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Studies on the Growth and Compositional Development

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Studies on the Growth and Compositional Development STUDIES ON THE GROWTH AND COMPOSITIONAL DEVELOPMENT OF ANTLERS IN RED DEER (CERVUS ELAPHUS) A thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy in the University of Canterbury by Paul David Muir Li nco 1n Co 11 ege 1985 Abstract of a thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy. STUDIES ON THE GROWTH AND COMPOSITIONAL DEVELOPMENT OF ANTLERS IN RED DEER (CERVUS ELAPHUS) by P.D. Muir The experiments described in this thesis investigated nutritional and physiological aspects of antler growth in red deer stags. The initial experiment (Section 3) examined the effects of winter nutrition on subsequent antler casting date and velvet antler weight. Mature stags on two different farm types (hill country, Farm H and irrigated lowland, Farm L) were offered three levels of winter nutrition, two levels of a concentrate supplement (~ libitum pellets and 1/2 ~ libitum pellets) and a basal hay ration. On both properties liveweight gains occurred in supplemented groups and liveweight losses in unsupplemented groups. At antler casting there were significant differences in liveweight of approximately 10 kg between fully supplemented and unsupplemented groups. Realimentation of winter liveweight losses subsequently occurred so that by the following rut the effects of winter undernutrition had been eliminated. On Farm H poor winter nutrition (hay only) resulted in a significant delay in casting date (13 days) and lower velvet antler yields (0.24 kg), than in stags offered the ad libitum concentrate ration. Stags on Farm H were 13 kg lighter at commencement of the trial than at Farm L and the differences in treatment effects obtained between farms may have been due to differences in body condition at commencement of the trial. An association was demonstrated between liveweight and date of antler casting, with heavier stags casting earlier than lighter stags. There was no effect of age of stag on casting date. Of the liveweights recorded, liveweight prior to the rut showed the best relationship with casting date, possibly because the seasonal nature of liveweight change meant that a weight recorded at this time gave the best indicator of the true frame size of a stag. Both age and liveweight significantly affected velvet antler weight, with increases of velvet antler weight of 0.26 kg between 3 and 4-year-old stags and of 0.30 kg between 4 and 5-year-old stags at the same liveweight. Within an age group velvet weight increased by 0.12 kg for each 10 kg increase in pre-rut liveweight. The experiments described in Section 4 comprised studies on antler growth and composition. In order to obtain data on antler growth and composition individual antlers were removed sequentially from mature red deer stags between 28 and 112 days after casting of hard antlers. Contralateral antlers were removed after stripping of velvet. Wide variation occurred in antler casting date (53 days) compared to date of velvet stripping (24 days). The duration of the period of antler growth may therefore be governed more by date of casting than by date of velvet stripping. Mean duration of the antler growth period was 164 days. Growth in length of the antler appeared to follow a sigmoid curve. However, between 28 and 112 days after casting, rates of elongation were close to linear. Mean length of hard stripped antlers was 0.71 m and between 28 to 112 days after casting mean rate of antler elongation was 0.62 cm/day. Over this period indivdual antlers increased in fresh weight at a rate of 13.7 g/d, with heaviest weight recorded 112 days after antler casting, at approximately 130% of final hard antler weight. Between 28 and 91 days of growth, volume of blood in the antler increased linearly at a rate of 194 ml/kg. Three phases of mineralization were demonstrated in developing antlers. Tips of growing antlers were cartilaginous and poorly mineralized. A zone of mineralization occurred 5.0 to 7.5 cm behind the antler tip which corresponded histologically to the transition from mineralized cartilage to trabecular bone. The second phase of mineralization occurred through continued accretion of trabecular bone in the antler shaft. The third phase, described as "terminal mineralization" in this study, appeared to be associated with a rapid increase in density of cortical bone in the periphery of the antler shaft. Terminal mineralization (between 91 and 112 days after casting of hard antlers) coincided with the slowing of growth in length, a decrease in relative blood volume in the antler and an increase in levels of plasma testosterone. These events occurred close to the summer solstice. At velvet stripping individual antlers had a mean weight of 1.12 kg and contained 81.1% dry matter (OM). Fat free organic matter (FFOM) and ash concentration in OM were 36.6 and 60.0%, respectively. Peak daily rates of FFOM and ash deposition occurred between 91 days and 112 days after casting, at rates of 1.4% of hard antler FFOM and 1.6% of hard antler