Age and psychogenic factors in response to food deprivation and refeeding in White Leghorn chickens

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1 Arch. Genügelk. 995, 59 (3), 75-8, ISSr Verlg Eugen Ulmer GmbH & Co Stuttgrt Age nd psychogenic fctors in response to food deprivtion nd refeeding in White Leghorn chickens Alter und psychisch bedingte Fktoren im Zusmmenhng mit Futterentzug und Wiederfütterung bei Weißen Leghorn-Hühnern. Zulkifli, E. A. Dunnington, P. B. Siegel Mnuskript eingegngen m 4. Juli 994 Introduction Mterils nd Methods Withdrl of feed for vrying periods of time is husbndry prctice tht my hve both physiologicl (e.g FREEMAN et l., 98; MAXWELL et l., 992; ZuLKIFLI et l., 993), nd psychologicl (DuNCAN, 97) consequences on n niml. While most feed restriction studies emphsize the importnce of physiologicl effects ith regrd to stress response, there is derth of informtion on the involvement of underlying psychogenic fctors. When n niml fils to fulfill its biologicl nd behviorl needs, frustrtion my be evoked (LEVINE, 985). Psychologicl stimuli contributing to thrting or frustrting situtions my be s potent s physicl insults in triggering both utonomic nd neuroendocrine responses (DANTZER nd MoRMEDE, 985). Psychogenic fctors my ctivte the hypothlmic-pituitrydrenl (HPA) xis despite the bsence of homeosttic perturbtion (SAPOLSKY, 992). In his revie, MASON (975) indicted tht plsm g lucocorticoid concentrtion declined in fsted primtes provided ith non-nutritive flvored plcebo. Thus, stisfction obtined from orl mnipultion of non-nutritionl substnce my ct s displcement ctivity, or frusttion outlet mong feed deprived primtes. Most reserch on effects of fsting hs emphsized physiologicl nd behviorl consequences tht occurred during the fst. Little informtion is vilble on the response, prticulrly leucocytic ltertions, folloing refeeding fter feed deprivtion. Experiments ith rts nd mice hve demonstrted tht the HPA xis my be ffected by ging (CmuEH et l., 98; SAPOLSKY et l 983; IDA et l., 984). Although, ged rts ere cpble of secreting glucocorticoid in response to vrious Stressors, they hd mrked inbility to terminte glucocorticoid elevtion t the end of stress s eil s delyed dpttion to mild Stressors. SA POLS KY (992 b) ttributed the phenomenon to loss of drenl corticoid negtive feedbck receptors, hich s ccompnied by loss of neurons. The purpose of the present study s to evlute the importnce of ge nd psychogenic fctors in response to fsting nd refeeding in White Leghorn chickens Genetic stock Animl nd Poultry Sciences Deprtment, Virgini Polytechnic Institute nd Stte University, Blcksburg, VA , USA Archiv für GeOügelkunde 3/ 995 Chickens used in Tril ere from White Leghorn popultion developed from lines divergently selected for ntibody response to SRBC (SIEGEL nd GROss, 98). The to selected lines ere crossed, then chickens from the F genertion ere bckcrossed to ech prentl line for three genertions. Only individuls from the high ntibody genetic bckground ith B 3 B 2 genotypes t the mjor histocomptibility complex (D unnington et l., 992) ere utilized in this tril. The chickens used in Tril 2 ere offspring of those used in Tril. Tril At htching (Dy ), chicks ere ingbnded, vccinted for Mrek's disese nd floor rered ith continuous lighting. At 56 d ys of ge, rhey ere trnsferred to groer pens ith ood shvings litter nd exposed to nturl lighting. Pullets ere moved on Dy 2 into individul cges (46 cm high, 3 cm ide nd 46 cm deep) ith verticl rods on the front nd horizontl brs on the bck nd sides. Cges ere in indoless room ith photoperiod from 6 to 2 h. Strter (2 % CP; 2,865 kcl ME/ kg), developer (4% CP; 2,827 kcl ME/ kg) nd l yer (6% CP; 2,76 kcl ME/kg) diets ere provided d libitum in msh form from Dys to 63, Dys 64 to 39, nd Dy 4 onrds, respectively. Feed s in trough ttched to the outside of the front of the cges. On Dy 37, 75 hens ere ssigned t rndom to one of four feeding regimens s follo: () d libitum feeding (AL); (2) food ithdrl for 48h, ith the provision of snd (SAND); (3) food ithdrl for 48 h (NO FOOD); or (4) food ithdrl for 48 h, imposed by blocking ccess to food by ire mesh hich lloed visul nd olfctory cues (WIRE). Ech group s mde up of 2 hens, except for AL, here there ere 5 hens. The AL hens ere housed t the upper decks to prevem hens in the other feeding regimes from seeing them becuse llelomimetic behvior (HUGHES, 97) nd socil fcilittion (FRANCHINA et l., 986) occur during feeding nd drinking. Prior experiments hve not shon tier effects for the mesurement criteri used in this study. In no cse could chickens fce ech other hen eting. Food ithdrl commenced t 8 h on Dy 329.

