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331. INFLUENCE OF THE ENDOCRINE SYSTEM ON GROWTH AND DEVELOPMENT R. J. GERRITS A. R. S., U. S. D. A. Introduction The field of endocrinology cannot be defined in a manner entirely acceptable t o all biologists or animal scientists. This is understandable as there are many points of view and many voids in our knowledge i n t h i s area. It is very hard t o give a precise definition for an endocrine gland because all cells possess some secretory capacity and therefore contribute t o the internal environment of the organism. For the most part the term "hormone" is probably applied too loosely and to a great variety of unrelated substances. Hormones can be identified as chemical agents which are synthesized by circumscribed parts of the body, generally specialized ductless glands, and are carried by the circulatory system to another part of the body where they evoke systemic adjustments by acting on rather specific target tissues or organs (Turner, 1966). regulate many processes such as growth, regeneration, reproduction, blood chemistry, metabolic rate, etc. Hormones act on the organs and tissues of the body by regulating the rate of specific metabolic reactions without contributing much at all t o the constituent cell. It can be said that ad- justments t o hormone levels require duration rather than speed, as opposed to the rapid coordinations of the body that are controlled by the nervous system. These biochemical adjustments are accomplished at the cellular level by virtue of their power t o augwnt or restrain special enzyme sys- tems. It is important that hormones be released at the right time and in the proper amounts of the normal organism if they are to accomplish their specific mission. Hormones are ineffective unless the target cells and tissues are capable of responding t o them. In general, hormones While it is neaxly impossible t o separate the neural and the hormonal components of regulatory processes, the complex system of endocrine glands in the vertebrates is quite clear. This system includes the pitui- tary, thyroid, parathyroid, adrends, gonads, pancreatic islets, and t h e hormone-producing part of the gastrointestinal tract. The material pre- sented in this paper will be limited to a very small segrnent of the endo- crine system and the effects on growth. More s p e c i f i c a l l y , I will attempt to discuss in this paper some of the important influences that are exerted by the hormone somatotrophin (STH growth hormone). selected material on the effect of testosterone (as an endogenous hormone) and of melengestrol acetate (a synthetic hormone) on growth. Also I will present some Hormones of the adenohypophysis Seven hormones are known t o have been released from the adeno- hypophysis, namely: (ACTH) ; thyrotrophin (TSH) ; prolactin (lactogenic hormone or luteotrophin) ; follicle-stimulating hormone (FSH) ; luteinizing hormone (LH or interstitial cell-stimulating hormone) ; and melanophore-stimulating hormone (MSH) . All somatotrophin ( STH or growth hormone) ; corticotrophin

Influence of the Endocrine System on Growth and Development

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331.

I N F L U E N C E OF T H E E N D O C R I N E S Y S T E M ON GROWTH A N D D E V E L O P M E N T

R . J . G E R R I T S A . R . S . , U. S . D. A .

Introduction

The f i e l d of endocrinology cannot be defined i n a m a n n e r e n t i r e l y acceptable t o all bio logis t s or animal s c i e n t i s t s . This i s understandable as there are many points of view and many voids i n our knowledge i n t h i s area. It i s very hard t o give a precise de f in i t i on for an endocrine gland because all c e l l s possess some secretory capacity and therefore contribute t o t h e in t e rna l environment of t h e organism. For t h e most pa r t t he term "hormone" i s probably applied t o o loosely and t o a great va r i e ty of unrelated substances. Hormones can be iden t i f i ed as chemical agents which are synthesized by circumscribed p a r t s of t h e body, generally specialized duc t less glands, and are car r ied by t h e c i rcu la tory system t o another par t of t he body where they evoke systemic adjustments by act ing on ra ther spec i f ic t a rge t t i s s u e s or organs (Turner, 1966). regulate many processes such as growth, regeneration, reproduction, blood chemistry, metabolic r a t e , e t c . Hormones act on the organs and t i s s u e s of t h e body by regulat ing t h e rate of spec i f ic metabolic react ions without contr ibut ing much at all t o t h e const i tuent ce l l . It can be said t h a t ad- justments t o hormone levels require duration r a the r than speed, as opposed t o t h e rapid coordinations of t h e body t h a t are control led by t h e nervous system. These biochemical adjustments are accomplished at t h e c e l l u l a r l e v e l by v i r t u e of t h e i r power t o augwnt or r e s t r a i n special enzyme sys- t e m s . It i s important t h a t hormones be released a t t h e r i g h t t i m e and i n t h e proper amounts of t h e normal organism i f they are t o accomplish t h e i r spec i f ic mission. Hormones are inef fec t ive unless the t a r g e t c e l l s and t i s s u e s are capable of responding t o them.

