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Studies of growth and nutrition in the Columbian black-tailed deer (Odocoileus hemionus columbianus) Bandy, P. J. 1955

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STUDIES OP GROWTH AND NUTRITION IN THE COLUMBIAN BLACK-TAILED DEER (Odocoileus hemionus columblanus) by P. J. Bandy A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS i n the Department of ZOOLOGY We accept t h i s thesis as conforming to the standard required from candidates for the degree of MASTER OF^^OLOGY Members of the Department of Zoology THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1955 Abstract E a r l y post-natal development was studied i n experimental Columbian b l a c k - t a i l e d deer (Odocoileus hemionus columbianus), with respect to c a l o r i c l e v e l s of Ideal d i e t s . The growth re-sponse, i n terms of weight, he a r t - g i r t h , height-at-withers and length of hind foot, showed that instantaneous r e l a t i v e growth rates were decreased by a decrease i n c a l o r i c consumption. How-ever, the growth pattern remained the same i n the two planes of n u t r i t i o n employed. The assessment of growth showed that weight and heart-g i r t h s i m i l a r l y r e f l e c t n u t r i t i v e condition and are af f e c t e d to the greatest extent by c a l o r i c intake. Height-at-withers also r e f l e c t e d the planes of n u t r i t i o n but the length of hind foot showed l i t t l e d i f f e r e n c e . Thus a method was outlined f o r evalua-ti n g the degree of fl e s h i n e s s or n u t r i t i v e condition by employ-ing the actual weight of the deer or the estimated weight from the h e a r t - g i r t h regression formula and the weight estimated from the hind foot. The index of condition was thus found to be one for normal deer, greater than one f o r above normal deer and less than one f o r deer i n poor n u t r i t i v e condition. Blood samples were taken from 20 to lj.65> days of age showing the changes i n some blood constituents with increasing age. A d d i t i o n a l l y , the blood constituents were analysed with respect to the two planes of n u t r i t i o n . With the exception of blood glucose and fibrinogen, no demonstrable differences between the two planes occurred. However, a l l the constituents studied, with the exception of plasma gl o b u l i n , changed with increasing age. The blood picture presented by the data from these deer are thought to represent a normal condition and may thus be used as a standard i n evaluating blood l e v e l s of wild Columbian black-t a i l e d deer. Table of Contents. Page L i s t of Tables (2) L i s t of Figures (J|) Acknowledgements (6) Introduction . 1 Methods and materials k (a) Experimental animals k (b) Hearing pens 5 (c) Feeding and care 6 (d) Feeding 9 (i) U.B.C. Ration #15-52 . . . . . 9 ( i i ) Milk r a t i o n 10 (e) Weights and measurements 11 (f) S a c r i f i c e of the experimental deer 11 (g) Blood samples; c o l l e c t i o n and preservation . . . . . 12 (h) A n a l y t i c a l procedures f o r blood samples 12 (1) Packed c e l l volume 12 2. Erythrocyte sedimentation rate . 13 3* Hemoglobin 13 k. Blood glucose 13 5 . Non-protein nitrogen llj. 6. Total proteins, fibrinogen, albumin and globulin l k Results and Discussion 16 (a) Computations of c a l o r i c requirements f o r maximum and r e s t r i c t e d growth 16 (b) Adequacy of the diets 19 (c) Actual food consumption 20 1. The milk r a t i o n 20 2. U.B.C. ra t i o n #15-52 21 (d) Success of the planes of n u t r i t i o n 23 (e) Relationships of i n d i v i d u a l growth curves to the planes of n u t r i t i o n 25 (f) Instantaneous r e l a t i v e growth rates and the planes of n u t r i t i o n 29 (g) Individual age relationships of l i n e a r growth . . . 3h (h) Relationships of l i n e a r growth to body weight . . . IJX) (i) An index to n u t r i t i v e condition 1+3 (J) The blood chemistry of the b l a c k - t a i l e d deer . . . . k9 1. Packed c e l l volume 1+9 2. Erythrocyte sedimentation rate 53 3* Hemoglobin 5U k. Blood Glucose 59 5 . The non-protein nitrogen of blood (N.P.N.) . . 65 6. Plasma t o t a l proteins 68 7. Plasma albumins, globulins and fibrinogen . . . 72 Summary and Conclusions 81 Page Literature c i t e d 85 Appendix . . . . . . . . . 91 (a) The composition of rations . . . . . . . . (I) (b) Some observable differences i n response to the . plane of n u t r i t i o n . . . . . . . . . . . . . . . . . ( i i ) (c) I l l n e s s , symptoms, treatment,and response . . . . . . (v) 42) LIST OF TABLES Table Page I The ea r l y h i s t o r y of H group 4 II The instantaneous r e l a t i v e growth rates i n r e l a t i o n to the plane of n u t r i t i o n . i 31 III Instantaneous r e l a t i v e growth rates for Columbian b l a c k - t a i l e d deer (Cowen and Wood, 1954) 34 IV Absolute l i n e a r growth rates i n r e l a t i o n to the plane of n u t r i t i o n 36 V The approximate percentage of the adult measure-ments achieved at b i r t h 38 VI The index of c a r d i t i o n f o r deer 1H and 3H 45 VII Packed c e l l volumes i n percent of whole blood f o r 1954 fawns ( J group) and yearlings (H group) . . . . 5 l V i l a Packed c e l l volumes i n percent of whole blood f o r the yearlings (H group) 52 VIII Erythrocyte sedimentation rates f o r fawns and yearlings 53 IX Hemoglobin values i n gras per 100 ml. blood f o r the 1954 fawns ( J group) 56 X Hemoglobin values i n gms per 100 ml. blood f o r the yearling deer (H group) 56 XI Blood glucose l e v e l s i n mgm per 100 ml. whole blood for 1954 fawns ( J group) i n comparison to H group (yearlings) 61 XII Blood glucose l e v e l s i n mgm per 100 ml. whole blood for the yearlings (H group) 61 XIII Non-protein Ng i n mgm per 100 ml. blood f o r the Fawns (J group) 66 XIV Non-protein Ng i n mgm per 100 ml. blood f o r year-lings (H group) 66 XV Plasma t o t a l proteins as gans protein per 100 ml. of plasma f o r the fawns (J group) 70 XVI Plasma t o t a l proteins as gms protein per 100 ml. of plasma f o r the yearlings (H group) 70 (3) L i s t of Tables - Continued Table Page XVII Plasma albumin i n gms protein per 100 ml. plasma f o r the fawns ( J group) 74 XVIII Plasma albumin i n gms protein per 100 ml. plasma f o r the yearlings (H group) 74 XIX Plasma globulins i n gms protein per 100 ml. plasma f o r the fawns ( J group) 76 XX Plasma globulins i n gms protein per 100 ml. plasma for the yearlings (H group) 76 XXI Blood fibrinogen i n gms protein per 100 ml. blood f o r the fawns ( J group) 78 XXII Fibrinogen i n gms protein per 100 ml. blood f o r . . yearlings (H group) 78 Appendix XXIII The composition of U.B.C. ration #15-52 (I) XXIV The approximate composition of U.B.C. r a t i o n #15-52 ( i ) XXV The approximate composition of P a c i f i c Evaporated Milk and Nurse Cow replacement (i) XXVI The composition of doe's milk (K i t t s et a l , 1955) • (i) XXVII The dressing percentage of experimental deer based on bled and eviscerated carcasses with the head and h i d remaining attached ( i i ) (k) L i s t of Figures Figure Page 1. Feeding standard for U.B.C. r a t i o n #15-52 9a 2. Feeding standard f o r f a t added milk r a t i o n . . . . . 10a 3. Standard curve f o r hemoglobin determination 13a k. Standard curve f o r glucose determination l k a 5. The c a l o r i c consumption of deer 5 J 21a 6. The c a l o r i c consumption of deer 6J 21b 7. Growth i n weight of deer 5 Jo" 25a 8. Growth i n weight of deer 6Je" 25b 9. Growth i n weight of deer lkJ9 25c 10. Growth i n weight of deer 15J9 25d 11. Growth i n weight of deer lHo* 25e 12. Growth i n weight of deer 2Ho* 25f 13. Growth i n weight of deer 10H9- 25g l k . Growth i n weight of deer 11H9 25h 15. Growth i n he a r t - g i r t h f o r 5J 3 5 a 16. . Growth i n hea r t - g i r t h f o r 6J 35b 17. Growth i n height-at-withers of 5 J 35c 18. Growth i n height-atrrwithers of 6J 35d 19. Growth i n length of hind foot of 5 J 35>e 20. Growth i n length of hind foot of 6J 35f 21. Growth of body weight i n r e l a t i o n to age and the plane of n u t r i t i o n of a l l experimental deer 36a 22. Growth of he a r t - g i r t h i n r e l a t i o n to age and the -plane of n u t r i t i o n of a l l experimental deer 36b 23. Growth of height-at-withers i n r e l a t i o n to age and the plane of n u t r i t i o n of a l l experimental deer . . . 36c (5) L i s t of Figures - Continued Figure Page 2I|. Growth of the length of hind foot i n r e l a t i o n to age and the plane of n u t r i t i o n of a l l experimental , deer 36d 25. The p r i o r i t y of nutrients for maintenance or production of tissues (from Hammond, 1952) 39 26. Regression of weight and h e a r t - g i r t h f o r a l l J and H deer lj.Oa 27. Regression of weight and height-at-withers f o r high plane male J and H deer Ij.la 28. Regression of weight and height-at-withers for low plane male J and H deer 1+lb 29. Regression-of weight and height-at-withers f o r a l l male J group (low plane1) l|2a 30. Regression of weight and height-at-withers f o r 2H (high plane 1) . l\2b 31. Regression of weight and height-at-withers for a l l deer 1(2 c 32. Regression of weight and length of hind foot f o r high plane male J and H deer • . I+2d 33. Regression of weight and length of hind foot f o r low plane J and H deer Ij2e 3k* Regression of weight and length of hind foot f o r low plane female J and H deer I|2f 35. Regress! on of weight and length of hirid foot for high plane female J and H deer . Ij2g 36. A graphic method for the evaluation of the index of condition 1+8 (6) ACKNOWLEDGEMENTS. The writer wishes to extend h i s sincere gratitude to Dr. I. MoT. Cowan, Department of Zoology, U n i v e r s i t y of B r i t i s h Columbia, f o r his many suggestions, his help and his encouragement throughout t h i s i n v e s t i g a t i o n . The writer also wishes to g r a t e f u l l y acknowledge the assistance of Dr. A. J . Wood and Dr. W. D. K i t t s i n supplying space f o r the experimental animals, laboratory space and equip-ment and for t h e i r suggestions and help throughout the experi-ment . I would also l i k e to thank a l l the graduate students of the Department of Zoology and the Department of Animal Husbandry who assisted i n so many ways. The writer i s also g r a t e f u l to the National Research Council of Canada f o r t h e i r f i n a n c i a l support i n the form of a research assistantship throughout the summer of 1954 and to the University of B r i t i s h Columbia f o r additional f i n a n c i a l support throughout the in v e s t i g a t i o n . I would also l i k e to thank members of the B. C. Game Commission, Mr. D. Robinson and Mr. C. Esslen who as s i s t e d i n the c o l l e c t i o n of experimental animals. 1. INTRODUCTION. Nu t r i t i o n and growth are fundamentally important to the understanding of adaptive physiology of a l l animals. Numerous studies on domestic animals have produced standards of growth and n u t r i t i o n from which deviations can be measured and analysed. Physiological adaptation to the environment and to envi-ronmental stresses are dependent p r i m a r i l y on n u t r i t i o n . Accord-in g l y wild ungulates r e f l e c t changes i n environmental conditions i n the growth of individuals and populations. Since population growth i s dependent upon the growth of reproducing u n i t s , i t i s important that the basic growth patterns of indiv i d u a l s be thoroughly understood, p a r t i c u l a r l y i n r e l a t i o n to n u t r i t i o n . Studies of growth and n u t r i t i o n i n the Columbian black-t a i l e d deer, Odocoileus hemionus columblanus, were i n i t i a t e d at the University of B r i t i s h Columbia under the d i r e c t i o n of Dr. I. McT. Cowan and Dr. A. J. Wood (Cowan and Wood, 1951+). The present study i s the continuation of t h i s p r o j e c t . For purposes of c l a r -i t y the objects of these experiments may be enumerated as follows: 1. To study growth and the r e l a t i o n s h i p of the c a l o r i c plane of n u t r i t i o n . 2. To study the assessment of growth and the plane of n u t r i -t i o n . 3. To study c e r t a i n aspects of the blood chemistry of the b l a c k - t a i l e d deer i n r e l a t i o n to growth and the c a l o r i c plane of n u t r i t i o n . Six fawns born i n 1953 and fourteen fawns born i n 195U (hereafter r e f e r r e d to as H group and J group, respectively) were 2. reared on two planes of n u t r i t i o n d i f f e r i n g only i n the quantity of food. H group consisted of four males and two females which were ra i s e d "by t h e i r dams i n c a p t i v i t y . J group consisted of fawns captured shortly a f t e r b i r t h on Vancouver Island, B.C., and subsequently b o t t l e - r a i s e d at the University of B r i t i s h Columbia. The planes of n u t r i t i o n were i n i t i a t e d subsequent to wean-ing i n H group but were started early i n the suckling period i n J group. The "high plane" f o r both groups was designed to provide a s u f f i c i e n t l y high c a l o r i c intake, of a well balanced r a t i o n (Ration U.B.C. #1^-52) to permit nearly maximal growth, on an l s o c a l o r i c feeding b a s i s . The "low plane" i n d i v i d u a l s i n both H and J groups were fed the same r a t i o n , at a l e v e l s l i g h t l y above that calculated f o r maintainance. Thus growth continued i n the low plane i n d i v i d u a l s but at a minimal rat e . Controlled feeding was accomplished by o f f e r i n g only the calculated amount and re-weighing the remaining food every twenty-four hours. Fluctuations In the d a i l y intake were c h a r a c t e r i s t i c of the high plane indiv i d u a l s whereas low plane animals t o t a l l y consumed the i r d a i l y r a t i o n * Accordingly the planes of n u t r i t i o n resulted i n d i f f e r e n t i a l growth r a t e s . Body-weight, hea r t - g i r t h , height-at-withers and length of hind foot were measured at i n t e r v a l s as an assessment of growth. Blood samples were also taken at i n t e r v a l s and analysed f o r packed c e l l volumes, erythrocyte sedimentation rate, hemoglobin, blood glucose, non-protein nitrogen, t o t a l plasma prot e i n , fibrinogen, albumin and globulin. The anatomical and chemical measurements were analysed 3. with reference to the stage of growth and the plane of n u t r i t i o n . Blood analyses also demonstrated monthly variations In the l e v e l s of c e r t a i n constituents which may be attributed to environmental changes such as temperature. METHODS AND MATERIALS. (a). Experimental Animals. The animals used i n these experiments were the Columbian b l a c k - t a i l e d deer, Odocoileus hemionus columbianus. H group con* s i s t e d of fawns born i n 1953 which were r a i s e d at the University of B r i t i s h Columbia or were presented to the University i n the f a l l of 1953* Table I shows the hi s t o r y of this group. TABLE I THE EARLY HISTORY OP H GROUP Animal No. Sex Date of B i r t h Reared Si r e Dam 1H o* June 17/53 By doe, U.B.C. Blaokie Jumpy 2H o" June 17/53 By doe, U.B.C. Blackie Diana 3H o* June 15/53 By b o t t l e , Stanley Park Wild Wild o" June 15/53 By b o t t l e , Stanley Park Wild Wild 10H 9 June lk / 5 3 By b o t t l e , Port Hardy Wild Wild 11H 9 July 3/53 By doe, U.B.C. Brownie Hardie The s i r e s and dams recorded for the fawns r a i s e d a t the Univers i t y of B r i t i s h Columbia can be considered to have been on a completely adequate d i e t during gestation and throughout the suckling period. The n u t r i t i o n a l h i s t o r y of the dams and fawns donated to U.B.C. from Stanley Park and Port Hardy i s not known. However, p h y s i c a l comparisons showed no marked differences i n thei r condition from that of the deer r a i s e d at U.B.C. 5. The 1954 fawns ( J group) were captured sh o r t l y a f t e r b i r t h at Courtenay, Vancouver Island, B.C. Nothing i s known con-cerning the n u t r i t i o n of the dams during gestation. However, a l l the adult deer seen during the c o l l e c t i o n of the fawns appeared to be i n healthy condition, r e f l e c t i n g r e l a t i v e l y good n u t r i t i o n , at l e a s t during the l a t t e r part of gestation. J group o r i g i n a l l y consisted of eleven males and seven females. D i f f i c u l t y i n adapting the fawns to b o t t l e feeding resulted i n the loss of one male and three females. Further n u t r i t i o n a l d i f f i c u l t i e s during early suckling r e s u l t e d In the death of three males and two females. Accordingly, complete records are available for seven males and two females i n J group, (b). Rearing Pens. H group deer were ra i s e d i n pens 6» x 6' x 4 ' constructed of plywood, during the winter of 1953-1954. The fawns were h a l -ter-broken and t i e d , two i n each pen with separate food and water containers f o r each i n d i v i d u a l . The pens were situat e d i n a shed-like p o r t i o n of a barn which was exposed only to the south. In early spring these animals were moved to i n d i v i d u a l pens constructed of 2" x 4" r a i l s . These pens were 30' x 4 ' x 8' with the f i r s t f i v e f e e t forming covered pens exposed to the south. Food, water, bedding were placed i n the covered porti o n . J group fawns were r a i s e d i n i n d i v i d u a l pens 5 ' x 3s* x 4 1 * constructed of plywood. These pens were situated i n an unheated b u i l d i n g but were f u l l y protected from the weather. The cleaning of a l l pens was f a c i l i t a t e d by the use of wood shavings to the depth of approximately one inch. Paper 6. excelsior was t r i e d but required more time and more effort, i n cleaning the pens. ( c ) . Feeding and Care. A p e l l e t e d r a t i o n (U.B.C. #15-52), designed by Dr. A. J . Wood of the Department of Animal Husbandry, was used as the major food source following the weaning of both the J and H groups. (The composition of this r a t i o n i s l i s t e d In the appendix.) Good q u a l i t y a l f a l f a hay was used as a roughage which was replaced by fresh grass cuttings when a v a i l a b l e . Both the p e l l e t e d r a t i o n and roughage were offered i n i n d i v i d u a l feeding trays f o r each animal. S a l t i n brick-form was also .placed i n the feeding trays separating the roughage from the p e l l e t e d r a t i o n . The p e l l e t e d r a t i o n was fed on an i s o c a l o r i c basis em-ploying calculated standards of feeding. The quantity of p e l l e t s to be fed to each deer was weighed and the remaining p e l l e t s were re-weighed a f t e r twenty-four hours. In contrast, roughage was offered at the rate of one quarter pound per day regardless of age of the deer or the type of roughage. The low c a l o r i c values of roughage contributed l i t t l e to the t o t a l c a l o r i c intake, con-sequently, unconsumed roughage was not re-weighed and d i d not enter into the calculated c a l o r i c requirements. I n i t i a l l y , J group was fed a d i l u t e s o l u t i o n of P a c i f i c Evaporated milk. Digestive d i f f i c u l t i e s , which were a r e s u l t of feeding d i l u t e d milk, were s a t i s f a c t o r i l y a l l e v i a t e d by feeding concentrated evaporated milk with added f a t i n the form of corn o i l . The analyses of doe's milk showed a t o t a l s o l i d s content of approximately 25$ and a f a t content of 10.5$ (For the composition 7 of deer milk and evaporated milk, see appendix.) Addition of eight grams of corn o i l f o r every 100 ml. of 1:1 milk and water, ra i s e d the f a t content of the r a t i o n to that of deer's milk. On finding the t o t a l s o l i d s of deer's milk to approximate 2$%f the fat-added, evaporated milk was fed i n concentrated form without correcting the f a t content. The f a t content of the r a t i o n fed was thus erroneously elevated to 15.8$. However, l i t t l e d i f f i c u l t y with feeding this d i e t was encountered and the r a t i o n proved sat-i s f a c t o r y . Milk was fed from regulation baby b o t t l e s and nipples. Openings i n the nipples were enlarged ±n order to f a c i l i t a t e feeding. . In addition, warming the milk to body temperature was found to be necessary to encourage feeding during the e a r l y suck-l i n g period. Three feedings per day were necessary during the early suckling period In order to provide the calcu l a t e d c a l o r i c Intake. During the l a t t e r stages of the suckling period two feedings per day were used which was l a t e r reduced to one, during weaning. Mineral d e f i c i e n c i e s were prevented by adding f e r r i c c i t -rate and copper sulfate to the milk and by o f f e r i n g Ringer's solution with added magnesium ch l o r i d e . During the weaning process, Ringer's solu t i o n was replaced with s a l t i n a brick-form (Lesl i e Salt-Range b r i c k ) . One raillilitre of a s o l u t i o n containing 10 grams of f e r r i c c i t r a t e and one gram of copper sulfate per $$0 ml. of s o l u t i o n was added to every l± l i t r e s of fat-added milk. The Iron and copper successfully prevented anemic conditions r e s u l t i n g from 8. the extraction of 20 ml. of blood every ten days. The addition of magnesium chloride (MgC^'oHgO) i n the amount of. 0,1% i n Ringer's s o l u t i o n was found to a l l e v i a t e symp-toms of magnesium deficiency which res u l t e d from feeding evapo-rated milk. Vitamin d e f i c i e n c i e s r e s u l t i n g from prolonged rumen development were mitigated by the administration of two drops of Pilehardene o i l every two days and by i n j e c t i o n of Solu-Zyme when necessary. Accordingly, vitamin D d e f i c i e n c i e s d i d not a r i s e and vitamin B d e f i c i e n c i e s were promptly a l l e v i a t e d by i n j e c t i o n s of the Solu-Zyme. Scouring, r e s u l t i n g from changes i n the d i e t , was quickly mitigated by the administration of crude aureomycin. Dosages ranged from j^ -1 tablet per day depending upon the size of the animal. Weaning proved d i f f i c u l t under r e l a t i v e l y aseptic condi-tions imposed on the fawns by the method of r e a r i n g . A milk re-placement, "Nurse-Cow," was t r i e d i n an e f f o r t to wean the J group fawns. Small amounts were added to the milk i n increasing quantities. However, distaste f o r the milk replacement was e v i -dent and i t was discontinued. Grass and p e l l e t s were placed In each pen and milk feeding was continued. Voluntary acceptance of grass and f i n a l l y of p e l l e t s allowed f i r s t the f a t content and then the milk to be decreased i n the amount fed. Weaning was thus achieved at a r e l a t i v e l y l a t e date. I l l n e s s , symptoms, treatment and response of the experi-mental animals i s recorded i n the appendix. 