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OCR for page 441
Sodium Chloride
In ancient times, the distribution of the population centers was predi-
cated essentially by three factors the availability of salt (NaCl), water,
and food (Batterson and Brodie, 19721. Salt, therefore, was among the
first of the specific nutrients recognized to be essential for animal
nutrition and health. Salt is widely distributed in nature, where it occurs
not only in the sea and other saline waters but also in dry deposits as
rock salt. The concentration of salt in seawater averages 2.68 percent.
Whereas table salt is the most common use of salt, it finds application
in literally thousands of commercial processes that yield products con-
taining either sodium or chlorine (Stander, 1970~. In 1976 domestic salt
production approached 40 million metric tons, of which about 2 million
metric tons were used by the feed industry (U.Se Department of the
Interior, 19761. Interest in the biological effects of salt is high because
of its importance in nutrition, and much literature has been published
concerning salt deficiencies and excesses, its relationships with potas-
sium and other minerals, and its potential impacts on human health.
ESSENTIALITY
Sodium and chlorine have been found to be essential constituents of
diets fed to all animals and recommendations for salt supplementation
have been prepared and published by the National Academy of
Sciences/National Research Council (1974) and by the Salt Institute
441
OCR for page 442
442 MINERAL TOLERANCE OF DOMESTIC ANIMALS
(Anonymous, 19741. Sodium and chlorine, along with potassium, in
proper concentration and balance are indispensable for a number of
important physiologic processes. Sodium, as the chief cation of the
extracellular fluids, is the most important ion in maintenance of osmotic
pressure, body fluid balance, and hydration of the tissues. Heart action
and nerve impulse conduction and transmission are highly dependent
upon proper proportions of sodium and potassium. These cations are
also essential for the operation of certain enzyme systems and the
maintenance of blood pH. Chloride functions mainly to ensure the
proper fluid-electrolyte balance.
Signs of a combined sodium and fluorine deficiency in cattle, sheep,
swine, and horses include a salt craving evidenced by the animals'
licking soil, rocks, wood, and other objects. Eventually, there is a loss
of appetite and productive parameters are adversely affected. Cattle
and horses take on an unthrifty appearance, and their hair coats
roughen. In sheep, wool growth is greatly reduced. In poultry, ac-
companying the reduction ire productive performance, there appear
nervous signs and dehydration. The end point of long-term salt defi-
ciency for all animals is death. Generally, the dietary requirements for
sodium in animals of economic importance approximate 0.2 percent,
with the lowest value (0.10 percent) being required by growing beef
calves and the highest Value (0.35 percent) being required by horses.
Chlorine requirements are less known, but it is apparent that salt sup-
plementation to satisfy the sodium requirements will also satisfy the
chlorine requirements.
METABOLISM
Comprehensive reviews on the metabolism of sodium and chlorine
have been published by Forbes (1962), Cotlove and Hogben (1962), and
by Tracor-Jitco, Inc. (1974~. Meneely and Battarbee (1976) have re-
viewed the interrelationship between sodium and potassium. These
reviews testify to the voluminous literature on sodium and chlorine
metabolism, and thus the following information, adapted from Church
and Pond (1974), comprises only a brief summary of our knowledge.
Sodium and chloride ions are absorbed by animals principally from
the upper small intestine. Approximately 80 percent of the sodium and
chloride entering the gastrointestinal tract arises from internal
secretions such as saliva, gastric fluids, bile, and pancreatic juice.
Thus, large variations in salt intake have relatively small effects on the
total amount of sodium and chloride entering the gastrointestinal tract.
OCR for page 443
Sodium Chloride
443
The regulation of body concentrations of sodium and chloride ions, as
well as potassium ion, is narrowly controlled by, as yet, incompletely
defined mechanisms. It is known that increased intakes of each element
are accommodated by ready excretion in the kidneys. Plasma levels of
sodium are controlled, in part, by aldosterone, which functions to in-
crease sodium reabsorption from the kidney tubule. Other control is
exercised by the antidiuretic hormone of the posterior pituitary, which
is responsive to changes in osmotic pressure of the extracellular fluid.