I n general, hormones

While it i s neaxly impossible t o separate t h e neural and t h e hormonal components of regulatory processes, t he complex system of endocrine glands i n t h e ver tebrates i s qui te c lear . This system includes the p i t u i - t a ry , thyroid, parathyroid, ad rends , gonads, pancreatic islets, and t h e hormone-producing pa r t of t h e gas t ro in t e s t ina l t r a c t . The material pre- sented i n t h i s paper w i l l be l imited t o a very small segrnent of t h e endo- c r ine system and t h e e f f e c t s on growth. More spec i f ica l ly , I w i l l attempt t o discuss i n t h i s paper some of t h e important influences t h a t are exerted by t h e hormone somatotrophin (STH growth hormone). selected material on t h e e f f e c t of tes tosterone (as an endogenous hormone) and of melengestrol acetate (a synthetic hormone) on growth.

Also I w i l l present some

Hormones of t h e adenohypophysis

Seven hormones are known t o have been released from the adeno- hypophysis, namely: (ACTH) ; thyrotrophin (TSH) ; prolac t in (lactogenic hormone o r luteotrophin) ; fo l l ic le -s t imula t ing hormone (FSH) ; lu te in iz ing hormone (LH or i n t e r s t i t i a l cel l -s t imulat ing hormone) ; and melanophore-stimulating hormone (MSH) . All

somatotrophin ( STH or growth hormone) ; cort icotrophin

332.

of these hormones are proteins or peptides, and three of them (FSH, LH, and TSH) contain carbohydrate i n addition t o amino acids.

Somatotrophin (STH)

It has been well es tabl ished t h a t STH i s secreted by the acidophil c e l l s of the pars d i s t a l i s and t h a t the pr incipal act ion of growth hormone i s on growth of bone and muscle. Growth hormones i so la ted from d i f f e ren t species exhib i t d i f f e ren t physiochemical propert ies . The molecular weight of STH hormone ranges from 21,500 t o 48,000 i n man, pig, beef, and sheep. The growth hormones with lower molecular weights are more acidic t h h n those with higher weights. Also, it i s important t o know t h a t body growth response of STH hormone i s d i f f e ren t among t he various ver tebrates and t h a t t he r a t responds t o somatotrophin from numerous mammalian species. The most common tes t employed i s the t i b i a t e s t . mone, t he increase i n width of t he proximal epiphyseal ca r t i l age of t he t i b i a of t h e hypophysectomized female rat or mouse i s determined. STH pro- duces continuous growth and widens the ca r t i l ages i n proportion t o t h e amounts given.

-

After administration of t h e hor-

Mechanisms of hormone act ion

There i s a considerable amount of knowledge about t he chemistry of hormones and t h e i r i n vivo and i n v i t r o e f f e c t s . However, su f f i c i en t re- search, espec ia l ly i n ver tebrates , has not been conducted t o disclose f u l l y how any hormone ac tua l ly performs at the molecular l eve l . Levine and Gold- s t e i n (1955) and Smith -- e t al. (1961) have demonstrated t h a t insu l in promotes the transfer of glucose i n t o t h e c e l l s of ce r t a in t i s sues , such as muscle and f a t . Three general points, each with varying modifications, have been proposed t o explain the mechanisms of hormone action, namely:

-- --

1. Hormones exert a d i r e c t e f f ec t on in t r ace l lu l a r enzyme systems.

2 . Hormones ac t t o control permeability re la t ionships at the c e l l surface or elsewhere and hence ind i r ec t ly control condition enzymic reac t ion.

3 . Hormones may produce t h e i r e f f e c t s d i r e c t l y by act ivat ing or suppressing par t icu lar genes.