9 (d). Feeding. ( i ) . U.B..C. Ration #15-52. The feeding standards used In these experiments were based on the equation, Basal energy metabolism = 70.5 * Cal./Kgra/day for adult, non-pregnant, non-lactating mammals (Brody, 1945)• Kleiber's p r i n c i p l e , Maximal food energy consumption = 5 Basal energy metabolism was u t i l i z e d to calculate the upper parameter of the standards. Thus 70.5 x W ° ' 7 3 x 5 Metabolizable Energy of the r a t i o n (Cal./lb) represents the maximal quantity which the deer could eat, i f the Columbian b l a c k - t a i l e d deer follow the same pattern as shown by many domestic animals. In order to prevent fluctuations of i n -take r e s u l t i n g from feeding maximal amounts, the quantity fed to the high plane groups was lowered to 75$ of maximal intake. Figure 1 shows the high plane standard which was based on the assumption that the metabolizable energy content of the U.B.C. r a t i o n #15-52 was 1200 Cal/lb when fed with one quarter pound of roughage. The low plane standard for U.B.C. r a t i o n #15-52 was a r b i t r a r i l y set at 60% of the high plane standard. Figure 1 shows that the resultant plane of n u t r i t i o n i s s l i g h t l y above a main-tenance l e v e l based on the assumption that Maintenance food energy consumption • 2 (Brody, 1945) Basal energy metabolism Accordingly, growth was not arrested by the low plane of n u t r i t i o n 10. but proceeded at a very slow rate. In contrast, the high plane standard permitted a t h e o r e t i c a l growth response of 75$ of maxi-mal growth since the plane of n u t r i t i o n was set 75$ below maximal Intake. . ( I I ) . Milk r a t i o n . Feeding standards for the milk r a t i o n were calculated i n a s i m i l a r manner to that used for U.B.C. r a t i o n #15-52. However, the 0.67 power of body weight (W) was used In the calculations f o r the fawns as i t has been shown that basal energy metabolism varies with the 0.67 power of body weight i n young animals In contrast to \P*73 i n adults. Thus the maximal food intake can be expressed as: 70.5 x W ° * 6 7 x 5  Metabolizable energy of the r a t i o n (Cal./fl.oz.) Assuming f a t to represent 9 Cal./gm., protein k Cal./gra. and car-bohydrate k Cal./gra. in terms of metabolizable energy, the f a t -added milk r a t i o n was estimated to contain 952.56 C a l . / l b . or 63.5 C a l . / f l . o z . This estimation does not seem unreasonable since i t represents approximately 90$ of the gross energy of the r a t i o n calculated on the bases that f a t represents h287 C a l . / l b . , protein 2563 C a l . / l b . and carbohydrates i860 C a l . / l b . •1 The high plane standard f o r t h i s ration^corresponds to the calculated maximal intake. Figure 2 shows that the low plane was again a r b i t r a r i l y set at 60$ of the high plane standard, r e s u l t i n g i n a plane of n u t r i t i o n above the maintenance l e v e l . The milk r a t i o n and U.B.C. r a t i o n #15-52 were provided d a i l y on an i s o c a l o r i e b a s i s . However, loss i n weight was neg-11. lected i n determining the amount to be fed, since a decrease i n the food provided would further depress loss i n weight. The experimental deer were randomly selected f o r the planes of n u t r i t i o n . Thus 2H, 3H and 10H were placed on high plane and 1H, 1+H and 11H on low plane. Odd numbers i n J group correspond to the high plane of n u t r i t i o n and even numbered represent the low plane. (e) . Weights and Measurements. Body weights and measurements were recorded at i n t e r v a l s f o r the purpose of assessing growth. Pawns were weighed i n a weighing crate s i m i l a r to those used f o r pigs. H group deer out-grew t h i s method and were l a t e r weighed on platform scales. Body measurements consisted of heart g i r t h , height-at-withers and length of hind foot and were measured by the use of a p l a s t i c tape marked i n centimeters. Heart-girth consists of the minimal chest circumference, approximately one inch behind the fore-legs. Height-at-withers was measured from the f l o o r to the highest point of the body above the shoulders while the deer stood erect. Length of hind foot was measured from the t i p of the calcaneus to the point of the hoof. (f) . S a c r i f i c e of the experimental deer. A l l experimental deer except 2J, I4.J, 6J and 10J were s a c r i f i c e d at the conclusion of the axperimental period on December k» 195>4» The remaining deer w i l l be kept f o r another year on a low plane of n u t r i t i o n . The deer marked f o r s a c r i f i c e were weighed, shot and b l e d . The carcasses were hung, eviscerated and reweighed. Organs were weighed separately and the skeletons 12 were saved f o r study. The relationships of organ weights and s k e l e t a l analyses w i l l be reported at a l a t e r date; however, dressing percentages are recorded i n the appendix. (g) . Blood samples; c o l l e c t i o n and preservation. Blood samples of 20 ml. were taken from each deer at i n t e r v a l s between July 5/51+ and September 23/51+. A 20 ml. syringe was used with 1^" s i z e 22 needles f o r venepuncture of the fawns. As the deer grew, larger sized needles were used to expedite the sampling procedure. The deer were held by assistants while 20 ml. of blood were withdrawn from the recurrent t a r s a l vein employing aseptic techniques. The blood samples were then flushed i n t o test-tubes containing 0.2 ml. of a 10$ potassium oxalate solution which had been oven-dried. Samples were then stoppered and stored at 6°C f o r a n a l y s i s . (h) . A n a l y t i c a l procedures'for blood samples. 1. Packed C e l l Volume. Hematocrit tubes were used tb determine the packed c e l l volume. Oxalated blood was centrifuged i n an International c l i n i c a l centrifuge at half-speed f o r f i v e minutes (approximately 1000 r.p.m.). The speed of centrifugation was then increased to approximately 2000 r.p.m. f o r f i f t e e n minutes. The l i d of the centrifuge was l e f t open during centrifugation i n order to prevent heating. Hematocrit values were recorded and the samples were recentrifuged at f u l l speed for another f i v e minutes. I f no change had occurred, the f i n a l hematocrit values were recorded but if. a change were noted, the samples were respun u n t i l no 13. further change occurred i n the packed c e l l volume. 2. Erythrocyte sedimentation r a t e . The Hellige blood sedimentation tube, with an inside diam-eter of 2.5 mm. was used i n this analysis. The procedure was modified from that reported by D1 Amour e_t a l , (19i+8) by d i l u t i n g the blood four parts to one with Ringer's s o l u t i o n instead of 3$ sodium c i t r a t e s o l u t i o n . Ringer's solu t i o n was used because potassium oxalate had previously been added to the blood to prevent coagulation. Sedimentation tubes were placed i n a perpendicular p o s i -t i o n and the rate of sedimentation was recorded i n mm. at the end of one hour. 3. Hemoglobin. The method of Wong (1928) was followed without modification for the quantitative determination of hemoglobin. A Coleman, Model 11A, spectrophotometer was used and a standard curve pre- ' pared (Figure 3 ) , i n order to d i r e c t l y translate the spectrophoto-meter readings to hemoglobin i n grams percent. Assuming the hemo-globin of deer's blood to contain 0.336$ Iron produced accurate r e s u l t s , as non-hemoglobin Iron accounts f o r only 1 to 2$ of the t o t a l blood i r o n . k. Blood glucose. Protein-free blood f i l t r a t e was prepared by the method of F o l i n (Hawk et a l , 195l)• This method was modified by the use of a centrifuge Instead of f i l t r a t i o n , to obtain the clear super-natant s o l u t i o n . Benedicts colorimetric method for the determination of fiiniUiiiiiiiiiiiliiii^^ •••••••••••••••••••••••••••••a •••••••••••••MHNII • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • a • • • • • • • • • • • • • • • • • • • • • • • • • a i i i i i i i i i i i i l i i i i i i i i i l l i i i i i i i i i i i i i i i i i i ! HKlUMIItMlllMK n::: : : : : : : : : : : : : : : i : :s: : : : :Ec:: :S:M: llHIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII I | | | | H i l l III HIIL. llHIIHIHIIIIIIHIIIIIIIlllllllll l l l l l l l l l l l l l l l I H i l l I II I II . IIIIIIIIIIIIIHIIIIIIIIIIIIIIIIIII l l l l l l l l l l l l l l l l I H i l l I I I I H I I I II I II II II II I I I l l l l l l l l l l l l l l l l I llllllllllllllll l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l IIIII i i Hill II I::::::::::::::::::::::: ::::::::: ::::: :: I : K S : ::::: :::Es:ssisssH:s::F : : : r : : : r " : : : : : : : : : K : : : : : Ill III 1 1 III III III m i l l III Ultlit III II II I I II III IIIII i i i i i HUH m i l l ijn: is:!::::::::::::::::::: n i l i i i i i i i i i l i i i i i i i i i i i i "-lis ::::::::::::::=:::::::::: 1 . . « « . . . . . . . . . . . . . . . a . . . . . :j|=::::::::::: =========|=^ E^^ E^ =^^ ^^ EE ^ ^^ ^^  fEEEEEp^ E^  EEEH^ i^^ ^^  ^^ ^^ ^^  ^ ^ ^ E ^ ^ E ^ ^ ^ ^ ^ ^ ^ ^ ^ ^^ ^ • • • • • • « • • • • • nun i nun • • • • • • • • • • • • • • • • • • • • • • • • I luiiiiiiiiiiNiim I r s : : H K I I I I I m i mm .."•••Hill I I I I I I I I I I H I M I I I I I I I I I I I I I I I I I I - •II l l l l l l l l l l l l l l l l iiiiiiii IIIII II II III IIIII III II II III IIIII III II III IIIII III III I •••••••••••*•••••• >m m i l I I I I I HIM I I H I I I I I I I I H I I I I I I I I I I I I :::::::::::: SSI I I I I I I I I I I " " " M I I I •lllllllllllllllllllllllltmiMIIIIHIIHt l 'Mil IIHII l l l l . ; " M | | H i l l l l l l l l l l l l l l l l l H I I I I I I U H IIIII III IIIIIIII I I U ; : * IIIIIIII m u m II m m m i i i i i m i H i m i i i i i m i m i i i m i i m m :=;:: ==•===!== i n I I H I I I I H I I HUH i i i i i . : v i u i i i IIHIIIIHIIIIIIII MilHl iiillilliiililBgHigilliilUIIII imil!gliliiilimimiliiiiiii^^ii!ill!liliy£iml S3ss3::ss:s iiiimini :===:::;:=:=:==: :: I llltllllMIIIIIIIIMIIMMItlllllMtlllB •••••••IIIMIIMMI ••••••••••••••HIIIIMIIIIIIIHIIMNIIMf ••••••lltllllltlllllllMIIIIIIIUlM • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • I 14. sugar (Hawk et a l , 19i>l), was used with one modification. B o i l -ing f o r s i x minutes was found to be i n s u f f i c i e n t to complete the reduction of copper by the glucose. Accordingly, samples were b o i l e d f o r twenty minutes and a standard curve was prepared (Figure 4) . A Coleman, model 11A, spectrophotometer was used i n the determinations of the reduced copper. The samples were read within I4.O minutes after adding the molybdic acid reagent. 5>. Non-protein nitrogen. The non-protein nitrogen of- deer blood was determined by the digestion method of F o l i n and Wu employing the F o l i n and Wu protein free blood f i l t r a t e . Modifications consisted of changing the colorimetric method of F o l i n and Wu to a t i t r a t i o n method, following digestion. A micro-Kjeldahl s t e a m - d i s t i l l a t i o n apparatus, modified by the S c i e n t i f i c Glass Apparatus Co. from the Parnas and Wagner model, was used f o r the d i s t i l l a t i o n . Reagents and methods employed for the s t e a m - d i s t i l l a t i o n and t i t r a t i o n are those reported by Steyermarky 19f?l. 6. Total Proteins, Fibrinogen, Albumin and Globulin. Micro-Kjeldahl techniques were used i n the determination of the t o t a l proteins, fibrinogen and albumins of blood. The reported gl o b u l i n l e v e l s r e s u l t from the difference between f i b -rinogen - albumin and t o t a l proteins. Methods used f o r the analysis of proteins are those reported i n Hawk et a l , 193>1, modified to a t i t r a t i o n method. Following digestion the samples were ste a m - d i s t i l l e d into 1$ b o r i c llliilliiilllilillll=l!!iiililliillilll|iiiiiiiiiiiiiiiiiiiiiiiiiiiH mmmmmmm\m\\mmimmmmm ::s::::s:::si.:- :::::::::::::::::::::::::::::::::::::: •Hi:!::: : !liii::::aL:s5:::s:i • • « • • • • • » • • « • • • • • • • • • • • • • • • • • • • • • a • a * . -• • • • • • • • • • • • •••••• •••••••••••• \mmm\m\m\mm E E : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : ::;::::::::::::::::::::::::::::: ::::::::::::::::::: • • • • • • • • • • • • • • • • • • « • • • * • • • • • • • • • • • ••< •••••••••••••••••••••••••••••(••••••••••••••••••I**••••••••• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •••••••••*••< • • • • • • • • •••> • •••••••••••I • • • • • • • • • • • a • • • • • • • • • • • • • • • • • • • • • • • ••••••••••••••••••••••••••••••••••••••I_. •••••••••a••••••••••••••!•••••••••••••••«• • • • • • • • • • • a • • • • • • • • • • * • • • • • • • aaiaiaa] • • • • • a i • l l t l l l l l M I I I H • • • • a i M i a i m o a a « i a t • • • l a i i M a i i M • I••••••••aaaaaaa••*«•»•••• IMIMNIMINIIINII •••••••••a !•••« • •••••••••• ••••• ••••• iMMMaai ===========s==s====s=ssEES: EEEEEi^ EEE SSSEEEEESE E=s^=Hzl^^ ^£ES££EESS i sssssss••••• ••••«••••••• • a a a a • • • • • a a a a i - - - ) • • • • • • * « • • • • • « I aaaaaaa aaaaaaiajaBBB •••••••••••••••••••in • ••••••••••••••Hi •••a ••••••••••••••••! aaaa ••••• •••••• I • • • • • • • • • I • • • • • • • • • ^ ; u " Inn n i I U in 111! m i i i i m i i i i i m i m i i i i 111 m i • IIIHIIIIIIII l l l l l l l l l i l l l l l l l l l l l l IBM I 11811 I i l l ! 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S B iiiiimiii l l l l l l l l l l l l l l l l llllll mn Kill "11 !!!! :::: l l l l l l i i i i i i i i i i i i m i i i i i i i i i i i i i i i a i i i i i i ii i i i i i i i i i i i i i i i i i i s i i i i i a •••••••••••••••••••••••••••••••••••••••I IIIIIHI l l l l I l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l 1 l l l l l l l l l l l l l l l l ::::::::::::: . -'""=========== i i i i i i a P ..Hill l l H i l l f f l ISmssss SSSSS •••••••••• SSSSSSSSn SSSSSSi ill • , • : . • u p • • • • • • • • • • • • • • I • fEEEssssEs EEEEsssssr ssssEsEEEs|ss|E=sEsElrE=sESSs S|S« a • • • • • « * • • • • • • • • • • • • • • :::::::::=:=::: l::::::::::::H»::::::::::H:::HH» |:::::::::::::::::::::::::::::::::::::::::::::::::: :::::::::::::: I:::::::::::::::::::::::::: . . . .....«.•••»• •! in mmm •• =—IIM l l l l l l l l l l l l l • I I I I I I I I I I I I I I I I I I I I I I I I I I I l l l l l i i i i i i i i i i i i i i i i i i i i i i m i i i i i i i i i i n i M i i i i i i i •• m i .. l i i i M i i i i i i i i i i i i i i i i i i i i i i i n i i i i i i i i a i i i i i i i i II i II i i i i i i i i i i • m i H I • • • • • • • i i i i i i n IIIIIII • • • • • • • • i m n i i i i i l a i i i i i m n i 11 IIIIIIIII I I IIII I N I I l i i H l i r ' lllllUIKHIIIIIIIIiaillllSIIHIIIIIIIHIIIIIIII ia • •• i i n i i m i • m i •• • • •IIIIIII. IIIHIIIIIIIIIIIIIIIIHIIIIIIIIIII l l l l l l l l II I •••••••IIIII I I I II II I I IIIIIIIII • m i n i m i i i i i i i i m m i i m i i i a i i i m i i n 111 IIIIIIIII I I I II II • i i i i m i m m m J •IIIIIIIII m i i i m i i m i i c t u i i i i J i i i inn mn IIIII IIIIIIIIIIIIIIIIIIIIIIIIIIII minimum IIIIIIIIIIIIIIIIIIIIIIIIIIII i i i i i i i m i i m 111 >]iii immiiii i: i in IIIIIIIII IIIIIIIII II ii!: :E s::s :::::: ssss :::: :::::: :::: I I I I I l l l l S | 11! 9811 I II I l l l l m m m i jiiiiyiiiiiiiiii m i m i n i l m i n i l • i • i n m i • • •III l l l l I I II II II • i l l ttiia •• •••• •••• •• l l l l l l l l l l l l l l l l • III Mil m II 1116 I K I I H I II i l l II I I : : : : : : : : : : : : : : : . . . . . --••.-•••••••••••••.»••••••••••••••••• ••••••.w I . . . . . . . . . i i i m i m m u M • • » m m | !i I i i i i i i i i I l N i i i l W • • • • •I n :::: •IIIIIII l l l l l l l l l l l l l l III! l l l l l l l l l l l l II I l l l l l l l l l l l l l l l l m i m i ~ II l l l l l l l l II III • II IIIIIII Iffl l l l l l l l l S3 SSS IIIIIII I l l l f l l l I ill l l l l l l IIIIIIIII l i m i t . ^ i»a« l l l l l l l l l l l l l l l l l I I H I I I I * M | l l l l l l l l •••••••I l l l l mi ll l l l l l l l I I I U mi IIIIIIIIIimi I I I IIIII! :::::: ::::::::::::::::::: ! • • • • • • • • • • • • • • • • • • • • • • • • • : : : : : : : : : ::::::::::HH::::::::::::m^ ••imiMIMllMIIMIimllllMWIMIMMIMH.IMimmmBl HI U I HWIIlMIII III IWII MlllllllMMWIli III III III III IIIIMIIIII I I I I I I I I I I I H l l l l l l l l IIIII mn iiii i i i i i imimiiiii i imi II II i II IIIIIIIII I H I II II I I IIIIIIIIIII H I IHL . mimm miiimmiiiimiiiii i i i mimiim i II u 1 1 miiiimim IIIII IIIII iimimi H mi mi mi i imiiii i i imii mm I I I B IIIIIIIIIII •••••••••••••••IIIIIII II ••••••••it ••••• •• • • I I I I I I I i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i I I I I I I I I I I I • • • • • • i i i i i i i i i m i i i i n i i i i i i i n i i n n II • • • • • • • i i i i i i i i i i i i i i i i i i i i i i i i m i i i i i m i i i i i n l • I Mil •••• l l l l I IIIIIII IIIII IIIIIII •••••••••••••••••••••••I n m i m i m i i i m i m i i i i i i i i i n u n i i i i m m m i i i i i i i i i i l m n m n i i m i m i m m m i i m m i i ] • ••IIMIIIMIMIIIIIII I  l l l l l l l l l l l l l l l l l l l l l l " m n II IIIII II l l l l l l i iiiiiimi minimi •III i i i i i IIIIIIIII III! H i l l l l l l l l l l l l l l III l l l l llll l l l l l l l l l i l l i i m i : l l l l I l l l l I l l l l l l III l l l l l l l l l l l l l l l l l l l l l l l l l l l l U t W f H I I I I I U l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l II II l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l l iimiiimmmii III IIIIIII l l Illlllllllllllllll IIIII llllllll 15.. acid as reported i n Steyerraark, 1951, using the modified Parnas and Wagner s t e a m - d i s t i l l a t i o n apparatus manufactured by S c i e n t i f i c Glass Apparatus Co. Digestion was allowed to proceed u n t i l the sample turned clear and the dense white fumes given o f f during digestion were reduced i n volume. The shortened digestion period produced accurate r e s u l t s within the l i m i t s of error of the whole experi-ment. Recovery tests using egg albumin produced an average exper-imental error of * 2.22$. Data r e s u l t i n g from the blood analysis was averaged f o r H and J groups, f o r the samples representing d i f f e r e n t ages and , with respect to the plane of n u t r i t i o n . Averages were tested for s i g n i f i c a n t differences by the method outlined by Snedecor (1946). 