Both hormones act to maintain a constant ratio of sodium to potassium
in the extracellular fluid. Chloride metabolism is controlled in relation
to sodium so that excess kidney excretion of sodium is accompanied by
chloride. Chloride excretion is also influenced by bicarbonate ion, with
a rise in plasma bicarbonate resulting in the excretion of a comparable
amount of chloride.
SOURCES
In general, foodstuffs do not contain sufficient sodium to provide for
optimum productive performance in livestock and poultry. Meyer et al.
(1950) found that most plant and plant products contain relatively small
amounts of sodium in comparison to animal products. In their survey,
oats contained only 0.008 percent sodium, whereas condensed fish
solubles contained 2.52 percent. With regard to chlorine, the broadest
range of values occurred between brewer's dried grains (0.03 percent)
and condensed fish solubles (4.63 percent). Sodium and chlorine con-
tents, respectively, were for corn (0.004 and 0.06 percent), clover hay
(0.14 and 0.15 percent), alfalfa hay (0.07 and 0.19 percent), soybean
meal (0.02 and 0.04 percent), and timothy hay (0.008 and 0.14 percent).
TOXICOSIS
LOW LEVELS
The toxicity of salt in animals has been reviewed by Tracor-Jitco, Inc.
(1974), the National Research Council (1974), and the Salt Institute
(Anonymous, 19741. The effects of salt administration are summarized
in Table 32.
The effects of relatively low levels (5 percent and less in the feed and
2 percent and less in the drinking water) have been measured in rumi-
nant species. Demott et al. (1968) provided lactating dairy cows with 0,
OCR for page 444
444 MINERAL TOLERANCE OF DOMESTIC ANIMALS
1, 2, and 4 percent salt in the grain (fed at a rate of 1 kg for each 2 kg
of fat-corrected milk) for a 2-week period. No adverse effects were
noted in the general health, milk production, or average body weight of
the treated cows. Jasteret al. (1978) studied the effects of saline drink-
ing water in dairy cows. In those studies, high-producing cows were
provided with drinking water with and without 2,500 ppm salt for a
28-day period. Although no changes were noted in feed intake and
digestibility and milk and blood concentrations of sodium, potassium,
calcium, magnesium, chloride, and phosphorus, the treated cows pro-
duced less milk and consumed greater quantities of water. In cattle,
Weeth et al. (1960) and Weeth and Haverland (1961) could produce a
toxicosis by administration of salt via the drinking water at levels which
ranged from 12 g per liter to 20 g per liter. The signs noted included a
severe anorexia, decreased water consumption, anhydremia, weight
loss, and collapse. The serum potassium and sodium concentrations
were elevated significantly, while both serum magnesium and urea
concentrations were depressed. In sheep, the effects of dietary salt
administration have been studied by Meyer and Weir (1954), Wilson
(1966), Jackson et al. (l971), and Kromann (Washington State Univer-
sity, personal communication, 1978~. Thus, levels of salt of 5 percent
and below were without adverse effects on weight gain, empty body
weight, carcass composition and energy gain in growing lambs, and
feed consumption in older sheep. Likewise, this level of salt did not
affect the productive performance of ewes maintained during growth,
fattening, breeding, gestation, and early lactation (253 days), nor did it
affect hematocrits, serum albumin and sodium concentrations, or milk
protein, sodium, and potassium concentrations.
In a series of studies by Peirce (1957, 1959, 1960, 1962, 1963,
1968a,b), it was found that sheep treated with 2.0 percent salt in the
drinking water lost weight continuously throughout the experimental
period and were found to be weak and listless. Diarrhea was occa-
sionally observed as was an increase in serum chloride concentration.
