Zurich Open Repository nd Archive University of Zurich Min Lirry Strickhofstrsse 39 CH-8057 Zurich www.zor.uzh.ch Yer: 2014 Differences in periprtl plsm prmeters relted to clcium homeostsis of diry sheep nd gots in comprison with cows Wilkens, Mirj R; Liesegng, Annette; Richter, Juli; Frser, Dvid R; Breves, Gerhrd; Schröder, Bernd DOI: https://doi.org/10.1017/s002202991400020x Posted t the Zurich Open Repository nd Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-100531 Journl Article Accepted Version Originlly pulished t: Wilkens, Mirj R; Liesegng, Annette; Richter, Juli; Frser, Dvid R; Breves, Gerhrd; Schröder, Bernd (2014). Differences in periprtl plsm prmeters relted to clcium homeostsis of diry sheep nd gots in comprison with cows. Journl of Diry Reserch, 81(3):325-332. DOI: https://doi.org/10.1017/s002202991400020x
1 2 3 4 5 6 7 8 9 10 11 Differences in periprtl plsm prmeters relted to clcium homeostsis of diry sheep nd gots in comprison with cows Mirj R. Wilkens *, Annette Liesegng,c, Juli Richter, Dvid R. Frser d, Gerhrd Breves, Bernd Schröder Deprtment of Physiology, University of Veterinry Medicine Hnnover, Foundtion, Hnnover, Germny Institute of Animl Nutrition, Vetsuisse Fculty Zurich, University of Zurich, Zurich, Switzerlnd c Center for Applied Biotechnology nd Moleculr Medicine, University of Zurich, Zurich, Switzerlnd d Fculty of Veterinry Science, University of Sydney, Sydney, Austrli 12 13 14 15 16 17 18 19 * Corresponding uthor: Mirj R. Wilkens, Deprtment of Physiology, University of Veterinry Medicine Hnnover, Bischofsholer Dmm 15/102, 30173 Hnnover, Germny. Tel.: + 49 511 856 7628; fx: + 49 511 856 7687. E mil ddress: mirj.wilkens@tiho-hnnover.de 20 21 22 Key words: trnsition; clcium homeostsis; close-up; dry period; ruminnts; one metolism; 23 24 Short title: Periprtl clcium homeostsis in ruminnts 25 26 1
27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 Summry: Recently it hs een demonstrted tht there re differences etween sheep nd gots in respect to the dpttion to clcium restricted diet. It ws the im of the present study to evlute whether species-specific peculirities lso occur when clcium homeostsis is chllenged y lcttion. Therefore, we investigted the time courses of plsm prmeters relted to clcium homeostsis (clcium, phosphte, clcitriol, the one resorption mrker CrossLps nd the one formtion mrker osteoclcin) during the trnsition period in multiprous nimls of oth species nd compred the results to dt from former study crried out with diry cows. Like in cows, plsm clcium nd the rtio of one formtion to one resorption decresed t prturition in gots while plsm clcitriol incresed. On the 10 th dy postprtum the one prmeters of gots reched preprtum vlues gin which ws not the cse in cows. Sheep were found to experience chllenge of clcium homeostsis lredy 10 dys efore prturition, reflected y very low rtio of one formtion to one resorption which ws not ccompnied y n increse in plsm clcitriol. Additionlly, sheep nd gots which hd een in milk for 3 months were smpled, dried-off nd smpled gin 6 weeks lter. In dried-off nimls there were no differences in prmeters of one metolism detectle. In conclusion we could show tht the contriution of one moilistion to the compenstion for the enhnced clcium demnd due to lcttion differs etween the three ruminnt species. 45 46 2