I n our attempt t o understand the mechanisms of hormone act ion we must r e f e r t o work t h a t has been conducted with insec ts . Microscopic ex- amination of t he giant chromosomes of chironomous larvae has shown t h a t t he administration of ecdysone causes a puffing of ce r t a in genes. i s t h a t the act ivated genes (DNA) begin t o synthesize spec i f ic RNA which i s t ransfer red t o t h e cytoplasm where it a c t s a s template f o r t h e synthesis of a pa r t i cu la r protein, e.g., enzymes (Karlson, 1962). The work of Liao and Williams-Ashman (1962) suggests t h a t ce r t a in ver tebrate hormones may ac t i n a s i m i l a r manner. It w i l l be most in te res t ing t o see how f a r t h i s concept can be substantiated i n processes other than development i n m a n n a l s .

The inference

333.

Biological act ion

A t one t i m e it w a s assumed t h a t STH had only an e f f e c t on general body growth, p a r t i c u l a r l y t h e skeleton, and when e x t r a c t s r i c h i n STH pro- duced other ac t ion they were a t t r i b u t e d t o contaminating f a c t o r s . There i s now s u f f i c i e n t evidence t o show t h a t STH plays an important ro le i n the metabolism of proteins, f a t s , and carbohydrates and also serves as a synergist t o enhance t h e e f f e c t s of other hormones.

The e f f e c t of STH on pro te in metabolism

Amino acids not u t i l i z e d by animals are normally converted t o urea and eliminated through t h e ur ine . It has been ably demonstrated t h a t STH encourages t h e animal t o r e t a i n amino acids which are t h e e s s e n t i a l building blocks f o r pro te in and t h a t t h e body weight increase normally observed after treatment with STH i s an a c t u a l increase i n t i s s u e pro te in .

Knobil (1961) states t h a t one important aspect of STH act ion i s t o promote t h e t r a n s f e r of e x t r a c e l l u l a r amino acids across c e l l membranes, p a r t i c u l a r l y i n t o muscle c e l l s .

Milman and Russel l (1950), L i -- e t al. (1949), and Luck -- e t al. (1954) have demonstrated t h a t t h e administration of growth hormones lowers blood amino acid nitrogen. Swislocki and Szego (1965) also demonstrated t h i s e f f e c t i n hypophysectomized ad l i b fed rats. I n t h e la ter study blood-amino nitrogen (BAN) l e v e l s i n t h e growth hormone t r e a t e d rats w e r e s i g n i f i c a n t l y lower than t h e controls a t one and f i v e hours a f t e r treatment with STH.

I n a recent study Beach and Kostyo (1968) showed t h a t when ra ts were injected d a i l y in t raper i tonea l ly with bovine growth hormone f o r seven days t h e amount of DNA i n t h e muscles r e c t u s femoris, gastrocnemius, p e c t o r a l i s -' major and diaphragm w a s increased s igni f icant ly . When STH w a s administered f o r one day it had no e f f e c t on t h e DNA of these muscles. However, following seven days of t r e a t n m t , with t h e hormone, t h e amount of DNA i n each of t h e muscles w a s markedly elevated. Since there w a s a con- comitant increase i n muscle weight, t h e concentration of DNA i n terms of micrograms per mg. of w e t muscle w a s a l te red s igni f icant ly . These d a t a show t h a t growth hormone can increase t h e amount of DNA i n s k e l e t a l muscles of hypophysectomized ra ts . Thus, it seems reasonable t o assume t h a t t h e increase i n muscle DNA which occurs during t h e normal course of growth i n rats i s dependent i n some measure on p i t u i t a r y growth hormone.

Ef fec t of STH on l iDids metabolism

The experiments of Welt and Wilhelmi (1950) suggested t h a t growth hormone administration leads t o a reduction of l i p i d synthesis. Their i n i t i a l s tud ies have subsequently been supported by several workers.

Hypophysectomy r e t a r d s t h e mobilization of depot f a t and tends t o ameliorate ke tos i s i n t h e d iabe t ic subject . STH encourages t h e movement of unes te r i f ied f a t t y acids from f a t reserves, consequently decreasing body f a t and increasing t h e l i p i d content of t h e blood plasma and l i v e . I n support of t h i s , Swislocki and Szego (1965)showed t h a t elevated l e v e l s of plasma nonester i f ied f a t t y acids occurred a f t e r t h e administration of growth hor- mones t o hypophysectomized ra ts .