Certain of Peirce's studies ( 1959, 1960, 1962' 1963) involved treatments
with salt combined with magnesium chloride, sodium sulfate, calcium
chloride, and sodium carbonat~bicarbonate, respectively. At an in-
take of total salts of 1.3 percent or less, little adverse effects were noted
in health, feed intake, or body weight. Certain of the combinations
(including magnesium chloride) did cause occasional diarrhea and
reduced feed consumption, but none affected wool production. Next,
Peirce used penned (1968a) or grazing (1968b) ewes and their lambs to
assess the effects of synthetic drinking waters composed to resemble
underground waters found in Australia. The water compositions tested
OCR for page 445
Sodium Chloridle
445
were 1.3 percent (mostly salt), 1.0 percent (mostly salt), and
0.5 percent (half salt and half sodium bicarbonate). No adverse
effects were observed on the health, feed consumption, or
wool production of the penned animals, although in one trial a
poorer reproductive performance was evidenced by the ewes
receiving the 1.3 percent and the 0.5 percent salt treatments. In
the grazing animals, 1.3 percent salt decreased body weight gains
in lambs and the reproductive rate in ewes, caused diarrhea, and
increased mortality in one of the two experiments done. The other
treatments either decreased rate of gain and wool production (1.0 per-
cent) or the percentage of ewes that lambed (0.5 percent), but had
no other effects. Wilson (1966) also found that salt provided at 2 percent
in the drinking water decreased feed consumption. Finally, in sheep,
Potter and McIntosh (1974) found that the addition of 1.3 percent salt
to the drinking water of pregnant ewes caused neonatal mortality in the
resultant lambs. The ewes receiving the saline drinking water had
significantly higher levels of plasma potassium and chloride and sig-
nif~cantly lower levels of plasma calcium and magnesium.
The single study located for swine was performed by Done et al.
(1959) and demonstrated that dietary salt at 3 percent combined with
drinking water restriction for an 11-day period was without adverse
effect.
A group of workers (Quigley and White, 1932; Barlow et al., 1948;
Kare and Biely, 1948; Paver e! al., 1953; Mohanty and West, 1969) has
assessed the effects of relatively low levels of salt administration in
chickens. The reported signs of salt toxicosis included decreased rate
of weight gain, increased mortality, diarrhea, edema, increased heart
size, nervousness, and degenerative changes in kidney, liver, spleen,
adrenal, heart, lung, central nervous system, and the gastrointestinal
tract. Similarly, the signs noted in turkeys by Matterson et al. (1946),
Roberts (1957), Robblee and Clandinir~ (1961), Harper and Arscott
(1962), and Morrison et al. (1975) were increased mortality, edema, loss
of body weight, increased incidence of ascites, pendulous crop, diar-
rhea, and gross pathologic lesions of the heart, kidney, and lungs.
HIGH LEVELS
Literature is also available that demonstrates the effects of salt admin-
istration to livestock and poultry at relatively high levels (in excess of
5 percent via the diet and 2 percent via the drinking water).
Meyer et al. (1955) demonstrated that fattening steers could tolerate
dietary levels of salt of 9.33 percent with few adverse effects. Daily
OCR for page 446
446 MINERAL TOLERANCE OF DOMESTIC ANIMALS
gains and dressing percentage were similar for the treated and control
steers, although there was a small difference in carcass grade. In sheep,
Meyer and Weir (1954) found that a dietary concentration of 13.1 per-
cent salt caused increased weight loss during lactation and a decreased
number of lambs raised, as well as an increase in the blood and milk
chloride concentration.
In swine, Bohstedt and Grummer (1954) were able to induce salt
poisoning by offering diets containing 6~ percent salt with a restriction
in the availability of drinking water. Several days posttreatment, the
signs of salt poisoning included nervousness, staggering, weakness,
and paralysis. Blindness was observed in one pig with liver changes
appearing as the only gross anatomical change. Done et at. (1959) and
Todd et al. (1964) also studied salt toxicosis in swine precipitated by
high dietary salt and restricted water intake. Signs noted were muscular
tremors, incoordination, convulsions, prostration, coma, and the typi-
cal lesions of meningoencephalitis.
In chickens, Blaxland (1946) estimated the lethal dose of salt ad-
ministered via the crop at about 4 g per kilogram of body weight.
Otherwise the toxic signs of salt poisoning in both chickens and turkeys
are as they were discussed under Low Levels.
In other poultry, Scott et al. (1960) have performed toxicity studies
in pheasant and quail. Dietary salt at 7.5 percent in both of these species
causes decreased rate of weight gain and increased mortality.
With regard to salt toxicosis in other animal species, the table of
acute effects prepared by Tracor-litco, Inc. (1974), is particularly
pertinent and is herein adapted as Table 33.