47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 Introduction Smll ruminnts re often used s models for cows to study sic spects of clcium (C) homeostsis. However, there re differences in homeosttic mechnisms etween gots nd sheep tht need to e considered efore ny results cn e extrpolted to diry cows, especilly with respect to the trnsition period. On ody weight sis the milk yield of diry gots is comprle to tht of diry cows nd hypoclcemi occurs in oth species usully t the onset of lcttion, while sheep develop hypoclcemi more often during lte gesttion (Oetzel, 1988). But when compring feeding ehviour, cttle nd sheep cn e regrded s grzers, while gots in their nturl hitt select high energy (concentrte) feed (Hofmnn, 1989). These differences in feed type might influence gstrointestinl minerl sorption s it hs een shown tht gots dpt more efficiently to dietry C restriction y up-regulting intestinl C trnsport thn sheep (Wilkens et l., 2011; Wilkens et l., 2012). Interestingly, it hs lso een reported tht reduction in C supply did not led to n increse in pprent digestiility of C in lctting cows (Tylor et l., 2008). Comprtive dt on the response of different ruminnt species to the chllenge of C homeosttic mechnisms t the onset of lcttion re limited. In studies investigting one metolism in gots nd sheep it hs een demonstrted tht round prturition sheep tend to hve greter concentrtions of one resorption mrkers nd more pronounced depression of one formtion mrkers in lood, compred to gots (Liesegng et l., 2006, 2007). As milk fever in diry cows is ssocited very closely with prturition, it ws the im of the present study to investigte chnges in plsm prmeters of C homeostsis during the periprtl period of sheep nd gots y tking smples in shorter intervls thn it hs een done in the studies mentioned ove nd compre them to respective dt from cows. Bsed on our former studies we hypothesise tht there re differences in the response to the periprtl chllenge of C homeostsis which cnnot e simply explined with the higher milk yield 72 usully oserved in gots nd cows. 3
73 74 75 76 77 78 79 80 Besides ionised nd totl C nd phosphte plsm, concentrtions of clcitriol, the iologiclly ctive metolite of vitmin D, were determined. Furthermore, the one mrkers were ddressed. Bone turnover or remodelling mens concurrent one formtion nd resorption (Allen, 2003). As solute vlues of iochemicl mrkers of one metolism re influenced y vrious fctors like ge, circdin rhythms nd seson (Hnnon & Estell, 2000), the rtio of osteoclcin to CrossLps ws used to ssess one turnover. The rtio of one formtion mrkers to one resorption mrkers hs een used efore to estimte the sttus of one remodelling in horses nd cows (Lepge et l., 1998; Liesegng et l., 2000). 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 Mteril nd methods Animls, diets, milking pttern nd lood smpling The study ws crried out with 5 to 6 multiprous nimls of ech species (sheep, gots nd cows), ged etween 2 nd 5 yers. All nimls were of reeds used for milk production in Centrl Europe: Est Friesin diry sheep, Snen type gots nd Holstein-Friesin cows. Averge ody weights were 66.3 ± 1.6 kg (sheep), 49.1 ± 1.7 kg (gots) nd 679 ± 25 kg (cows). Sheep nd gots were housed in free-stll rn t the Deprtment of Physiology of the University of Veterinry Medicine, Foundtion, Hnover, Germny. During the entire oservtion period, the nimls hd ccess to hy (Tle 1), minerls nd wter d liitum. The nimls were group-fed preprtum, receiving 75 g concentrte (Tle 1) per 10 kg ody weight per dy. Immeditely fter prturition, the kids nd lms were removed nd milk yield ws recorded for 20 consecutive dys. The ewes nd does were milked nd provided with concentrte three times dily t 0800, 1400 nd 2000 h. The mount of concentrte ws clculted for ech individul niml ccording to the milk yield of the previous milking (800 g per kg milk) to meet the requirements for diry sheep nd gots recommended y the Society of Nutrition 98 Physiology (GfE). 4