2 76 ZuLKIFLI et l., Age nd psychogenic fctors in response to food deprivtion Hen-d y ovultion percentge nd totl number of norml nd defective eggs (VAN MIODLEKOOP nd SIEGEL, 976) ere recorded for ech hen dily from Dys 37 to 342. Blood smples ere obtined from the ing vein ith EDT A s nticogulnt t 2, 24, 36, nd 48 h fter onset of food deprivtion. Five hens per tretment ere bled in ech period, except for the AL group, here four nd three hens ere smpled in the first three nd lst bleeding period, respectively. A durtion of 48 h s lloed before the sme hen s bled gin. Blood smers ere prepred using My-Grunld-G iems stin nd heterophils (H) nd lymphocytes (L) ere counted to totl of 6 cells (GRoss nd SrEGEL, 983). At 8 h on Dy 33, ll hens ere relesed to d Jibitum feeding nd similr bleeding procedures ere conducted t 2, 24, 36, nd 48 h fter refeeding. Wter s vil ble t ll times. Behviorl observtions ere mde during the onset of the feedin g regimens nd prior to ech bleeding period. Observtions consisted of recording the generl behvior of ech group of five hens hen tretments ere imposed nd henever the reserchers entered the room here the chickens ere mintined. During the period of food ithdrl, msh nd snd consumption of AL nd SAND gro ups, respectively ere mesured for groups of five hens dil y. Similr mesurements ere mde in ll feeding regimens during refeeding. Tril 2 At htch, 2 chicks (mles nd femles) ere ingbnded nd ssigned t rndom, in groups of 6, to 2 strter btteries ith ire floors nd electric het. Loction of the AL pens s the upper levels s described in Tril. Chicks ere fed Strter diet ith sme formultion s for Tril. Lighting s continuous nd ter s vilble throughout the experiment. At 8 h on Dy 2, chicks ere subjected to one of the four feeding regimens (AL, SA D, Fig.. Men heterophil to lymphocyte rtios t vrious stges of fst.ing by feeding regimen., b, c, d: Mens ithin durtion of fsting ith no comm on letters differ t P ::;.5. Pooled SEM =.2 (2 h);.4 (24 h);.5 (36 h) ;.7 (48 h) H /L -Verhältisse bei verschiedee Sittione einer Niichterung /erschiedlicher Art t :c.. :le.8 ::; ::r I.. : t;;.4 :c DURATION OF FASTING (HOURS) Fig. 2. Men heterophil to lymphocyte rtios t vrious stges of fsting t 2 nd 329 dys of ge., b: Mens ithin durtion of fstin g ith no common letters differ t P::;.5. See Figure for pooled SEM H /L-Verhiiltisse bei verschiedenen SitJtionen einer Niichtmmg bei 2 d 329 Tgen FOOD, nd WIRE) described in Tril. All chicks ere refed t 8 h on Dy 23. Similr blood smpling ( pen per feeding regimen for ech bleeding period), food intke mes urements (pen bsis), nd behviorl observtions (ech pen of chicks) s described in Tril ere rnrried out. S ttistici n!jses Dt from both trils ere used in nlysis of vrince ith fctoril rrngement of tretments in fixed effect model. Heterophil to lymphocyte rtios ere nlyzed seprtely for fsting nd refeeding ith ge nd feeding regimen s min effects. Feeding regimen nd ge ere the min effects for totl food nd snd intke nlyses, hile stge of refeeding s lso included in the model for dily food consumption. For Tril, egg production prior to nd fter onset of fsting s exmined seprtely. When interctions ere significnt, seprte nlyses ere crried out ithin ech min effect. When significnt differences mong. tretments ere observed, comprisons mong mens ere mde by Duncn's multiple rnge tests. Significnt differences ere ssumed t P s.5. L'lll M>l.OIT\M..., l:lll t 2. &Si Wfll!.5 :c.. :le ::; ::r I R esuits ''"""" (gj... : :c.5 t;; DURATION OF FASTING (HOURS) 48 H eterophii/!jmphoryte rtios During food ithdrl, there s no evidence of feeding regimen interctions ith other min vrü,lles for H /L rtios. There s no effect on H /L rtios 2 h fter subjecting chickens to the vrious feeding regimens (Figure ). By 24 h, hoever, rtios for SAND nd FOOD chickens ere elevted. By 36 h, responses ere sirnilr for SAND, NO FOOD, nd WIRE groups ith ll greter thn for AL. All g roups differed from ech other b y 48 h, ith the WIRE regimen hving the highest rtios folloed by FOOD, SA D, nd AL. Chickens fsted on Dy 329 (OLD) exhibited higher rises in H /L rtios by 2, nd 24 h Arch iv für Genügelkunde 3/995