16 RESULTS AND DISCUSSION. (a). Computations of caloric requirements for maximum and  r e s t r i c t e d growth. , Feeding standards f o r these experiments were based on the assumption that basal metabolism i n the Columbian b l a c k - t a i l e d deer follows the same pattern shown by many domestic animals. Thus basal metabolism = 70 .5 x W Cal/Kgm/day was used to prepare the standard, where 70 .5 represents the c a l o r i c requirements per u n i t weight and W represents body weight i n kilograms. Investigators such as Sarrus, Rameaux, Bergmann,' Rubner and Mech report the value of M b n i n the order of ^ whereas Kleiber reports "b" to equal j^. An average f o r numerous species shows "b M i n the order of 0 •731+ (Brody, 191+5). Since no work has been done on the basal metabolism of deer, the average f o r many species was employed. Thus Basal metabolism s 70.5 x Cal/Kgm/day was assumed to represent the pattern i n the Columbian b l a c k - t a i l e d deer. The above equation i s true only of mature animals, uncom-p l i c a t e d by growth, reproduction or l a c t a t i o n . During e a r l y growth, metabolism per W*3 shows a peak when the t o t a l metabolism Is pl o t t e d against weight on a log-log g r i d . P r i o r to t h i s peak, metabolism tends to be d i r e c t l y proportional to simple weight, W 1 , 0. and thereafter to W°«67 u n t i l growth i s complete (Brody, 191+5) • Various investigators have reported values f o r ^b" rang-ing from 0.15 to 1.17 f o r d i f f e r e n t species and f o r d i f f e r e n t phases of growth under various conditions. However, since the metabolic pattern has not been studied f o r the deer, the feeding 17. standards were s i m p l i f i e d by using 0.67 as the value of nb" p r i o r to weaning and 0.73 during post-weaning growth. The Nformer was based on the assumption that metabolism may be thermodynamlcally closer to an expression of the surface law during early stages of 2 growth, I.e., surface area varies with the j power of body weight. Kleiber reported that the maximum r e l a t i v e food capacity, i . e . maximal food energy consumption basal energy metabolism i s approximately 5 i n such diverse animals as c a t t l e , rabbits and chickens (Brody, 191+5). Since i t i s reasonable to assume that maximum food intake would produce maximum growth, 70.5 x Wb x 5 Metabolizable energy of food was regarded as the upper l i m i t of the feeding standards. In order to prevent digestive d i f f i c u l t i e s and fluctuations In con-sumption which might arise at t h i s l e v e l of intake, the high plane deer were fed 75$ of the i r calculated maximal food consump-tion as shown i n Figure 1 for U.B.C. r a t i o n #15-52. Results showed th i s l e v e l of feeding to produce rapid growth with con-s i s tant food consumption. However, i n an e f f o r t to produce max-imal growth i n J group while on the milk r a t i o n , the high plane fawns were fed the calculated maximal food intake l e v e l (Figure 2 ) . Digestive d i f f i c u l t i e s from over-feeding during the suckling period caused food consumption of the high plane J group to fl u c t u a t e . A d d i t i o n a l l y these deer were seldom able to consume the calcula t e d d a i l y food r a t i o n . This tends to suggest that the feeding standard was too high and thus "b" should be les s than 0.67, or 18. the maximum food consumption i s less than f i v e times basal during early growth. Towards the conclusion of the experiments, i t was decided to determine whether 70.5 x W0*^ x 5  Metabolizable energy of the r a t i o n adequately represented the maximal food intake f o r U.B.C. r a t i o n #15-52. Daily rations f o r 3H and 10H were gradually increased above the 75>$ l e v e l of the high plane standard u n t i l p e l l e t s r e -mained In the feeding trays a f t e r 2k hours. I t was thus found, f o r example, that 3H reached a maximum food intake of 1+.9 l b s . of the r a t i o n at 120 l b s . body weight. Figure 1 shows that the c a l -culated maximum food intake for this weight i s k.8 l b s . There-fore i t may be concluded that 70.5 x W°-73 x 5 i s a reasonable basis for estimating the maximal food intake f o r deer under these experimental conditions and at approximately one year of age. The low plane standards were calcul a t e d to provide 60$ of the food intake of the high plane deer, at the same body weight. Figures 1 and 2 indicate that the low planes of n u t r i t i o n were above the calculated maintenance standard f o r both the milk and the p e l l e t e d r a t i o n . Due to experimental d i f f i c u l t i e s i n v e s t i -gators have been unable to accurately e xpress maintefaance as a function of basal metabolism. However, i t has become apparent that Maintenance food energy consumption basal energy metabolism approximates two. Accordingly, maintenance requirements may be expressed by: v, 70.5 x WP x 2  Metabolizable energy of the r a t i o n 19. Thus the low plane of n u t r i t i o n for U.B.C. r a t i o n #15-52 w a s 11.27$ above the calculated maintenance l e v e l and f o r the milk r a t i o n , 33*38$ above maintenance. Growth rates on the low plane standard for U.B.C. r a t i o n #15-52 were minimal, as expected from the proximity of the low plane feeding standard to the maintenance l e v e l . This tends to suggest that maintenance can be expressed by the equation: Maintenance food requirements = 70.5 x W°* 73 g 2  Metabolizable energy of the ratio n during the post weaning growth phase. (b). Adequacy of the d i e t s . U.B.C. r a t i o n #15-52 was designed by Dr. A. J. Wood of the Department of Animal Husbandry, for the express purpose of providing a complete d i e t for deer. (The composition of this r a t i o n i s recorded i n the appendix.) Use of this r a t i o n during the present experiments and during the experiments conducted by Cowan and Wood (195U), proved i t to be adequate i n a l l respects as healthy conditions and rapid growth were maintained. The fat-added milk r a t i o n proved to be inadequate In some respects. The abnormally high f a t content (15.8$) plus the high l e v e l of feeding f o r the high plane fawns resu l t e d In blockage of the p y l o r i c sphincter and possibly a reduction i n the amount of milk the fawns were able to digest. Blockage and inflammation of the p y l o r i c sphincter were evident i n two fawns that died. Frequently, the remaining high plane fawns appeared bloated and l o s t t h e i r appetities. Massage of the stomach r e l i e v e d t h i s con-20. d i t i o n by f o r c i n g the serai-coagulated milk through the sphincter. Symptoms of magnesium deficiency arose during the course of the suckling period. Fresh cow's milk i s not d e f i c i e n t i n magnesium but concentration of the milk by evaporation reduced the concentration of magnesium i n solution to a c r i t i c a l l e v e l due to i t s low s o l u b i l i t y . Vitamin B def i c i e n c i e s were also encountered during milk feeding. However, these d e f i c i e n c i e s arose p r i m a r i l y from a pro-longed development of the rumen since these vitamins are synthe-sized by rumen microorganisms. ( c ) . Actual food consumption. 1. The milk r a t i o n . Actual food consumption of the milk r a t i o n by J group fawns fluctuated markedly between 2k-hour periods. Figures £ and 6 I l l u s t r a t e the c a l o r i c intake f o r deer 5J and 6 j . The f l u c t u a -tions may be attributed to d i f f i c u l t i e s In digestion a r i s i n g from an abnormally high f a t content of the r a t i o n and/or to force feeding a plane of n u t r i t i o n which was too high f o r early post-n a t a l development. Comparisons of the c a l o r i c intake f o r £j and 6 J show that the fluctuations of intake were greater and more frequent i n the food consumed by 5J (high plane). This tends to suggest that the plane of n u t r i t i o n was too high and, as previously stated, the equation for the c a l c u l a t i o n of intake requires r e v i s i o n . How-ever, minor fluctuations i n the food consumed by the low plane deer indicates minor digestive disturbances due to the r a t i o n i t s e l f . 21. Addition of a milk replacement (Nurse Cow) to the milk r a t i o n , i n an e f f o r t to promote weaning, resulted i n greater fluctuations as shown by the data f o r 6 j . Similar fluctuations r e s u l t i n g from feeding Nurse Cow occurred on the high plane group, but the previously f l u c t u a t i n g consumption of food abscures the fluctuations due to feeding Nurse Cow. Accordingly, Nurse Cow was discontinued when i t was found to be unpalatable. 2. U.B.C. r a t i o n #15-52. Fluctuations were markedly decreased i n the d a i l y food intake of the high plane J group fawns and eliminated e n t i r e l y from the intake of the low plane fawns of t h i s group when weaning was accomplished. Continuation of the fluctuations i n the high plane group tends to suggest that the feeding standard f o r U.B.C. r a t i o n #15-52 was too high for the immediate post-weaning period. Calculations of the standard employing 0.68 or l e s s for the power of "b" may have resulted i n a more consistant intake f o r this phase of growth. Similar fluctuations were encountered i n the food consum-ption of the H group during the post-weaning period. However, at the onset of spring, food consumption became more constant i n both high and low plane deer. The absence of marked fluctuations of the d a i l y food consumption for H group tends to suggest that the high plane of n u t r i t i o n was not too high. Thus 70.5 x W ° ' 7 3 x 5 Metabolizable energy of the r a t i o n may accurately represent the maximal food consumption during t h i s phase of growth. Increasing the d a i l y rations f o r the high plane H group deer above the high plane standard showed a close agree-2 2 . ment between the actual maximal food consumption and the theoret-i c a l maximal food consumption during t h i s phase of growth. Voluntary reduction i n food consumption occurred i n l a t e summer and early f a l l In a l l H deer except 11H. Deer number 3H, a high plane male, reduced his food consumption from k.9 lbs/day to approximately 3«0 l b s . from the middle of August to the end of the experiment on December k. Reduction i n food consumption for this animal coincided with loss of velvet from the antlers and the onset of reproductive behaviour. Low plane deer 1H and kH did not show s i m i l a r reductions of food consumption at t h i s time. However, changing the low plane deer to high plane on September 1 permitted rapid growth and sexual a c t i v i t y was beginning to be-come evident i n late November. Coincidentally, food consumption was v o l u n t a r i l y reduced i n these two animals by November 2$. This tends to suggest that sexual a c t i v i t y i s accompanied by a voluntary reduction i n food consumption i n the male deer. I t i s also apparent that the low plane of n u t r i t i o n may have prevented or delayed sexual a c t i v i t y had the low plane been continued. Changing the low plane males to high plane allowed sexual a c t i v i t y to become evident at a much l a t e r date than that shown by 3H which was on a continuous high plane. In both cases the Inception of sexual a c t i v i t y was accompanied by a voluntary reduction i n food consumption. Deer 10H, a high plane female, v o l u n t a r i l y reduced i t s food intake i n late November (about November 28) but 11H d i d not. The reduced food consumption of 10H coincides with the season of sexual a c t i v i t y i n the females. Deer 11H did not reach this 2 3 . stage due probably to the previous low plane of n u t r i t i o n . Further study i s necessary In order to evaluate the r o l e of n u t r i t i o n i n r e l a t i o n to sexual a c t i v i t y i n the Columbian b l a c k - t a i l e d deer. The present data indicates that a low plane of n u t r i t i o n may delay sexual a c t i v i t y or may prevent i t e n t i r e l y i n both males and females. This agrees with the work of Lutwak-Mann, 1 9 5 1 , who found n u t r i t i o n to influence the production, re-lease and s t a b i l i t y of hormones. However, i t i s also apparent that during the active r u t period there i s a voluntary reduction of the food consumption of males which may l a s t f o r the ent i r e r u t period. A d d i t i o n a l l y one female also showed a reduction i n food consumption f o r a short period of time, probably coincidental with phases of the oestrus c y c l e . (d). Success of the planes of n u t r i t i o n . The two planes of n u t r i t i o n r e s u l t e d i n d i f f e r e n t i a l growth rates between the two planes i n both H and J groups. How-ever, differences i n growth rate are smaller between the two planes of n u t r i t i o n In J group during the suckling p eriod. Planes of n u t r i t i o n were i n i t i a t e d on November 17/53 i n H group, at an average age of 151+ days. Previously these deer may be considered to have been on a high plane of n u t r i t i o n as they were fed on an ad l i b d i e t . Thus the growth rates shown fo r these deer on t h e i r age—body weight graphs, up to 151+ days, represents r e l a t i v e l y higher rates than would have resulted from either the high or low planes of n u t r i t i o n delineated by the feeding standards. L i t t l e difference i n growth responses between the two 2k. • planes of n u t r i t i o n was evident between November 17/53 and. May l / 5 k . The high plane deer consumed s l i g h t l y l e s s food than was indicated by the prepared standards f o r r a t i o n #15-52, thus a t t r i b u t i n g to the f a c t that the standard was too high f o r this period of growth as suggested previously. Af t e r May l / 5 k , the high plane H group showed a marked increase i n growth compared to that of the low plane (see Figure 2 1 ) . Thus i t appears that p r i o r to t h i s time the high plane deer were incapable of expressing an i n -creased growth response over that of the low plane even though the plane of n u t r i t i o n was adequate. This tends to indicate that growth was depressed during the winter as has been shown for c a t t l e (Joubert, 1951+) • I t i s suggested that the food consumed during the winter period may be u t i l i z e d f o r the maintenance of tissue and f o r thermal adjustments to lowered environmental tem-peratures at the expense of growth. However, a reduced food con-sumption, coincident with sexual a c t i v i t y , has been demonstrated, which may also act to depress growth. The planes of n u t r i t i o n during the suckling period i n J group did not produce a marked difference i n growth response (See figure 21). Fluctuations i n the d a i l y food intake of the high plane deer resulted i n an o v e r a l l lowering e f f e c t of the actual plane of n u t r i t i o n . However, high plane deer i n t h i s group tended vto show a s l i g h t l y higher growth response than did the low plane. A d d i t i o n a l l y , average growth rates during growth phase K-^  were 0.0187 f o r the high plane and 0.0132 f o r the low plane (see Table I I ) . Thus, the planes of n u t r i t i o n during the suckling period d i d not p a r a l l e l the difference which should r e s u l t from 25 the feeding standard but did r e f l e c t s l i g h t differences i n growth ra t e . Consumption of r a t i o n #15-52 by J group subsequent to weaning produced s a t i s f a c t o r y differences on the growth responses to the two planes of nutriton (see Figure 21). (e). Relationships of i n d i v i d u a l growth curves to the planes  of n u t r i t i o n . Numerous d e f i n i t i o n s of growth can be found i n the l i t e r a -ture yet few include a l l the ramifications of growth. The most simple and perhaps the most complete d e f i n i t i o n of growth i s " b i o l o g i c a l synthesis." This includes c e l l m u l t i p l i c a t i o n , en-largement and the incorporation of environmental materials. Growth, i n terras of the whole animal, during post-natal develop-ment, p r i m a r i l y r e s u l t s from c e l l enlargement and the incorpora-t i o n of materials taken from the environment. However, c e l l m u l t i p l i c a t i o n also occurs i n the production of blood c e l l s and ectoderm c e l l s . Thus "growth" has been used i n these experiments to represent an increase i n a l l three types of synthesis. Total body weight has been used as an index of growth throughout.these experiments. Loss i n body weight can occur which apparently contradicts the conception that growth i s i r r e -versible.. However, since loss i n weight r e s u l t s from changes i n the environment of a fortuitous nature, minor changes i n body weight maybe considered as mere f l u c t u a t i o n s . Accordingly, body weight may be used operationally to q u a n t i t a t i v e l y measure growth. Figures 7 - l k are examples of the growth curves of i n d i v i d u a l deer of both J and H groups. The data were smoothed Ill KiHiniiiiiiiiiiiiiiiiiiiiii 'HHP • I!!! • » jjii _______ 1IIIL __Er _^__*______c__B . •___________•_______•_§ in in II mm ••• •• • • • • H i l l I • i r r i i i i - ' i i i i - ' r i i _ IIIIIIIIIIIIIUIII m I I I " M I H " I I | T I H I I I H I I I l l l l l l l l l I I I 111 I__ ^taiiWIiiliiiliiiiiiliiilliiliiliiliiiiiilliiiii WW in ii! • iiiii isilfiiiiHiili! - ! \ \ -nnasssTK::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::: K:::::::n:::::::::::::;::::::::::::::::::::::::::::::::::::::::::: :: :::::::::::::::::::::::::::::; M!!l!!!l!!ii!ii!iil[l!!l!!i!!!!jl!!l ••Hj^ss'^M^r:"' lilliliilllilllllllilllilil lliliii illliilii iiHRHISl S S S s g i i a : iia£8 - S I ssss:s.ss-_ I H Slffiiil^iiS^^Siiiiiill l i B i i i i i i i i i i n i y i iiiii i l l l i i l i i Hlimlliliili!..... !::::.:::::::::::::::::::::: •mmi!:::iiii!!!!i!:iii!!:!:!!!::!!:i •••••••••••••III It..1111.. Ill l l l l l l l l l l l l l l l l l l - - • ( = L—r: = rf ~=irn-;z- = = - = - : b i l i l l i l l ======i===-??