See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/232010995

Acid-base balance in broilers

Article  in  World's Poultry Science Journal · March 2007

DOI: 10.1017/S0043933907001286

CITATIONS READS

47 416

3 authors, including:

Ana Vitória Fisher da Silva Alex Maiorka

Universidade Federal do Paraná Universidade Federal do Paraná
31 PUBLICATIONS   627 CITATIONS    153 PUBLICATIONS   1,239 CITATIONS   

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Amylase View project

Evaluation of pelleting parameters of broiler diets View project

All content following this page was uploaded by Alex Maiorka on 27 August 2015.

The user has requested enhancement of the downloaded file.

010028_Journal_4 01-02-2007 15:27 Pagina 73

DOI: 10.1079/WPS2005128

Acid-base balance in broilers

S.A. BORGES, A.V. FISCHER DA SILVA* and A. MAIORKA

Universidade Federal do Paraná – UFPR, 81530-000 Curitiba PR Brasil

Corresponding author: avitoria@ufpr.br

High ambient temperature could result in numerous physiological and metabolic

changes in broilers, that adversely impact broiler performance and immune
response. In addition to environmental control techniques that have been frequently
used to reduce the negative impact of heat stress on birds’ performance, other
measures have been studied. Lately, proper nutritional management has shown to be
effect as a preventive measure against heat stress because the function of the
thermoregulating system of broilers (heat production, evaporative and non-
evaporative routes for heat dissipation) can be influenced by diet. This applies
particularly to the establishment of proper electrolyte balances due to their
physiological importance in the heat stress mechanism. Thus, nutritional
mechanisms should be reassessed as a tool to control this metabolic dysfunction in
birds.

Keywords: blood parameters; broilers; electrolyte balance; heat stress

Introduction
The evolution of poultry production has resulted in an young broiler with a high efficiency
in converting different types of feed into animal protein. In spite of this, a number of
metabolic and management problems have emerged, among them heat stress. The
susceptibility of birds to heat stress increases as both relative humidity and ambient
temperature exceed the thermal comfort zone, thus making heat dissipation difficult and,
as a consequence, increasing the bird’s body temperature, adversely impacting
performance.
Some measures can be taken in order to minimize losses due to heat stress, including the
use of fans and nebulisers, management of protein and energy in the diet, acclimatization
of birds, use of anti-thermal agents, ascorbic acid, electrolytes, feeding management and
management of drinking water. One of the consequences of stress is the change in the
acid-base balance with the occurrence of respiratory alkalosis. Therefore, one of the
methods used to control heat stress is the chemical management of the acid-base balance
in birds by compounds such as sodium bicarbonate (NaHCO3), potassium chloride (KCl),
calcium chloride (CaCl2) and ammonia chloride (NH4Cl) in water and/or feed (Borges,
1997). The object of this paper is to discuss the detrimental effects of heat stress on birds
and some techniques that minimize them.

© World’s Poultry Science Association 2007

World’s Poultry Science Journal, Vol. 63, March 2007
Received for publication January 17, 2005
Accepted for publication June 6, 2006 73
010028_Journal_4 01-02-2007 15:27 Pagina 74

Heat stress physiology in birds: S.A. Borges et al.

Heat stress physiology in birds

Being homeotherms, birds have a thermoregulating centre located at the hypothalamus

that is able to control body temperature by physiological mechanisms and behavioural
responses by producing and releasing heat, thus determining the maintenance of normal
body temperature (Macari et al., 1994).
Among the compensatory physiological responses of birds when exposed to heat is
peripheral vasodilation, resulting in an increase in non-evaporative heat loss. Therefore, in
an attempt to increase heat dissipation, birds manage to increase their surface area by
keeping their wings spread away from their bodies, ruffling their feathers and intensifying
peripheral circulation. Non-evaporative heat loss can also take place with the increase in
urine output if this water loss is offset by greater cool water consumption.
Another physiological response is the increase in respiratory rate, resulting in excessive
carbon dioxide (CO2) losses. Therefore, partial CO2 pressure (pCO2) decreases, causing a
decrease in the concentration of carbonic acid (H2CO3) and hydrogen (H+). In response to
that, kidneys increase HCO3- excretion and reduce H+ excretion in an attempt to keep the
bird’s acid-base balance. This change in the acid-base balance is called respiratory
alkalosis.