3 ZuLKIFLl et l., Age nd psychogenic fctors in response to food deprivtion r.8.4 Oh d AD LIBITUM SAND NO FOOD WIRE FEEDNG AEGMEN 2. r.8 +2h c c h c Sh + 48h 2. c r.8.4 b AD LIBITUM SAND NOFOOD WlRE FEEOING AEGJMEN AD LJBITUM SAND NOFOOD WIRE FEEOING AEGMEN Fig. 3. Men heterophil to lymphocyte rtios t vrious stges of refeeding here ge (2 dys, solid line; 329 dys, dshed line) by feeding regimen interctions ere significnt (except for nd + 2 h)., b, c: Mens ithin n ge ich no common letters differ t P.5. * Significnt difference beteen ges. Pooled SEM =.7 ( h);.5 (+2 h);.4 (+24 h);.4 (+36 h);.6 ( +48 h) H /L Verhiiltnisse bei verschiedenen Sitttionen der Wiederfiittemng bei Unterschieden in Behdltng und Alter of food ithdrl thn those tht ere feed deprived t 2 dys of ge (YOUNG) (Figure 2). The difference beteen ge groups disppered by 36 h ith reversl of trend by 48 h. Although H /L rtios declined ithin 2 h folloing refeeding, rtios did not return to rhe level of AL for ny of the other feeding regimens (Figure 3). At this time the SA D regimen bd loer rtios thn FOOD nd WIRE chickens, hich ere similr. Age by feeding regimen interctions ere observed for the subsequent durtion of refeeding. The interctions resulted from delyed recovery mong YOU G chickens during relese to d libitj feeding. After 24 nd 36 h of refeeding, expcept for WIRE here H /L rtios of YOUNG chickens ere elevted, there ere no ge-relted differences in response mong feeding regimens. Among OLD hens, hoever, regrdless of prior tretment, ll hd similr H /L rtios folloing 36 h of refeeding nd these levels ere mintined therefter. On the contrry, ithdrl from fsting for 48 h resulted in mrked rise in H /L rtios of YOUNG chickens under SA D nd WIRE regimens. Egg prodction Prior to the onset of food deprivtion (D ys 37 to 328), percentge hen-dy ovultions s similr for ll tretment Archiv für Geflügelkunde 3/ 995

4 78 ZuLKIFL! et l Age nd psychogenic fctors in response to food deprivtion 9 eo 7 5 () 4 : "- AOLIBITUM SAND NOFOOD 6 WIRE AGE (DAYS) Fig. 4. Percentge hen-dy ovultion by feeding regimen. Feed s ithdrn (l) on Dy 329 nd returned (j ) on Dy 33. Prozentle tägliche Ov/tionsrte bei Ftterentzg mit 329 T gen tmd lf7iederfüttemng mit 33 Tgen Behvior Unless stted otherise, behviorl descriptions ere similr for both ge groups. AL chickens spent mjor proportion of their time eting nd resting du ring morning nd evening observtions, respectively. When presented ith snd during onset of fsting ( h), chickens initilly pproched the feeders, pecked t nd re some snd for minute or to. They then either engged in snd pecking or moved y from the troughs. Unlike their SA D counterprts. WIRE chickens pecked constntly t the ire mesh insted of moving y from feeders. Severl OLD hens, shoed "explortory" behvior by loering their heds to see the undersurfce of food throughs. At the time hen feeders ere removed, FOOD chickens did not exhibit ny specific rection. Hoever, fe OLD hens put their heds our from cges nd pecked t the ire grills here the feed troughs hd been locted. groups (Figure 4) nd no defective eggs ere produced. During the 48 h fsting period (Dys 329 to 33), production ptterns of the four groups remined similr. Hoever, commencing from Dy 33, there s drmtic decline in percentge hen-dy ovultions mong WIRE nd SA D hens hile those under NO FOOD hd moderte drop in production. There s rpid rise in ovultions mong SA D nd WIRE hens folloing 6 h of refeeding (Dy 334) ith lg in the rise in the FOOD group by fe dys. Percentge hen-dy ovultions from Dys 329 to 342 (post fsting period) ere higher for AL nd NO FOOD regimens thn