====iililli —;====s|||||||||||| ^ L y S a a S ^ S M i M H n S i i i i i M i H N u l E l " ! piiijjsiipiaiann • m i l l J I I I M I I I I I I I I I I I I I I m i i m i l l lllllllll! III! §iiyi!ii!illlilj B ' l ' i ' f l l I B i l l 2 6 . by the method of l e a s t squares and growth rates were calculated f o r each phase of l i n e a r i t y (Brody, 19k£). . Five primary phases of l i n e a r i t y are evident from b i r t h to one year of age when weight i s p l o t t e d against age on an a r i t h -log g r i d . Four of these phases are shown on Figures 7 - l k . A phase of growth between b i r t h and the s t a r t of phase K has been reported by Cowan and Wood, 195k. This early phase (designated K_ i n Table III) l a s t s f o r approximately two to three days and represents a growth rate i n the order of 8-9$. Since J group fawns were older than three days at the s t a r t of the experiment, phase K_ i s absent from the data f o r t h i s group. S i m i l a r l y this data i s missing f o r 3H, kH and 10H. Phase K was not treated separately i n the case of the doe-raised fawns 1H, 2H and 11H. However, data points on Figures 12 and l k show thi s rapid growth to be present during the f i r s t two to three days. The second phase of l i n e a r i t y (K) i n the growth curves l a s t s for approximately 10-12 days. Phase K and K represent the periods of most rapid growth due to the r e l a t i v e absence of growth I n h i b i t i n g forces. Reasons for the change i n rate between these two phases of growth are not known. However, true homeothermy i s not accomplished f o r the f i r s t few days of l i f e i n many mam-mals and i t i s therefore suggested, that t h i s f i r s t break i n the growth curve represents a metabolic change associated with the Inception of true homeothermy. Growth rates again decreased between 12-18 days of age to a t h i r d phase of l i n e a r i t y (K^). The break between K and K-^  phases of growth coincided with the change to fat-added milk i n 27. the d i e t of J group. However, t h i s break also occurred i n doe-r a i s e d fawns 1H, 2H and 3H, and therefore cannot be a t t r i b u t e d to changes In the d i e t . Temperature records, kept f o r noting deviations from normal, show an increase from approximately 101.0 to 102.0°F during this period. Thus, the break i n growth rates between K and i s thought to represent further p h y s i o l o g i c a l changes associated with thermoregulation. A d d i t i o n a l l y the change i n growth rate may be associated with an anatomical l i m i t a t i o n to further Increments of food intake. P r i o r to t h i s point I t i s suggested that the consumption of food was less than the anatomi-c a l capacity of the digestive system. Since the. r e l a t i v e increase i n the development of v i s c e r a l organs i s less than that of the body as a whole (Brody, 19kf>), successive increments i n consump-tio n must be l i m i t e d by the slower development of the v i s c e r a l organs. An increased d i s a b i l i t y of the high-plane J group to con-sume the calculated d a i l y food r a t i o n during this period tends to substantiate t h i s suggestion. The rate of growth was again retarded between phases K-^  and Kg. Figures 11, 12 and l k , f o r doe-raised fawns, show thi s break i n the growth curve to occur concurrently with the period of weaning. However, Figures 7, 8, and 10 show a phase of growth K^ & not represented i n the curves of doe-raised fawns. Differences between the two groups l i e s i n the prolongation of the suckling period i n J group due to aseptic conditions and to the preventing of consumption of small amounts of s o l i d food dur-ing the suckling period. Thus under more natural conditions phase K would not occur i n the growth curves f o r J group and 2 8 phase K^ would change d i r e c t l y into phase Kg. This tends to sug-gest that the change i n growth r a t e s at t h i s point i s related to development of the rumen for the b a c t e r i a l fermentation of n u t r i -ents associated with the period of weaning. Maintenance of r e l a t i v e l y high glucose l e v e l s during phase K^ft i n the J group tends to substantiate t h i s point as w i l l be shown l a t e r . Figures 11, 12, 13, and l k show the next change i n growth rates between phases K 2 and K^ to occur concurrently with the commencement of planes of n u t r i t i o n . However, Figure 12 represent-ing the growth curve of 2H shows that a break i n the growth rate occurred p r i o r to the commencement of planes of n u t r i t i o n . Thus, thi s change i n growth rate i s thought to represent the pubertal i n f l e c t i o n i n growth which i s obscured i n the remaining H group through a p p l i c a t i o n of planes of n u t r i t i o n . Figures 7t 9 and 10 show a si m i l a r break i n the growth curve which occurs at approxi-mately the same age of 1^0 days. Discovery of one r i p e ovarian f o l l i c l e i n l£j shortly afterwards, substantiates the f a c t that puberty had been reached, at l e a s t i n one animal. Behaviour of some of the high plane, J group males indicated that many, i f not a l l , had reached puberty. Thus the change i n growth rates between phases Kg and K^ may be a t t r i b u t e d to metabolic changes associated with puberty. Additional changes i n growth rates occurred through man-ip u l a t i o n of the planes of n u t r i t i o n . Growth rates were increased from phase K^ to K^ i n both high and low plane deer of H group. Change i n rate of growth of the high plane group, with the excep-tion of 2H, resulted from feeding above the standard i n an e f f o r t 29. to determine the maximal food consumption. Increased growth i n the low plane deer resulted from a change to the high plane of n u t r i t i o n i n order to determine whether they were s t i l l capable of an increase i n growth rate i f l i m i t a t i o n s of food supply were removed. Accordingly, an increase i n the rate of growth i s apparent i n both groups. Deer 2H died p r i o r to changes i n n u t r i t i o n (Figure 12), but an increase i n growth was recorded from phase to during the l a t t e r part of May, 195k. This increase i n growth i s thought to be a release from a growth depression due to environmental conditions associated with winter or sexual a c t i v i t y , as was previously suggested (see Figure 2 k ) . The low plane deer 1H, kH and 11H were not able to express ah increase i n growth a t this time due to the low standard of feeding. Individual growth curves f o r 3H and 10H do show small increases i n the rate of growth dur-ing early spring similar to that shown by 2H. Decreased growth rates were noted for a l l deer i n H group i n l a t e f a l l of 195k (approximately November 2 0 ) . Voluntary re-duction i n food consumption shown previously to be associated with the inception of sexual a c t i v i t y may be d i r e c t l y responsible for the decreased growth. However, since decrease i n food consump-t i o n proceeded decrease i n growth rate i n the high plane H group, an addi t i o n a l metabolic change i s indicated which may be associated with lowered environmental temperatures. ( f ) • Instantaneous r e l a t i v e growth rates and the planes of  n u t r i t i o n . The instantaneous r e l a t i v e growth rate (k) during the 30, s e l f - a c c e l e r a t i n g phase can be expressed by the formula dW/dt W where "W" represents the weight i n kilograms, at time ttt". Inte-gration of this formula provides a method f o r the mathematical evaluation of instantaneous r e l a t i v e growth. Thus In W? - In W, 1 1 (1) for the s e l f accelerating phase of growth (Brody, 19kf?). Similar-i l y , -k = dW/dt (2) A-W for the s e l f - i n h i b i t i n g phase of growth where "A" represents the f i n a l mature weight and nW" i s the weight at time " t . " Formula (1) was used f o r the c a l c u l a t i o n of a l l k-values i n the analyses of growth data as an accurate value f o r "A" was not known. In addition, estimations of "A" by graphic methods did not r e s u l t i n a reasonable f i g u r e . Accordingly formula (1) was used for both the s e l f - a c c e l e r a t i n g and s e l f - i n h i b i t i n g phases of growth. Data thus calculated i s valuable i n the comparison of indivi d u a l s although i t does not represent the true values for the l a t e r phase. Table II shows the instantaneous r e l a t i v e growth rate for a l l experimental deer. Planes of n u t r i t i o n were not imposed on H deer u n t i l the beginning of phase and on J group u n t i l the beginning of phase K-^ . Consequently, k-values for phase K may be compared i n order to i l l u s t r a t e differences between doe-raised and "bottle-raised fawns. Doe-raised fawns 1H, 2H and 11H demonstrate an instantan-eous r e l a t i v e growth rate of approximately 6% compared to 3$ for 31 TABLE II THE INSTANTANEOUS RELATIVE GROWTH RATES IN RELATION TO THE PLANE OP NUTRITION Animal k k l k 2 , k 3 Sex Plane of n u t r i t i o n 1H 0.06k2 0.0175 0.0079 0.0003 Low plane k& - - 0.0058 0.0008 0" Low plane 11H 0.0573 0.01k2 0.0035* 0.0009 9 Low plane Average 0.0607 0.0158 0.0057 0.0007 2H 0.06kl O.Olkk 0.0013* Q.OOI3 0* High plane 3 H - - 0.0075 0.0022 0" High plane 10H - - 0.0052 0 .0023 9 High plane Average 0.06kl O.Olkk . 0.00k7 0.0019 1J * O.OkOl 0.0220 - mm 0* High plane 3J 0.0kk7 0.015k 0.0067 0.0087 0* High plane 5 J 0.0656 0.0206 0.0118 0.006k a* High plane 7J 0.02k3 0.0182 - - 0* High plane 9J 0.0323 0.0216 - - High plane 15J 0.0259 O.Olkk 0.0097 0.006k 9 High plane 17J 0.02k5 - - - 9 High plane Average 0.0353 0.0187 0.009k • 0.0072 2J 0.0370 o . o i k 5 0.0123 - 0* Low plane k j " 0.0319 0.0115 0.0006 0" Low plane 6 j 0.0351 0.0132 0.011k 0.0015 o" Low plane 10 J 0.0232 0.013k 0.0155 0.0013 0* Low plane 18J - 0.0152 0.0125 0.0008 0" Low plane IkJ 0.035k 0.0107 O.OOkk 0.0008 9 Low plane 32 TABLE II - Continued Animal k k l k 2 k 3 Sex Plane of n u t r i t i o n 16J 0.0k21. 0.0137 - - 9 Low plane Average 0 . 0 3 k l 0.0132 0.0095 0.0011 Low growth rates probably due to the deer being very excitable and nervous animals Note 1. Values may be read as a percentage growth rate by multiplying the above figures by 100. Note 2. The double l i n e Indicates the approximate s t a r t of the planes of n u t r i t i o n . the b o t t l e - r a i s e d fawns. Lower growth rates f o r J group during t h i s phase may have resu l t e d from shock associated with capture and new environmental conditions. A d d i t i o n a l l y , slower growth may have r e f l e c t e d poor n u t r i t i v e conditions i n the wild does dur-ing pregnancy compared to the captive does or may have been due to an i n f e r i o r n u t r i t i v e condition i n iiie fawns r e s u l t i n g from the a r t i f i c i a l milk d i e t . Degrees of success i n adaptation to the new conditions resulted i n i n d i v i d u a l v a r i a t i o n s of growth rates during t h i s phase. In general, low plane animals i n both groups r e f l e c t e d comparatively lower instantaneous r e l a t i v e growth rates. However, k 2 values f o r J group show no difference between the two planes of n u t r i t i o n . This was due to the loss of control of cabric In--take during the weaning period. Ration #15-52 and grass were placed i n the pens during the l a t t e r part of the suckling period i n order to i n i t i a t e consumption of s o l i d foods. Milk r a t i o n s were reduced as p e l l e t s were eaten i n increasing quantities with-33. out regard f o r the planes of n u t r i t i o n . Thus, control of c a l o r i e intake was l o s t during t h i s period. Instantaneous r e l a t i v e growth rates were higher i n J group than i n H group during the K 2 phase. This i s believed due to a recovery type e f f e c t r e s u l t -ing from l i b e r a l food conditions associated with the method of weaning. Thus, given the opportunity, absolute growth l o s t through slow growth rates during phase K and K-^  was p a r t i a l l y recovered during phase Kg. The values of K^ show that the high plane H deer showed an I.R.G.R. almost three times higher than the low plane group while i n J group the difference i s almost 7 times higher f o r the high plane. A d d i t i o n a l l y , growth of the low-plane J group was more rapid than growth of the low-plane H group. This tends to suggest that the planes of n u t r i t i o n were r e l a t i v e l y higher i n J group than i n H group i n spite o f the fact that the same stand-ard and ration was used i n both eases. Thus, the feeding stand-ard f o r this period of growth was too high, as was suggested previously. In order to reduce the rate of growth i n the low plane J group the power of "b n should have been less than 0.73* Comparisons of Tables II and I I I show that e a r l y growth rates found i n these experiments are similar to those found by Cowan and Wood (195>k). However, the f i r s t short period of very ra p i d growth (K_) was omitted during the present experiments. Values of k and are intermediate b&tween the values of J and H groups suggesting that the milk r a t i o n used f o r J group was not greatly i n f e r i o r to does milk. Thus low values f o r k i n J group may be attributed to the shock of changes i n environment. 3 U . TABLE I I I INSTANTANEOUS RELATIVE GROWTH RATES FOR COLUMBIAN BLACK-TAILED DEER (COWAN AND WOOD, 1951+) Deer k a k k ^ Sex 1 0.0799 0.0300 0.0109 Twins 9 2 - 0.0532 0.0150 It d 3 - 0.01+95 0.0150 Twins d k - 0.01+95 0.01^2 n d 5 - 0.0L70 0.0108 Single d 6 0.1350 0.100** 0.0150 Single J 7 0.0980 0.0500 0.Q150 Twins d 8 0.0910 0.05^0 0.0137 n d Average 0.1010 . O.Oij.17 0.0137 The data was col l e c t e d at f a i r l y wide i n t e r v a l s during the l a t t e r stages of the growth. Thus the weaning break apparent i n the present study does not occur i n the data of Cowan et a l . Following the K phase of growth with a k value of 0.1350, there was a markea decline i n weight. Thus, the high value f o r k (0.100) may be the r e s u l t of recovery from a previous slump. Note: The above k values were rearranged i n order to compare growth rates with those of the present experiments. (g). Individual age relationships of l i n e a r growth. Growth, i n terms of mass, i s associated with increases i n M n M dimensions. Since the proportion of each dimension remains more or less constant throughout growth of the animal as a whole, rates of growth must vary f o r d i f f e r e n t l i n e a r dimensions. How-ever, c e r t a i n l i n e a r dimensions are a f f e c t e d to a greater or 35. smaller degree by growth promoting or retarding f a c t o r s . This provides a means of measuring the e f f e c t of various factors on growth. Linear growth tends to occur In an arithmetic progression and to p l o t l i n e a r l y on arithmetic paper. Thus, i t may be expres-sed by the formula Y = a + bX where n a K i s the Y-Intercept, and "b M i s the slope of the l i n e or rate of growth. "YM equals body mass and "Xw represents a l i n e a r measurement (Brody, 1945)• Heart-girth, height-at-withers and length of hind foot were used f o r the study of the growth i n terms of l i n e a r dimen-sions. Figures 15-20 represent data from these measurements for fawns 5 J and 6j as t y p i c a l examples for the whole group. A l i n e of best f i t was determined by the method of l e a s t squares, assum-ing that growth was l i n e a r f o r the e n t i r e period. Heart-girth shows the most r a p i d growth r a t e , increasing at a rate of approximately 16$. Height-at-withers and hind-leg length grew less r a p i d l y at 13$ and 6.5$ r e s p e c t i v e l y . The l a t t e r represents the growth r a t e of purely s k e l e t a l elements while h e a r t - g i r t h and height-at-withers include the development of a d d i t i o n a l tissues such as muscle and f a t . Growth of a l l these dimensions i s affected by the plane of n u t r i t i o n to a greater or l e s s e r degree. Table IV shows the re l a t i o n s h i p of the growth of three l i n e a r measurements with respect to the plane of n u t r i t i o n . Heart-girth, since i t has the most rap i d rate of growth, i s affected by the n u t r i t i o n a l l e v e l to the greatest degree. Height-at-withers i s also affected but to a le s s e r degree, compatible with a slower r a t e of growth. The ~ 40 t 20 « X Age in days (T) G r o w t h in h e a r t - g i r t h f o r 5 J 100 120 Age in days (T) F IG . 16 G r o w t h in h e a r t — g i r t h f o r 6 J 20 40 60 80 100 12 Age in days ( T ) • 17 G r o w t h in h e i a h t - a t - w i t h e r s of 5 J !0 140 160 6 0 2 FIG. 18 0 40 60 80 100 120 140 160 Age in days ( T ) G r o w t h i n h e i g h t - a t - w i t h e r s o f 6 J 60 • • 50 cms. ( 40 ind foot in 30 0 - 0 6 5 4 T + 23 *8 • CO o o> e OP 20 • — * 1 s m _ l 10 2 0 4< 3 6 Age 0 8 in days 1 0 IC T) )0 12 0 1A 0 16 .0 IE 10 FIG. 2 0 G r o w t h in l e n g t h o f h i n d f o o t o f 6 J 36. length of the hind foot is affected only slightly and comparative-ly, the length of the hind foot may be considered to be relatively unaffected by the plane of nutrition. TABLE IV ABSOLUTE LINEAR GROWTH RATES IN RELATION TO. THE PLANE OP NUTRITION Absolute Growth Rate i n Percent Linear Measurement High plane (5 J) Low plane (6j) Difference Chest g i r t h 17.08$ 15.59$ 2.49$ Height-at-withers 14.28$ 12.73$ 1.55$ Length hind foot 7.14$ 6.54$ 0.60$ Instantaneous relative growth rates, In terras of weight were previously shown to differ between the two planes of nutri-tion. However, the differences were mathematically small in comparison to differences between the planes of nutrition In terms of absolute, growth. Figure 21 shows the weights for a l l deer in relation to the two planes of nutrition. For example, one high plane male H deer weighed approximately 55 kgms at 410 days of age whereas the corresponding low plane male weighed approximately 29 Kgms at 412 days of age. This represents a dif-ference of 50$ whereas difference in weight at the beginning of the experiment was i n the order of 28$. Figure 22 shows that heart-girth is also materially affected by the plane of nutrition. Comparisons of Figures 21 and 22, which are drawn to the same scale, show that the magnitude of difference between absolute growth of heart-girth is less than that of weight. Since the growth of weight and heart-girth show FIG. 22 Growth of heart -g i r th in relation to age and the plane of nutrition of all experimental deer S t a r t o f n u t r i t i o n a l I p l a n e s f o r H - g r o u p I I * I l « . * >„ i S t o r t o f n u t r i t i o n a l p l a n e s f o r J - g r o u p E n d o f m i l k f e e d i n g • H i g h p l a n e J - g r o u p m a l e s o L o w p l a n e J - g r o u p m a l e s x H i g h p l a n e H - g r o u p m a l e s a L o w p l a n e H - g r o u p m a l e s A H - g r o u p f e m o l e s L o w p l a n e p l o c e d on h igh p l a n e FIG. 23 Growth of height-at-withers in relation to age and the plane of nutrition of all experimental deer S t a r t o f n u t r i t i o n a l p l o n e * f o r J - g r o u p S t a r t of n u t r i t i o n a l I p l a n e f o r H - g r o u p E n d o f m i l k f e e d i n g "o • x" • * | • H i g h p l a n e J - g r o u p m o l e s 0 L o w p l a n e J - g r o u p m a l e s x H i g h p l a n e H - g r o u p m a l e s 0 L o w p l a n e H - g r o u p m a l e s • H - g r o u p f e m a l e s A J - g r o u p f e m a l e s L o w p l a n e p l o c e d on h i g h p l a n e c FIG. 24 Growth of the lengh of hind foot in relation to age and the plane of nutrition of all experimental deer I I S t a r t o f n u t r i t i o n a l p l a n e t f o r J - g r o u p S t a r t of n u t r i t i o n o l p l a n e t f o r H - g r o u p E n d o f m i l k f e e d i n g ® . S> ® 0 -L o w p l a n e p l a c e d o n h igh p l a n e • H i g h a n d low p lane J - g r o u p ) x H i g h a low p l a n e H g r o u p m a l e t ® H i g h 8 low p l a n e H - g r o u p f e m a l e * O 4 0 8 0 1 2 0 1 6 0 2 0 0 2 4 0 2 8 0 3 2 0 3 6 0 4 0 0 4 4 0 4 8 0 A g e i n d a y t 37. similar responses to the planes of n u t r i t i o n i t should be possible to u t i l i z e h e a r t - g i r t h to estimate body weight. Height-at-withers reacts s i m i l a r l y to h e a r t - g i r t h with respect to the plane of n u t r i t i o n but to a smaller degree (F i g -ure 23), and hind-leg length (Figure 21+) r e f l e c t s the plane of n u t r i t i o n l e a s t of the three l i n e a r measurements. Since the rate of growth decreases i n the same order as does the degree to which n u t r i t i o n a f f e c t s the measurement, i t i s apparent that f a c t o r s r e s t r i c t i n g growth tend to affect the dimensions with the most rap i d rate of growth to the greatest degree. Accordingly, weight i s most s e r i o u s l y a l t e r e d by the planes of n u t r i t i o n while length of the hind foot i s l e a s t affected. As weight can vary to a much greater degree than can hind-foot length, a wide range of weights may be represented by the same length of hind f o o t . For t h i s reason, the l a t t e r measurement i s u s e f u l i n analyzing the condi-t i o n of the deer. Comparisons of the growth of these dimensions already achieved at b i r t h tend to substantiate use of h e a r t - g i r t h f o r weight estimation and hind foot length for the analysis of condi-t i o n . Table V shows that c a t t l e are r e l a