334.

E f fec t of STH on carbohydrate metabolism

The following general statements may be m a l e regarding t h e ac t ion of STH on carbohydrate metabolism when administered t o m a m m a l s (Turner, 1966) :

1. The hormone tends t o produce hyperglycemia, thils aggravating d i abe te s .

2 . It i n h i b i t s t he ac t ion of i n s u l i n and i n s u l i n e f f e c t .

3. It increases muscle glycogen when given t o hypophysec- tomized sub jec t s .

4. It produces a permanent d i abe te s mel l i t i s i n c e r t a i n spec ies when given over prolonged periods.

The la t te r e f f e c t probably r e s u l t s from t h e eventual des t ruc t ion of t h e 8 - c e l l s of t h e pancrea t ic i s l e t s which sec re t e i n s u l i n . It has been shown t h a t ra t s rece iv ing excessive carbohydrates by tube feeding develop tem- porary d i abe te s when given STH.

Studies on STH i n meat animals

Baird -- e t d. (1952) assayed t h e hypophysis of two s t r a i n s of p igs se lec ted f o r e igh t generations on t h e b a s i s of rap id and slow rate of ga in . The two s t r a i n s reached u l t imate s i z e a t s i g n i f i c a n t l y d i f f e r e n t ages. When t h e growth hormone content of t h e p i t u i t a r y glands of t h e two s t r a i n s w a s determined, it w a s found t h a t at d l ages at which cornpaxisons w e r e made t h e glands of t h e f a s t e r growing s t r a i n contained s i g n i f i c a n t l y more somatotro- phin per u n i t of gland t i s s u e than d id t h e glands of t h e slower growing s t r a i n . This study showed t h a t t h e amount of somatotrophin per u n i t of p i t u i t a r y t i s s u e w a s t h e same i n t h e youngest and t h e o ldes t animal and t h a t it d id not v w y s i g n i f i c a n t l y throughout t h e growth period.

Armstrong and Hansel (1956) also demonstrated t h a t t h e r o l e of STH f o r con t ro l of growth w a s t r u e i n Hols te in h e i f e r s . concentration of somatotrophin per un i t of p i t u i t a r y t i s s u e w a s t h e same i n young growing ca lves as it i s i n a n i m a l s which have reached t h e growth p la teau . t h a t r ap id growth occurs as long as t h e r a t i o of t h e c i r c u l a t i n g somatotro- phin per u n i t of body t i s s u e remains s u f f i c i e n t l y high t o st imulate p ro te in synthes is and t o cause bone and muscle growth. The t h r e e l a t te r s tud ie s c i t e d a l l support t h e " d i l u t i o n theory," t h a t is , when t h e s i ze of t h e animal increases , t h e amount of hormone ava i l ab le per uni t of weight i s "diluted." Thus, o lder , heavier animals r e t a i n t h e i r a b i l i t y t o r e p a i r t i s s u e but l o s e t h e i r capacity t o grow a t a r ap id r a t e .

They showed t h a t t h e

These s tud ie s , along with t h e study of Simpson -- e t a l . (1950), show

Related s tud ie s on Duroc swine se lec ted f o r high and low back.fat

H e t z e r and P e t e r s (1965) and Hetzer and Harvey (1967) reported on t h e e f f e c t of s e l e c t i o n on Duroc s w i n e f o r both high and low backfat t h i ck - ness. A non-selected con t ro l l i n e derived from t h e same source as t h e se lec ted l i n e w a s maintained. After 1 0 generations of s e l ec t ion , t h e high-

335.

and low-fat Duroc l i n e s d i f fe red i n backfat by 2.6 cm. or 68% of t h e i n i t i a l mean. Selection basis on backfat depth w a s e f f ec t ive i n both the upward and downward d i rec t ion . y ie lds of lean meat and less f a t than the l i n e selected f o r m a x i m u m depth of backfat .