those under SAND nd WIRE hich ere similr. For percentge of defective eggs (reltive to totl number of ovultions), there s no difference beteen AL (.74) nd FOOD (2.92). Hoever, only the former s superior to both SA D (9.87) nd WIRE (8) hile NO FOOD nd SAND shred similr vlues. Food nd snd consumption For ech ge group, consumption of feed nd snd s indexed reltive to the AL regimen hich s considered s %. No ge effect s found in percentge of snd consumed during food deprivtion hen mesured dily (YOUNG, 3.3%, OLD, 9.3%) nd s totl (YOU G, 8.6%, OLD, 26.6%). Regrdless of ge, chickens consumed more snd on the second (4.%) thn the first (8.2 %) dy of food ithdrl. Feeding regimen by ge interctions ere significnt for both dily nd totl feed intke hen chickens ere refed. Hoever, the interctions my be inherent becuse food intke of AL regimens t both ges ere considered s %. Upon relesing to d libitutn feeding, ll YOU G chickens consumed similr mount of feed, heres dily (to dys) nd totl feed consumption mong OLD chickens vried ccording to feeding regimen (Figure 5). When relesed to d libitutn feeding, chickens fsted t 2 dys of ge consumed more food on the second thn the first dy of refeeding. The mount of feed consumed by the OLD chickens s similr for both dys resulting in significnt ge by Stge of refeeding interction (Figure 6). 8 ::; E "' :::; Cl 2 bc ti :- "JJ;;... I. b... b....._ FEEDING REGIMEN Fig. 5. Men percentge dil y (top), nd totl (bottom) food consumption (reltive to d libitm intke) during refeeding here ge (2 dys, solid line; 329 dys, dshed line) by.fet;c;!ing regimen interctions ere significnt., b, c: Mens ithin il.ige ith no common letters differ t P.5. * Significnt differ'l'qce beteen ges. Pooled SEM=.3 (top); 3.6 (bottom) ' Prozentler tglicher nd Totl-Fterverbrh hrend der Wiederfütterungsperiode c b Archiv für Geflügelkunde 3/995

5 ZuLKrFLT Age-relted differences in response ere observed folloing 2 h of food ithdrl. Even though possible disturbnces ere minimized by hving these chickens isolted from ll others, ppernce of the observers initilly cused excitement nd flightiness mong OLD chickens. These rections ere folloed by pecking of ire nd snd for those groups. On the contrry, t this time YOU G chicks ere either resting or sleeping under the het source. After 24 h of fsting, ppernce of observers gin induced excitement nd flightiness ith the rection less intense in AL chickens. While NO FOOD chickens remined flightly nd excited, severl ctivities seen during the onset of fsting such s eting nd pecking of snd, nd occsionl ire pecking ere gin observed. Some OLD hens subjected to WIRE ere found fcing in the opposite direction of feed troughs. This s prticulrly cogent becuse the ter cups ere just djcent to the feeders. Observtions conducted t 36 nd 48 h ere nlogous to those t 2 nd 24 h for OLD hens, respectively. For the YOU G ones, hoever, fter 36 h of fsting, insted of resting under the het source, chicks stood round food troughs nd periodiclly engged in pecking of ire nd snd. When feed s returned, feeding responses ere similr for ll OLD chickens. Among YOUNG chicks, hoever, responses of those from the SAND nd WIRE groups ere sloer thn those of their NO FOOD counterprts. Observtions folloing 2, 24, 36 nd 48 h reveled tht, regrdless of prior tretment, chickens ere either eting or resting. Disrnssion The present findings dd to the groing body of evidence tht response to fsting my be physiologiclly nd psychologiclly modulted (see revies by D uncan, 97; MENCH, 992). After being food-deprived for 48 h, WIRE nd NO FOOD chickens hd similr durtions of physiologicl disruption but the WIRE chickens, probbly due to greter psychologicl stimultion, ere more distressed s mesured by H /L rtios. These observtions suggest the thrting nture of the WIRE tretment nd concomitntly the potency of underlying psychogenic fctors in eliciting the