HEAT STRESS VS. BLOOD PARAMETERS

The blood system is particularly sensitive to changes in temperature, being an important
indicator of physiological responses in birds to stressing agents. Quantitative and
morphological changes in blood cells are associated with heat stress and translated by
variations in haematocrit value, number of circulating leukocytes, erythrocyte content and
haemoglobin content erythrocytes.
There is an increase in haematocrit counts with heat stress, which can be explained by
an increase in the number of red blood cells. The heterophil/lymphocyte ratio is altered as
a result of the increase in heterophils and reduction in lymphocytes, with the
heterophil/lymphocyte ratio having been proposed as a sensitive measure of chronic stress
in broilers. Another response is the increase in glucose concentration as a direct response
to greater adrenaline, noradrenaline and glucocorticoids secretion (Borges, 1997; Borges,
2001).

HEAT STRESS VS. ELECTROLYTES

Electrolytes can be defined as chemicals that break down into their ionic constituents,
having as their main physiological function the maintenance of the body acid-base
balance. Sodium (Na+), potassium (K+) and chloride (Cl-) are essential ions for the
maintenance of osmotic pressure and acid-base balance of body fluids. Therefore, the
effects of ionic balance in the diet on the broiler performance can be related to variations
in the acid-base balance (Mongin, 1981).
K+ is the main cation of the intracellular fluid, while Na+ and Cl- are the main ions of the
extracellular fluid. Osmoregulation is achieved by the homeostasis of these intra- and
extracellular ions. Under optimal conditions, water and electrolyte contents are kept
within narrow limits. However, electrolyte (Na+ or K+) loss without any change in body
water content reduces the osmolality of these fluids.
K+ is involved in many metabolic processes, including the arginine-lysine antagonism,
nervous conduction, excitement, muscle contraction, synthesis of tissue proteins,
maintenance of intracellular homeostasis, enzymatic reactions, osmotic balance and acid-
base balance. As a consequence, changes in K+ homeostatis can affect cell function.
Studies with thermal dehydration followed by rehydration in humans have shown that the
degree of water deficit in intracellular fluid was associated with loss of intracellular K+ and

74 World’s Poultry Science Journal, Vol. 63, March 2007

010028_Journal_4 01-02-2007 15:27 Pagina 75

Heat stress physiology in birds: S.A. Borges et al.

the extracellular fluid deficit was linked to loss in plasma Na+. The degree of intracellular
rehydration was determined by restoring K+ (Nose et al., 1988).
In mammals, respiratory alkalosis causes a reduced competition between H+ and K+ for
urinary excretion, thus increasing K+ loss in the urine. Excess K+ ions compete with buffer
anions in the renal tubule fluid, preventing H+ removal, which in turn is reabsorbed, and
the result maybe acidosis (Bacila, 1980). When this mechanism is present in broilers, it
may increase K+ requirements during heat stress. Recent evidence shows that the
intercalated cells of the collecting duct, which secret acid, also secret H+ and this process
is enhanced by hypokalemia and seems to be a strong contributor to renal acidification
(Cunningham, 1999).
Plasma Na+, K+ and Cl- levels are affected by heat stress. K+ and Na+ concentration
decreases as temperature rises, (Borges, 1997), while Cl- increases (Belay and Teeter,
1993).
The increase in Cl- decreases H+ excretion and HCO3 reabsorption by the kidneys. This
might contribute to blood acidification which, in turn, seems to be an appropriate response
to alkalosis. However, stress time should be taken into account, since Salvador et al.,
(1999) observed a reduction in Cl- serum levels when broilers were subjected to chronic
stress for one week (42 to 49 days).
K+ serum levels are also influenced by heat. K+ excretion is influenced by hormonal
factors (aldosterone, antidiuretic hormone - ADH and deoxycorticosterone), acid-base
balance and cation balance. K+ excretion rate by urine is variable, being connected to Na+
serum concentrations and the bird’s hydration status, where losses can be caused by an
increase in water consumption, since the osmotic gradient favours the movement of
intracellular fluid water to the urine, which may carry K+ . The increase in K+ intake results
in greater urinary loss, with the bird having little capacity of conserving body K+. K+ serum
levels decrease during stress (Borges, 1997; Salvador et al., 1999). The decrease in K+
plasma levels is attributed to an increase in excretion of this ion during chronic stress and
an increase in intracellular K+, which is ordinarily found in acute stress. In humans,
hyperkalemia may result in metabolic acidosis, both due to the reduction in ammonia
excretion and to the limitation of carbonate reabsorption by the kidneys (Ait-Boulahsen et
al., 1995).