Pigs from the Duroc low backfat l i n e produced higher

Davey and Kincaid (1964) conducted i n i v i d u d feeding trials with pigs from high- and low-fat Duroc l i n e s . Carcasses were separated i n t o lean, fa t , bone, and skin at ages varying from 90 t o 400 days. The abso- l u t e weight of f a t and lean up t o 220 days of age showed l i n e a r t rends with age. Differences i n carcass composition for pigs fed normal d i e t s as estimated from regressions on age were: high Duroc, 23.2, 0; low Duroc, 17.3, 4.1; kg. of f a t and lean, respectively. The da ta show t h a t t h e high- fa t l i n e began t o l a y down more f a t than lean a t 18 weeks of age. However, t he low-fat l i n e did not begin t o l a y down more f a t than lean u n t i l 10 weeks l a t e r .

Gerrits -- e t al. (1964) found t h a t t h e an ter ior p i t u i t a r y weight of animals from the l i n e s of Duroc swine selected f o r high and low backfat d i f fe red s igni f icant ly from those of t h e control . This study a l so revealed t h a t t he heart , pancreas, and length of s m a l l i n t e s t ines from the selected l i n e s d i f fe red s ign i f i can t ly from the control . Thyroid weights between the l i nes were not s ign i f icant . cholesterol l e v e l s i n the Duroc selected f o r high backfat were s ign i f icant ly correlated with depth of backfat.

Johnson and Gerrits (1965) reported t h a t

Growth hormone study i n Duroc swine selected f o r h i& and low backfat

I n considering the f indings i n the previously c i t ed experiments and because of t he spec i f ic e f f e c t s known t o be mediated by the act ion of growth hormone and the responses manifested i n other species, a study w a s designed t o determine i f se lec t ion f o r high and low backfat had a l te red t h e concentration of growth hormone per - se.

The growth hormone content of an ter ior p i t u i t a r i e s from 30 Duroc swine balanced by l i n e and sex and selected on t h e bas i s of backfat depth w a s assayed using hypophysectomized female rats. The animals were d l slaughtered at a constant weight, and there w a s no s igni f icant difference i n the average age of t he animals at the t i m e of slaughter. There was a s igni f icant difference between all l i n e s with respect t o depth of backfat and percent of l ean cu ts . One ham f ron each animal w a s separated i n t o lean, fat , and bone. were s ign i f icant ly d i f f e ren t except f o r t h e percent of lean between control and low-line Duroc. The an ter ior p i t u i t a r i e s from f i v e animals of each sex i n each l i n e were pooled f o r t h e assay. The assay procedure used w a s t h a t of Greenspan -- e t a l . (1950). The an ter ior p i t u i t a r i e s from t h e control l i n e w e r e s ign i f i can t ly heavier than those from both of t h e selected l i n e s . Based on two assay c r i t e r i a , each l i n e contained the same amount of growth hormone per un i t of tissue. The amount of growth hormone contained per un i t of t i s s u e t e s t ed i n each a n i m a l w a s not d i f f e ren t as measured by t h e increase i n the growth of hypophysectomized female rats or i n s ize of epiphyseal ca r t i l age width. When t h e an ter ior p i t u i t a r i e s w e r e adjusted For weight, .the t o t a l STH potency ranged from highest t o lowest i n the control , low, and high l i nes , respect ively. Growth hormone concentration ner uni t of t i s s u e

The values from the three pa r t s of t h e ham between l i n e s

336.

between the three l i n e s w a s not s ign i f icant ly d i f f e ren t . This suggests t h a t genetic se lec t ion f o r high and low backfat has probably a l te red the a b i l i t y of t h e t i s s u e s t o respond t o STH.

Effect ~f endogenous and exogenous hormones on growth

It has been ably demonstrated t h a t t he implantation or addition of d i e thy l s t i l bes t e ro l ( a synthetic estrogen) t o t h e ra t ions of c a t t l e , mainly steers and he i fe rs , i s a common and benef ic ia l pract ice . Since many a r t i c l e s have been published on DES and it has been reviewed by previous speakers at t h e Reciprocal Meat Conference, I w i l l not discuss any speci€ic e f f e c t s of DES at t h i s t i m e .