HPA xis. In erlier studies, D uncan nd WooDGUSH (97; 972) proposed from behviorl observtions tht presenting food under visible cover cused frustrtion mong fsted chickens. KOENE (993) confimed these observtions nd found differences in vocliztion mong hite nd bron egg lyer stocks. Suppression of H /L rtios mong WIRE chickens folloing 24 h of food ithdrl my be medited by visul stimuli hich ct s cue for them to expect the forthcoming event, refeeding. Similrly, HARVEY et l. (983) noted tht fsted chickens exhibited decline in plsm concentrtion of corticosterone hen simply shon food. Hoever, in the present study, nd s noted by HARVEY et l. (983), the suppression of stress response s only trnsient. They suggested rectivtion of the HP A xis if visul stimuli ere not folloed by metbolic reinforcement nd ttributed the phenomenon to "hope-disppointment syndrome". In discussing the importnt role of psychogenic fctors in eliciting stress response, MASON (975) postulted the possibility of minimizing ctivtion of the HPA xis due to physicl Stressors by modifying the involved emotionl Archiv für Geflügel kunde 3/ et l., Age nd psychogenic fctors in response to food deprivtion b. CU z ::;; E & CO :J <( u.. (.'.) z 4 l AYSOLD () 329 DAYS OLD ex: Cl. 24 HOURS 48 HOURS OURATION OF REFEEDING Fig. 6. Men percentge dily food consumption (reltive to d libiti intke) during refeeding here ge by stge of fsting interctions ere significnt., b: Mens ithin n ge ith no common letters differ t P.5. * Significnt difference beteen ges. Pooled SEM =.22 Prozentuler tiiglicher Ftterverbrch viihred der Behdlg rousl. Studies ith pigs (D ANTZER nd MORMEDE, 98), primtes (MASON, 975) nd rts (SAPOLSKY, 992) support this thesis nd suggest the importnce of displcement ctivities or frustrtion outlets in meliorting the psychologicl threts. Our dt provide dditionl support to these findings, becuse H /L rtios of SAND ere higher thn for AL but loer thn for NO FOOD nd WIRE chickens. Aprt from the ctul consumption of the nutritionlly inert snd, the displced behvior of redirected pecking nd "snd plying" my hve been importnt in helping them cope ith psychologicl components of fsting nd eventully ttenute physiologicl responses. The OLD hens in o ur ex periment did not exhibit impired recovery from disruption of homeostsis due to ge hich is inconsistent ith erlier studies ith rts (CHIUEH et l., 98; SAPOLSKY et l., 983; lda et l., 984). In terms of " physiologicl ge", our OLD hens my hve been different from the rts used in previous ork nd the reported ge-ssocited hippocmpl degenertion (SAPOLS KY, 992b) my hve not occurred extensively mong the ged hens used in this study. The inconsistency my lso be ttributed to ge-relted vrition in feedin g behvior during refeeding, here the lighting progrm nd reproductive sttus (SYKES, 983) could be responsible. Feed intke of refed OLD hens vried ccording to fsting regimen nd remined the sme on both dys of refeeding, heres ll YOUNG ones consumed similr mounts of food ith higher intke on the second thn first dy of relese to d libititll feeding. ZuLKIFLI et l. (993) reported tht chicks exhibited trnsient rise in H /L rtios hen ithdrn from more severe fsting. Rpid replenishment folloing ter deprivtion my lso be stressful (MARSDEN et l., 965), suggesting tht grdul releses from environment l insults m y enhnce termintion of poststress response. A third possible explntion is relted to erly experiences nd response to stress. Becuse OLD hens ere exposed to vrious lighting progrms nd types of housing, they my hve hd more stressful erly experiences thn their YOU G

