NationalLibraryOfScotland/medical-history-of-british-india huggingface dataset:
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Phosphorus deficiency.—Naturally occurring deficiencies of mineral elements
have been shown to affect fertility. Of these, possibly the most important is
deficiency of phosphorus since it is of widespread occurrence in natural pastures
throughout the world. It has been shown by Theiler and his associates in South
Africa, by Henry in Australia, and by Eckles, Becker and Palmer and Hart in the
United States, that deficiency of phosphorus results in abortions and sterility.
In observations on 200 native scrub cows, half of which received a supplement of
bone meal, Theiler, Green and Du Toit (34) found that 80 per cent. of the bone-
meal lot calved normally, while only 51 per cent. of the controls calved. Du Toit
and Bisschop (35) report an experiment continued for three years with 109 bone-
meal-fed cows and 20 controls in which the bone-meal-fed group produced 87.3
per cent. of the possible number of calves, while the control group produced only
56.5 per cent. of the possible number. Eckles, Becker and Palmer (36) also found
that feeding a phosphate supplement improved fertility. They attributed the low
fertility associated with phosphorus deficiency to infrequency of ovulation. Hart
(37) finds that, in certain range areas in the States, where the calf crop is low,
the pasture, although quantitatively sufficient, is deficient in phosphorus and pro-
tein and the calcium/phosphorus ratio is high. On such pastures cows become
emaciated and œstrus does not occur. Laboratory experiments on rats, designed
by Hart to throw light on the causation of this sterility, have shown that under-
nutrition stops the normal cycle of ovarian changes, and that a diet low in pro-
protein and phosphorus, or with a high calcium/phosphorus ratio, interrupts the
cycle or greatly lengthens the interval between ovulations. Hart describes
trials in supplementary feeding in which a supplement of cotton-seed cake, rich
in protein and minerals, gave excellent results, 50 per cent. of the cows being
re-bred within a month after calving.
Calcium deficiency.—It is not generally considered that deficiency of calcium
occurs to any great extent under natural conditions. It is, however, known to
occur in some areas, for example, the Falkland Islands and hill pastures in Scot-
land (38). In most cases observations on such areas are complicated by the co-
existence of general under-nutrition. We have, therefore, no data regarding the
effect of calcium deficiency on fertility under natural conditions.
In experiments in which dairy cows were fed greatly restricted rations, con-
fined practically to a single fodder-plant oat, wheat or maize, Hart (39, 40, 41, 42)
and his co-workers showed that addition of calcium was essential to secure repro-
duction. The results were further improved by addition of butter or cod-liver
oil and salt. Hart also records that cows fed on hay and straw grown on acid
soils and containing less than the normal percentage of calcium were unable to
produce normal healthy calves. In an experiment using a ration of low calcium
NUTRITION IN RELATION TO REPRODUCTION
203
content Meigs (43) found that cows were very difficult to get in calf, while con-
trols with a supplement of calcium carbonate gave much better results. Kennedy-
found that animals on a diet deficient in calcium were sterile, although ovula-
tion occurred. The uterus was pale, flaccid and in a state of inactivity. Sim-
monds (44) found, in extensive experiments on rats, that a minimum percentage
of calcium in the ration was necessary for successful reproduction, and that in-
crease above the minimum gave improved reproduction with better condition of
the parents. An interesting observation by Küst is quoted by Frei (45), namely,
that rickets and nymphomania occur together in the same herds. Davidson (46)
has shown that protracted deficiency of calcium in the ration of breeding sows
leads to a very considerable increase in the number of pigs born dead. The effect
is cumulative, becoming more pronounced in the second and third generations.
A salt mixture representing the mineral elements of blood meal, had a marked
effect on the occurrence of œstrus after weaning. Withholding this mixture
delayed the appearance of œstrus. From the composition of the ration, it
is inferred that the effect was due either to potassium or to iron.
Other observations in which the chief deficiencies concerned were probably of
calcium or phosphorus or both, are the following. Rettger, White and Chapman
(47), in an experiment designed to produce contagious abortion in young heifers
by feeding suspensions of cultures of Br. abortus, fed their experimental animals
first on pasteurized milk and later on a ration of corn meal, groundnuts, linseed
oil meal, bran and sodium chloride, with green food only occasionally. No conta-
gious abortion developed, but great difficulty was experienced in breeding, the
control group being even more affected than those fed the cultures. After three
months on a ' well-balanced ration' they came in heat, and were successfully bred.
Kossmag (48) observed that the occurrence of abortions was prevented when a
silica-rich, base-poor diet was replaced by one poor in silica and including sodium
chloride and calcium chloride.
