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The Physiology of Growth by S Postgrad Med J: first published as 10.1136/pgmj.26.298.417 on 1 August 1950. Downloaded from 417 THE PHYSIOLOGY OF GROWTH BY S. LEONARD SIMPSON, M.A., M.D., F.R.C.P. Consultant Endocrinologist, St. Mary's Hospital, etc.; Formerly Research Wor7Aer, Lister Institute Although clinical and experimental evidence postulated an antagonism between growth and had previously indicated that skeletal growth sex-stimulating hormones. Further observations depended upon the pituitary gland, it was not suggested that corpora lutea were formed before until I92I that Evans and Long, at the University ovulation occurred, and that his emulsions con- of California, demonstrated that a saline emulsion tained gonadotrophic luteinizing hormone. Al- made from the anterior pituitary lobe of fresh though Evans' original giant rats were apparently glands of cattle contained a growth hormone. The adipose, later work (Lee and Schaffer, 1934), hypophyses were thrown into 40 per cent. ethyl showed that there was a retention of nitrogen and alcohol, agitated with a glass rod, transferred to a relative excess of body protein. This was also two changes of sterile normal saline, and triturated true of the excess of tissue deposited in and with force and speed with ocean sand, diluted with around the abdomen. Evans stated in a recent saline, and decanted. The layers of sand, cell discussion in London that rats treated with pure fragments and opaque pink fluid permitted the growth hormone, unlike those in the initial latter to be decanted. This emulsion, when experiments with crude extracts, were not injected daily in doses of I to i ml. intraperiton- adipose. eally into young rats, accelerated their skeletal Another fundamental step was the development growth rate, compared with controls, and con- by P. E. Smith (I926) of a technique of completecopyright. tinued this growth for 20o days, or more, after hypophysectomy on the rat, without interfering growth normally ceases. Later, improved chemical with the hypothalamus. There resulted ' an methods permitted the preparations of alkaline almost complete growth stasis and a rapid regres- extracts which contained the growth hormone sion in the size of the adrenals (involving chiefly in purer form. It is interesting and important to the cortex), the thyroid and the sex apparatus ' note that Evans (1923-4) recorded that success, (Fig. i). At the Institute of Biology at Berkeley in mammals, with parenteral injections of beef University, California, rats are completely hypo- hypophysis extracts was only obtained ' after physectomized without complications in about failure in a long series of massive oral administra- two minutes for each operation, and there is http://pmj.bmj.com/ tions '. There is no doubt that in mammals the always a plentiful supply of hypophysectomized groWth hormone is ineffective when given by rats for experimental work. Such rats may be mouth, although this well-established experi- used for the assay of growth hormone, either by mental fact is still ignored by some clinicians. The weight increase, or by tail growth (Freud et al., rat is a particularly suitable animal for testing 1935, I939). Smith and MacDowell (I930) growth hormone because its epiphyses remain studied a strain of black silver dwarf mice, brought open until late in life. from England by Professor L. C. Dunn in I928, The effect of extracts containing pituitary in whom the characteristic of dwarfism was shown on October 1, 2021 by guest. Protected growth hormone has also been dramatically by Snell to be recessive. These mice cease to demonstrated in hypophysectomized rats and in grow after the end of the second week, but, congenitally dwarfed mice. Evans (1923) termed although there is some delay in sexual develop- his normal rats injected with pituitary emulsions ment, reproduction takes place. The essential ' hypophysis giants', and stated that they were physiological defect is absence of the growth 'twice as heavy as the largest individuals known hormone, and the essential histological defect in to us from our own and published records for this the pituitary gland of these animals is a complete animal species'. They were fat animals ; 'the absence of the eosinophil cells. Implantation of skeleton, most of the viscera especially the heart the pituitary glands of these dwarf mice into and lungs, liver, kidneys, and alimentary tract immature female mice of a normal strain produces showed a true overgrowth, but not so the repro- no acceleration of growth but accelerates sexual ductive tract, the uterus and oviduct remaining development. The injection of growth hormone infantile'. He also observed that the growth and will produce growth in these dwarf mice. Furtier maturation of ova were-impaired' or prevented, a'nd experimenta' evidence to the effect that ''the Postgrad Med J: first published as 10.1136/pgmj.26.298.417 on 1 August 1950. Downloaded from 4I8. POSTGRADUATE MEDICAL JOURNAL August I 950 eosinophil cells are the source of the growth Is there a Specific Growth Hormone ? hormone was provided by Smith who showed that Pituitary extracts containing