Nutritional measures to control heat stress

USE OF SALTS
The use of salts in drinking water or in the feed is an alternative that is frequently used
by broiler growers to reduce losses resulting from heat stress. Among the main salts used
are potassium chloride (KCl) and sodium bicarbonate (NaHCO3).
K+ is abundantly present in most ingredients of animal diets, unlike Na+, which is present
in nutritionally inadequate amounts in natural food used for animal nutrition.
KCl supplementation in the feed and/or drinking water for birds has been proposed as a
way to minimize the consequences of high temperatures on performance (Smith and
Teeter, 1993). By supplementing 0.50 and 1.00% of KCl in the feed of broilers raised in
summer, Borges (1997) concluded that there was a 3.5% improvement in weight gain
(P<0.05).
NaHCO3 has been used by the poultry industry in an attempt to minimize losses caused
by heat stress, particularly during summer time. Performance results show that a 0.5 and
1.0% supply of NaHCO3 added to the feed of broilers subjected to temperatures varying
from 39 to 41ºC and 34 to 36ºC led to a trend of improved feed consumption, weight gain
and feed conversion (Fischer da Silva et al., 1994). Borges (1997), By supplementing 0.5;

World’s Poultry Science Journal, Vol. 63, March 2007 75

010028_Journal_4 01-02-2007 15:27 Pagina 76

Heat stress physiology in birds: S.A. Borges et al.

1.0 and 1.5% of NaHCO3 in the feed of broilers during summer time and did not see any
effect on performance.
Weight gain in broilers is improved after KCl, NaCl and K2SO4 supplementation in
water or in the feed, although the acid-base balance in the birds is altered. The addition of
NH4Cl to restore blood pH to normal values had not effect on birds performance (Teeter
and Belay, 1996).

APPLICABILITY OF THE ELECTROLYTE BALANCE THEORY

The acid-base balance is influenced by the environment and the diet. Several reports
point to the effects of electrolyte balance in the diet on birds’ productive performance. The
maintenance of this balance can be an important measure to improve the performance of
broilers raised under high temperatures and to overcome the harmful effects of respiratory
alkalosis resulting from heat stress.
Diets formulated with high Cl contents (NH4Cl, HCl and CaCl2) decrease blood pH in
broilers, adversely affecting their growth under thermoneutrality. However, acid or base
intake, electrolyte balance, the environment, their interactions and implications on the
performance of birds still have to be defined.
In addition to carrying away a large amount of organic acid, heat stress may be
associated with electrolyte losses through cell membranes, (Fischer da Silva et al., 1994).
Prevention of electrolyte unbalance can be achieved by incorporating cations and anions
to the diet, being usually expressed in mEq/kg (Mongin, 1981). However, electrolyte
availability may be influenced by intestinal and renal homeostatic regulation and by the
greater absorption of monovalent ions.
Some authors have described equations to explain the relationship between cations and
anions and the acid-base balance. For Melliere and Forbes (1966) this interrelationship
can be described by the following equation:

Relative level = mEq cations = Ca + Mg + Na + K

mEq anions PO4 + Cl + SO4
However, the ions that are essential for the maintenance of the acid-base balance are:
Na+, K+ and Cl-. In addition to the minimum amounts birds need in their feed to meet their
nutritional requirements, it is essential that the proportion among these ions is optimal to
maintain the acid-base homeostasis and to get the best performance from the birds
(Mongin, 1981). In order to maintain the acid-base balance, birds have to regulate acid
intake and excretion. There are differences in the intake and excretion of anions and
cations in the diet. However, acids produced in metabolism (endogenous H+) also
contribute to the acid-base balance. The following equation can be described for situations
where birds are in a constant acid-base balance, without either acid or base excess or
deficiency:

ingested (Anions - Cations) t + endogenousH+ = excreted (Anions - Cations )

According to Mongin (1981) the result of the acid power of Na+ + K+ – Cl- intake is equal
to the difference of excreted cations and anions (excreted (cations – anions)) plus the
production of endogenous acid (endogenousH+), plus excess bases (BEecf) or alkaline reserves.
The optimal electrolyte intake, in terms of acid-base balance can minimize the presence of
BEecf, tending to zero. The optimal requirement of electrolyte balance was defined in
terms of mEq(Na+ + K+ - Cl-)/kg of feed around 250 mEq/kg.

(Anions - Cations) = excreted (Anions - Cations ) + endogenousH+ + BEecf

ingested
or,

76 World’s Poultry Science Journal, Vol. 63, March 2007

010028_Journal_4 01-02-2007 15:27 Pagina 77

Heat stress physiology in birds: S.A. Borges et al.

ingested (Na+ + K+ - Cl-) = excreted (Anions - Cations ) + endogenousH+ + BEecf

mEqNa + mEqK – mEqCl = 250

Example: 0.35%Na x 10000 / 23 = 152 mEq Na

0.68%K x 10000 / 39.1 = 174 mEq K
0.30%Cl x 10000 / 35.5 = 84 mEq Cl
152 + 174 – 84 = 242 mEq/kg of feed

In these equations, some factors have to be considered:

a) In the equation it is assumed that only the minerals Na, K and Cl have an impact on
the acid-base balance, without considering the way they are ingested. Sodium and
potassium supplementation increases pH and blood HCO3-, while chloride addition
decreases these parameters (Hurwitz et al., 1973). However, there is evidence that
metabolizable anions have an influence on the acid-base balance. Gorman and Balnave
(1994) concluded that the weight gain associated with sodium carbonate and sodium
bicarbonate was significantly different in diets with the same electrolyte balance, thus
concluding that heat stress can lead to a metabolic requirement for the bicarbonate ion;
b) The equation does not take into account either the specific effects of each ion or the
individual requirements for these ions, which may limit its use. We should bear in mind
that not always these ions are quantified in the ingredients, with K being abundantly
present in most ingredients that compose diets for birds, whereas Na is present in small
amounts. Nutritional recommendations for these electrolytes for broilers also vary, NRC
(1994) recommends 0.30, 0.20 and 0.20; 0.30, 0.15 and 0.15% for K, Na and Cl from 0 to
3 and 3 to 6 weeks of age, respectively;
c) Although other cations and anions also take part in the acid-base balance, they are not
taken into account in this equation due to their secondary importance. Therefore, taking
into account the electrolytical potential of elements we can classify them in terms of their
importance for the body acid-base balance. For example, K, Na and Cl have a greater
electrolytical potential than Mg, S, P and Ca, with the electrolytical potential of the latter
being greater than that of Fe, Mn, Zn, Cu, Se, Mo, Co and I. Trace elements have the
ability to function as electrolytes, but they are present in small amounts in feed and in low
concentrations in bird tissues, which naturally reduces their impact on the acid-base
balance and on the electrolyte balance in birds. Therefore, the complete electrolyte
equation would be: (Na+ + K+ + Ca++ + Mg++) - (Cl- + SO4= + 2PO4- + HPO4-) (Borges,
2001).
d) In relation to the elements that are not considered in the summarized equation by
Mongin (1981) we could say the following: bivalent cations are not as rapidly absorbed as
monovalent cations; magnesium (Mg) is commonly supplied in feeds; phosphate is hard
to be quantified because it comes from various sources; calcium absorption rate is
controlled by the endocrine system; sulphate is present in small amounts, being related to
methionine catabolism prevention.
e) The interrelationship between mineral ions and other nutrients, such as between Na
and Cl and the arginine: lysine ratio in heat-stressed broilers (Brake et al., 1998). In
broilers aged 3 to 7 weeks raised under heat stress, the optimal arginine: lysine ratio was
1.34 when Na and Cl were kept according to NRC (1994) recommendations. However,
when Na and Cl contents in the diet were increased, the arginine:lysine ratio was below
1.05, thus showing that under heat stress the adjustment of the electrolyte balance for
maximum bird performance can be related to the composition of amino acids in the diet.
Hurwitz (1981), unlike Mongin (1981), proposed that the balance between Na+ and Cl-
is what primarily determines HCO3- concentration and plasma pH. K+, Ca++ and Mg++