Cotton and Buck (15) state that the abortion rate in a herd numbering over
200, and negative to contagious abortion, was from 5 to 13 per cent. They be-
lieve that some dietetic deficiency must be the cause, although the ration fed
was, in their opinion, adequate. McGilvray (49) also holds the view that mineral
deficiency lowers resistance to abortion disease and increases sterility. This
would be particularly liable to occur in herds stall-fed on restricted rations.
Graves (50) found that foxes reared in captivity failed to breed until the meat in
their ration was reduced and green vegetables were supplied.
Iodine deficiency.—In areas of endemic goitre throughout the world, breeding
difficulties in domestic animals tend to occur (51). It was first shown by Ennis
G 2
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Smith (88) that premature birth and imperfect development of the young in pigs
could be prevented by administration of iodine to the sows. Stiner (89) and
Schropp (90) have both found that administration of iodine, either as an inorganic
salt or in pasture manured with iodine, had a beneficial effect on fertility, the
number of services required being reduced. Muller-Lenhartz and v-Wendt (91)
found similar results to follow the feeding of a mineral mixture containing cal-
cium, phosphorus, iron and iodine.
It has also been suggested that conditions of relative iodine deficiency might
arise in areas where deficiency of iodine is not thought to occur, under conditions
of intensive feeding. The results of a four years' experiment at the Rowett In-
stitute suggest that in stall-fed cows there may be a relative deficiency of iodine
adversely affecting reproduction (communicated).
Fluorine excess.—Experiments are accumulating to show that an excess of
fluorine in the diet may have a detrimental effect on fertility. Since excess of
fluorine occurs under natural conditions in at least two known areas (Morocco and
some districts in U. S. A.), the experimental findings may be of some practical
significance. In Morocco, excess of fluorine in the drinking water gives rise to
pathological conditions in sheep (52, 53, 54) and in the U. S. A. a similar excess is
held responsible for abnormal dental enamel in man (55).
McCollum and his co-workers (56) found that feeding fluorine as sodium
fluoride at a level of 0.05 per cent. of the ration had no effect on fertility.
Similarly, Schulz and Lamb (57) found no impairment of reproduction from feed-
ing 0.05—0.1 per cent. of the ration as sodium fluoride, but Bergara (58), feeding
5 mg. sodium fluoride per rat per day, failed to get reproduction. Del Castillo
(59) and Chaneles (60), also found injurious effects. Velu (61), on the basis of
experiments in which he has shown that continued administration of fluorine
to rats causes suppression of the cestrous cycle, suggests that similar effects
may be produced in domestic animals from long-continued consumption of
fluorine-containing water.
Protein deficiency and excess.—In natural grazings a deficiency of phosphorus
is usually correlated with a low nitrogen content of the pasture, especially in the
mature herbage and under drought conditions. Hart (37) from his observations
on range pastures, lays stress on the dual deficiency. In many cases it would be
impossible at present to judge what part of the effect of such grazing in reducing
fertility is due to under-nutrition and what part to deficiency of phosphorus or
protein.
Numerous experiments on small animals have been made which show the
great importance of both the amount and the quality of the protein supplied for
NUTRITION IN RELATION TO REPRODUCTION
205
breeding efficiency. In a series of experiments on rats, Slonaker and Card (62,
63, 64) showed that the age at puberty, the length of the reproductive span and
the size of litters were all affected by the quality of the protein fed. Rats
receiving animal protein matured earlier than those receiving only vegetable
protein; the intervals between litters were about two months in the former
and varied up to four months in the latter. The period of reproductive
activity on animal protein was seventeen months ; on vegetable protein only two
to seven months. Sterility was high on vegetarian diet; the number of young
in litters was greatly reduced and the effect was cumulative, the vegetarian rats
being completely sterile in the third generation. Simmonds (44) showed, that a
diet in which only vegetable protein was supplied greatly reduced fertility, while a
high proportion of animal protein increased it. Wan and Wu (65) also found a
significant difference in litter size in favour of omnivorous, as compared with
vegetarian, rats. Later, Slonaker (66) showed that there is an optimum protein
level for reproduction at 1. 4 per cent. of the ration (animal protein), higher or
lower levels giving diminished litter size and increased sterility. This level was also
best for length of reproductive span. A large excess of animal protein over the
optimum level causes pathological changes in the ovary and promotes sterility (67).
The difference in value between animal and vegetable protein for reproduction
may possibly be due to deficiency of essential amino-acids in the latter. Courrier
and Raynaud (68) have shown that rats fed on a diet deficient in lysine showed no
sexual cycle. Males on the same diet had relatively poorly developed testicles.
In poultry feeding it is well recognized that sufficiency of calcium and phospho-
rus and an adequate supply of good quality protein are essential for egg production.