Recent s tudies have shown t h a t t he feeding of melengestrol acetate (MGA), an act ive progestin or ig ina l ly t e s t ed t o synchronize the estro-ds cycle, improves r a t e of gain and promotes growth i n he i fe rs . Bloss -- e t a l . (1966) reported increases i n rate of gain and feed e f f ic iency when KA-fed he i f e r s were compared t o cont ro ls . -- e t a l . (1968), crossbred he i f e r s fed .3 mg. of E A i n t h e feed per head per day f o r 140 days gained s igni f icant ly f a s t e r (21%) and made 11% more e f f i c i e n t use of t h e i r feed than he i f e r s receiving the same feed without MGA. I n t h e l a t te r study, carcass grades, marbling score, and dressing percentage of %A-fed h e i f e r s and controls conformed subs tan t ia l ly with the r e s u l t s of Bloss e t aL. (1966). unaffected by E A i n t h e feed; however, no spec i f ic measurements taken with regaxd t o the amount of f a t or l ean i n the carcass were reported.

I n a recent study reported by O'Brien

I n the study of O'Brien -c e t a l . (1968) dressing was --

N o changes i n general body conformation or vaginal prolapses, as reported t o occur i n estrogen-treated he i fe rs , w e r e observed among any of t he EA-treated h e i f e r s (Bloss -- e t al., 1966). Zimbelman and Smith (1966) showed t h a t E A dos-.s of approximately .2 mg/heifer/day w i l l prevent e s t r u s i n some he i fe rs , and doses of .5 mg/day w i l l prevent e s t r u s i n almost a l l he i f e r s . a n i m a l s but r a r e l y i n MGA-treated animals. Heifers which were found t o be sexually mature and then spayed grew more slowly and had poorer feed ef - f i c i ency than mature in t ac t he i fe rs . The feeding of E A t o spayed he i f e r s did not improve growth r a t e or feed eff ic iency s igni f icant ly . These re - sults, along with results on f o l l i c u l a r s i z e , substant ia te t he hypothesis proposed by Bloss L- e t al. (1966) t h a t E A caused increased weight gain by allowing continual endogenous estrogen secret ion. The failure of s t e e r s t o respond t o MGA seems t o be consis tent with t h i s hypothesis.

They also reported ovulation occurred regular ly i n untreated

Effec t of tes tosterone on growth

It i s w e l l known t h a t tes tosterone promotes pro te in anabolism and decreases t h e urinary l o s s of nitrogen without increasing the non-protein nitrogen of t h e blood and t h a t it produces at least a temporary increase i n body weight. This suggests t h a t t h e hormone causes a t r u e storage of nitrogen i n t h e form of t i s s u e protein. It should be noted t h a t androgens increase t h e body weight of young hypophysectomized r a t s ; but most of t h i s ga in i s accounted f o r by the increased mass of t he genital. coml;lex, ap- parent ly consequent upon protein re ten t ion (Turner, 1966). The changes i n body weight produced by tes tosterone vary with t h e species and a l s o depend on the nu t r i t i ona l s t a t u s of t he animal.. Karlson (1963) and Liao and

337.

Williams-Ashman (1962) have demonstrated t h a t the microsomes from the pros ta te gland of t he tes tosterone-treated r a t have a greater a b i l i t y t o incorporate sa l ine C-14 i n t o protein than microsomes f rom cas t ra ted sub- j e c t s . Since nucleic acids are int imately associated with the synthesis of proteins, t he growth processes i n i t i a t e d by androgens and various t a r g e t t i s s u e s could be mediated at t h e l e v e l of the gene.

Bra tz le r e t al. (1954) reported t h a t barrows cas t ra ted at 180 l b s . and boars had a higher percentage of l ean i n the lo in , a lower dressing per- centage, and less backfat than pigs castrated a t 70 days and 100 l b s . Zobrisky e t al. (1959) and Teague e t al. (1962) also reported a s igni f icant ly higher yield of l ean meat from the carcasses of boars. showed t h a t t he carcasses of boars and g i l t s were longer and had less fa t covering over t h e shoulder and back area of l o i n and a higher iodine number than those cas t ra ted la te . I n t h i s study, acceptab i l i ty t e s t s showed t h a t t he age of cas t ra t ion did not a f fec t f lavor , odor, o r tenderness of meat.