6 8 ZuLKlFLf et l Age nd psychogenic fctors in response to food deprivtion counterprts tht ere kept in the s me pens ith constnt photoperiod throughout the experiment. Erl y stressful experiences.my hve considerble impct on response to subsequent perturbtions of homeostsis (ZuLKIFLI et l., 993, 994, b). Mechnisms involved fo r the errtic nd robust fluctutions of H /L rtios during refeeding mong YOUNG chickens subjected to WIRE nd SA D re uncler. lt is possible, hoever, tht the phenomenon could lso be linked to feedin g behvio r durin g refeeding. As ex pected, food ithdrl suppressed ov ultion nd incresed incidence of defective eggs. Elevtion of plsm concentrtion of drenline, due to stress, hich cused mong o ther fctors, dely in oviposition, cesstion o f uterine motility nd retention of eggs (SYKES, 955 ; H ughes et l., 986) my hve cco unted for the inferior egg yield of fsted hens. There is lck of greement on methods to ssess elfre or degree of stress in livestock. D espite skepticl vies nd inconsistencies mong reported findings, productivity is considered s one o f the typicl indictors of elfre (see revies by D uncan, 98; M ENC H, 992). Our dt suggest tht productivity nd physiologicl indexes do not necessrily correlte eil. For exmple, ltho ugh NO FOOD hens hd higher H /L rrios thn those under SAND nd AL regi mens, their hen-dy ovultions ere greter thn nd similr to the former nd ltter, respectively. Psychologicl events, such s in thrting situtions, my provide stimuli tht cn influ ence ggressive behvior (see revies by D u CAN, 97; ME 'CH, 992). Frustrtioninduced ggression hs been nored in bboons (SAPOLSKY, 99), pigs (D A 'TZER et l., 98), nd domestic fo l (D uncan nd Wooo-GuSH, 97). Despite the thrting nture of the WIRE gro up, o ur experimentl procedure s such tht physicl spects of ggression ere precluded. This s becuse chicks used in T ril 2 ere onl y 2 dys of ge, prior to estblis hment of pecking rights (D AsoN nd SIEGEL, 967), nd res ults shoed no ggressive behvio r. As fo r the OLD hens, physicl contct mong individuls s precluded becuse they ere cged individully. Excitement nd fli ghtiness exhibited by the fsted chickens hen observers ppered suggest tht presence o f humns provide cue signlling the chickens to except presenttion offeed. Fish (DA vrs nd BARDACH, 965), nd finches (CLIFTON, 979) ere ctive during nticiption of foo d. This rection, hoever, my further excerbte the stressfulness of fsting du e to "hope-disppointment syndrome" (MoRER, 96 ; s cited by HARVEY et l., 983). lt s uncler hether the tendency o f redirected pecking t ire mesh s detrimentl to the ongoing frusrrtion, beneficil to the physiology of the chickens s in displcement ctivities or neither. When lyers ere denied food by covering feed trou ghs ith opque bords, th us ttenuting thrting, pecking of covers s lso noted (PRESTO, 987). In the present experiment, it is unlikel y tht ire pecking could be linked to socil fcilittion or llelomimetic feeding behvior becuse the AL chickens ere out of their sight. The opposite o rienttion of severl WIRE hens my be considered s conspicuous disply of frustrtion. The behvior could be interpreted s "looking for n exit" ction or voidnce from thrting situtio ns (D uncan, 97). Emotionl rousl such s exposure to novel environment elicited locomotor ctivity nd escpe ttempts by pigs {DANTZER nd MORMEDE, 985). There is evidence from severl sources (see D ANTZER nd MORMEDE, 985; MENC H, 992) suggesting tht MASON's (975) thesis regrding elimintion of emotionl components nd ttenution of detrimentl physio logicl responses could offer some promising solutions for stress-relted mngement procedures. Our dt suggest the possibility of reducing ctivtion of the HPA xis in food-deprived chickens by providing nutritionlly inert substnce, hich modifies the psychologicl threts of the procedure. Su!llmry Physiology, behvior, nd egg production ere mesured in White Leghorn chickens (2 nd 329 dys of ge) su bj ected to () d libitu!ll feeding (AL) ; (2) food ithdrl for 48 h, ith the provision of snd (SA D ); (3) food ithdrl for 48 h (NO FOOD); or (4) food ithdrl for 48 h, imposed by blocking ccess to food by ire mesh