Optimum quantities and ratios are being determined by experiments at all the
main centres of research (69, 70, 71). According to the most recent results,
plant protein, adequately supplemented with calcium and sodium chloride, may give
as good results as animal protein. There are, however, no data to show whether
this is true for mammals.
Vitamin deficiency.—The vitamin most clearly connected with reproduction is
vitamin E (72). This vitamin is fat-soluble, and occurs in nature, chiefly in green
vegetables, notably lettuce, and in the embryos of certain cereals, especially wheat.
It is found in milk and egg-yolk, and is present in muscle, fat, pituitary, placenta,
and to a less extent in intestine and liver. It does not occur in plant oils, and is
apparently absent from cod-liver oil, kidney, brain and testis. A diet free from
vitamin E does not prevent maturation or shedding of ova in rats, and implanta-
tion is normal. A first litter may even be born, but as depletion of reserves pro-
ceeds, a characteristic condition with death and absorption of the foetus results.
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In males, spermatozoa become non-motile, then disappear, while germinal epitheli
um degenerates and testis weight is reduced.
More recently, it has been shown that a deficiency of vitamin A also causes
degeneration of the germinal epithelium, and complete sterility (73, 74, 75). This
effect is more rapidly produced than that of vitamin E deficiency.
From time to time it has been suggested that other vitamins were necessary for
reproduction. It is doubtful, however, whether deficiency of vitamin B has any
specific effect on fertility. The present position (76) appears to be that the effect
on fertility of deficiency of vitamin B is due, not to lack of the vitamin itself, but
to the inanition characteristic of the condition. Vitamin D appears, at least under
certain conditions, to be necessary for normal reproduction in birds, but this is
again probably only a secondary effect associated with the effect of vitamin D on
mineral metabolism and the high mineral requirements for growth of the egg.
It is uncertain whether deficiency of vitamins affecting fertility occurs under
natural conditions. It has been suggested by Velu (77) that deficiency of vitamin
E may be the cause of a condition in sheep in Morocco in which pregnancy occurs,
followed by death and absorption of the foetus. The point does not appear to
have been experimentally investigated. It has also been thought that vitamin E
might be deficient in the rations of farm animals, particularly those stall-fed on
poor rations. Experiments have therefore been made by Graves and Miller (78),
and by Rogers (79) in the United States, and at the Department of Animal Gene
tics in Edinburgh (communicated) to test whether the feeding of sprouted oats
would influence sterility in cases not attributable to any definite pathological condi
tion. The results, although suggestive, are inconclusive. (There appears to be
some doubt as to whether sprouted oats contain any considerable amount of
vitamin E.) On the other hand, Vogt-Mφller and Bay (80) claim that a single intra
muscular injection of an ether extract of wheat germ cured sterility in cows kept
under poor conditions on a restricted feed.
Glands of internal secretion. At least two glands of internal secretion have
definite correlations with, and effects on, the ovary ( 81, 82 ). These are the thyr
oid and the anterior lobe of the pituitary. Either a deficiency or an excess of thy
roid secretion will interfere with ovarian function, causing cessation of ovulation
and sterility. The anterior lobe of the pituitary produces two hormones; one
appears to be necessary for maturation changes in the ovary, the puberty hormone,
and the other promotes luteinising changes. Injection of the former into immature
animals causes early puberty; of the latter, excessive production of lutein with
inhibition of follicular development and sterility. It has been found that extracts
of anterior pituitary may replace vitamin E in a deficient diet, and that the anter.
NUTRITION IN RELATION TO REPRODUCTION
207
ior pituitary ceases to be active in animals fed on a vitamin E deficient diet. Some
correlation, therefore, appears to exist between vitamin E and the hormones of the
anterior pituitary. It is also known that under-activity of the thyroid (simple
goitre, cretinism) is, in some way, correlated with iodine metabolism, and that
over-activity may, to some extent at least, be controlled and modified by diet (83).
Sterility due to disturbance of the functions of these glands may, therefore, ulti-
mately be due to lack of dietary balance of a kind not yet fully understood.
Interesting observations have been made on the effect of diet on the production
of spermatozoa. Polowzow and Naguzew (84) have shown that production in the
stallion is definitely related to quality of food, and that eggs and bran have a
stimulating effect. Parchutin (85) records exactly similar observations.
Nordby and Bollen (86) describe a case of temporary sterility in a boar fed on
Canadian field pea screenings. A change of diet to barley, oats, tankage, oil meal,
alfalfa and sodium chloride resulted in complete restoration of fertility in six weeks.
At the time of the change in diet the pH of the semen was 7.61, and motility of the
spermatozoa was ' very unsatisfactory '. On the altered diet the pH fell to 7.35
and motility improved along with the reduction of pH.