--

-- -- Charette (1961) also

The study of Frescot t and Lamming (1964) showed t h a t ca s t r a t ion consis tent ly depressed growth r a t e i n c a t t l e and sheep. They fur ther point out i n t h e i r study t h a t t h e carcasses of ca s t r a t e s contain much more f a t and less l ean and bone than those of t h e i n t a c t animals. Boars a l so yielded a much leaner carcass containing 19 l b s . less f a t and 7 l b s . more l ean than the carcasses of hogs a t 260 l b s . of liveweight.

Collectively, these s tudies point out t h a t the use of t h e i n t a c t male holds considerable poten t ia l with present demands f o r more e f f i c i e n t production and t h e production of more l ean m e a t .

Concludim remarks

On t h e foregoing pages, I have considered only a handful of t he issues on problems germane t o the e f f ec t of the endocrine system on growth and development. Without doubt, many other considerations could or should have been included. ta in ing t o the f i e l d of endocrinology w i l l understand t h e d i f f i c u l t i e s i n t ry ing t o present a comprehensive discussion of t h e top ic i n a l imited t i m e . On the other hand, it i s hoped t h a t the cen t r a l theme has been presented. Surprisingly l i t t l e work has been conducted on the spec i f ic e f f e c t s and bene f i t s t h a t might be real ized i n meat animals through s tudies i n endo- crinology. Also, we as animal s c i e n t i s t s have been pa r t i cu la r ly slow i n applying basic discoveries i n biology t o all aspects of m e a t animal prcduc- t i o n . Resolution of t h e problems and r ea l i za t ion of t he bene f i t s w i l l r e - quire m a x i m u m e f f o r t as well as e f f ec t ive u t i l i z a t i o n of t a l e n t s from many s c i e n t i f i c d i sc ip l ines working together . Thus, i n meeting our responsi- b i l i t i e s t o society as animal s c i en t i s t s , we are faced with a demanding but exc i t ing challenge for the fu ture .

Those familiar with the vas t amount of literature per-

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Baird, D. M., A. V . Nalbandov, and H. W. Norton. 1952. "Some physiological causes of g e n e t i c a l l y d i f f e r e n t r a t e s of growth i n swine." J. An. Se i . 11:292.

Beach, R . K. and J. L. Kostyo. 1968. "Effect of growth hormone on t h e DNA content of muscles of young hypophysectomized ra ts . I '

Endocrin. 82: 882.

Bloss, R . E., J. I. Northam, L. W. Smith, and R . G . Zimbelman. 1966. "Effec ts of oral melengestrol ace t a t e on t h e performance of f e e d l o t c a t t l e . " J. An. Se i . 25:1048.

Bra t z l e r , L. J. , R. P. Soule, Jr., E. P. Reineke, and P. Paul. 1954. "The e f f e c t of t e s tos t e rone and c a s t r a t i o n on t h e growth and carcass c h a r a c t e r i s t i c s of swine. J. An. Sc i . 13: 1 7 1 .

Charette, L. A. 1961. "The e f f e c t s of sex and age of male a t c a s t r a - t i o n on growth and carcass q u a l i t y of Yorkshire swine." Canad. J. An. Sc i . 41:30.

Davey, R. J. and C . M. Kincaid. 1964. "Response of swine se lec ted f o r backfat th ickness t o d i f f e r e n t energy in takes ." J . An. Se i . 23: 1197. ( Abst .)

Gerrits, R . J., L. A. Johnson, H. 0. Hetzer, and C . M. Kincaid. 1964. "Some anatomical and physiological measurements on swine se lec ted f o r high and low backfat." J. A n . Sci . 23:905. (Abst.)

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Hetzer, H. 0. and W. R. Harvey. 1967. "Selection for high and low f a t n e s s i n swine." J. An. Se i . 26:1244.

Johnson, L. A. and R. J. G e r r i t s . 1965. "Biochemical and hematological measurements on swine se lec ted f o r high and low backfa t . J. An. Sc i . 24:1217. (Abst.)

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Knobil, E . 1961. "The p i t u i t a r y growth hormone: some physiological considerations." Systems. Basic Books, Inc. , New York.