t i v e l y more mature at b i r t h than are deer by v i r t u e of the development achieved at b i r t h . However, i t i s evident i n both c a t t l e and deer that weight i s the least developed at b i r t h and that hind foot length i s developed to the greatest extent. I t i s then reasonable to expect that l i m i t a t i o n s of the post-natal environment can most serio u s l y a f f e c t those measurements which have the greatest development yet 38 TABLE V THE APPROXIMATE PERCENTAGE OP THE ADULT MEASUREMENTS ACHIEVED AT BIRTH Cattle Deer Deer Measurement (Brody, 191+5) (250 lbs) (100 lbs) Weight 4 8 3 6 Heart-girth 1+1+ 25 27 Height-at-withers 55 1+0 1+2 Length of hind foot 68 (leg length) 5 l 1+5 to achieve. Conversely, growth-limiting fac t o r s can only s l i g h t l y a f f e c t those measurements which have completed a large proportion of their f i n a l s i z e at p a r t u r i t i o n . The computed values f o r deer show a s i m i l a r r e l a t i o n s h i p between the measurements of growth to those reported f o r c a t t l e by Brody. Differences between c a t t l e and the Columbian b l a c k - t a i l e d deer indicate that height-at-withers has completed a smaller percentage of i t s f i n a l development at b i r t h i n the deer. Thus, where Brody uses height-at-withers as an index f o r assessing the plane of n u t r i t i o n i n c a t t l e , t h i s measurement would not provide accurate r e s u l t s i n the deer. Length of hind foot i n deer has completed a comparable percentage of i t s f i n a l development i n deer to that of height-at-withers i n c a t t l e . Accordingly length of hind f o o t should provide an accurate index of condition for the deer. Differences i n the above responses to the planes of n u t r i t i o n may be explained by reference to Figure 25. The d i a -gram shows that bone has a higher p r i o r i t y f o r blood nutrients than have either muscle or f a t . I f the supply of nutrients i s 39. decreased one arrow may be removed from each t i s s u e . Thus, f o r one u n i t of nutrient decrease, there w i l l be a 100$ decrease on the nutrients going to f a t , $0% to muscle, 33$ to bone and 20% to the c e n t r al nervous system. Since weight i s p r i n c i p a l l y . t h e r e s u l t of muscle and f a t accumulations, a decrease on the plane of n u t r i t i o n w i l l greatly a f f e c t weight. In contrast, bone i s r e l a t i v e l y unaffected since a unit decrease i n nutrients removes only 33% of the supply to t h i s t i s s u e . Therefore a s k e l e t a l measurement would be r e l a t i v e l y unaffected by n u t r i t i o n compared to measurements invo l v i n g muscle and f a t . Heart-girth represents muscle, f a t and skeleton and there-fore may then be used i n estimating body weight since i t i s affected s i m i l a r l y to body weight by growth-limiting or growth-promoting f a c t o r s . In contrast, hind leg length has been shown to change r e l a t i v e l y l i t t l e i n growth rate as a r e s u l t of a low plane of n u t r i t i o n . For t h i s reason i t can be used i n measuring condition i n the Columbian b l a c k - t a i l e d deer. (h). Relationships of l i n e a r growth to body weight. In animal production i t i s important to know the l i v e weight of an animal f o r c a l c u l a t i n g food requirements, energetic e f f i c i e n c i e s and other functions and f o r measuring growth. Ani-mal producers do not always have weighing scales to accurately determine l i v e weight. Thus, a method which i s suitable f o r estimating body weight without scales i s advantageous. Investiga-tors i n w i l d l i f e management have c o l l e c t e d semi-dressed weights at numerous checking s t a t i o n s . However, the weight thus c o l l e c t e d i s of l i t t l e value unless i t can be corrected to a l i v e weight b a s i s . I t i s with these uses i n mind that estimation of l i v e weight by the use of body measurements was explored. Figure 26 shows the regression of weight and h e a r t - g i r t h . Since a l l the data from high and low plane deer are p l o t t e d on this graph, i t i s apparent that a s a t i s f a c t o r y c o r r e l a t i o n exists between weight and hea r t - g i r t h , regardless of the plane of n u t r i -t i o n . Thus, this regression may be used for the graphic estima-t i o n of body weights. P l o t t i n g a l i n e a r measurement such as h e a r t - g i r t h , against body weight on arithmetic-coordinate paper r e s u l t s i n a parabola (Brody, 19i+5)« but when plot t e d on log-log paper a st r a i g h t l i n e i s achieved. The regression may thus be represented by the :=!===: =====: H I H H L f i S i i i i i i i i l l l i i • • • • • • • < :::::::::::: i i i i i i i i i i i i i i l l l i " i u i i i i i i i i i i i i i i i l i I H I I i l i m i m i i R i i =======: •========== • • MM MM :::::: BE Ear •R4rw<B|H • mn mn i !!!!!!!!!•!! :====: iiiiiiHiiii • PI j i i i i i i i i i i i i i i i i i i iaiBiaii I riwl UVIIIII I «¥£» l i ! ! ! ! ! ! ! ! __555i_ i l f S i m i g i i i ^ B H I IliillllHIII • • • • • • • I I A i l • • tmm jf <*«. •• _ ir« i i ii •• • •• »• Rffl Wffi r.M i :: • i II l l l l iiii iiii HE "I iiii IIIII •Hit: •••• II —•• II iiii mi i • II mi " "1! 1 HI ::::::••::•:•::!======= I I I I I H I M I I I I I I I I M H H M i a i H H H • III III.— »l HIM — — — — • — —» —• — — l l l l l l l l l i i i i i i i i i aaia>fl i iBMii i i i i iMi • ••••••••••IIIIIIIII ••••••••••••IIIIIIIII imam ill:::':': •••••i •!•»•••• ••••••in ••••••••• ••••••III •••••••II • l l l l l l l l I ••••IIIII I lllllllll IIIIIIIII •••uiaiMi mi II m i l m i • • • B i i i n a i •••Hilar— s s s s s s s : •• • • • • • • • • a • • • • • • • • •I llll I i i i i Ii I i i i i I T S : - -III III III ••PI I'ii'i 11 HI l l l l l l l l l l l l l l IIIIIIHIIII IIIIIII II l l l l IIIIIIHIIII jiiiiiiiiiii .1':; nilil,H:iliU: IIIIIIHIIII IIIIII IIII - -1 III l l l l l l l l l l l I IIIIIIHIIII • SSSSSSSSggg::: n a i i a i M a i M « a « * n a i i i H rwrw it . . •Si 5 i J iJI II I I I IIIII III Hill • • • • • III IP I llll IIIII I I I imiii i H i m m HI in. formula log M = a +• log G x b where " M w i s the body weight i n k i l o s , "a" i s the Y-intercept, G Is the l i n e a r measurement, i . e . , heart g i r t h i n cms, and "b" i s the slope of the l i n e . Data represented In Figure 2 6 were smoothed by the method of le a s t squares and the standard error of estimate was computed. The formula f o r the l i n e log M - 1+.5708 + log G 2.61+52 may then be used to estimate the weight of a Columbian b l a c k - t a i l e d deer once the h e a r t - g i r t h i s known, or the weight may be g r a p h i c a l l y determined from Figure 2 6 . The slope of the regression l i n e f o r h e a r t - g i r t h and body weight i s 2 . 6 f o r the deer compared to 2 . 7 5 - 2 . 9 1 f o r d i f f e r e n t breeds of c a t t l e (Brody et a l , 1 9 3 7 ). _The slope f o r deer i s i n the same order of magnitude as that f o r c a t t l e suggesting that i t can be used- i n the estimation of the l i v e weight of deer as i t can i n c a t t l e . Height-at-withers p l o t t e d against body weight produces a slope of approximately 3 . 6 i n deer compared to 1+.25 to 1+.1+3 for several breeds of dairy c a t t l e (Figures 27-29). Thus height-at-withers shows a much steeper slope than does h e a r t - g i r t h when regressed against body weight. Since height-at-withers was l e s s affected by the plane of n u t r i t i o n than was h e a r t - g i r t h , i t seems l i k e l y that the slope of the l i n e would d i f f e r f o r the two planes of n u t r i t i o n . Figures 27 and 2 8 show that differences In slope between the two planes d i d occur. A slope of 3.7I4.5I1 was found f o r the high plane male deer compared to 3.5209 f o r the low plane. On this basis height-at-withers would be unsatisfactory f o r the estimation of body weight* However, d i f f i c u l t i e s i n measuring 5 6 7 8 9 1 J•••!•••!!• • I S m r S ) ' S i m B II I I I I M B I H I i i i iR r i i P i i i i i i i I H i i l p i l i l l l l m u m Ssssssss ssisss"" ====== i i i i i i i i i i i i i i i i IIIIIIII mir •SB =11111 =11111 •SSSiS n u n 1 1 ssss: jsssssss 1SSSS : : : : : : EE: : : : : K H H a i s.;:i! •••in i i imi i i i . nun m i l IIIIIIIIIIIII3IIU <t ••••••••••••IIIIIIII IIIII IIIII IIIII iiiiiii IIIII ilIIiiiiliiiillllilHIIIi! isiIs»iiiiiiii»!iH§ii! Ill t r a i n 5 b 7 8 9 IHI iliiiiiEiaiiiiiiiHieiHyiiBlBiiHHHii^i^HlliniHHl Iiiiiii • S S B S S s s s s s : l i i i i s s E i i i i i i l IIIII IIIII ••••II I I I I I I llllllllllll IIIIIIIIHIIIINIIirjMIIIIII iiiiiiiiimiiiiiaiHH miiiiii'if HlWi'f IfilllfJill lllllliilli!,:!! I H I : : : :-r ' : ! :! :! — J . f J W I jljij- Illijiiiijijjjiijjijjiijlii I : iil!!!!i I i i i i i i i mm iiiiiiii! t r a i l == nsssxssi at",: , ===== 25:3:::::Hif jj »:> :::!S!:;:>m:HS::::::::$ •* I * tm • ai* • fill • III ii: M I f i l m i n J I H mi i i n m -iUMgll l l "~ •••••••IIIIIII ••••IIIIIIIIII III lllllllllllllll ••lllll l l l l l l l llllllllllll IIIIIIIIII E3*IIIIIIK]<]IIIIIU ji!ji!iij!j===l|l| ::::::::!::: : : : : : i : : i : s s B s a iiiii IIIIIIIIIIII ::;:: iiiiiii! liiiii mil _ IMIIIIIHMIIIIIIIIIIIMIIII mi iiiiimiiaii•••mininun mi iimiiiiiiiiiiiiiiiiiii mi iiiiiiiiiiiiiiii imur III! IIIIIIII linn IHI llllllllllllllllllll mi iniiiiiimiiiii iniiiiliMiii iiiiiiiiiimiiiiiiimiiiii mil !!! i l l ! : Sssssssss : s : : : : : : : : : : ! : s s s s 3 s s s : s s : : : : : : : : : i i i i i i l l l l l l l l | i | I H H i inSIMiT* l i l i lHi i i ip i i i i i i i ip igig I IliilBiMS HiiiiHiiiithitiiMi iloliHSIIiilli ••••!!••!!! !Sii!iiS!!iSiil!liliSH •••"!•!"!!!! II iiii i iiiiii ! •iiiitiiiiiiitiniiaii ii ii in II IIIIIIIIII ••• . IIIIIIIIII mil I llllllllllllllllllilH • ••••nun iiiiiini I I I I I I I I I I•••• iiiii • • l l l l l l l l l l l l i i i i i i i i n i i m i i i i i i i i iiiiHHiiiiiiiiiiii i i i n i i i i i i i i i iiiiiiiiiiiii! i i IIIIIIIIII Iliiiiiiiliiil sssiss::: I B • •• mmm mm mm mm mm mm m . i i !••••••• i l l ! iiiiliiNi IHMUi iHN iiiii iiiii IHH iHH====^^ i^iI i i i i i i i i i i " " ••••in 7 8 9 1 3 4 5 fc 7 8 9 1 " 1 1 1 1 -iiiiiii Iiiiiii! IIIIIIIIIIIIIIIIII IIIIIIIIIIIIIIIIIIIIIIIII l l l l l l l l l l l l l l l l l l l l 3 4 5 6 7 8 9 I I I I I I I I I I IIIII I I I I I I I I I am ::::: I IIIII! :::: : : i i : : : : i u n a u r n IIIII •••••IIIIIIII IIIIIIII IIIII I I I I I I I I IIIII iiinmiiHisniiiiiioiiiiiiii b S T B S SSS! j l l l l l l l l l l l l l l l l l l j l l l j j j ! I i ; ' : ! i 3 i ! a i i l l l l l = i l ! i ; i I l l i a i i i i i i i l i i i i I M i l l IIIII IIIII • • • • • • • • • • • • • ;;;::;;::i:::!i==sssssssss:s Hill 4 "IIIHIBiBi H i i i i i i i i l i i i i i i i i i i j i i iy i ============== ::: U l l i i i i i i Mm |niniiniiii=ri^^S==IliiliIIiiiiiiIiini!H ~ i | = l l l i l i r ~ ' llll! H i l i l i i i i ^ i i l i i i i i i i i i i l i i •mm' Iiiii i i i i i i i i i i l l ii llll 7 8 9 1 =111111111 ======:: :::::: ' S S B B S S S : : :::::::: ii'i'i ii •ran •• il II II • I II M i i i H i i i •••iiiiiiti II •••••IIIIIIIII lllllllllllllllllllllll • i r -Blilillllilliiiiiliilliii '••••mi * I I M i i i i i i i i i i i mn — — — — — — — !:!:!!!!!!::I!!!:::!::::!::::!::::::SSBSSSS I I E IIIIIII iiiiiiiiiiiiiiiiiiiiiiimiiiiiiiiiHMf I f f l B 3 4 5 6 7 8 9 1 3 4 5 6 T 8 9 I 42 t h i s l i n e a r dimension resulted i n large standard error of estimate which may In turn have produced greater differences i n slope be-tween the planes of n u t r i t i o n than a c t u a l l y e x i s t . Figures 29 and 30 show that l i t t l e difference a c t u a l l y exists between two n u t r i t i o n a l planes i n terms of height-at-withers when data i s selected so as to reduce the standard error of estimate. Considering a l l male J group to represent low plane* and deer 2H to represent high plane* on the b a s i s of Figure 21, resulted i n s i m i l a r slopes for height-at-withers against body weight f o r both regressions. Thus, 3*6996 represents the slope fo r 2H (high plane*) and 3.6420 represents the slope f o r a l l male J group fawns (low plane ). On this basis height-at-withers may be used s i m i l a r l y to h e a r t - g i r t h i n estimating body weight. Figure 31 represents a l l the data f o r height-at-withers. Very large standard errors of estimate are evident a t t r i b u t i n g to the inaccuracy of using height-at-withers as a measurement f o r e s t i -mating body weight. It.was shown previously that a s k e l e t a l measurement i s less l i k e l y to be affected by the plane of n u t r i t i o n than heart-g i r t h or height-at-withers. Figures 32-35 show the regression of a s k e l e t a l measurement, length of hind f o o t , to body weight. Un-fortunately, t h i s measurement was not taken during early growth of the J group deer. Accordingly, a r e l a t i v e l y large proportion of growth had already been made p r i o r to the commencement of the feeding standards and differences between high and low plane are not as marked as they should be. I f the low plane of n u t r i t i o n affects weight to a greater I l l llll IIIIIIIIII 7 - 8 9 1 5 fc 7 8 9 ) l::;:::Ritiniii i B j g f i i i i i i i i i i l l i l i iiiii i i i i l l l l i m i i i i i i i i i i i I i i i i i i i i i ^ i i i i i i i i i i . . i l i l i i l l l l l l l § I • •••Ml I ••••• ••••• m i l • : : : : : : : : : : : : : : :BBBBS r-*»J IIIIIIIIIIIIIIII itr=a iiiiiimiiit-i l\\m i i i i i i i i ssssss: ssssas • i i i n H I III Si! • m i H i l l M i l l IIIII ::::: •••• •••• i i ipiii iai • r i l l i i i i i J i m II IIIII II i i SSSS! •••• IIIII l l l I I I I I II I IIIII II I I I I I I II I IIIII :'A Kill!! ii i ' if,'iiiii i if II r.aw 'ten Mil 11 in" i i if II in Mini i n i m i i • ii iiir l l l l l l l l l l l l l l ====== i i i i i i i i iiiiiii l i i i i i i l i i l l l i iiiiiii mm I i i i i i i i i i III i i i i i i i i i i n i i i i i i i i i i S S S S ! ======= I i i i i i i i i i i i i i i i : i i i i i i i i i i i i i i i i i i i i i i i I S S S J S S B B B S S S S S S S B S S ; :============:=:= i s i i s s i i i i i i i i i i i • ( • • • • • • • • • • • • • • • I I ;;;;;;;;;:;::;:==; I i n n inn mil • • • • i IIIMIIIII |mi HI i i i i i n m i i i IIIIIIII III! l l l l l l l l l l l l l l l IIIIIIHNH1' i r n i i i i in in nvt. m i l ni l !! - - — S - - - - " " 5 i i i i i I II • • • • • • • • • • • • • • • • M i l • ••••a • • • • • • • • • • l l l l l l l l l l l l II • • • • I I I I I I I I II l l l l l l l l l l l l II l l l l l l l l l l l l "III IIIIIIII II l l l l l l l l l l l l II I H I :::: Hiii.iSiiii ••••••••••I • • • • • • • • • • • • i i i i i i i ••••••••••••••••••I •••••••••••••iiiuii • • • • • • • • I I I I I I I I I I ••IIIIIIIIIIIIIIII •••• m i l ••••• nir • • • • • • i n IIIII mi n •••••••••! M i l l mi IliiliP" I l l l i i i i i i i i i i i i i i i i i i i n n i i i i j u i i imii i l i i imisni i i i i i i ! i l l 1 1 ! T ! i i i i i i i ! ! : : : : : : : : • • • • • • • • I • • • • • i M l i i l l i i i i i m i i u a a a a •If — !!! •iiimniiiii •IIIIIIIIII'" • • • • • • I • • • • • • « •••••••  •III IIIIIIIIII lllllllllllllll • • i i i i i i i i i m i i i i i i i t i i i i i i t i i i i i a i i i i i i i i m • l l l l l i l l l l l l l i i l l l l l l l i i l i i i i i i i i l i i l l l l l l • iiiiiiMsiiiiiMiiHiliiipiiiiiii ==::: : : : : : : : : : : r=:::::::::::::i::i:i:::i is::::::: :::::::: iiiiiiii i t a ; ^ | | | | i ; g ; f i i i i i i i Siilii I i i i i i i i i i ! iiiiiiiiiiiiiiiiiiiiif HI I l i l i i l i i i i i I S S B B S J S a S l ! \mwmm sss ssx: ::::: EE: :::: ::::: ••>••• • • • • • • • i n • a • • • • •••*••>•••*! ••••••••>• I H M N H H H l :::::::::::: :::::::::::: illlllliliilii :::: i i ! • ••••••IIIIIIK lllllllllllllll IIIIIII II Iiiiiiiiii III iii i i i i i i i i ••• I B • • • I I I Hill I llll I llll IIIII I llll llll IIIII I llll IIIII IIII mi 7 - 8 9 1 i i i i i i i i i i i i 5=5::::: : [ l i l i i i i i i i i i H I i i J l l i i i i i i i i l l i l l i i :========: ^ r-' i i i i i i i i i i i i i i i i m :SEsS||5SE|iS||E: llllll •I#J ilrli i j i i I « , H S u n n II • l I* II Fl •III Si l l uii'jr II nun llllll __ nun •• 11(111 • • llllll •II l l l l 111111111 l l l l III ••••••  -: ! ! ! ! ! ! ! ! E E = E E S S B : Bi£SSSSl . . i i i i i i i i i l i l l l Iliiilllii i i i i i i i i i i Iiiii ilUIIIIii IIIIIIII1MHMUUM i i : : i : : : i s s s s s a s s : ••I • • • • • • • • • I I M I I I H l H M H « H M a 1111 minim :: iiiiililll l l l l l E I I i i l l i l l l l f l IKjflllllllll 1111  I i i i i i i i i i i i i i i l l l i J . V : ; I : : : : : : mtLk ft ] » H I I B I KiUB \inun p i l l I i i i i i i i — i tan •••• "1 H i -l l IS E:: iii!!:i:!ssssa:asss:s i H i r r r r i m ] i IHIHIIIIII IIHIIIIIIjlI I l l l l l i l S ! mi • • • •• • miiiii I III III III •• i in II in H in in II in ii • • • • • • • • • M III! ••••••» l l l l ( II •I i i i i i i sli!• ••••• : :: • • • • • • • • • ••••• ! •••••••• I M i l • ••••••• I I I I I :::::: llllll • H i l l • • • • i n •• IIIII - mir in; ii • • • • II • H I H i r r r r l imn IIIII III; llllll II !! in in i i i i i i i i • • i i m i . . • • • I I I I I I I I II • • • • Eiii •in in P H I II im i i i in mn i HI iiiiiiiiiliiiillilHj IlissiiiiiiHIillllllli mn mini lllllll :::: IIIiiiiiiiiiiIilIIllIiII!IiIiiiiiiiiiiiiiIiiiI!III!===i=li :=s==a=ssai:t ======= • i II • . i l l . . . . • • • k •• ••••••• l l l l l l l l l l l _ •• • • • • • • • • I I I • •• l l l l l l l l l l l • •• I I •••••HIM • I i i i i l i l l l ssssss:::; i i iiiiiiiiin iiiiiiiiiii II lElIIIIIIIV • •••••• IIIlL • • • • • • • I I I I I I I III II • • • • • I ._ •••••• • •••••• •••••• • • • • • • • • • • • • ! •••llllllll • • • • I I I I I I I lllllllllll lllllllllll lllllllllll lllllllllll lllllllllll Iiiiililll •Iimi lllllllll in I I I I I I I II !======I§I i i i i i i •I • II i ii i S B B I sssssss: ::=: ...!• iiiiiliiiillliilUHilnlill!!!! 5 6 7 8 9 1 S S f S S S S S S _ a a > B a a a S I i i i i i i i i i i i i i i i t mm 1======= i i i i i i i ! r-4*ii f i l l 111=1=1 •SssSs . J S S S S S =========== i i i i i i i i i i m m i i i i i i i i H HI li!l!f^!!!!!!i!!!i[iiil iliFfi'iiii = I I i i i i l l l l i i I I I I I ! M H H m l==IIIIIIIIlIIillIIII jKjjjiEiiii! • I I 111 IIIIIIMIIIIIIIIIIII... ••• lllllllllllllll I I I I I I I I Hi! itiiimmiij ill/jllllllliiliillllliliiiii . iirrrii|iiiiiiiiiiiiiiiiiiiini i » mil mi I ii 'iiiiiiiiiiill I V, IIIIIIIIIIINIIIIIIHIIIIIII llllllllllllllllltlllllllllll lllllllllllllll !ll!!'IIIII E H E l I B a i l ::::::::: iiiiiii::! I lk M mii i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i IIIIIIIIIIIIIII IIIIIIIIIIIIIII •5 5 I i i i i i i i i i i i i i i m mm n55555aHHaa^f isa iaa i ia s i i l i i i i i i i BBS 1 - ! 1 l ' ! ! i l S : : : . 1 ; ! ! ! ! ! i : . ! . . ' . ! ! ! ! l EEs mmEimmm\m\ i i i i i i i Iliiiillllfiliilfil ; i i========: IIIIIIII! • • • • • • • • • • • • H H i i l i l l i i i i i i i i i i i i mi mi illililli i i = S S = = = = = = = = = = i : = = = = = s = = : : ========== r mm iSBSSSSSSSS ^ i i i i i i l i v ; ' i i i i i i H H I H L 1 1 9 m sill _. II I I!IC;!|! IIIII IIIIIII II IHI ...IIIII IHHI ii! S S H I S S i i S i l K i i i l • • • • • • •••••••••• II • • • • • • • • • • < • • • • • II • • • • • • • • • • • • • • l l l l I I H I I • • • I I I I I I I II • • l l l l l l l l l l l l l l l l . • I J I I I I I I I I I I I I I I I I • ••••••••IIIIIIIIII ~iBSSS *555i •#•• ========= == ::::::::: •IIIIIIII " i i i i i i i ! E E : : : : : : : i i i i i i i i i i i i i i i i : ! : ! IIIIIIIIII IIIIIIIIII IIIIIIIIII ill l i i i i i i i H i i i i i i i i i l u ^ s I I i i mw\\\mmm=m%=m • • • • • H'IIIIIIIIIIIHI M I Si!:!!!!!!!!!!!!!!!!::! • • • • • • • • IIIIIIIII n • •••••••••••••in IIIIIII i • • • • • • • • • • • • • •II I IM I I I •iiiiiiiiniiiiiiiiiiii • ••••••••••••IIIIIIIIII l l l l l l l l l l l l l l l l l l l l l l l l l l mam ••••• IIIIIIIIII iiiiiiiui ~ ~ ||Kl>"li:V'Tin. • ••••• • • • • • • ^ • i i i -iiiiiiiiiiii !!