parigi Bini, R., Xiccato, G., Cinetto, M. and Dalle Zotte, A. (1992) Energy and
protein utilization and partition in rabbit does concurrently pregnant and
lactating. Animal Production 55, 153–162.
Partridge, G.G. and Allan, S.J. (1982) The effects of different intakes of crude protein
on nitrogen utilization in the pregnant and lactating rabbit. Animal Production
35, 145–155.
Partridge, G.G., Garthwaite, P.H. and Findlay, M. (1989) Protein and energy retention
by growing rabbits offered diets with increasing proportions of fibre. Journal of
Agricultural Science 112, 171–178.
Pascual, J.J., Cervera, C., Blas, E. and Fernández-Carmona, J. (1996) Milk yield and
composition in breeding rabbit does using high fat diets. In: Lebas, F. (ed.)
Proceedings of the 6th World Rabbit Congress, Toulouse, Vol. 1. Association
Française de Cuniculture, Lempdes, pp. 259–262.
Pettigrew, J.E. (1995) Amino acid requirements of breeding pigs. In: Haresign, W.
(ed.) Recent Advances in Animal Nutrition. Butterworths, London,
pp. 241–256.
Sawin, P.B., Deneberg, V.H., Ross, S., Hafter, E. and Zarrow, M.X. (1960) Maternal
behavior in the rabbit: hair loosening during gestation. American Journal of
Physiology 198, 1099–1102.
Taboada, E., Méndez, J., Mateos, G.G. and de Blas, J.C. (1994) The response of
highly productive rabbits to dietary lysine content. Livestock Production Science
40, 329–337.
Taboada, E., Méndez, J. and de Blas, J.C. (1996) The response of highly productive
rabbits to dietary sulphur amino acid content for reproduction and growth.
Reproduction, Nutrition and Development 36, 191–203.
Villamide , M.J. and Fraga, M.J. (1998) Prediction of the digestible crude protein and
protein digestibility of feed ingredients for rabbits from chemical analysis.
Animal Feed Science and Technology 70, 211–224.
Xiccato, G., Parigi Bini, R., Cinetto, M. and Dalle Zotte, A. (1992) The influence of
feeding and protein levels on energy and protein utilization by rabbit does.
Journal of Applied Rabbit Research 15, 965–972.
Xiccato, G., Parigi Bini, R., Dalle Zotte, A., Carazzolo, A. and Cossu, M.E. (1995)
 Protein Requirements 143

Effect of dietary energy level, addition of fat and physiological state on

perfomance and energy balance of lactating and pregnant rabbit does. Animal
Science 61, 387–398.