World’s Poultry Science Journal, Vol. 63, March 2007 77

010028_Journal_4 01-02-2007 15:27 Pagina 78

Heat stress physiology in birds: S.A. Borges et al.

make up the other cations and HCO3-, proteins and other anions in low concentrations
(phosphate, sulphate, lactate and piruvate) are plasma anions. HCO3- and proteins
(including hemoglobin) constitute basic buffers and like Beecf are considered metabolic
components of the acid-base balance. Beecfs Express the amount of acids or bases which,
when added to one litter of blood, return pH to normal. The basic buffer is considered as a
union between the acid-base balance and the electrolyte balance. The H+ ion is not present
in the diagram since it has influence in smaller concentrations than Cl-, K+ and other ions.
With variation in Na+ and Cl- levels in the feed, an improvement was observed in the
growth of birds when the Na Cl ratio was around 1:1 with the use of feeds containing
(Na++ K+ - Cl-) at 200mEq/kg (Hurwitz et al.,1973). Borges et al., (1999) found that the
best electrolyte balance varied according to the manipulated electrolyte. The authors
recommend that extreme Cl- (0.15 and 0.71%), K+ (0.98 and 1.21%) and Na+ levels (0.15
and 0.60%) should be avoided in pre-starter diets. Rondón et al., (2000) have also found a
variation in the best electrolyte balance depending on the manipulated ion. The best
electrolyte balance was of 250 mEq/kg, when Na+ levels varied and 319 mEq/kg when the
manipulated ion was K+. Changes in the acid-base balance and unbalances in (Na+ + K+ -
Cl-) supplementation caused loss of appetite, with reduction in weight gain, affecting feed
conversion, decreasing egg production and when unbalances are not compensated for
there is an increase in mortality rates (MONGIN, 1981). In birds with alkalosis, blood
concentrations of the electrolytes (Na+, K+) are decreased. A reduction in the alkalosis
status occurs when the Na:Cl ratio decreases, with the addition of 0.5 and 1.0% of CaCl2,
with an 8.0% improvement in bird performance (Teeter et al., 1985).
Johnson and Karunajeewa (1985) concluded that an electrolyte balance in the diet lower
than 180 mEq/kg and higher than 300 mEq/kg decrease the weight of the birds when
assessed at 42 days. An optimal electrolyte balance was found for feeds containing from
250 to 300mEq/kg. Likewise, Hulan et al. (1987), researching the effect of feeds
containing Na+ + K+ - Cl- in different ratios and still varying the calcium level, found that
the worst and the best weight gain were achieved when the “Mongin number” was 174 and
215 mEq/kg, with 1.38 and 0.95% calcium, respectively.
The protein source used in the feed can affect the electrolyte and acid-base balance,
because certain sources, particularly from animal by-products, increase the production of
organic acids and reduce Na and K contribution, increasing the relative amount of Cl
(Portsmouth, 1984). The supply of feeds constituted basically of soybean meal which has
low Na contents and high K contents, showed a significant response in the development of
broilers supplemented with 0.5 and 1.0% NaCl (March, 1984).
The impact of the cation/anion ratio on the acid-base balance in broilers, blood pH and
growth rate was assessed by Hurwitz et al. (1973). Broiler growth rate was greatest when
blood pH was 7.28, with a decrease in growth being found when pH values were greater
than 7.30 or lower than 7.20. The electrolyte ratio of the diet for the maximum growth
varied from 226 to 260mEq/kg. However, if the response is totally due to changes in pH
or to other electrolyte or metabolic effects is still to be defined. Teeter et al. (1985) have
shown that during panting pH values greater than 7.25 depress growth rate and feed
efficiency. Increases in blood pH can be reduced by a reduction in the respiratory rate of
birds. These increases can occur with acclimatization temperatures, but during acute stress
the respiratory rate can be reduced and heat loss efficiency can be improved by the
maintenance of high water consumption (Belay and Teeter, 1993). Mortality during heat
stress can be inversely related to water consumption (Branton et al., 1986).
Borges (2001) concluded that the response to electrolyte balance in the diet depends on
ambient temperature. The electrolyte ratio in the diet affects bird performance, with the
optimal ratio varying from 186 to 250mEq/kg. A high electrolyte balance (340 and 360
mEq) can result in metabolic alkalosis. In heat stress birds retain more electrolytes (Na, K