==== •••••••••a mil M M Kliill lll a • • I I I I I I I I I I • •aiiii-iiiiiiia llllllllllllllll llllllllllllllll — ••••llllllllllll • III llllllllllllllll ~ IIIIIIIIIIIIIIII llllllllllllllll 7 8 9 '••••PI iiiiiii in S S S S — i S S : : : : : ==——======: s s s s r ^ s s s r - u - « =LJ--==E=EE:=..--..!::.: iiiiiiiiii I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I IIIJIIIIII IIIIIIIII llllllllllll ill i i i i i i i i i i m m i HI llll • S i U f f l l H iiiiiii IIIJJiL • •Bai • • • • • • ••I • H I ••IIIIIIIIII mil llllllllllll iiiiiiiiiiii •••IIIIIIIIII IIIIIII ..JIU... iiiiiiiiiiiin ill llllll Illlili mmm II II i n i n •II l l l l l l l l llll llll • a n ••••mi m i llll llll IIIIIII llll IIIIIIII IIIIIIII IIIIIIII IIIIIIII ffi :!:: illi 5 €> 7 8 S> 1 3 4 5 6 7 8 9 7 - 8 9 1 I I s i i i i l i i l l l l l H I I I H l i r ! ! i i i i i i i i i i i i l i i i i i i i i H i i i i i i ::::::: ::::::: I i i i i i i i i i§ ====== i S I IIIIIII I III MUIMil I III II i - l l !1 !>««•• I IIIIIII I III i H i : , i IIRI I H i l l i l ! 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Figures 32-35 show a difference i n slope of 0.1226 between the two planes f o r males and 0.0022 f o r females. These small differences were due to the actual s i m i l a r i t y of the two planes of n u t r i t i o n i n J group. Had the two growth responses been more widely separated larger differences i n slope would probably have been recorded. Comparisons of the slopes of a l l three l i n e a r measurements shows that a 1% increase i n heart-girth, height-at-withers and length of hind foot corresponds respectively to a 2.6$, 3»6$ and 4.0$ increase i n body weight. Thus, h e a r t - g i r t h corresponds most c l o s e l y with body-weight while length of hind foot i s r e l a t e d to body weight changes to a r e l a t i v e l y small degree. On t h i s basis h e a r t - g i r t h may be used In the estimation of body weight while length of hind foot may be used to determine the degree of fleshiness or condition of a deer. (1). An index to n u t r i t i v e condition. Knowledge of the condition of deer i n various areas i s necessary i n order to adequately evaluate n u t r i t i v e conditions. De s c r i p t i v e l y , t h i s i s d i f f i c u l t but the provision of a quantita-t i v e gauge of condition can be of value i n i n t e r p r e t i n g the e f f e c t of environmental factors., Raising deer under known n u t r i t i v e conditions has provided a standard by which comparisons with wild members of the species can be made. Linear growth measurements of the captive deer show that quantitative analysis of n u t r i t i v e conditions are p o s s i b l e . 1 * . Heart-girth has been shown to be a usable measurement i n the determination of l i v e body-weight. In contrast, length of hind foot represents a wide range i n body weights and i s r e l a t i v e -l y unaffected by the n u t r i t i v e plane. Therefore, use of Figures 26 and 32 f o r estimating body weight w i l l usually r e s u l t i n d i f -ferent values. The value r e s u l t i n g from h e a r t - g i r t h measurements w i l l approximate actual weight whereas weights r e s u l t i n g from the use of hind foot measurements w i l l represent the weight of the standard (captive) deer f o r that measurement. Thus, i n a "normal" deer, the weight estimated by both methods should be equal whereas i n an undernourished deer, the weight estimated from h e a r t - g i r t h w i l l be less than the weight estimated from hind foot length. Mathematically, t h i s may be expressed by the formula: I = W^ g from h e a r t - g i r t h and Vil^ i s the weight estimated from the hind foot length. weight from the hind foot length depending upon which plane of n u t r i t i o n i s to be used as a standard. Use of the high standard (Figure 32) w i l l tend to accentuate the index of condition i n undernourished deer. By using t h i s standard "normal deer" w i l l correspond to the high plane of n u t r i t i o n , whereas by using Figure 33 the "normal deer" w i l l correspond to the experimental low plane. Accordingly, the reference base must be chosen which w i l l s u i t the intended purpose. Table VI shows the index of condition f o r two male deer calculated at d i f f e r e n t stages of growth and by the two references where I i s the index of condition, W, c Either Figure 32 or 33 van be used i n estimating the 1*5. TABLE . THE INDEX OP CONDITION Series Deer No. Plane of n u t r i t i o n Actual wt • Kgms• Heart-g i r t h (cm) Est. wt. from heart g i r t h A 1H Low 3U.01+ 7 5 . 5 31*.5 3H High 3l* .o l* 71+.0 33.0 B 1H Low 32 .67 75.0 31*.0 3H High 68.71* 97.0 66.0 t i*6. FOR DEER 1H AND 3H Length Est. wt. from hind Index of of hind foot using condi-bion using foot (cms) P i g . 33 Pig. 32 Aver. . P i g . 33 Pig. 32 Aver. 1*2 . 36 1*1* ho 0.96 0.79 0.86 39 28 33 30 1.18 1.00 1.08 1*2 36 1*1* l*o 0.95 0.77 0.85 1*2 36 1*1* l*o 1.81* 1.50 1.65 1+7. bases Figures 32 and 33. Series A consists of data chosen so that the actual weights were the same f o r both deer. This occurred shortly a f t e r establishment of the planes of n u t r i t i o n . Series B represents data chosen f o r i d e n t i c a l hind foot lengths which occurred near the end of the experimental period thereby representing wide differences i n actual weights. I t i s apparent that the index of condition.tends to be 1.00 or greater f o r the high plane of n u t r i t i o n and less than 1.00 f o r the low plane. I t should be noted that the use of Figure 32 (high plane regression of body weight against hind foot length) produced an index of n u t r i t i o n of 1.00 for 3H i n Series A tending to suggest that deer 3H was i n a normal condition. The index of n u t r i t i o n calculated by using the same regression produced a value of 1.50 i n Series B. This may be due to large depositions of f a t i n the animal which would tend to give a r e l a t i v e l y large heart-g i r t h i n r e l a t i o n to hind foot length and thereby a r e l a t i v e l y large estimated weight. A graphic method can also be used i n the evaluation of n u t r i t i v e condition. I f the weight estimated from the h e a r t - g i r t h equals the weight estimated from the hind foot length, the index of condition w i l l be 1 .0. Thus, i f weight estimated from heart-g i r t h i s p l o t t e d against weight estimated from hind foot length as shown in Figure 36, the normal deer would be represented by point "C.M I f "C" i s joined to the o r i g i n by a s t r a i g h t l i n e the slope of the l i n e i s 1.0;. Therefore the normal* deer i n the graphic method i s also represented by a value of 1 .0. Thus f i e l d data 1*8. may be p l o t t e d which may f a l l on either side of the "normal" l i n e , i . e . , points "a" and "b." By joining points "a" and "b" to the o r i g i n , the slope of the l i n e may be determined i f a mathematical value f o r condition i s desired. The value thus obtained w i l l be the same as that produced d i r e c t l y by the formula I Q = ^hc* Weight estimated from length of hind foot FIG. 3 6 . A graphic method for the evaluation of the Index of condition The standard used i n this method of evaluating condition apply only to the Columbian b l a c k - t a i l e d deer. However, the method i s applicable to other, species provided that reference bases are constructed f o r the regression of h e a r t - g i r t h against body weight and length of hind foot against body weight. These references may be constructed from f i e l d data, but more s a t i s f a c -tory r e s u l t s would be obtained by using experimental animals 1+9. under con t r o l l e d n u t r i t i v e conditions. ( j ) . The blood chemistry of the b l a c k - t a i l e d deer. Due to the number of determinations which were c a r r i e d throughout the experiment, duplicate analyses were not possib l e . Thus, the standard deviations of the means shown i n the following tables represent v a r i a t i o n s due to experimental error and i n d i v i d -u a l variations i n the deer. However, standardization of a l l meth-ods used, reduced the experimental errors to a minimum and there-fore, i n d i v i d u a l v a r i a t i o n forms the greater part of the standard deviations. , Differences i n blood lev e l s associated with sex was over-looked i n the following a n a l y s i s . The r e l a t i v e l y few females would not a f f e c t the averages to a s i g n i f i c a n t degree and there-1 fore both males and females are included i n the averages. The following analyses of blood l e v e l s should indicate n u t r i t i o n a l stress associated with the low plane deer as compared with the high plane animals. However, animals tend to react i n such a way as to' minimize or undo the e f f e c t s of the stress by lowering t h e i r c a l o r i c requirements and decreasing c a l o r i c expend-i t u r e (Anon, 1951)• For this reason, the ef f e c t of two planes of n u t r i t i o n which are both above the maintenance l e v e l , can have but l i t t l e e f f e c t on c r i t i c a l l e v e l s of the constituents of blood, provided that no actual s p e c i f i c deficiency e x i s t s . On the other hand, age and the associated metabolic pathways at d i f f e r e n t stages of growth does a f f e c t blood constituents. 1. Packed c e l l volume. Changes i n packed c e l l volume of blood r e f l e c t s many 5 o . types of p h y s i o l o g i c a l changes. The volume i s increased by such factors as high a l t i t u d e , muscular exercise, heightened environ-mental temperatures and numerous pathological conditions. C e l l volume i s decreased at high barometric pressures and low environ-mental temperatures as well as being the r e s u l t of some pathologi-c a l conditions (Best et a l , 1 9 5 0 ). In a d d i t i o n , age of the a n i -mal affects the packed c e l l volume, p a r t i c u l a r l y i n early post-n a t a l development. Red c e l l s are larger (macrocytic) i n f o e t a l blood than i n adult blood (Parkes, 1 9 5 2 ). This tends to elevate the packed c e l l volume i n the very young animals, as was found by McLeroy ( 1 9 5 2 ) , and Wise e_t a l (191*7). However, the volume soon decreases to a l e v e l below that of the adult, due to a change i n blood c e l l s to the microcytic type and to the high t o t a l blood volume displayed by a l l young animals (Hansard et a l , 1 9 5 3 )• As the production of adult red blood c e l l s increases with the growth of hemopoietic tiss u e s , the red c e l l volume gradually increases to the adult l e v e l . Table VII shows that the packed c e l l volume was already below that of the adult l e v e l at the time of the f i r s t a n a lysis. S i g n i f i c a n t increases occurred throughout the experi-mental period, but the packed c e l l volume of J group was s t i l l below that of the yearlings at 100 days of age. Rosen et a l , (1952) and Rusoff et a l (1951*), showed that hematocrit values varied between months i n mule deer and dairy c a t t l e , r e s p e c t i v e l y . Both investigators report peaks to have i occurred i n the red blood c e l l l e v e l i n May and June which they associated with heightened environmental temperatures. However, the present data showed no s i g n i f i c a n t difference between monthly TABLE VII PACKED CELL VOLUME IN PERCENT OP WHOLE BLOOD FOR 1951+ FAWNS (J GROUP) AND YEARLINGS (H GROUP) 1951+ Fawns (J Group) H Group Date of blood sample July 5 July 12 July 22 Aug. 3 Aug. 31 Sept. 23 July 1 2 -Aug. 31 Age In days 20 27 37 1*9 77 100 392-1+65 Mean red c e l l v o l . % 3l*.l* 3 5 . 2 3 6 . 2 1*1*. 3 1*3.5 1+9.8 5 8 . 2 Standard Deviation ± 5 . 2 ± 1*.9 ± 5 . 2 ± 8 . 8 • 8.1* ± 7 . 5 ± 7 . 0 Comparison to (H Group) P = < 0 . 0 1 <0.01 < 0 . 0 1 < 0 . 0 1 < 0 . 0 1 <0.01 Range 21+. 3-1*0.0 31*.0-66.0 3 0 . 5 - 6 5 . 0 3 0 . 0 - 5 5 . 0 1*0.5-63.5 1*1*. 0 - 5 9 . 0 1*1*.0-70.0 Mean P.C.U. (H Group) 58.9±1*.9 5 6 . 9 + 5 . 3 6 7 . 2 ± 2 . 5 51*.1*±5.8 5 2 . 1 + 5 . 8 >o.o5 <o.oi <o.01 >o.o5 5 2 . values. In addition, Rosen e_t a l , (1952) showed that the average packed c e l l volume d i f f e r e d between areas of poor and good n u t r i -t i o n a l conditions. Table VI]& shows that under the conditions of the present experiment, the Columbian b l a c k - t a i l e d deer showed no s i g n i f i c a n t difference between the two planes of n u t r i t i o n . TABLE V i l a PACKED CELL VOLUME IN PERCENT OP WHOLE BLOOD FOR THE YEARLINGS (H GROUP) Plane of n u t r i t i o n High plane H Low plane H Both high and low plane Dates of analysis July 12-Aug. 3 i July 12-Aug. 3 July 12-Aijg. . Age i n days 392 - 1*65 392 - 1*65 392 - 1+65 Mean 5 9 . 1 5 7 . 5 5 8 . 2 S. D. + 5 . 8 t 7 . 9 4 7 . 0 Range 5 0 . 5 - 6 9 . 0 i+l+.O - 7 0 . 0 l+i+.o - 7 0 . 0 P >o . o 5 Haden (1932) and Kaplan (1951+) report sex differences i n the packed c e l l volume, the lower values corresponding to females. The present study showed no sex differences i n the packed c e l l volume of H group f o r the Columbian b l a c k - t a i l e d deer, 58.h% and 5 7 . 9 5 $ representing male and female, r e s p e c t i v e l y . The values obtained i n these analyses are higher than those reported f o r c a t t l e by Rusoff et a l (1951+), and f o r goats, cats, r a b b i t s , dogs and man by Emmons, 1 9 2 7 . These high values are thought to represent the true l e v e l s f o r the Columbian black-53 t a i l e d deer since centrifugation was c a r r i e d out f o r 20-30 minutes at approximately 2,000 r.p.m. Schlenker (1952) found these condi-tions to produce complete packing and accurate r e s u l t s . 2. Erythrocyte sedimentation rate. Sedimentation rates i n the experimental deer varied wide-l y between indivi d u a l s and between samples from the same i n d i v i d -u a l . Thus no c o r r e l a t i o n with the plane of n u t r i t i o n could be demonstrated but a s i g n i f i c a n t difference did e x i s t between the fawns and yearlings (Table VII I ) . Rosen et a l (1952) also found wide variations i n the erythrocyte sedimentation rates i n mule deer. TABLE VIII ERYTHROCYTE SEDIMENTATION RATES FOR FAWNS AND' YEARLINGS J Group (fawns) H Group (yearlings) Mean - S.D. 11.63 • 12.7 6.19 ± 7.91+ Range 0 .5 - 59.0 0.5 - 3U.0 P <0.05 Numerous factors a f f e c t the rate of sedimentation. Sp e c i f i c gravity of the plasma, size of the corpuscles, clumping of c e l l s , shape of the erythrocytes, temperature, l e c i t h i n - c h o l -e r t e r o l r a t i o of plasma and the red c e l l concentration a l l a f f e c t the rate of sedimentation (Parkes, 1952). In addition, pathologi c a l conditions such as anemia, malignant tumours and inflamatory conditions have been shown to increase the sedimentation r a t e whereas decreased rates are associated with a l l e r g i c states, 51*. peptone shock and hemolytic anemias (Parkes, 1952). Other i n -vestigators have also shown a r e l a t i o n s h i p between basal metabol-ism and the rate of erythrocyte sedimentation (Baisset et a l , 191*9) . Pathological conditions may be r u l e d out as causing the variations i n sedimentation rates i n the present experiments. However, differences i n the development of hematopoietic tiss u e s , i n water balance, si z e of the corpuscles and the temperature dur-ing analysis undoubtedly served to produce the wide variations found i n the young deer. However, since blood volume i s high i n young animals (Hansard ejb a l , 1953), "the sedimentation rate proved to be higher i n the fawns than i n the yea r l i n g s . 3. Hemoglobin. The hemoglobin content-of blood i s affected by numerous factors which tend to decrease the hemoglobin l e v e l from normal. Age, sex, climate, a l t i t u d e , loss of blood, increased blood de-st r u c t i o n and impaired bldod formation, a l l a f f e c t the rate of hemoglobin production (Best e_t a l , 1950) • Since the rate of hematopoeisis i s i n d i c a t i v e of n u t r i t i o n a l d e f i c i e n c i e s , the blood picture of normal Columbian b l a c k - t a i l e d deer reported here may be used as a standard f o r field'comparisons. However, the e f f e c t of age, sex and cl i m a t i c factors must be accounted for before true comparisons can be made with the standard. Hawk, et a l (1951) report f o r humans, that the hemoglobin l e v e l remains between 11-12 ragra.$ during the f i r s t six months of l i f e , followed by a gradual increase u n t i l the age of 16 years i s reached. Since ruminants are r e l a t i v e l y more mature at b i r t h 55 than are humans, the length of time for a t t a i n i n g the adult l e v e l i s r e l a t i v e l y short. Thomas, _et a l (1951+) found the hemoglobin le v e l s of dairy calves to be r e l a t i v e l y high at b i r t h and to de-c l i n e shortly a f t e r b i r t h , reaching a minimum value at 1+0-60 days. According to t h i s i n v e s t i g a t o r , the hemoglobin l e v e l then gradu-a l l y increased and reached the adult l e v e l during weaning. Wise, et a l (19U7) substantiates these changes i n hemoglobin leve l s but indicates that the minimum value i s reached at 21 days. Pigs show a minimal l e v e l at approximately ll+ days (Vestal, e_t a l , 1938). Table IX shows that a s i m i l a r decline i n hemoglobin l e v e l s i s found i n the b l a c k - t a i l e d deer. However, the minimum value occurred at approximately 37 days. The decrease i n hemoglobin l e v e l i s associated with a change from f o e t a l hemoglobin to the adult type. The f o e t a l hemoglobin d i f f e r s from that of the adult i n i t s a b i l i t y to become saturated at lower oxygen tensions (Prosser, et a l , 1950). A d d i t i o n a l l y , f o e t a l hemoglobin i s manu-factured mainly i n the l i v e r while adult hemoglobin i s produced i n the bone marrow. Hence a change i n t o t a l hemoglobin l e v e l occurs during the change from f o e t a l to adult hemoglobin. I t i s also suggested that the high hemoglobin l e v e l s found s h o r t l y after b i r t h i s associated with colostrum intake during the f i r s t few days of post-natal l i f e . Colostrum i s high i n p r o t e i n , p a r t i c u l a r l y i n the globulin f r a c t i o n , which would provide b u i l d -ing materials for hemoglobin, thus ensuring r e l a t i v e l y high hemoglobin production during the ea r l y suckling stage. Adult hemoglobin leve l s were attained at approximately 100 days of age 56. TABLE HEMOGLOBIN VALUES IN GMS PER 100 ML. Date of blood sample July 5 July 12 July 22 Age i n days 20 27 37 High plane means 11.01+ 11.51 10.09 Low plane means 11.68 11.02 8.96 Mean for a l l fawns 11.36 t 3.11 • 11.27 + 1.53 9.53 t 3.92 P >Q, 05 <o 02 Range 8.31 - 15.00 9.08 - 11*.63 8.28 - 13.51 TABLE HEM0GL( ©IN VALUES IN C JMS PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age i n days High plane means Low plane means Means f o r a l l year-lings P Range 392 18.57 18.67 18.62 +-0.83 1 , 1+02 15.22 11+.50 11+.86 +r-tv23 1+11+ 15.1+9 .11*. 1+2 11*.96 + 0.97 <o 17.26 - 19.71+ 01 >o 12.65 - 16.08 05 11+.29 - 16.80 57 IX BLOOD FOR THE 1951+ FAWNS (J GROUP) Aug. 3 Aug. 31 Sept. 23 July 5-Sep .23 1+9 1 0 . 8 8 1 0 . 7 9 1 0 . 8 3 + 3*25 77 11.21+ 1 0 . 3 6 1 0 . 8 0 * 1 .51 100 10.91 10.21+ 1 0 . 5 0 + 1 .57 >o.o5 . >o 7 . 7 2 - 1 3 . 1 3 05 >o. 8 . 3 1 - 13.31+ )5 8 . 3 1 - 1 2 . 6 0 20 - 100 /10.91+ • 2.1+8 [\l0.1+l+ • 2.1+8 1 0 . 7 1 ± 2.1+8 P > 0 . 0 5 7 . 7 2 - 1 5 . 0 0 X BLOOD FOR YEARLING DEER (H GROUP) Aug. 31 Sept. 23 July 12 - Sept.. 2 ; 1+1+2 1 5 . 0 0 1 3 . 0 4 1 3 . 8 2 f 1 .11 1+65 1 2 . 5 3 1 1 . 1 0 1 1 . 6 7 * 1 .25 392 - 1+1+2 1 5 . 6 1 t 2 . 0 8 ^ 1 4 . 3 5 t 2 . 6 1 / 11+.93 + 2.1+2 9 . 8 0 - 19.71+ P = > 0 . 0 5 >o.o5 <o 1 2 . 6 0 - 1 5 . 1 5 ,02 9 . 