This last observation suggests a possibility that the supposed effect of infect-
ions of the female genital tract as due to ' spermatoxins' and ' spermalysins'
might, in reality, be the effect of a change in pH. It has been shown that such
an alteration, for instance, in regard to wound infections, may completely change
the nature or pathogenicity of the organisms present, and changes in the pH of
body fluids and secretions may easily be effected by diet. Hence the infections
themselves and the supposed toxic effect on spermatozoa might both be due to an
abnormal pH.
Conclusion. This brief review of the literature shows that under-nutrition or
malnutrition due to imbalance of the constituents of a diet, vitamins, proteins or
minerals, may seriously interfere with normal reproduction. The influence of such
faulty diets has been clearly demonstrated in experimental work with laboratory
animals. It is known that similar defective diets occur, with similar results under
extensive conditions of animal husbandry, and they may be reproduced under
intensive conditions on artificial rations. No data exist to indicate to what extent
such factors do actually affect fertility in practical, intensive or semi-intensive,
farming, but they may be of considerable economic importance.
In addition to this direct influence of diet on fertility, there is the other, possib-
ly even more interesting and important, indirect effect though its modifying action
on susceptibility to infections. It is known that malnutrition practically deter-
mines susceptibility to certain infections, and there is evidence that it may be an
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important predisposing factor to infection with Br. abortus and other organisms
which interfere with reproduction. It is perhaps unlikely that an organism of the
apparently high virulence of Br. abortus should become innocuous under the most
perfect dietary conditions, but the relative importance, as regards subsequent
sterility, of malnutrition of the host and virulence of this or other infecting orga-
nism has not yet been established. There is obviously urgent need for further
extensive investigations on the whole subject of nutrition in relation to reproduc-
tion and the various infections which interfere with conception and the normal
development of the foetus.
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(1) Quoted in Vet. Rec., 1931, 11, 1050.
(2) Garrad, G. H. Vet. Rec., 1931,11, 525-527
(3) Private communication.
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209
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(37) Hart, G. H. ' The Scientific Principle of Nutrition with particular reference to Percentage.
Calf Crop in Range Herds '. 11th Int. Vet. Congr., London, 1930.
(38) Orr, J. B. ' Minerals in Pastures,' H. K. Lewis, London , 1929.
(39) Hart, McCollum, Steenbock and Humphrey. Res. Bull. No. 17, Agric. Exp. Stat.,
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(40) Hart, McCollum, Steenbock and Humphrey. J. Agri. Res., 1917, 10, 175-198.
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(43) Meigs, E. B. Proc. World's Dairy Cong., 1924, p. 1046.
(44) Simmonds, N. Amer. J. Hyg., 1924, 4, Supplement.
(45) Küst, Frei, W. Schw. Arch. f. Tierheilk., 1931, 73, No. 11.
(46) Davidson, H. R. J. Agric. Sci., 1930, 20, 233-264.
(47) Rettger, L. F., White, G. C. and Chapman, L. M. Storrs Agric., Exp. Stat. Rep., 1920-23.
Pub. Doc. No. 34, pp. 59-88.
(48) Kossmag, M. Tierärztl. Rundschau, 1931, 37, 386-387.
(49) McGilvray, C. D. Ontario Dept. Agric. Bull. No. 304, 1925.
(50) Graves, E. F. J. Amer. Vet. Med. Assoc., 1930, 77, 354-361.
(51) Orr, J. B., and Leitch, I. ' Iodine in Nutrition '. M. R. C. Spec. Rep. Series, No. 126, 1929.
(52) Velu, H., and Balozet, L. Bull. Acad. Vet. de France, 1931, 4, 373-376.
(53) Velu, H. C. R. Soc. Biol., 1931, 108, 635-637 .
(54) Velu, H. Ibid., 750-752.
(55) Churchill, H. V. J. Dent. Res., 1932, 12, 141-148.
(56) McCollum, E. V., Simmonds, N., Becker, J. E., and Bunting. R. W. J. Biol, Chem., 1925,
63, 553.
(57) Schulz, J. A., and Lamb, A. R. Science, 1925, 61, 93.
(58) Bergara, C. C. R. Soc. Biol., 1927, 97, 600-602.
(59) Del. Castillo, E. B. Ibid., 1928, 99, 1405.
(60) Chaneles, J. Rev. Soc. Argent. Biol., 1929, 5, 317-396.
(61) Velu, H. Note to Soc. Pathol. Exotique, Dec., 1931.
(62) Slonaker and Card. Amer J. Physiol., 1923, 64, 35-43.
(63) Slonaker and Card. Ibid., 167-180.