78 World’s Poultry Science Journal, Vol. 63, March 2007

010028_Journal_4 01-02-2007 15:27 Pagina 79

Heat stress physiology in birds: S.A. Borges et al.

and Cl) in an attempt to maintain the acid-base balance. The amount of electrolytes
excreted in the urine depends on their concentration in the feed and on ambient
temperature. Water consumption depends directly on bird age and on the Na+K-Cl ratio in
the feed, where an increase in water consumption caused by the greater Na+K-Cl ratio has
a direct impact on litter moisture and reduces rectal temperature in birds.
Borges et al. (2003 a) evaluated effects of dietary electrolyte balance [DEB; Na+K-Cl,
mEq/kg)] under summer conditions in Brazil. The DEB of 240mEg/kg gave best weight
gain and feed conversion ratio. Ideal DEBs predicted by regression analysis, based on
weight gains and feed conversion ratios, were 236 and 207 mEq/kg of feed from 0 to 42
days, respectively (Figure 1).
Male broiler chicks on new litter were used to evaluate effects of environmental
temperatures and various dietary electrolyte balances [DEB; Na+K-Cl, mEq/kg)] in
Brazil. Two temperature regimens were: 1) thermoneutral and 2) cyclic daily simulated
natural heat stress. The DEB treatments, made from a corn-soybean meal and soy oil basal
mash diet were: 40, 140, 240, and 360mEq/kg in starter and grower feeds. The response to
DEB depended on temperature. In thermoneutral temperature, the Pmax points for 21 and
42 day weight gains were at DEBs of 250 and 201 mEq/kg, respectively, with highest 42-
d feed intake at 220mEq/kg. Independent of ambient temperature and live performance,
the most normal acid-base balance (blood pH and base concentration) and heterophil:
lymphocyte ratio were obtained with feeds containing between 140 and 240 mEq/kg. By
treatment main effects, the DEB of 240 mEq/kg with 0.35% Na and 0.366% and 0.294%
Cl in starter (0.745% K) and grower (0.666% K), respectively, gave best 42-d weight gain
and feed conversion ratio (Table 1) (Borges et al., 2003 b).