8 0 - 13.31+ 58. as shown by a comparison of Tables IX and X. The data f o r the y e a r l i n g deer (H group) shows a peak hemoglobin l e v e l f o r July 12 followed by a decreased, but s t i l l elevated l e v e l from July 22 to August 31* The f i n a l value f o r September 23 r e f l e c t s a much lower hemoglobin l e v e l . Similar chan-ges i n hemoglobin leve l s have been reported f o r d a i l y c a t t l e and mule deer by Rusoff, et a l (1954), & n d Rosen (1952), r e s p e c t i v e l y . These investigators a t t r i b u t e d the changes i n blood lev e l s to changes i n environmental temperatures. However, since the summer i s a period of ample food supply and u s u a l l y good growth conditions, the changes In lev e l s of hemoglobin may be associated with hormonal regulations of growth and metabolism. Tables IX and X show that no s i g n i f i c a n t differences occurred i n hemoglobin l e v e l s between high and low plane deer. Since the planes of n u t r i t i o n were simply differences i n c a l o r i c intake and the low plane deer were f e d above the maintenance l e v e l , i t i s suggested that the r e s t r i c t i o n of c a l o r i c intake alone does not affect the rate of hemoglobin formation. However, Bueckler (191+9) found that a r e s t r i c t i o n i n the quantity of food protein i n t e r f e r e s more s e r i o u s l y with hematopoiesis than does a r e s t r i c t i o n of ccalories. A d d i t i o n a l l y , Rousseau, e_b aJL (1951+) found that hemoglobin l e v e l s decreased i n calves f e d a low c a l o r i c Vitamin A d e f i c i e n t r a t i o n but remained normal on a high c a l o r i c Vitamin A d e f i c i e n t r a t i o n . This tends to suggest that a low c a l o r i c l e v e l may act concurrently with cert a i n s p e c i f i c n u t r i t i o n a l d e f i c i e n c i e s to lower the hemoglobin l e v e l . However, d e f i c i e n c i e s of metalic Ions, necessary f o r hematopoiesis, would a f f e c t the 59 hemoglobin l e v e l s regardless of the c a l o r i c plane of n u t r i t i o n . The values reported herein may be regarded as normal since the c a l o r i c plane was not below maintenance and since no s p e c i f i c vitamin or mineral d e f i c i e n c i e s were produced by the diet which were not immediately a l l e v i a t e d . I4.. Blood Glucose. Blood glucose was determined by the method of Benedict, on a P o l i n and Wu protein free f i l t r a t e . The Benedict method i s reported to y i e l d values 20-30 mgm$ higher than the true glucose l e v e l (Hawk, et a l , 1 9 5 l , and Reid, 1950). The elevated l e v e l r e s u l t s from the presence of non-glucose reducing substances (saccharoides). However, this f r a c t i o n appears to be r e l a t i v e l y constant and therefore i t s presence does not unduly influence the i n t e r p r e t a t i o n of variations In blood glucose lev e l s obtained by th i s method. Venous blood, taken from the recurrent t a r s a l vein, has a lower blood sugar l e v e l than a r t e r i a l blood (West, et a l , 195>1), thereby p a r t i a l l y correcting the values obtained by Benedicts method. Table XI shows that the blood glucose l e v e l i s high a t b i r t h but decreases r a p i d l y a f t e r approximately t h i r t y days. At 100 days of age, the blood glucose lev e l s of J group were s t i l l s i g n i f i c a n t l y d i f f e r e n t from those of H group. This i s believed to be due to the prolonged weaning period. The high blood sugar levels of young ruminants has been noted by many workers (Snook.et a l , 1938; McCandles et a l , 1950; Passraore et a l , 1938; and Reid, 1953)• These investigators 60 record that the decrease i n blood glucose l e v e l i s coincident with the onset of rumination. For this reason i t was believed that the decrease resulted from metabolic changes associated with the inception of b a c t e r i a l fermentation of carbohydrates. How-ever, Jacobson, e_fc aJL (1951) reported that delayed rumination due to the prolonged feeding of milk i n young calves, d i d not a f f e c t the pattern of changes i n blood glucose lev e l s shown by normal calves. The data shown i n Table XI indicates that both views may be represented. The high I n i t i a l l e v e l indicates that glucose i s the main source of energy exchange i n the newborn. A f t e r b i r t h , I t i s u n l i k e l y that such aseptic conditions could occur so as to e n t i r e l y prevent the accumulation and growth of rumen micro-organ-isms within the suckling fawn. These organisms may then ferment some glucose, r e s u l t i n g from the s p l i t t i n g of lactose. However, most of the glucose Is absorbed together with the galactose f r a c -tion and transported to the l i v e r where the galactose i s converted to glucose, ^ h e a b i l i t y of the digestive system to absorb glu-cose d i r e c t l y i n the young ruminant has been demonstrated by Reid (1952) and Holmes (195227* Therefore, these two sources of g l u -cose serve to maintain a high blood sugar l e v e l . The decrease i n blood glucose from the immediate post-n a t a l l e v e l to that recorded at approximately t h i r t y days of age may be due to fermentation of the glucose by small populations of b a c t e r i a . Werkman, et a l (1951) state that glucose fermenting c e l l s w i l l aquire competance on prolonged exposure to l a c t o s e . Thus, fermentation of carbohydrates may a c t u a l l y begin i n the 61. TABLE BLOOD GLUCOSE LEVELS IN MGM. PER 100 ML. WHOLE BLOOD H GROUP Date of blood sample July 5' July 12 July 22 Age i n days 20 27 37 High plane means 95.0 t 16.2 8J.-1 *: ,13.3 49.6 ± 9.1 Low plane means 85.4 ± 13.6 70.2 ± 9.2 46.9.4 11.2 Means f o r a l l fawns 90.2 4 14.1 76.6 4 . 1 4 . 0 48.3 ± 9.8 P 05 <o .05 Range 63.0 - 121.0 57.5 - 116.0 25.5 - 67.0 High values for high plane mean believed due to feeding Nurse Cow milk replacement to the high plane fawns. TABLE BLOOD GLUCOSE LEVELS IN MGM. PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age i n days 392 402 414 High plane means 51.3 + 1.1 .34.7 + 7.0 40.0 + 7.7 Low plane means 35.7 i 11.5 30.8 4 3.0 38.5 + 9.0 Means for a l l year-l i n g s 43.5 ± 7.3 32.8 + 4.8 39.8 ± 7.4 P * 0 05 <o, 05 Range 28.0 - 52.0 27.5 - 41.5 29.0 - 45.5 'One deer only, #2H - a highly nervous animal **Low plane switched back to high plane as of Sept. 1. No s i g n i f i c a n t difference between planes of n u t r i t o n (P 0.05) for the Sept. 23 sample. 62 XI FOR 1954 FAWNS ( J GROUP) AND IN COMPARISON TO (YEARLINGS) Aug. 3 * Aug. 31 Sept. 23 July 12-Sep .23 49 6 3 . 1 ± 4 2 . 1 4 9 . 4 i 1 3 . 2 5 6 . 3 + 3 1 . 0 77 4 6 . 2 i 1 2 . 0 4 1 . 6 ± 1 5 . 2 4 3 . 9 ± 1 2 . 6 100 4 3 . 3 * 7 . 2 4 4 . 6 ± 4 . 5 4 3 . 9 * 5 . 4 392 - 465 4 0 . 7 ± 1 0 . 6 3 3 . 0 • 7 . 5 3 6 . 4 ± 8 . 8 > 0 . 0 5 >o 2 7 . 0 - 1 5 3 . 0 05 >o. 2 7 . 0 - 6 5 . 0 05 <o 3 6 . 0 - 5 i . o 05 2 1 . 5 - 5 4 . 0 XII WHOLE BLOOD FOR THE YEARLINGS (H GROUP) Aug. 31 442 5 4 . 0 * 2 5 . 3 ± 5 . 3 3 9 . 7 t 5 i i >0.05 <0 2 1 . 5 - 5 4 . 0 Sept. 23 465 2 7 . 8 + 3 . 1 3 2 . 2 + 2 . 4 3 0 . 0 t 2 . 2 05 2 5 . 5 - 3 2 . 0 July 12 - Sept. 23 392 - 465 4 0 . 7 + 1 0 . 6 - 3 3 . 0 * 7 . 5 • 3 6 . 0 t 8 . 8 2 1 . 5 - 5 4 . o p <.o.o5 63. suckling ruminant and thereby decrease the amount of sugar being absorbed d i r e c t l y Into the blood stream from the rumen. However, as the fermentation of lactose occurs i n the absence of growth on the part of the micro organisms, the population w i l l remain small, thus preventing fermentation from proceeding to completion. There-fore less glucose i s being absorbed d i r e c t l y , but the blood l e v e l would s t i l l remain above the adult l e v e l due to the incompleteness of fermentation. Further study i s necessary to c l a r i f y the exact mechanisms of the decrease i n blood glucose l e v e l s . Table XII shows that fluctuations of the blood sugar l e v e l occurred, producing a peak on July 12 and a s i g n i f i c a n t decrease on Sept. 23 (P<O.Of>). These seasonal differences are thought to represent metabolic changes associated with increased growth dur-ing the summer months, followed by decreased growth associated with hormonal r e g u l a t i o n . Since the deer were fe d the same r a t i o n both i n summer and winter, the increase i n blood l e v e l s i n July r e f l e c t e d favourable environmental conditions. This tends to suggest that temperature and/or l i g h t may a f f e c t the metabolism of the deer at t h i s time. However, the decreased blood sugar le v e l s occurred at the time of sexual a c t i v i t y and the voluntary reduction i n food intake which was noted previously. Therefore the decreased sugar l e v e l s may be associated with increased energy expenditure r e s u l t i n g from the hormonal regulation of sexual a c t i v i t y . The yearling deer (H group) showed s i g n i f i c a n t differences i n the blood glucose l e v e l i n r e l a t i o n to the plane of n u t r i t i o n . I t should be noted that the low plane deer were placed on high 61+. plane on September 1 thereby increasing the blood sugar l e v e l s so that there was no s i g n i f i c a n t difference between high and low plane deer by September 23. Table XI shows that no s i g n i f i c a n t differences i n blood glucose l e v e l s occurred i n J group. F a i l u r e of separation was probably due to f l u c t u a t i o n s i n d a i l y food intake which tended to equalize the two planes of n u t r i t i o n . On the basis of the data f o r the yearlings, i t may be possible to u t i l i z e blood glucose l e v e l s to evaluate the condition of carbohydrate metabolism i n wild animals. However, i t would appear that only prolonged major differences between two groups would r e s u l t i n d i f f e r e n t blood glucose l e v e l s . Since glucose i s r e l a t i v e l y less important as a source of energy to a ruminant than a non-ruminant, (Reid, 195l), fluctuations i n n u t r i t i o n are more l i k e l y to af f e c t the v o l a t i l e f a t t y acid l e v e l s than the glucose l e v e l s . I t has been found that f a s t i n g f o r 21+ hours i n ewes, produced no change i n blood glucose l e v e l . In addition treatment with i n s u l i n also f a i l e d to depress blood glucose le v e l s of sheep and goats beyond a point s l i g h t l y above the hypo-glycemic shock l e v e l (Passmore et a l , 1938)• The u t i l i z a t i o n of glycogen from the l i v e r and the formation of glycogen from amino-acids of tissue proteins under n u t r i t i o n a l stress provides a ho:m eostatic mechanism preventing major fluctuations i n glucose leve l s as long as the necessary materials are present. However, prolonged n u t r i t i o n a l stress does produce a s i g n i f i c a n t difference i n blood sugar l e v e l s as shown by the data f o r H group. 5 . The non-protein nitrogen of blood (N.P.N.). 6 5 . 5 . The non-protein nitrogen of blood ( t t . P . H . ) The non-protein nitrogen levels of blood result from ammonia, urea, uric add, creatine, creatinine, amino acids, glutathione and others. Urea contributes 4 5 % of the nitrogen to the N . P . U . Thus, variations i n U . P . U . are due largely to variations in the urea content of blood, (Hawk et, a l , 1 9 5 1 ) , ard reflect impaired renal function or excessive tissue cata-bolism such as would occur during starvation or severe malnu-tr i t i o n , (Best et. a l , 1 9 5 0 ) . Anderson, et. a l , ( 1 9 3 0 ) and Braun, ( 1 9 4 6 ) , found that the N . P . H . level of bovine blood tends to decrease with age. However, i t i s apparent that these workers were dealing with changes in adult blood levels with increasing age, as young L ruminants show an increase i n J Y . P . N . ' with increasing age, (Table X I I I ) . Successive analyses were not significantly d i f -ferent but a comparison of the mean values .'for a l l fawns of July 5 and September 2 3 show them to be highly significant ( P 0 . 0 1 ) . Table X I I I shows that wide fluctuations occurred in the U . P . 2 J . levels of the yearling deer. A high degree of varia-b i l i t y has also been reported for beef cattle and dogs by Colby et a l , ( 1 9 5 0 ) and Costa Val, ( 1 9 5 0 ) respectively. Accordingly, the blood W . P . N , level i s of l i t t l e value in measuring the effect of a factor on a group of animals since the individual variation obscures the results. However, the direction of variations in relation to the normal or average has been used to detect renal malfunction, (Hawk, et a l , 1 9 5 1 ) . 66. TABLE NON-PROTEIH Ug ITS MGM PEE 100 ML. Date of blood sample > July 5 July 12 July 22 Age i n days 20 27 37 High plane means 27.65 2.52 31.27 9.03 32.28 12.98 Low plane means 31,01 8.80 24.76 4.79 33.00 6.76 Means for a l l fawns 29.33 5.99 . 28.01 6.94 32.64 9.94 P O.i 05 .„ O.i 05 Ranges 24.64 - 43.68 15.68 - 45.56 13.44 -•47.04 TABLE HOB-PROTEIN JT2 Iff MGM PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age in days 392 402 414 High plane means 35.00 10.84 39,11 11.32 29.96* Low plane means 35.15 4.82 42.55 8.30 50.40* Mean for a l l yearlings36.07 7.51 45.82 8.88 40,19 14.45 P 0,05 0.05 Ranges 23.52 - 45.08 31.36 - 61,32 29.96 - 50.40 *1 animal only. 6 7 . XIII BIOOD FOR THE FAWNS (J GROUP) Aug. 3 Aug. 31 Sept. 23 Jul y 5^Sept. 23 49 77 100 20 - 100 33.12 12.00 51.43 .7.30 45.50 6.12 35,11 9.19 38.22 14.86 45.50 15.94 44.52 6.09 34,71 9.04 35.47 12.81 48.04 12.03 44.91 5.75 34.91 9.03 0.05 0.05 >^ o. 05 18.20 - 55.72 23.24 58.24 36.96 - 55.44 13.44 - 58.24 XIV BLOOD FOR YEARLINGS (H GROUP). Aug. 31 Sept. 23 . July 12-Sept. 23 442 465 392-465 48.81 4.65 43.42 15.21 58.24* 39.62 0.19 45.69 9.41 58.24* 45.14 3.29 44.06 12.47 0.05 _ 39.48 - 53.76 23.52 -> 61.32 68. Tables XIII and XIV indicate that there i s no significant difference between planes of nutrition with respect to the N.P.H. level of blood. Since H.P.lf. appears to act as a buffer system in the maintenance of a nitrogen balance, i t i s unlikely that stable differences would exist between planes of nutrition, except under extreme nutritional stress. Morgulis et a l , (1924), found in dogs, that the N.P.2J. increases at an early stage of fasting and then remains at a more or less fixed level u n t i l the extreme stage of starvation i s reached. At this time a new and greater increase occurs i n blood Jl.P.lff. However, no such changes could be expected in the present experiments as a l l the deer were above maintenance feeding levels and were not sub-jected to fasting. The overall average blood non-protein nitrogen for fawns and yearlings, 38.85 - 9.89 mgm. % i s thought to represent a normal level for the Columbian black-tailed deer up to 465 days of age. This figure may thus be used as a standard for the in-terpretation of variations*that may be found under f i e l d condi-tions. Both males and females were included as no differences between the two sexes were evident, as i s the case in dogs, (Leichsenring, et a l , 1939). 6. Plasma total proteins The plasma proteins primarily function i n the maintenance of a normal distribution of water between blood and tissues. A secondary function appears to be a buffer action in the acid-base balance (Hawk, et_ a l , 1951). Thus plasma proteins vary 69. within the individual in order to maintain the water and acid-base balance. However, greater variations are found between species than between individuals within a species. Under ordi-nary conditions of health, the total proteins remain relatively constant between individuals of the same species (Best e_t a l , 1950). However, i t has been found that severe malnutrition, particularly associated with variations in protein intake, af-fects the level of total proteins i n the plasma, (Goldsmith, 1949; Chakravarti, 1952; J.olllfe et a l , 1950; Weeoh, et. a l , 1935; Klosterman, et a l , 1950 and Sansi, 1946). Age and sea: are also reported to affect the concentra-tion to total plasma proteins, (Howe, 1925; Dalgaro, et, a l , 1950; and Fischer, at a l , 1954). However, Iteiohsenring, et_ a l , (1939) found no essential sex differences i n dogs. Data from the pre-sent experiments shows no essential sex differences but the affect of gestation and lactation may alter the results. Howe, (1925), Dalgaro, et a l , (1950), and Fischer, et a l , (1954), have reported low concentrations of total protein in the blood of foetuses and newborn in sheep, rats and other young animals. Table XV shows that no significant changes occurred in the total protein level from 20 to 49 days of age. Samples taken at 49 and 100 days of age show a significant difference which was thought to be due to partial starvation imposed on the fawns i n order to encourage the consumption of solid food. How-ever, the averages for the fawns were significantly lower than the averages for H-group. Table XVI shows that no significant monthly variations 70. T A B L E PLASMA T O T A L P R O T E I N S A S GrMS P R O T E I N P E R 100 M L . Date of blood sample July 5* July 12 July 22 Age i n days 20 27 37 High plane means 7.86 5.67 6.10 Low plane means 6.12 5.56 5.61 Mean for a l l fawns 7*08 2.14 5.61 1.48 6.01 1.16 P ( 3.05 Range 4.27 - 10.54 3.66 - 8.65 5.04 - 8.51 *Serum. TABLE PLASMA TOTAL PROTEINS AS GrMS PROTEIN PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age in days 392 402 414 High plane means 6.08 6.33 6.08 Low plane means 7.16 6.90 6.40 Mean for a l l yearlings 6.62 1.55 6.61 0.48 6.24 0.61 P 0.05 0.05 Range 4.57- 9.10 5.97 - 7.24 5.98 - 7.03 71. XV OF PLASMA FOR THE FAWNS fJ GROUP) Aug. 3 Sept. 23 July 5-Sept.23 ** 0.05 49 100 20-100 5.71 5.11 5.71 0.89 6.09 4.78 5.60 1.27 5.90 0.84 4.91 0.36 5.65 1.10 0.05 0.01 4.78 - 7.57 4.33 - 5.45 3.66 - 10.54 XVI OF PLASMA FOR THE YEARLINGS (H GROUP) J GROUP (FAWNS) Sept. 23 July 12-Sept.23 P July 5-Sept. 23 465 392-465 20-100 6.18 6.16 0.62 5.71 0.89 0.05 6.51 6.74 0.91 5.60 1.27 6.38 0.22 6.47 0.76 0.01 5.65 1.10 0.05 6,11 - 6.71 4.57 - 9.10 3.66 - 10.54 7 2 . were found i n the y e a r l i n g deer and/no v a l i d difference occurred between the two planes of n u t r i t i o n . Since both planes of n u t r i -t i o n were above the maintenance l e v e l , tissue breakdown and loss of weight did not occur i n general. (Thus marked changes i n the l e v e l of t o t a l proteins of blood could not result from t h i s con-d i t i o n . However, had starvation conditions been imposed, p a r t i -c u l a r l y i n terms of protein, the t o t a l proteins of blood would decrease. Bansi, ( 1 9 4 6 ) found i n humans suffering from hunger edema that the t o t a l serum proteins had decreased to 3 5 % of the normal value. Zlosterman, e_t a l , 1 9 5 0 also found i n sheep, that a decrease i n protein intake from 0 . 3 3 to 0 . 1 8 l b s . of d i g e s t i b l e protein caused a change i n serum t o t a l proteins from 6 . 9 5 gms. % to 6 . 7 5 gms. %. However, at the l e v e l s fed during the present experiment, no such differences i n t o t a l protein l e v e l s of plasma were evident between the two planes of n u t r i t i o n i n either J or H groups. 