99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 Dt from cows were otined from previous study (Wilkens et l., 2012). The cows were kept on commercil diry frm locted in Sxony, Germny. They were fed totl mixed rtion d liitum efore nd fter prturition (C content.p.: 7.8 g/kg DM; C content p.p.: 10.0 g/kg DM). The clves were removed within 24 h postprtum, fter which milking ws done twice dily. In the present study, the periprtl plsm prmeters of 6 cows of the former control group were compred with respective vlues from the smll ruminnts. From clculted gesttionl dy 140 (sheep nd gots) or dy 175 (cows), two lood smples (lithium heprin nd EDTA) were collected from the jugulr (sheep nd gots) or coccygel vein (cows), every other dy t 0900 h until dy 10 postprtum. Additionl smples were tken immeditely fter prturition nd t 6, 12 nd 24 h postprtum. The preprtum smples were re-clssified ccording to the exct dy of prturition. Smples tken on dy 2 nd 1 preprtum were clssified s smple dy -2, smples tken on dy 4 nd 3 s smple dy -4, etc. A further 6 gots nd 5 sheep, of the sme ge nd reed were smpled fter three months of lcttion. They were then dried off nd smpled gin six weeks lter. The protocol of niml tretment ws pproved y the Animl Welfre Commissioner of the University of Veterinry Medicine, Foundtion, Hnover, Germny, nd its conduct ws supervised ccording to Germn Animl Welfre Lw. 116 117 118 119 120 121 122 123 Smple nlysis After mesuring ionised C in whole lood with lood gs nlyser (Chiron Dignostics, RpidL TM 348), smples were centrifuged (2,000 g, 15 min) t mient temperture nd plsm ws collected nd stored t -18 C. The concentrtions of phosphte nd totl C in plsm of sheep nd gots were mesured colorimetriclly y stndrd spectrometric techniques (Srkr & Chuhn, 1967; Kruse-Jrres, 1979), while totl C nd phosphte concentrtions in the plsm smples from cows were 5
124 125 126 127 128 129 130 131 determined y stndrd dignostic methods in the lortory of the Clinic for Cttle of the University of Veterinry Medicine, Foundtion, in Hnnover. Plsm clcitriol of sheep nd gots ws mesured with commercil rdioreceptor ssy (Immundignostik AG, Bensheim, Germny). Quntifiction of the one formtion mrker osteoclcin nd the one resorption mrker CrossLps ws done using commercil enzymelinked immunosorent ssy kits ccording to the mnufcturer s instructions (MicroVueTM Osteoclcin EIA Kit, Quidel Corportion, Snt Clr, USA; Serum CrossLps, IDS Ltd, Frnkfurt m Min, Germny). 132 133 134 135 136 137 138 139 140 141 142 143 Dt presenttion nd sttisticl nlysis Dt re presented s men ± SEM. The dily milk yield of sheep nd gots is corrected for the nimls ody weights. All sttisticl nlyses were performed using GrphPd Prism 5.0 (GrphPd Softwre, Sn Diego, USA). Gussin distriution ccording to Kolmogorov-Smirnov ws confirmed for ll dt. Comprison of periprtl ionised nd totl C, phosphte, CrossLps nd osteoclcin plsm concentrtions ws crried out y nlysis of vrince (ANOVA) for repeted mesurements with sturted model of the fixed fctor species. Bonferroni post-test ws pplied to revel species differences t ech time point nd differences over time within ech species. In ll cses, P vlues < 0.05 were considered sttisticlly significnt. 144 145 146 147 148 149 Results Milk yield Three of the 5 ewes nd 5 of the 6 does produced twins, while the remining smll ruminnts nd ll the cows hd single offspring. Milk yield ws not ffected y the numer of offspring. There ws no significnt difference in milk yield etween the sheep nd gots over the first few 6