ash. For a stag producing 2.24 kg of hard antler mean rates of FFOM and ash deposition over this period were 9.3 and 18.3 g/d, respectively. On a whole antler basis calcium concentration in antler ash remained constant, at around 35%. Therefore peak rate of antler calcium deposition would be 6.4 g/d. In the final experiment (Section 5) mature stags were offered a maintenance ration of greenfeed oats during the period of peak calcium requirement for antler growth and the kinetics of calcium metabolism were examined using a radio-isotope (45Ca ). Rates of faecal endogenous loss were low and at approximately 6.4 mg/kg BW per were half the estimated requirements of ARC (1980) for sheep and cattle. Availability of calcium from greenfeed oats was low (mean, 37%) and less than 30% of total calcium requirements were derived from the diet. Poorly mineralized skeletal bones indicated that the shortfall in antler calcium was derived from the skeleton. In spite of a severely negative calcium balance stags were capable of maintaining high and apparently normal rates of antler calcium deposition (mean, 44 mg/kg BW per day). Antlers appear to be acting as a sink with calcium being irreversibly deposited in the antler and lost to the animal's body. On the assumption therefore that antler calcium behaves like calcium lost during lactation a kinetic model of calcium metabolism in the stag was developed. TABLE OF CONTENTS Page SECTION 1 - Introduction 1 SECTION 2 - Literature review 2 Introduction 2 Antler growth and development 2 1) Pedicles 2 2) Antler casting 3 3) Antlet morphology 3 \~y) Early development 3 ,b) Ossification 5 \ " 'tJ Velvet 5 d) Blood supply 6 e) Innervation 6 '4) Sequence of antler development 6 5) Antler composition 7 a) Sources of minerals for growth of antler bone 9 Factors affecting antler growth 9 1) Age 9 2) Nutrition 10 a) Pedicles 10 b) Casting 10 c) Antler growth 11 3) Photoperiod 12 4) Hormones 13 a) Androgens 13 b) Oestrogens 14 c) Other hormones 14 Page SECTION 3 - Effects of winter nutrition on antler development 16 Introduction 16 Experimental methods 17 Results 22 1) Liveweight changes 22 2) Feed intake 22 3) Casting date 25 4) Antler growth 27 Discussion 33 SECTION 4 - Antler growth and composition 38 Introduct ion 38 PART A 40 Experimental methods 40 1) Experimental outline 40 2) Radio-isotope preparation 40 a) 51Chromium labelled red blood cells (51Cr-RBC) 40 b) Iodinated human serum albumen (125I-HSA) 41 C) Injection and blood sampling procedure 41 3) Antler removal, measurement and sectioning 41 a) Antler removal 41 b) Antler measurements 41 c) Antler sectioning 42 4) Blood volume 42 5) Chemical analysis 45 6) Histology 45 Results and discussion 48 1) Antler measurements and sectioning 48 2) Blood volume 48 3) Mineralization 50 Conclusions 52 Page PAin B Experimental methods 53 1) Experimental outline 53 a) Nutritional treatments 53 b) Antler sampling 54 2) Antler removal, measurements and sectioning 55 3) Blood volume 56 4) Chemical analysis 56 a) Antler samples 56 b) Feed samples 57 5) Histology 57 6) Blood sampling"and testosterone assay 57 7) Data analysis 58 Results 59 1 ) Effects of treatment 59 2) Growth of the antler 59 a) Antler casting to velvet stripping 59 b) Tine development 62 c) Increase in length 62 d) Increase in weight 65 e) Antler regrowth 65 3) Antler blood volume 67 a) Distribution in the antler shaft and brow tine 67 b) Relative blood volume in the antler shaft and brow tine 69 c) Relative blood volume in the whole antler 69 4) Antler mineralization 69 a) Antler shaft 69 b) Brow tine 77 c) Total mineralization in the antler shaft , brow tine and whole antler 78 d) Rate of organic matrix deposition 80 e) Rate of ash deposition 82 Page 5) Antler histology 82 a) Growth of the antler tip 82 b) Zone of mineralization 85 c) Mineralization of the antler shaft 85 6) Plasma testosterone 88 Discussion 91 1) Treatment effects 91 2) Growth of the antler 92 3) Antler blood volume 94 4) Antler mineralization 96 5) Mineral requirements for antler growth 100 SECTION 5 - Calcium metabolism in stags during peak antler mineralization 102 Introduction 102 Experimental methods 105 1) General 105 2) Low calcium diets 105 3) Balance Trials 107 4) Sample preparation and analysis 108 5) Radio-isotope counting 108 6) Data analyses 109 7) Model of calcium metabolism 110 Results 112 1) Nutrient intake 112 2) Antler composition 114 3) Calcium concentration in plasma and rates of excretion of calcium in urine and faeces 115 4) Faecal endogenous loss and rate of calcium absorption 117 5) Skeletal composition 117 Discussion 119 1) Antler mineralization 119 2) Plasma calcium 121 3) Faecal endogenous loss and urinary excretion of calcium 121 4) Calcium absorption 122 5) Model of calcium metabolism in the stag 123 Page SECTION 6 - General discussion and summary 129 Practical aspects and conclusions 134 ACKNOWLEDGEMENTS 137 REFERENCES 139 1 SECTION 1 I nt roduct ion Red deer (Cervus elaphus) have been farmed in New Zealand since 1969 (Coop and Lamming, 1976), producing high per hectare returns from antlers, venison and sale of live animals.
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