hich lloed vis ul nd o lfctory cues (WIRB). Fstingelicited elevtion in H /L rtios s evident by 24 h. Folloing 48 h of fsting, WIRB chickens hd the highest H /L rtios folloed by SAND, NO FOOD, nd L. Within 36 h of refeeding, older chickens hd recovered from fsting-induced stress but H /L rtios of yo unger ones from WIRB nd SAND tretments shoed errtic fluctutions. When relesed to d libittm feeding, food intke of older chickens vried ccordin g to prior tretment nd remined the sme during bo th dys of replenishment, heres ll you nger o nes consumed similr mounts of feed ith n intke hig her on the second thn first dy o f refeeding. Fsted chickens displyed prominent behviorl responses ith occsionl ge-relted discrepncies. Chickens subjected to WIRE shoed the most frustrtion. Provision of snd ppered to reduce "emotionl" spects of feed deprivtion. Alter und p.rychisch bedingte Fktoren illl? usnnmnhng lllit Futterentzug und Wiederfiitterung bei Weißen Leghorn-Hühnern I. Zulkifli, E. A. Dunnington und P. B. Siegel Z usm!llenjssmg Psychologie, Verhlten und Legeleistung urden bei Leghornhühnern im Alter von 2 und 329 Tgen gemessen, enn fütterungstechnisch d-libitum-fütterung (I), Futterentzug für 48 Stunden mit E rstz durch Snd (II), gänzlicher Futterentzug für 48 Stunden (lll) oder Futterentzug durch Verhinderung des Trgzugnges mit einem Drht (IV), lso bei noch visuellem bz. Geruchskontkt mit dem Futter, ngeendet urden. Ds durch Fsten nsteigende H /L-Verhältnis (Hetero phile/lymphozyten) r nch 24 Stunden feststellbr. Die Tiere der Gruppe IV htten die höchsten H /L-Werte, gefolg t von denen der Gruppen ll, lll und I. Nch 36-stündiger Wiederfütterung htten sich die älteren Tiere vom Entzugsstreß erholt, ährend die H /L-Werte der jüngeren von Gruppe IV und ll schnkten. Wenn genüchterte Tiere ieder zur Fütterung zugelssen urden, vriierte der Futterverbruch der älteren Hühner ents prechend ihrer früh eren Fütterung, und ds duerte 2 Tge, ährend lle jüngeren Tiere et dsselbe frßen, und zr m 2. T ge mehr ls m ersten. Die hungernden Tiere zeigten bechtliche Verhltensstörungen mit gelege ntlichen ltersbedingten Abeichungen. Die Hühner der Gruppe IV ren m meisten frustiert. Der Erstz durch Snd (Gruppe ll) reduzierte offensichtlich die "emotionelle Sitution" eines Futterentzuges. Arch iv für Genügelku nde 3/995

7 ZULKIFLI et l., Age nd psychogenic fctors in res ponse to food deprivtion S tichvorte Huhn, Alter, Fütterung, Fütterungsteehnik, Nüehterung, Verhlten, Frusttion, Legeleistung References CHIUEH, C., S. NESPOR, S. RAPOPORT, 98: Crdiovseulr, sympthetie nd drenl eortiel responsiveness of ged Fiseher-344 rts to Stress. Ne urobiol. Aging, CLIFTON, P. G 979: Responses to regulr deprivtion of food. Anim. Behv. 27, D ANTZER, R., M. ARNONE, P. MoRMEDE, 98: Effeets of frustrtion on behvior nd plsm eortieosteroid levels in pigs. Physiol. Behv. 24, - 4. DA TZER, R., P. MORMEDE, 98: Pituitry-drenl eonsequenees of d junetive etivities in pigs. H orm. Behv. 5, D ANTZER, R., P. MüRMEDE, 985: Stress in domestie nimls : A psyehoendoerine pproeh. Pges 8-96 in: Anim! Stress. G. P. MOBERG, ed. Wverly Press Ine., Bltimore, MD. DAVIS, R. E., J. E. BARDACH, 965: Time eo-ordinted prefeeding etivity in fish. Anim. Behv. 3, D AWSON, J. S., P. B. SIEGEL, 967: Behvior ptterns of ehiekens to ten eeks of ge. Pou ltry Sei. 46, D uncan, I. J. H., 97: Frustrtion in the fol. Pges 5-3 in : Aspeets of Poultry Behviour. B. M. FREEMAN nd R. F. GoRDON, eds. British Poultry Seienee, E dinburgh. D uncan, I. J. H., 98: Anim! rights-niml elfre: A seientist's ssessment. Poult. Sei. 6, D uncan, I. J. H., D. G. M. WooD-G us H, 97: Frustrtion nd ggression in the domestie fo l. Anim. Behv. 9, D uncan, I. J. H., D. G. M. WooD-GuSH, 972: Thrting of feeding behvior in the domestie fol. Anim. Behv. 2, D unn INGTON, E. A., C. T. LARSEN, W. B. GRoss, P. B. SIEGEL, 992: Antibody