150 151 152 dys fter prturition. However, dily milk production of the gots then incresed to mximum of 0.085 l/kg BW, wheres milk yield of the ewes remined more or less constnt t level of 0.04 l/kg BW per dy (Fig. 1). 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 Mcro minerls in lood nd plsm Whole lood nd plsm concentrtions of ionised nd totl C nd phosphte mesured in sheep, gots nd cows round prturition re given in Figures 2, 3 nd 4. Compred to dy 2 preprtum, ionised C mesured in whole lood of gots showed significnt decrese t prturition nd lso on the 10 th dy postprtum. There ws no similr chnge in ionised C ssocited with lming in ewes. The trnsient decrese oserved in cows with minimum t 12 hours postprtum ws not sttisticlly significnt ecuse of high individul vrinces, ut compred to sheep nd gots postprtum ionised C in cows ws significntly lower. However, the gret chllenge of C homeostsis in cows t prturition ecomes ovious when the plsm concentrtions of totl C re considered. In ll species, smll, ut not sttisticlly significnt decrese in plsm phosphte ws oserved t prturition. Intriguingly, pronounced increse in plsm phosphte ws found on the 10 th dy postprtum in gots. 168 169 170 171 172 Clcitriol The profile of plsm concentrtions showed tht in gots, clcitriol ws highest t prturition while sheep hd no significnt chnges nd cows showed the highest clcitriol levels on the 2 nd dy postprtum (Fig. 5). 173 174 Mrkers of one formtion nd resorption 7
175 176 177 178 179 180 The rtio of the one formtion mrker osteoclcin to the one resorption mrker CrossLps is given in Figure 6. In sheep plsm, very low rtio ws found throughout the entire oservtion period, while gots nd cows hd significntly greter rtios during the lst gesttionl dys which then decresed t prturition. On the 10 th dy postprtum sl vlues were found gin in gots, while the cows osteoclcin to Crosslps rtios remined s low s those of sheep. 181 182 183 184 185 186 187 188 189 190 191 Effects of terminting lcttion Comprisons of plsm concentrtions of totl C, phosphte, clcitriol nd the rtio of osteoclcin to CrossLps in ewes nd does fter three months of lcttion nd six weeks fter the end of lcttion re presented in Tle 2. Plsm concentrtions of totl C were not ffected y lcttion sttus. However, lower concentrtions were found in oth groups of sheep compred to gots. Both species showed smll increse in plsm phosphte fter lcttion hd cesed. While no effects of lcttion sttus were found with clcitriol concentrtions, the rtio of osteoclcin to CrossLps incresed mrkedly in oth species fter lcttion hd cesed. In contrst to periprtl vlues, there were no species differences in these one mrker rtios in non-lctting nimls. 192 193 194 195 196 197 198 199 200 Discussion There is sudden increse in the outflow of C from lood t the onset of lcttion compred to the lst dys of gesttion (Horst et l., 2005). The resulting trnsient decrese in plsm C t prturition nd shortly fter hs een descried for cows in numerous studies (Reinhrdt et l., 2011). The discrepncy etween ionised nd totl C in cows fter prturition in the present study could e explined y fctors cused y the negtive energy lnce lmost lwys reported for cows in erly lcttion (Butler & Smith, 1989). For exmple, ltertions of plsm lumin 8
201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 or ph might hve ffected the rtio etween ionised nd totl C. In contrst to the group fed diry cows, the smll ruminnts were feed individully ccording to their milk yield. The sence of such depression of C t prturition in sheep hs lredy een reported (Liesegng, 2008). This might suggest tht in contrst to gots nd cows there is no prticulr chllenge to the mechnisms of C homeostsis t lming due to the secretion of C in milk. However, s there were only minor differences in milk production etween sheep nd gots immeditely fter prturition, it is unlikely tht the C demnd for milk production during the first 24 hours p.p. ws very much higher in gots compred to sheep. The dily plcentl trnsfer in ewes etween dy 133 nd 140 of gesttion hs een shown to mount for 255 mg per kg foetl ody weight (Durnd et l., 1983). Assuming C concentrtion of 2.5 g/l in colostrum (Liesegng 2008), for the sheep in the present study the loss of C into the milk hs een pproximtely 4 g on the first dy p.p. compred to n