FACTORS INFLUENCING TOXICITY
The major factor that influences salt toxicosis in animals is the avail-
ability of drinking water. In the presence of an adequate supply, ani-
mals can tolerate relatively large quantities of dietary salt.
TISSUE LEVELS
Several studies are available concerning the tissue levels of sodium and
chloride resulting from excess dietary salt in food-producing animals.
In cattle, drinking water concentrations of 1.0 and 1.2 percent salt
resulted in serum sodium concentrations of 345 and 356 mg/dl, respec-
tively (Weeth and Haverland, 19611. In swine under conditions of water
restriction, Todd et al. (1964) reported the following values for various
OCR for page 447
Sodium Chloride
447
tissues (sodium and chloride, respectively in parts per million fresh
weight): cerebrum, 1,886 and 2,769; cerebellum, 1,817 and 2,769; liver,
1,748 and 2,840; kidney, 2,231 and 3,728; heart, 1,265 and 3,195; and
spleen, 1,795 and 2,910. Finally, in chickens, Barlow et al. (1948) have
provided these values for chloride derived from chickens fed 10 percent
dietary salt (in parts per million fresh weight): tendon, 3,596; lung,
3,010; liver, 1,924; leg muscle, 983; and kidney, 2,098.
MAXIMUM TOLERABLE LEVELS
Maximum tolerable levels of dietary salt in animals were established as
follows: a level of 4 percent was set for lactating cows, since this level
was the maximum level tested by Demott et al. (19681; a level of 9.0
percent was set for other cattle and sheep based on the studies of Meyer
et al. (1955) and Meyer and Weir (1954), respectively; a level of 8
percent was set for swine due to a lack of documentation of toxicosis
in swine when given adequate supplies of fresh dunking water; a level
of 2 percent was set for poultry based on the studies of Barlow et al.
(1948) in chickens and Matterson et al. (1946) in turkeys; the level of 3
percent for horses and rabbits was obtained by extrapolation.
SUMMARY
Salt is widely distributed in nature, and its components have been
found to be indispensable nutrients for all animals. Sodium and chlorine
function in the body to control osmotic pressure, fluid balance, heart
action, and nerve impulse conduction and transmission. Salt toxicoses
are characterized by increases in water consumption, anorexia, weight
loss, edema, nervousness, paralysis, and a variety of signs dependent
upon animal species. In many cases, provision of fresh water is effec-
tive in reducing the severity of the signs, but there is no doubt that
reducing the dietary salt load (in poultry) is also required to mitigate the
toxicity..
OCR for page 448
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456 MINERAL TOLERANCE OF DOMESTIC ANIMALS
REFERENCES
Amberg, S., and H. F. Helmholx. 1915. The fatal dose of various substances on intra-
venous injection in the guinea pig. J. Pharmacol. Exp. Ther. 6:595.
Anonymous. 1974. Salt for Livestock, Poultry and Other Animals. Salt Institute, Alex-
andria, Va.
Barlow, J. S., S. J. Slinger, and R. P. Zimmer. 1948. The reaction of growing chicks to
diets varying in sodium chloride content. Poult. Sci. 27:542.
Batterson, M., and W. W. Brodie. 1972. Salt the Mysterious Necessity. The Dow
Chemical Company, Midland, Michigan.
Behrens, B. 1924. Studies on the mechanism of sodium chloride poisoning. I. Significance
of osmotic processes. Arch. Exp. Pathol. Pharmacol. 108:39.
Blaxland, J. D. 1946. The toxicity of sodium chloride for fowls. Vet. J. 102:152.
Bohstedt, G., and R. H. Grummer. 1954. Salt poisoning of pigs. J. Anim. Sci. 13:933.
Boyd, E. M., and M. N. Shanas. 1963. The acute oral toxicity of sodium chloride in albino
rats. Arch. Int. Pharmacodyn. 144:86.
Church, D. C., and W. G. Pond. 1974. Basic Animal Nutrition and Feeding. D. C.
Church. Corvallis, Oregon.
Cotlove, E., and C. A. M. Hogben. 1962. Chloride, pp. 109-157. In C. L. Comar and F.
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Representative terms from entire chapter:
drinking water
