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1 Wgeningen UR Livestock Reserch Prtner in livestock innovtions Report 335 The effect of optimized lighting conditions on fether pecking nd production of lying hens Het effect vn geoptimliseerd kunstlicht op verenpikken en productie vn leghennen April 2

2 Colophon Publisher Wgeningen UR Livestock Reserch P.O. Box 65, 82 AB Lelystd Telephone Fx E-mil Internet Editing Communiction Services Copyright Wgeningen UR Livestock Reserch, prt of Stichting Dienst Lndbouwkundig Onderzoek (DLO Foundtion), 2 Reproduction of contents, either whole or in prt, permitted with due reference to the source. Libility Wgeningen UR Livestock Reserch does not ccept ny libility for dmges, if ny, rising from the use of the results of this study or the ppliction of the recommendtions. Wgeningen UR Livestock Reserch nd Centrl Veterinry Institute of Wgeningen UR, both prt of Stichting Dienst Lndbouwkundig Onderzoek (DLO Foundtion), together with the Deprtment of Animl Sciences of Wgeningen University comprises the Animl Sciences Group of Wgeningen UR (University & Reserch centre). Single numbers cn be obtined from the website. ISO 91 certifiction by DNV emphsizes our qulity level. All our reserch projects re subject to the Generl Conditions of the Animl Sciences Group, which hve been filed with the District Court Zwolle. Abstrct Fether pecking is one of the mjor problems in commercilly kept lying hens. The current reserch considers the relevnce of colour of light in the fether pecking problem. Keywords Lying hens, ultrviolet light, fether pecking, fer, production Reference ISSN Author(s) M.A.W. Ruis B.F. Reuvekmp H. Gunnink G.P. Binnendijk Title The effect of optimized lighting conditions on fether pecking nd production of lying hens Het effect vn geoptimliseerd kunstlicht op verenpikken en productie vn leghennen Report 335

3 The effect of optimized lighting conditions on fether pecking nd production of lying hens M.A.W. Ruis B.F. Reuvekmp H. Gunnink G.P. Binnendijk April 2

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5 Voorwoord Voor u ligt de Engelstlige rpportge vn het onderzoek nr het effect vn licht op het gedrg en de productie vn leghennen. Het onderzoek is uitgevoerd in opdrcht vn Philips Lighting en het Productschp Pluimvee en Eieren. Het is uitgevoerd door onderzoekers vn Wgeningen UR Livestock Reserch op twee proefccommodties in Lelystd. Het onderzoek drgt bij n de inspnningen die verricht worden om verenpikken bij leghennen tegen te gn met het oog op het toekomstige verbod op snvelbehndelingen. Deze rpportge beschrijft de chtergrond, de methodes, resultten en conclusies vn drie opeenvolgende dierproeven, in de periode Specil voor dit doel heeft Philips Lighting nieuwe lmpen ontwikkeld. Excte technische detils en (kleur)eigenschppen vn de nieuwe lmpen wren niet bekend bij de onderzoekers en zijn vertrouwelijk. We willen Philips Lighting en het Productschp Pluimvee en Eieren grg bednken voor het toegezegde vertrouwen om het onderzoek door ons te lten uitvoeren.