Conclusions
The exposure of broilers to high ambient temperature results in respiratory alkalosis,
causing a drop in production performance. Feed formulation based on the electrolyte
balance concept, as well as the addition of salts to water and/or feed are practices that can
be implemented to correct distortions in the acid-base balance resulting from heat stress.

References
AIT-BOULAHSEN, A., GARLICH, J.D. and EDENS, F.W. (1995) Potassium chloride improves the
thermotolerance of chickens exposed to acute heat stress. Poultry Science 74: 75-87.
BACILA, M. (1980) Bioquímica Veterinária. Varela , São Paulo, 534 pp.
BELAY, T. and TEETER, R.G. (1993) Broiler water balance and thermobalance during thermoneutral and high
ambient temperature exposure. Poultry Science 72: 116-124.
BORGES, S.A. (1997) Suplementação de cloreto de potássio e bicarbonato de sódio para frangos de corte
durante o verão. MS Thesis, Universidade Estadual Paulista, Jaboticabal, São Paulo, Brasil.
BORGES, S.A. (2001) Balanço eletrolítico e sua interrelação com o equilíbrio ácido-base em frangos de corte
submetidos a estresse calórico. PhD Thesis, Universidade Estadual Paulista, Jaboticabal, São Paulo, Brasil.
BORGES, S.A., ARIKI, J., SANTIN, E., FISCHER da SILVA, A.V. and MAIORKA, A. (1999) Balanço
eletrolítico em dieta pré-inicial de frangos de corte durante o verão. Revista Brasileira de Ciência Avícola 1:
175-179.
BORGES, S.A., FISCHER da SILVA, A.V., ARIKI, J., HOOGE, D.M. and CUMMINGS, K.R. (2003a)
Dietary electrolyte balance for broiler chickens under moderately high ambient temperatures and relative
humidities. Poultry Science 82: 301-308.
BORGES, S.A., FISCHER da SILVA, A.V., ARIKI, J., HOOGE, D.M. and CUMMINGS, K.R. (2003b)
Dietary electrolyte balance for broiler chickens exposed to thermoneutral or heat-stress environments. Poultry
Science 82: 428-435.
BRAKE, J., BALNAVE, D. and DIBNER, J.J. (1998) Optimum dietary arginine:lysine ratio for broiler

World’s Poultry Science Journal, Vol. 63, March 2007 79

010028_Journal_4 01-02-2007 15:27 Pagina 80

Heat stress physiology in birds: S.A. Borges et al.