estimted dily demnd of 2 g for lte gesttion. But in contrst to does nd cows, the osteoclcin to CrossLps rtio in ewes ws lredy very low t the end of gesttion. Since in non-lctting nimls there were no differences etween gots nd sheep, this low formtion to resorption mrker rtio is most proly not chrcteristic for sheep in generl nor cused y vritions in the sensitivities of the ELISA kits. The time course of the osteoclcin to CrossLps rtio round prturition more proly suggests tht in ewes the C homeosttic mechnisms hve lredy een chllenged efore lming nd there my hve een greter contriution of the skeleton to mintennce of C homeostsis thn in the other two ruminnt species. This is in line with the clinicl oservtion tht ewes often develop hypoclcemi during the lst dys efore lming when the foetl skeleton is minerlised (Oetzel, 1988) nd tht the periprtl plsm C concentrtions re lower in twin-ering thn in single-ering ewes (Roofi et l., 2013). However, ll of the gots except one in the present study lso produced twins. As it hs lredy een demonstrted tht gots hve greter ility to dpt to dietry C restriction thn sheep 9
227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 (Wilkens et l., 2012), it my e tht the does were dpted etter thn sheep to the enhnced C demnd of foetl one growth in lte pregnncy. The onset of lcttion induced smll decrese in ionised C ccompnied y concurrent increse in clcitriol in does. A decrese in ionised C leds to secretion of prthyroid hormone (PTH) which thn stimultes the renl 1α-hydroxylse, the enzyme needed for the trnsformtion of 25-hydroxyvitmin D to 1,25- dihydroxyvitmin D, the ctive hormone clcitriol (Frser & Kodicek, 1970). Assuming tht C homeostsis in ewes hd lredy een severely chllenged efore prturition, one would expect clcitriol response occurring even erlier in these nimls. However, this ws not the cse. Lower clcitriol concentrtions in sheep compred to gots experiencing the sme dietry C restriction hve lredy een demonstrted (Wilkens et l., 2010; Wilkens et l., 2012). As there is no suitle ssy ville for the determintion of PTH in smll ruminnt species, it is difficult to revel the resons for this phenomenon. In cows nd gots one resorption seemed to e induced not efore prturition. But while gots reched pre-kidding vlues t the 10 th dy postprtum gin, cows nd sheep s osteoclcin to CrossLps rtios remined very low. This might point to re more rpid stimultion of gstrointestinl C sorption in gots llowing the nimls do reduce one moilistion gin. For sheep it hs een shown tht the mximl efficiency of gstrointestinl C sorption ws reched 2 months fter prturition, lthough one resorption hd lredy een stimulted efore lming (Brithwite et l., 1970). Comprle lnce studies crried out with gots re not ville. In cows significnt increse of gstrointestinl C sorption ws found to tke plce one week postprtum (vn t Klooster, 1976) or etween the 10 th nd the 20 th dy fter prturition, respectively (Rmerg et l., 1970). It should e tken into ccount tht these dt were otined some time go when milk production in diry cows ws not s high s nowdys. Aginst the ckground of the low osteoclcin to CrossLps rtios found in cows on the 10 th dy postprtum it might e speculted tht one moilistion ws more prolonged in cows of 252 the present study. 10