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28. W e l t , I. D. and A. E. Wilhelmi. 1950. "The e f f ec t of adrenalectomy and of t he adrenocorticotropic (ACTH) and growth hormones on the synthesis of f a t t y acids." Yale J. B i o l . Med. 23:99.

29. Zimbelman, R . G . and L. W. Smith. 1966. "Control of ovulation i n c a t t l e with melengestrol ace ta te . 11. Effec ts on f o l l i c u l a r s ize and ac t iv i ty . " J. Reprod. F e r t i l . 11:193.

30. Zobrisky,S.E.,D.E. Brady, J. F. Lasley, and L. A. Weaver. 1959. "Significant re la t ionships i n pork carcass evaluation. I. Lean cu ts as c r i t e r i a for l i v e hog value." J. An. Sci. 18:420.

D. A. CRAMER: Thank you very much, D r . Gerrits, and I w i l l thank all of you gentlemen f o r a s e r i e s of very f i n e papers. I th ink you wodd all make top-notch meats men. Fortunately, t he group ups t a i r s i s also running a f e w minutes behind, so we do have time f o r a f e w questions. I th ink we can run u n t i l about two o'clock for questions. I won't take time out from our discussion t o introduce Dave Topel, because most of you know him, anyway. 1'11 j u s t l e t him get s t a r t ed with t h e questions.

D. G. TOPEL: Thank you, Dave. I ' m sure you w i l l a l l agree t h a t a2.1 of the people on t h i s program are very in te res ted i n composition, and maybe not d i r e c t l y with meats, but cer ta in ly i n the end product t o 8 ce r t a in degree. A t t h i s t i m e I would l i ke . t o ask f o r questions regarding the papers. Are there any questions?

PAUL LEWIS, Arkansas: I would l i k e t o d i r e c t t h i s t o the second speaker--how are you analyzing for sodium and potassium?

DR. EWAN: O u r meat analyzing i s by photometric methods.

LEWIS: Do you worry about any interferences?

EWAN: We haven't investigated t h i s point as yet . I believe there are differences i n response depending on the l e v e l s of sodium and potassium t h a t aze present, and t h i s i n pa r t i cu la r . I n other words, when you go t o look between t i s sues , with a system t h a t i s geared t o work with plasma or serum, you would have t o a l t e r your system t o be sure t h a t you are get t ing accurate measurements, because of t he r e l a t ive enhancements.

DR. HENDRICKSON, Oklahoma: I would l i k e t o address a question t o Gene Allen. Fa t i s deposited i n connective t i s sue and I am wondering a s the fa t bui lds up, i s there an increase i n the connective t i s sue?

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DR. A L B N : I n thinking back, i n dressed beef animals, I don' t t h i n k we have shown a l a r g e difference i n f a t t y t i s s u e i n some of t h e mus- c l e s we have looked a t . These changes have been more i n t h e composition, but as far as I am aware of , I would say no.

HENDRICKSON: You don ' t f e e l t h a t there i s a generation of con- nective t i s s u e as l i f e proceeds?

ALLEN: Not much.

DR. TOPEL: If there are no f u r t h e r questions, I w i l l t u r n it back t o D r . Cramer.

DR. C R M R : Thank you, D r . Topel. This has been an excel lent session. We are running a b i t behind schedule and are due back i n t h e Main Auditorium f o r a paper which you all w i l l want t o hear on Changes i n Standards f o r Pork Carcasses and Slaughter Hogs by John Pierce and Peter W i l l i a m s of t h e U.S.D.A. So l e t ' s adjourn t h i s session and proceed t o t h e Main Auditorium.

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STATISTICAL COMMITTEE W N AUDITORIUM GEORGIA CENTER

H. W. OCKERMAN: I ' l l c a l l t h e meeting t o order and since we're a b i t behind schedule I ' l l not name t h e other Embers of t h e S t a t i s t i c a l Committee. They are l i s t e d i n t h e back of t h e program. We are for tunate t o have two i n t e r e s t i n g papers t h i s afternoon and two highly qual i f ied speakers: George Brissey of Swift on "Sampling" and C . F . Parker of Ohio on "Biological Variation". George will be OUT f irst speaker.