chickens is altered during heat stress in association with changes in intestinal uptake and dietary sodium
chloride. British Poultry Science 39: 639-647.
BRANTON, S.L., REECE, F.N. and DEATON, J.W. (1986) Use of ammonium chloride and sodium
bicarbonate in acute heat exposure of broilers. Poultry Science 65: 1659-1663.
CUNNINGHAM J.G. (1999) Tratado de Fisiologia Veterinária. Guanabara, Rio de Janeiro, 454 pp.
FISCHER da SILVA, A.V., FLEMMING, J.S. and FRANCO, S.G. (1994) Utilização de diferentes sais na
prevenção do estresse calórico de frangos de corte criados em clima quente. Revista Setor de Ciências
Agrárias 13: 287-92.
GORMAN, I. and BALNAVE, D. (1994) Effects of dietary mineral supplementation on the performance and
mineral retentions of broilers at high ambient temperatures. British Poultry Science 35: 563-572.
HULAN, H.W., SIMONS, P.C.M. and VAN SCHAGEN, P.J.W. (1987) Effect of altering the cation-anion
(Na+K-Cl) and calcium content of the diet on general performance and incidence of tibial dischondroplasia of
broiler chickens housed in batteries. Nutrition Reports International 33: 397-408.
HURWITZ, S., COHEN, I. and BAR, A. (1973) Sodium and chloride requirements of chick: relationship to
acid-base balance. Poultry Science 52: 903-909.
HURWITZ, S. (1981) Requirements and interaction of monovalant ions in nutrition. Proceedings of the Annual
Minerals Conference, pp. 27-35.
JOHNSON, R.J. and KARUNAJEEWA, H. (1985) The effects of dietary minerals and electrolytes on the
growth and physiology of the young chick. Journal of Nutrition 115: 1680-1690.
MACARI, M., FURLAN, R.L. and GONZALES, E. (1994) Fisiologia Aviária Aplicada A Frangos de Corte.
Funep, Jaboticabal, 246 pp.
MARCH, B.E. (1984) Sodium chloride supplementation of all plant protein broiler diets. Poultry Science 63:
703-705.
MELLIERE, A.L. and FORBES, R.M. (1966) Effect of altering the dietary cation-anion ratio on food
consumption and growth of young chicks. Journal of Nutrition 90: 310-314.
MONGIN, P. (1981) Recent advances in dietary cation-anion balance: applications. Proceedings of the Poultry
Nutrition Society, pp. 285-294.
NOSE, H., MACK, G.W., SHI, X. and NADEL, E.R. (1988) Role of osmolality and plasma volume during
dehydration in humans. Journal Applied Physiology 65: 325-331.
NATIONAL RESEARCH CONCIL (1994) Nutrient requirements of poultry. 9th Revision Edition. National
Academy Press, Washington.
PORTSMOUTH, J. (1984). Changes needed in nutrient input data relating to leg problems in poultry. Feedstuffs
56: 43-52.
RONDÓN, E.O.O., MURAKAMI, A.E., FURLAN, A.C. and GARCIA, J. (2000) Exigências nutricionais de
sódio e cloro e estimativa do melhor balanço eletrolítico da ração para frangos de corte na fase pré-inicial (1-
7 dias de idade). Revista Brasileira de Zootecnia 29: 1162-1166.
SALVADOR, D., ARIKI, J., BORGES, S.A., PEDROSO A.A. and MORAES, V.M.B. (1999) Suplementação
de bicarbonato de sódio na ração e na água de bebida de frangos de corte submetidos ao estresse calórico. ARS
Veterinária 15: 144-148.
SMITH, M.O. and TEETER, R.G. (1993) Carbon dioxide, ammonium chloride, potassium chloride, and
performance of heat distressed broilers. Journal of Applied Poultry Research 2: 61-66.
TEETER, R.G. and BELAY, T. (1996) Broiler management during acute heat stress. Animal Feed Science and
Technology 58: 127-142.
TEETER, R.G., SMITH, M.O., OWENS, F.N., et al. (1985) Chronic heat stress and respiratory alkalosis:
occurrence and treatment in broiler chicks. Poultry Science 64: 1060-1064.

80 World’s Poultry Science Journal, Vol. 63, March 2007

010028_Journal_4 01-02-2007 15:27 Pagina 81

Heat stress physiology in birds: S.A. Borges et al.

Table 1 Weight gain and feed conversion of broilers raised under two ambient temperature regimens and
receiving different dietary electrolyte balances (DEB; Na+K-Cl, mEq/kg).

Weight gain/bird (g) Feed conversion

DEB, mEq/kg 0 to 21 d 0 to 42 d 0 to 21 d 0 to 42 d

Thermoneutral room
40 737b 2,415b 1.521 1.770
140 743b 2,451ab 1.489 1.768
240 796a 2,583a 1.529 1.740
340 761ab 2,410 b 1.563 1.798
Pooled SEM 7.19 24.9 0.01 0.01
Heat stress room
40 732 2,378 1.507 1.758
140 731 2,334 1.511 1.774
240 750 2,422 1.527 1.775
340 746 2,390 1.546 1.784
Pooled SEM 5.76 27.6 0.01 0.01
abMeans within a column and treatment grouping lacking a common superscript differ (P>0.05).

Figure 1 Effects of different dietary electrolyte balances (Na+K-Cl, mEq/kg) on predicted weight gains
and feed conversion ratios of broiler chickens from 0 to 42 days of age.

World’s Poultry Science Journal, Vol. 63, March 2007 81

View publication stats