253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 The reson for the slight decrese in plsm phosphte oserved in ll species is usully ssumed to represent loss of phosphte to the milk (Roertson et l., 1956). Furthermore, it hs een shown tht decrese in plsm C induces concurrent increse in cortisol. Cortisol ws demonstrted to hve n impct on plsm phosphte (Horst & Jorgensen, 1982). No stisfying explntion could e found for the increse in gots plsm phosphte concentrtions on the 10 th dy postprtum. As the lood concentrtion of ionised C showed second decrese on the 10 th dy, too, these ltertions might e ssocited with the steep rise in milk production oserved in does etween the 7 th nd the 12 th dy fter prturition. The role of phosphte in lctting nimls is not yet fully understood. In contrst to clcium, plsm concentrtions of phosphte re not tht strictly controlled nd the exchnge etween the different pools is more complicted. Phosphte is not only moilised from one, ut lso from soft tissues nd in ruminnts lrge mounts of phosphte secreted vi the slivry glnds nd resored gin in the gstrointestinl trct hve to e tken into ccount (Brithwite, 1983). However, the effect of drying-off on plsm phosphte underlines tht it might e influenced y the hormonl regultion mechnisms involved in lcttion nd one metolism. 268 269 270 271 272 273 274 275 Conclusion We could demonstrte tht in line with former studies sheep seem not to respond with n dequte stimultion of clcitriol synthesis to chllenges of C homeostsis. Furthermore, striking differences regrding periprtl one metolism were oserved etween the three ruminnt species investigted. To evlute whether sheep or gots re the etter model for studying C homeostsis in the diry cow, further studies under more controlled feeding regimes of the cows nd lrger numer of nimls re needed. 276 277 The finncil support of the Deutsche Forschungsgemeinschft (SCHR 342/8-2) is grtefully 278 cknowledged. 11
279 12
280 Tles 281 282 Tle 1 283 284 285 Ingredients of hy nd concentrte fed to sheep nd gots expressed s percentge of originl sustnce; n.d.: not detectle using stndrd procedures. 286 Hy Concentrte Crude protein 8.21 20.0 Crude ft 1.52 2.60 Crude fire 26.5 11.0 Crude sh 7.30 8.70 Clcium 0.41 0.90 Phosphorus 0.32 0.55 Sodium 0.11 0.20 Energy [MJ per kg] 7.50 10.2 Vitmin D [IU/kg] n.d. 1,000 287 288 13
289 Tle 2 290 291 292 293 294 Totl clcium (mmol/l), phosphte (mmol/l), clcitriol (pg/ml) nd the rtion etween osteoclcin (ng/ml) nd Crosslps (ng/ml) in plsm of gots nd sheep mesured fter 3 months in milk nd 6 weeks fter hving een dried-off, respectively. Results of 2wy ANOVA (species, effect of lcttion nd interction). Mens ± SEM. 295 Totl clcium Phosphte Clcitriol Rtio osteoclcin to CrossLps Gots Lctting (6) 2.56 ± 0.06 1.69 ± 0.09 12.8 ± 0.86 46 ± 5.8 Dried-off (6) 2.63 ± 0.03 1.87 ± 0.14 12.8 ± 0.82 62 ± 5.2 Sheep Lctting (5) 2.20 ± 0.06 1.59 ± 0.08 13.4 ± 3.35 35 ± 5.3 Dried-off (5) 2.18 ± 0.09 2.01 ± 0.16 8.1 ± 0.90 62 ± 12 ANOVA Species P < 0.001 n.s. n.s. n.s. Lcttion n.s. P < 0.05 n.s. P < 0.05 Interction n.s. n.s. n.s. n.s. 296 14
297 Figures 298 299 300 301 302 303 304 Figure 1 Dily milk yield of gots nd sheep normlised to the nimls ody weight. RM ANOVA reveled significnt effects of time (P < 0.001), species (P < 0.001) nd n interction of time nd species (P < 0.001). Significnt differences etween species reveled y Bonferroni posttest for ech dy re mrked with sterisks (P < 0.05: *; P < 0.01: **; P < 0.001: ***). Mens ± SEM. 305 306 307 308 309 310 311 Figure 2 Blood concentrtions of ionised clcium mesured in gots, sheep nd cows round prturition. RM ANOVA reveled significnt effects of species (P < 0.01) nd n interction of time nd species (P < 0.05). Different letters represent significnt differences within one species. The symols # nd $ indicte significnt differences etween cows nd gots (#) or sheep ($), respectively. Mens ± SEM. 312 313 314 315 316 317 Figure 3 Plsm concentrtions of totl clcium mesured in gots, sheep nd cows round prturition. RM ANOVA reveled significnt effects of time (P < 0.001) nd species (P < 0.001). Different letters represent significnt differences within one species. The symols # nd $ indicte significnt differences etween cows nd gots (#) or sheep ($), respectively. Mens ± SEM. 318 319 320 321 Figure 4 Plsm concentrtions of phosphte mesured in gots, sheep nd cows round prturition. RM ANOVA reveled significnt effects of time (P < 0.001) nd n interction of time nd species 15