responses to combintions of ntigens in White Leghorn ehiekens of different bekground geno mes nd mjor histocomptibility complex genotypes. P oultry Sei. 7, FRANCHINA, J. J., A. B. DYER, s. J. ZACCARO, A. H. SCHULMAN, 986: Soeilly feilited drinking behvior in ehieks ( Cllus domestics): Reltive effeets of dri ve nd stim ulus meehnisms. Anim. Lern. Behv. 4, FREEMAN, B. M., A. c. c. MANNING, I. H. FLACK, 98: The effeets of restrieted feeding on drenl eortiel etivity in the immture domestie fol. Br. Poult. Sei. 22, GROSS, W. B., H. S. SIEGEL, 983: Evlution of heterophil/lymphoeyte s mesurements of stress in ehiekens. Avin Dis. 27, HARVEY, S., H. KLANDORF, Y. PINCHASOV, 983: Visu.l nd metbolie Stimuli euse drenoeortiel suppressn m fsted ehiekens during refeeding. Neuroendoerinology 37, H ug HES, B.., 97 : Allelomimetie feeding in the domestie fol. Br. Po ult. Sei. 2, H ughes, B.., A. B. GILBERT, M. F. BROWN, 986: Ctegoriztion nd euses of bnorml egg shells: Reltionship ith stress. Br. P oultry Se. 27, foa, Y., M. TANAKA, A. TsuDA, 984: Recovery of stres.sindueed inerese in nordrenline turnover s delyed m speeifie regions of old rts. Life Sei. 34, Archiv fü r Geflügelkunde 3/995 8 KüENE, P 993 : Emotionl expressions nd fru strtion in domestie fols. Pges in: Proeeedings of the Fourth Europen Symposium on Poultry Welfre. C. J. SAVORY nd B.. H ughes, eds. Edinburgh, UK. LEVI E, s 985: A definition of Stress? Pges 5-69 in: Anim[ Stress. G. P. MoBERG, ed. Wverly Press Ine., Bltimore, MD. MARSDEN, S. J., G. S. McKEE, M. J. CRANDALL, 965: Wter depri v tion nd replenishment in poults. Poultry Sei. 44, MASON, J. W., 975: A historiel vie of the stress field. J. Hum. Stress, MAXWELL, M. H P. M. HOCKING, G. W. RüBERTSON, 992: D ifferentil leueoeyte responses to vrious levels of food restrietion in broilers, turkeys nd dueks. Br. Poult. Sei. 33, MENCH, J. A., 992: The elfre of poultry in modern produetion systems. P o ult. Sei. Rev. 4, PRESTON, A. P., 987: Restrieted feedin g time nd the behvio r of eged J yin g hens. Br. Poult. Sei. 28, SAPOLSKY, R. M., 99: Stress in the ild. Sei. Am. 262, SAPOLSKY, R. M., 992 : Neuroendoerinology o f the stressresponse. Pges in : Behviorl Endoerino logy. J. B. BECKER, S. M. BREEDLOVE nd D. CREWS, eds. MIT Press, Cmbridge, MA. SAPOLSKY, R. M., 992b: Stress, the Aging Brin nd the Meehnism of euron Det h. MIT Press, Cmbridge, MA. SAPOLSKY, R. M., L. C. KREY, B. S. M ceen, 983: The drenoeortiel stress-response in the ged mle rt: Impirment of reeovery from Stress. Expt. G erontol. 8, SIEGEL, P. B., W. B. GRoss, 98: Produetion nd persistenee of ntibodies in ehiekens to sheep erythroeytes.. Direetionl seleetion. Poultry Sei. 59, - 5. SYKES, A. H., 955 : The effeet of drenline on oviduet motility nd egg produetion in the fol. Poultry Sei. 34, SYKES, A. H., 983: Food intke nd its eontrol. Pges - 29 in : Physiology nd Bioehemistry of the D omestie Fol. B. M. FREEMAN, ed. Aedemie Press Ine., London. VAN MmDLEKOOP, J. H., P. B. SIEGEL, 976: Clssifietion of bnorml ehieken eggs. Po ultry Sei. 55, ZuLKIFLI, I., E. A. D unnington, W. B. GRoss, A. S. LARSEN, A. MARTIN, P. B. SIEGEL, 993: Responses of drf nd norml ehiekens to feed restrietion, Eimeri tenell infeetion nd sheep red blood eell ntigen. Poultry Sei. 72, ZuL KIFLI, I., E. A. D unn INGTON, W. B. GRoss, P. B. SIEGEL, 994 : Food restrietion erly o r lter in life nd its effeet on dptbility, disese resistnee nd immunoeompetenee of het-stressed drf nd nondrf ehiekens. Br. Poult. Sei. 35, ZuLKIFLI, I., E. A. D unnington, W. B. GRoss, P. B. SIEGEL, 994 b: Inhibition of drenl steroidogenesis, food restrietion nd eclimtion to high mbient tempertures in ehiekens. Br. Poult. Sei. 35, Kontktdresse : Pul ß. Siegel, Anim! nd Poultry Scicnces Depnmeot, Virgini Polycec hnic Institute nd Srtc Univcrsity, Blcksburg, VA , USA

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