growth hormones the peripheral part of the bovine gland contains were found by earlier observers to contain also eosinophil cells, and, when stripped from the adrenotrophic and thyrotrophic hormones as well central basophil portion, will yield growth hor- as prolactin. Apart from any direct effect on mone. Clinically, gigantism is often associated skeletal growth attributable to hormones of the with an eosinophil adenoma. adrenal cortex and thyroid and possibly to pro- It is also of interest to note that the thyroid and lactin, it is probable that these hormones have adrenal cortex of Dunn's dwarf mice were hypo- effects on metabolism which indirectly favour plastic, suggesting that the eosinophil cells are the growth, especially in hypophysectomized animals source of the thyrotrophic and adreno-cortico- in which these glands are hypoplastic or atrophic. trophic hormones. Riddle and Bates (I938) suggested therefore that certain pituitary extracts owe their action upon Measurement and Site of Action of Growth growth to a balanced combination of prolactin, Hormone thyrotrophic and adrenotrophic hormones rather Since the epiphyses of normal rats remain open than to a special growth hormone. Earlier experi- in adult life, earlier methods of assay of growth ments, however, indicated that the growth hor- hormone depended upon observations on the mone was a specific entity. Thus Evans (I924) re- body weight of normal rats after ' a plateau' in cords that Smith and Allen showed that beef their weight curve was established, indicating hypophysis is effective by mouth in producing cessation of growth. After the technique of growth in hypophysectomized tadpoles, but that it hypophysectomy was mastered, the effect on the does not produce metamorphosis or repair the weight of the hypophysectomized rat appeared to atrophied adrenal cortex (interrenal body) or the have a greater exactitude and specificity. Freud, thyroid. Smith, in 1930, demonstrated that whereas Levie and Kroon (I939) used the tail of the intramuscular transplantation of living pituitary hypophysectomized rat as a more exact measure- glands from adult rats into dwarfed hypophysec- ment of the effect of the growth hormone (Fig. 2). tomized rats restored the size and function of the copyright. They also made radiographic and histological thyroid, adrenals and gonads as well as producing studies of fundamental importance. Using an growth, injections of a saline suspension of fresh almost purified growth hormone, they demon- gland affected growth only. It would therefore strated its specific chondrotrophic effect. In the appear that although extracts of growth hormone normal growing rat, the epiphyseal zone shows as usually prepared contain adrenotrophic and cells arranged in columns, little cartilaginous thyrotrophic hormones which have some effect on matrix, vascular connective tissue and a fine growth, there is a specific growth hormone which network of cancellous bone. In the hypophysec- can be chemically separated. This is in keeping tomized rat (Fig. i), the regular columns of with the existence of clinical conditions in which http://pmj.bmj.com/ cartilage cells characteristic of growing cartilage the only defect appears to be that of growth are undeveloped and irregularly placed, the hormone, although other clinical conditions con- longitudinal arrangement is lost, mitosis is firm that the thyroid, the adrenals and the gonads minimal, the matrix has increased and the fine themselves have an important influence on growth. primary cancellous bone has disappeared. Growth We shall see later that further purification of the hormone immediately repairs this histological growth hormone led to more conclusive proof of picture, which reverts to normal, and shows active its specificity. growth of cartilage. There is no alteration in the on October 1, 2021 by guest. Protected development of bone tissue itself, and the growth The Thyroid and Growth hormone is seen to have a biologically typical point In the strain of congenitally dwarfed mice of attack at the proliferating cartilage. Freud, referred to above, thyroid extract alone produced Levie and Kroon (I934) found that the epiphyses an increase of growth, although not as much as of the tail of the rate soon become closed after pituitary growth hormone (Bates et al., 1935). hypophysectomy. Thus after four weeks a trans- The rate of growth of young normal mice is verse bony plate closes the epiphysis, from which a consistently accelerated by thyroxine injected number of massive bony processes project towards daily, but the period of this acceleration lasts for the diaphyseal marrow cavity. This can be en- only five weeks after which time the control mice tirely prevented by the early use of growth hor- overtake the'treated animals and the ultimate size mone after hypophysectomy, but it cannot be is the same. The skeleton of rats thyroidectomized reversed after it has *occurred. Smith (I930), at birth does not develop beyond the skeletal age of however, found replacement therapy renewed iS days, even with pituitary growth extract, unless growth three months after hypophysectomy.
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