322 323 324 (P < 0.001). Different letters represent significnt differences within one species. The symols # nd indicte significnt differences etween gots nd cows (#) or sheep ( ), respectively. Mens ± SEM. 325 326 327 328 329 Figure 5 Plsm concentrtions of clcitriol mesured in gots nd sheep round prturition. RM ANOVA reveled significnt effects of time (P < 0.05) nd species (P < 0.05). Different letters represent significnt differences within one species. Mens ± SEM. 330 331 332 333 334 335 336 337 Figure 6 Rtio etween plsm concentrtions of osteoclcin nd CrossLps mesured in gots, sheep nd cows round prturition. RM ANOVA reveled significnt effects of time (P < 0.001), species (P < 0.001) nd n interction of time nd species (P < 0.001). Different letters represent significnt differences within one species. The symols #, $ nd indicte significnt differences etween gots nd cows (#), sheep nd cows ($) or sheep nd gots ( ), respectively. Mens ± SEM. 338 339 16
340 341 342 343 344 345 346 347 348 349 350 351 352 353 354 355 356 357 358 359 360 361 362 363 364 References Allen MJ 2003 Biochemicl mrkers of one metolism in nimls: uses nd limittions. Veterinry Clinicl Pthology 32 101-113 Brithwite GD, Glscock RF & Rizuddin S 1970 Clcium metolism in pregnnt ewes. British Journl of Nutrition 24 661-670 Brithwite GD 1983 Clcium nd phosphorus requirements of the ewe during pregnncy nd lcttion. British Journl of Nutrition 50 723-736 Butler WR & Smith RD 1989 Interreltionships etween energy lnce nd postprtum reproductive function in diry cttle. Journl of Diry Science 72 767-783 Durnd D, Brithwite GD & Brlet JP 1983 The effect of 1 lph-hydroxycholeclciferol on the plcentl trnsfer of clcium nd phosphte in sheep. British Journl of Nutrition 49 475-80 Frser DR & Kodicek E 1970 Unique iosynthesis y kidney of iologicl ctive vitmin D metolite. Nture 228 764-766 Hnnon R & Estell R 2000 Prenlyticl vriility of iochemicl mrkers of one turnover. Osteoporosis Interntionl 11 S30-44 Hofmnn RR 1989 Evolutionry steps of ecophysiologicl dpttion nd diversifiction of ruminnts: comprtive view of their digestive systems. Oecologi 78 443-457 Horst RL, Goff JP & Reinhrdt TA 2005 Adpting to the trnsition etween gesttion nd lcttion: differences etween rt humn nd diry cow. Journl of Mmmry Glnd Biology nd Neoplsi 10 141-156 Horst RL & Jorgensen NA 1982 Elevted plsm cortisol during induced nd spontneous hypoclcemi in ruminnts. Journl of Diry Science 65 2332-2337 Kruse-Jrres JD 1979 Klinische Chemie Vol II: Spezielle klinisch-chemische Anlytik, Fischer, Stuttgrt Germny 17
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Milk yield in L per kg ody weight nd dy 0.10 *** 0.08 0.06 0.04 * ** 0.02 0.00 5 10 15 20 Dys fter prturition Gots (6) Sheep (5) Fig. 1
C 2+ in mmol/l 1.50 1.25 $ # 1.00 0.75 10 dys.p. 2 dys.p. t prturition 6 hours p.p. 12 hours p.p. 1 dy p.p. 2 dys p.p. 10 dys p.p. Gots (6) Sheep (5) Cows (6) Fig. 2
C t in mmol/l 3.0 2.5 2.0 $ # c # $ # # c c c # # c c 1.5 10 dys.p. 2 dys.p. t prturition 6 hours p.p. 12 hours p.p. 1 dy p.p. 2 dys p.p. 10 dys p.p. Gots (6) Sheep (5) Cows (6) Fig. 3
P i in mmol/l 3.5 3.0 # 2.5 2.0 1.5 1.0 0.5 10 dys.p. 2 dys.p. t prturition 6 hours p.p. 12 hours p.p. 1 dy p.p. 2 dys p.p. 10 dys p.p. Gots (6) Sheep (5) Cows (6) Fig. 4
Clcitriol in pg/ml, gots nd sheep 30 20 10 0 10 dys.p. 2 dys.p. t prturition 12 hours p.p. 2 dys p.p. 10 dys p.p. Gots (6) Sheep (5) Fig. 5
Rtio osteoclcin to CrossLps 80 60 40 $ $ # 20 c c c c c 0 10 dys.p. 2 dys.p. t prturition 6 hours p.p. 12 hours p.p. 1 dy p.p. 2 dys p.p. 10 dys p.p. Gots (6) Sheep (5) Cows (6) Fig. 6