7 . Plasma albumins, globulins and fibrinogen The factors a f f e c t i n g the t o t a l protein of plasma also af f e c t the l e v e l s of the component fr a c t i o n s albumins, globulins and fibrinogen. However, d i f f e r e n t f r a c t i o n s respond d i f f e r e n t l y to various s t i m u l i . Thus a change i n one f r a c t i o n may cause a v a r i a t i o n i n t o t a l protein l e v e l s while the other f r a c t i o n s re-main constant. Tables XVII to XXII show that over a period of time, only albumins are s i g n i f i c a n t l y d i f f e r e n t i n both fawns and yearlings. Thus changes i n t o t a l protein associated with age are due mainly 73. to changes i n the albumin f r a c t i o n . This i s i n agreement with the work of Fischer, et a l , (1954), and Howe, (1925). Two other investigators, Gainer, (1952) and Braun, (1946) report that glo-bulins d i f f e r to a greater degree with increasing age than does albumin. However, i t appears that albumins are the more impor-tant variants i n the young animals while globulins play a more important r o l e i n increasing age of adult animals. Table XVIII shows that the difference between the l e v e l s of albumins i n the fawns and the yearlings i s s i g n i f i c a n t whereas no v a l i d d i f -ference i s shown for the globulins or fibrinogen. Under severe n u t r i t i o n a l stress i t has been noted that the t o t a l proteins of blood deorease. This i s due mainly to a decrease i n the albumin fraction- (Chakravarti, 1952; Howe, 1925; Xlos.terman, et, a l , 1950 and Weeoh, et_ a l , 1935). Since the deer were not subjeoted to severe n u t r i t i o n a l stress no v a l i d d i f -ference was shown between the planes of n u t r i t i o n with respect to t o t a l protein of blood. Accordingly there i s no s i g n i f i c a n t d i f -ference between the two planes of n u t r i t i o n i n terms of either albumins or globulins (Tables XVII to XX). However, Table XXII shows that a s i g n i f i c a n t difference d i d occur between the two planes with respect to fibrinogen l e v e l s i n H. group. Sinoe fibrinogen contributes r e l a t i v e l y l i t t l e nitrogen to the t o t a l protein nitrogen, t h i s difference i s obscured and does not af-fect the t o t a l protein l e v e l s s u f f i c i e n t l y to produce a d i f -ference between the planes of n u t r i t i o n . Foster, et, a l , (1921-22) showed that " f i b r i n " l e v e l s vary with a v a r i e t y of s t i m u l i i n -cluding the d i e t . Since the quantity of f i b r i n i s dependent on 74. TABLE PLASMA ALBUMIN IN GMS PROTEIN PER 100 ML. Date of blood sample July 5* J u l y 1£ July 22 Age i n days 20 27 37 High plane means 2.97 4.18 2.76 Low plane means 2.24 3.77 2.99 Means for a l l fawns 2.62 0.87 3.98 1.04 2.87 0.63 P 0.01 Range 1.60 - 4.46 2.7.3 - 6.01 1.97 - 4.01 *Serum. **Data of July 5 ommitted from c a l c u l a t i o n s . TABLE PLASMA ALBUMIN IN GMS PROTEIN PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 e i n days 392 402 414 High plane means 4.86 3.44 4.16 Low plane means 4.63 4.34 3.89 Means for a l l year-l i n g s 4.75 0.85 4.12 0.81 4.02 0.40 P 0.05 0.05 Range 4.07 - 6.20 3.44 - 5.12 3.62 - 4.62 75. XVII PLASMA FOR THE FAWNS (J GROUP) Aug. 3 Sept. 23 July 5-Sept.23** 49 100 20-100 3.24 3.74 3.48 0.95 3.34 3.26 3,36 0.76 3.29 0.73 3.46 0.41 3.42 0.84 0.05 0.05 2.15 - 4.54 2.73 - 3,86 1.60 - 6.01 0.05 XVIII PLASMA FOR THE YEARLINGS (H GROUP) J GROUP Sept. 23 July 12-Sept.23 P July 5-Sept, 23 465 392-465 20-100 4.81 4.46 0.82 3.48 0.95 0.05 4.27 4.28 0.58 3.36 0.76 4.49 0.43 4.36 0.67 0.01 3.42 0.84 0.05 3.99 - 5.09 3.44 - 6.20 1.60 - 6.01 76. TABLE PLASMA GLOBULINS IN GMS PROTEIN PER 100 ML. Late of blood sample July 5* July 12 July 22 Age i n days 20 27 37 High plane means 3.64 0.89 1.47 1.26 2.87 1.08 Low plane means 5.57 2.34 1*42 1.67 2.51 1.23 Means for a l l fawns 4.69 1.74 1.44 1.47 2.69 1.11 P 0.05 Ranges 2.01 - 8.94 0.10 - 4.14 1.21 - 5.13 *Serum. **Data of July 5 not included i n calculations. TABLE PLASMA GLOBULINS IN GMS PROTEIN PER 100 ML. Bate of blood sample July 12 July 22 Aug. 3 Age in days 392 402 414 High plane means 0.86 1.09 2.22 0.73 1.65 0.44 Low plane means 2.06 2.01 2.22 0.49 2.21 0.77 Means for a l l year-lings 1.46 1.43 2.22 0.56 1.93 0.56 P 0,05 0.05 Ranges 0.04 - 4.39 1.40 - 2.81 1.25 - 2.95 77, XIX PLASMA FOR THE FAWNS (J GROUP) Aug. 3 Sept. 23 July 5-Sept.23** P 49 100 20-100 2.11 1.22 0.92 0,41 2.05 1.24 0.05 2.31 1.36 1,21 0.41 2.07 1.30 2.21 1.24 1.08 0.38 2.06 1.26 0.05 0.05 0.23 - 3.75 0.48 - 1.88 0,10 - 8,94 XX PLASMA FOR THE YEARLINGS (H GROUP) J 6R0PP Sept, 23 July 12-Sept.23 P July 5-Sept. 23 465 392-465 20-100 1.09 0.36 1.49 0.83 2.05 1.24 0.05 1.76 0.38 2.06 0.97 2.07 1.30 1.49 0,32 1.79 0.89 0.05 2.06 1.26 0.05 0.83 - 2.20 0.04 - 4.39 0.10 - 8.94 78, TABLE BLOOD FIBRINOGEN IN GMS PROTEIN PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age i n days 27 37 49 High plane means 0.473 0.053 0.447 0.117 0.358 0.121 Low plane means 0.369 0.108 0.474 0.164 0.446 0.271 Means for a l l fawns 0.407 0.088 0.460 0.129 0.398 0.191 P 0.05 0.05 Ranges 0.277 - 0.537 0.315 - 0.803 0.170 -0.870 TABLE FIBRINOGEN IN GMS PROTEIN PER 100 ML. Date of blood sample July 12 July 22 Aug. 3 Age in days 392 402 414 High plane means . 0.315 0.056 0.288 0.036 0.270 0.046 Low plane means 0.468 0.085 0.349 0.103 0.340 0.053 Means for a l l year-lings 0,391 0.064 0.319 0.068 0.305 0.044 P 0.05 0.05 Ranges 0.270 - 0.553 0.240 - 0,441 0,230- 0.400 79 XXI BLOOD FOR THE FAWNS (J GROUP) Sept. 23 July 12-Sept. 23 100 0.435 0.107 0.344 0.138 0.584 0.110 0.05 0.235 - 0.558 27-100 0.425 0.104 0.05 0,410 0.170 0.417 0.151 0.170 - 0.870 XXII BLOOD FOR THE YEARLINGS (H GROUP) ' 3 GROUP Sept. 23 July 12-Sept, 23 P Jul y 12 - Sept.23 465 392-465 27-100 0.277 0.086 0.288 0.048 0.425 0.104 0.01 0.478 0.264 0.409 0.144 0,410 0.170 0.398 0.196 0.351 0.107 0.05 0,417 0.151 0,05 0.196 - 0.719 0.196 - 0.719 0.170 - 0.870 80. the amount of fibrinogen present, as well as other factors, the effect of diet cannot be c l e a r l y depicted. However, Table XXII indicates that the fibrinogen l e v e l i s higher i n the low plane group than i n the high plane deer. 81. Summary and Conclusions Two groups of Columbian b l a c k - t a i l e d deer were raised i n c a p t i v i t y on two d i f f e r e n t planes o f n u t r i t i o n . One group had reached the pubertal break i n growth on an ad l i b d i e t p r i o r to the inception of the planes of n u t r i t i o n whereas the other group was separated on the two planes from shortly a f t e r b i r t h to 168 days of age. A fat-added, synthetic milk diet similar to doe's milk was found to produce rapid growth In the high plane fawns and low instantaneous r e l a t i v e growth rates i n the low plane fawns. However, growth i n both planes was retarded during the suckling period due to f a i l u r e to i n c i t e rapid growth o f the rumen micro-organisms. A p e l l e t e d r a t i o n designed to meet a l l the n u t r i t i o n a l requirements of the deer was used during the post-weaning growth period. The d a i l y food-ration was estimated, on an iso c a l o r i c basis from an average expression o f basal metabolism, (basal meta-bolism a 70.5 x Wb Cal/kgm./day), The high plane deer were fed 75% of the estimated maximal food intake (70.5 x x 5) and the low plane deer were fed 60% of the high plane r a t i o n (45% of maximal calculated intake). Growth rates i n terms of both weight and l i n e a l dimensions were s i g n i f i c a n t l y d i f f e r e n t as a r e s u l t of the two planes of n u t r i t i o n . However, the difference was smaller i n the case of the suckling fawns, due probably to using the incorrect value of "b n. The value of "b? was Interpreted as 0.67 during the 8 2 . suckling period and 0.73 during the post-weaning growth period. However, digestive d i f f i c u l t i e s and the f a i l u r e to consume the calculated d a i l y rations proved that the value of "b" i s l e s s than 0.67 during the suckling period and should probably have been approximately 0.67 from weaning to about one year of age. A value of 0.73 was found to be accurate i n the case of a 120 l b . deer, suggesting that maximal food energy consumption may be accurately estimated by the formula 70.5 x w0'75 x 5 Metabolizable energy of the r a t i o n for a deer of 100-plus l b s . S i m i l a r l y , the very slow growth rate of the low plane deer during the post-weaning period showed that 70.5 x W x 2  Metabolizable energy of the r a t i o n may be used to adequately represent a maintenance l e v e l of n u t r i t i o n . Instantaneous r e l a t i v e growth rates varied from 10% shortly a f t e r b i r t h to 0.03% following weaning thus forming f i v e phases of l i n e a r i t y i n the age-weight growth curves. In general the plane o f n u t r i t i o n affected a l l phases of growth and the growth rates were lower for the low plane deer. Linear measurements, such as heart-girth, height-at-withers and length of hind leg i n r e l a t i o n to age, were treated as being l i n e a r throughout the experimental period. Thus equa-tions were calculated for each measurement using the formula Y=ax + b. It was shown that the plane of n u t r i t i o n affected each of these measurements to some degree as was indicated by 83. differences i n the slope of the l i n e s representing the regression of the l i n e a r measurement against age. Thus heart-girth was affected i n a s i m i l a r manner to body weight whereas length of hind leg-age regressions showed l i t t l e change between the planes of n u t r i t i o n . These re l a t i o n s h i p s were used i n the regression of l i n e a r measurements against body weight. Thus heart-girth was shown to be a useful measurement i n estimating body weight while length of hind leg can be used to determine the condition etc degree of fleshiness of the animal. A. method for the d i r e c t and graphical evaluation of con-d i t i o n was proposed on the basis of the difference i n estimated weights using heart-girth and hind l e g length as c r i t e r i a . Beer of good to excellent condition tend to have an index of condi-t i o n greater than 1.0 whereas animals i n poor condition have an index of condition of l e s s than 1.0. This method of evaluation should prove usef u l i n comparisons of the n u t r i t i v e conditions of various areas as. r e f l e c t e d by the conditions of the deer l i v i n g within that area. Analyses of several blood constituents were car r i e d out i n r e l a t i o n to growth and to the planes of n u t r i t i o n i n order to provide standards for future comparisons. Ho differences i n the l e v e l s of blood constituents were discernable i n the fawns (J group) due to the small difference between the two actual planes of food consumption. However, data f o r the yearlings (H group) indicated that blood glucose l e v e l s and fibrinogen were affected s i g n i f i c a n t l y by the two planes of n u t r i t i o n . The other deter-minations showed no difference betv/een the two planes, but i t i s 84. suggested that a plane of n u t r i t i o n below the maintenance l e v e l would undoubtedly affeot the l e v e l s of a l l constituents. Addi-t i o n a l l y , s p e c i f i c d e f i c i e n c i e s would also cause changes i n the l e v e l s o f ce r t a i n blood constituents. Packed c e l l volume, sedimentation rates, hemoglobin, glu-cose, N.P.N, t o t a l p r o t e i n fibrinogen and albumin l e v e l s of blood were shown to vary with age. Only the globu l i n f r a c t i o n of blood remained constant throughout the development of the fawns. Sea-sons for the changes i n blood l e v e l s of these constituents were suggested. Seasonal changes were noted i n the l e v e l s of hemoglobin and glucose. It was suggested that suitable conditions for rapid growth ocour during the summer months which are not asso-ciated purely with the plane of n u t r i t i o n . Thus hemoglobin and glucose l e v e l s are elevated during the summer. The decrease noted i n early f a l l i s thought to be associated with hormonal regulation and sexual a c t i v i t y since a voluntary reduction of food consumption was also noted at the inception of sexual be-haviour. In general, the blood constituents fluctuate s l i g h t l y i n order to minimize environmental s t r e s s . 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( i ) T a b l e i ^ r f The composition o f U.B.C. r a t i o n #15-52. Ingredients. l b s . A l f a l f a meal* 460 Whole corn. 640 Wheat bran. 280 Beet pulp. 200 Molasses. 140 Soybean meal. 140 Copra (coconut o i l meal) 100 Calcium Carbonate. 20 Sa l t - i o d i z e d . 20 Calcium limestone. 20 Total weight 2020 This r a t i o n was fed as a l / 5 " dry p e l l e t . T a b l e ^ i Y The approximate composition of U.B.C. r a t i o n #15-52. % of yet weight. % of dry weight. Moisture. ! 10.4 Protein. 15.4 17.3 Fat. 4.0 4.5 Fibre. 8.4 9.5 N.F.E. 53.6 60.0 Ash. 7.7 8.7 X X V Table.TV. The approximate composition o f P a c i f i c Evaporated Milk and Nurse Cow replacement. P a c i f i c Evap. Milk Nurse Cow Protein. — - 25$ Fat. 7.8% 20% Fibre. 1.1 Total s o l i d s . 25.5% TableVV.. The composition of doe's milk* ( K i t t s et a l , 1955). Sample I Sample II' Total s o l i d s . 25.08 1 22.04 Water. 74.92 1 77.96 Fat. 10.50 10.50 Protein. 9.56 8.96 Lactose. 3.91 4.67 Ash. 1.56 1.30 1. Constituents as percentages of whole milk. 2. Milk drawn from the doe before death. 3. Milk drawn from the doe immediately a f t e r death by excis i o n ( i i ) of the udder. 4. The fawn was born on May 27, 1954 and the milk was drawn on June 19, 1954. Table.... The dressing percentage o f experimental deer based on bled and eviscerated carcasses with the head and hide remaining attached. l i v e weight (lbs.) Bled and Dressing % Evisoerated Wt.flbs.) 27.6 19.5 70.65 32.0 22.0 68.75 41.1 31.0 75.61 46.5 35.0 75.27 83.0 65.5 78.91 84.0 70.5 71.43 102.5 80.5 78.54 111.0 85.5 77.02 148.0 114.0 77.02 In s u f f i c i e n t data precludes representation i n graphic form. Dressing percentage tends to increase with increasing weight but varies depending upon the condition of the deer. Some observable differences i n response to the plane of  n u t r i t i o n . A. H-group. 1. Pellage. Shortly a f t e r the commencement of the two planes of n u t r i t i o n , the hair on both sides of the low plane deer broke off so that the proximal grey portion was l e f t . l a t e r i n the summer of 1954 these short grey hairs became sparse and minute lesions appeared on the s k i n . The pellage i n the high plane group remained normal. 2. Antler development f i i i ) Antler development was retarded i n the low plane group. At the conclusion o f the experiment, the low plane male deer possessed spike-antlers of approximately two inches i n length. Beer number 3H had forked antlers approximately nine inches i n length, at the end of the experiment. The other high plane deer (EH) broke the growing t i p s early i n the summer but s t i l l produced three large, s i x inch spikes f o r each antler, by the time i t died of a broken neck. Thus, the plane of n u t r i t i o n oan markedly a f f e c t antler development. However, the mechanisms which allow normal s k e l e t a l development but poor a n t l e r growth are not understood and further work i s necessary to c l e a r t h i s point. It "would appear that n u t r i t i o n may a f f e c t hormonal regu-l a t i o n and thereby the growth of a n t l e r s . 3. A c t i v i t y and Behaviour. The plane of n u t r i t i o n markedly affected behaviour and a c t i v i t y . In general, the high plane deer were more vigorous and active and l e s s nervous than the low plane deer. 4. Moult. Accurate records o f the beginning of moulting were not kept for H group. It was apparent, however, that moulting ooourred much l a t e r i n the low plane deer than i n the high plane group. B. J-group. 1. Pellage. During the suckling period, the pellage of the low plane fawns tended to be dry and without l u s t e r . A f t e r weaning (iv) these c h a r a c t e r i s t i c s became more pronounced g i v i n g a " f l u f f y " appearance to the head and body. Some of the low plane animals started to lose h a i r from the inside of the front legs p r i o r to the conclusion of the experiment. In contrast, the high plane fawns possessed an abundant pellage with a high l u s t e r . 2. Antler development. Antler development was f i r s t noted on September 28/54. In general, the high plane fawns had v i s i b l e a n t l e r but-tons at t h i s time while the low plane fawns had buttons which could be f e l t but not seen. Antler development was probably retarded i n a l l of J-group due to the low growth response i n comparison to doe raised fawns. 3. A c t i v i t y and behaviour. The behavioural differences between the two planes of n u t r i t i o n became very apparent aft e r weaning and was s i m i l a r to that reported for H-group. However, a greater degree of nervousness was evident i n the low plane fawns which may have been due to d e f i c i e n c i e s i n the vitamin B complex. 4. Moult. Moulting started i n most of J-group fawns i n e a r l y September and was p r a c t i c a l l y complete i n 6J, 7*7, 10J, 14J and 15J by September 30. Fawns numbered 2J, 3J and 4J had started to moult at t h i s time but the fawn-spots were s t i l l d i s t i n c t l y v i s i b l e . This was thought to be due to i l l n e s s which retarded moult i n these three fawns, Due to s i m i l a r i t y i n the plane of f v ) n u t r i t i o n between high and low planes at the time of weaning/ there was no observable difference i n the time and rate of moult between the planes. 5. Sexual a c t i v i t y . " Deer S3 and 5J showed a marked sexual i n t e r e s t i n the high plane female, 15J. However, none of the low plane male fawns showed any interest i n her, nor did the high plane males show any intere s t i n the low plane female, 14J. This tends to suggest that sexual maturity had been reached by the high plane males and females but not by the low plane deer. On December 4/54, fawn 15J had a ripe Graafian f o l l i c l e i n the right ovary. The low plane female 14J showed no such a c t i v i t y of the ovaries. I l l n e s s , symptoms, treatment and response. 1. A discharge from the eyes occurred i n a number of fawns due possibly to splashing milk on the eye while learning to bottle-feed. The eye l i d s were bathed with opthalmic boric acid and opthalmic aureomtoin ointment was rubbed on the eyelids. This treatment soon oleared the condition. 2. Scouring probably due to n u t r i t i o n a l disturbances was encountered, p a r t i c u l a r l y before the f a t content of the milk was increased, and while the evaporated milk was s t i l l being d i l u t e d . During scouring, the fawns became weak and l o s t t h e i r appetite. Milk was forced down the throat with a pipette and doses of crude aureomycin tablets were administered, depending upon the size of the animal. I f the fawns weakened from scouring and l o s s fvi) of appetite, a heat lamp was supplied. When severely weakened an attempt was made to restore the water balance and provide energy by the administration of 10% glucose by intravenous d r i p . For example 6J was treated i n t h i s manner oh June 18 and the fawn survived to the end of the experiment. In some cases, glucose was administered o r a l l y i n solution* This treatment was used when the fawns showed l o s s of appetite but were not serious-l y weakened. 3. Magnesium d e f i c i e n c i e s were encountered i n 16J and 18J. Slight muscular seizures were apparent and the head was thrown backwards over the neck or held to one side. The fawns were given 10 oo of Ringer's s o l u t i o n containing 0.1% magnesium ohioride. Both fawns displayed normal conditions within three hours a f t e r administration. 4. Blockage of the p y l o r i c sphincter occurred i n a num-ber of the fawns due possibly to the high fat content of the milk or due to foroe feeding at a high l e V e l . The symptoms of t h i s blockage were aocompanled by a bloated appearance of the v i s c e r a which sometimes caused the flanks to be stretched very t i g h t l y . A d d i t i o n a l l y the fawns con t i n u a l l y ground t h e i r teeth and would not accept the b o t t l e when attempting to feed them. In advanced stages a mucous-like exoretion replaced the normal feces showing that no food was leaving the stomach. It was found that massaging the v i s c e r a l region with the hand, using a strong pressure from the diaphragm p o s t e r i o r l y , caused the food to pass through the p y l o r i c sphincter, thereby reducing the discomfort, (Vii) 5. Vitamin B d e f i c i e n c i e s were suspected i n a number of the fawns although no t y p i c a l symptoms were apparent u n t i l August 27/54 when 10J showed c h a r a c t e r i s t i c symptoms of pantothenic acid deficiency. Symptoms consisted of a p a r t i a l p a r a l y s i s of the l e f t side and a "goose-stepping" motion of the l e f t hind l e g while walking. The head was held i n a t i l t e d p o s i t i o n to the right and the fawn showed extreme nervousness. One ml. of a mixture of B-vitamins (Solu-Zyme) was administered by intra-?muscular i n j e c t i o n . Symptoms had com-p l e t e l y disappeared s i x days a f t e r treatment. 6. H-group fawns developed a bronchial congestion during the winter of 1954-54. A rasping cough was successfully treated with aureomycin-triple-sulfa tablets, the dosage being deter-mined by the size of the deer, as s p e c i f i e d by the manufacturer. \ 

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