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7 Smenvtting Achtergrond en hypothese Vn verenpikken is bekend dt het vk te herleiden is nr een rme omgeving met weinig prikkels en mogelijkheden tot het uiten vn ntuurlijk gedrg. Wgeningen UR Livestock Reserch onderzocht in opdrcht vn Philips Lighting en het Productschp Pluimvee en Eieren (PPE) in hoeverre verenpikken teruggedrongen kn worden door npssingen in lichtkleur, optiml fgestemd op het kippenoog. De chterliggende gedchte ws dt kippen meer kleuren kunnen onderscheiden dn mensen, en dt ze in tegenstelling tot mensen ook ultrviolet A (UV-A) licht kunnen zien. Het is bekend dt dit UV-A licht, nwezig in dglicht een grote invloed heeft op het ntuurlijke gedrg vn kippen. Met UV-A licht zien kippen meer detils in de omgeving en herkennen ze soortgenoten beter. Dit stimuleert ntuurlijk gedrg zols bodempikken, bodemkrbben en stofbden en vermindert de intentie om nr elkr te gn pikken. Bij voorkeur zl een kip deze ctieve gedrgingen bij lichtsterktes boven de 5 lux willen uitvoeren. Het belng vn licht voor het ntuurlijke gedrg en het kunnen zien vn kippen is nuwelijks bekend in de prktijk. De verlichting in pluimveestllen - vk witte TL-verlichting bevt nuwelijks tot geen UV-A licht. Drnst worden in de prktijk reltief lge lichtsterktes gehnteerd, meestl tussen de 5 en 3 lux. Drie dierproeven In de periode zijn drie opeenvolgende dierproeven uitgevoerd met Lohmn Bruin (LB) en Lohmn Selected Leghorn (LSL) leghennen. De snvels vn de hennen bleven intct. De hennen werden ls eendgskuikens in grondhokken gepltst. Nst het gedrg is het gewicht vn de hennen en de kwliteit vn het verenkleed nuwlettend gevolgd. Op bsis vn voortschrijdend inzicht werden in elke proef twee nieuw ontwikkelde lmpen getest, en vergeleken met de resultten onder stndrd witte verlichting. De nieuwe lmpen zijn ontwikkeld door Philips Lighting, en er is gebruik gemkt vn hoogfrequente PL lmpen. In de nieuwe lmpen is met nme gevrieerd met de hoeveelheid en excte golflengte vn het UVA licht. De excte technische detils en (kleur)eigenschppen vn de nieuwe lmpen wren niet bekend bij de onderzoekers. In proeven 1 en 2 werden de leghennen in groepjes vn gehuisvest, op bodems ingestrooid met houtkrullen. Beide proeven duurden tot het einde vn de opfokperiode (17 weken). De lichtsterkte ws 2 of 4 lux in proef 1, en 4 lux in proef 2. In proef 3 werden de hennen gehuisvest in groepjes vn 3, bij 4 lux. Er werd gestrt met bodems zonder strooisel, om de omstndigheden minder optiml te mken. Uiteindelijk kregen LSL hennen op een leeftijd vn 6 weken en LB hennen op een leeftijd vn 14 weken beschikking over strooisel. Strooisel werd verstrekt op het moment vn uitbrken vn verenpikken bij de respectievelijke merken. In deze proef werden de hennen ook in de legperiode gevolgd, tot een leeftijd vn 5 weken, om ook een effect op technische kengetllen te bestuderen. Pikgedrg Zols bleek in de eerste proef hd lichtsterkte nuwelijks effect op de resultten. Alleen wren de hennen bij 2 lux wt minder ctief. Op verschillende momenten in de opfokperiode vn proef 1 werd met een nieuwe lmp meer bodempikken en minder zcht verenpikken wrgenomen, vergeleken met de stndrd lmp. Dit ws ook het gevl in proef 2, mr lleen bij de testlmp met de hoogste UV-A output. Vermoedelijk is het beter is ls er minder zcht verenpikken voorkomt, ngezien dit gedrg over kn gn in beschdigend verenpikken. Genoemde positieve effecten wren overigens soms lleen bij één merk zichtbr. Beschdigend verenpikken werd in proeven 1 en 2 niet wrgenomen. In de derde proef ws het beeld beduidend nders. In de legperiode werd onder de testlmpen bij LB hennen meer verenpikken gezien dn bij de stndrd lmp en ws de veerschde n strt en rug op 5 weken ntoonbr groter. Bij LSL hennen werd nuwelijks verenpikken wrgenomen in de

8 legperiode. Deze hennen hdden l in de vroege opfok strooisel gekregen, en dit verklrt wellicht het verschil. Overig gedrg en technische resultten Als rectie op een persoon of op een onbekend voorwerp toonden leghennen vn één of beide merken in lle proeven minder mijdingsgedrg bij één vn beide testlmpen. Dit ws indictief voor minder ngst. Er is in de proeven 2 en 3 ook ntoonbr meer comfortgedrg gezien, vergeleken met stndrd verlichting, zols stofbden en poetsen. Over het lgemeen hdden de nieuwe lmpen geen effect op de technische resultten in de opfok- en legperiodes. Bij sommige testlmpen ws de uniformiteit lger. In de legperiode vn proef 3 ws de voerconversie hoger bij één vn de testlmpen. Bij de ndere testlmp werden meer grondeieren gevonden. De oorzk is onduidelijk. Discussie en vervolg De resultten lten zien dt kunstlicht dt ook ultrviolet licht uitstrlt bij kn drgen n het stimuleren vn het gewenste pikgedrg. De hennen worden dn gestimuleerd om meer nr de bodem en minder nr elkr te pikken. Vermoedelijk doorslggevend in een positief effect vn geoptimliseerd licht is de tijdige nwezigheid vn voldoende prikkels en fleiding. Omgekeerd beredeneerd werkt een optimle verlichting vn een kle omgeving verenpikken eerder in de hnd. Dit geeft n dt een goede verlichting lleen niet voldoende is om ongewenst pikgedrg tegen te gn. Eenml ngeleerd ongewenst pikgedrg in de opfok lijkt overigens moeilijk f te leren, ook niet ls de omstndigheden in de lte opfok en legperiode beter zijn. Gezien de smenhng met en fhnkelijkheid vn ndere fctoren, zols strooisel en wellicht voer, zl Philips Lighting nog geen ngepste lmp op de mrkt brengen. Als in de toekomst een lmp wordt ngeboden, zl dit gebeuren in combintie met een strooisel- en voerdvies. Voor dit n te bevelen "pkket" is nog vervolgonderzoek nodig. Dit onderzoek kn in de prktijk pltsvinden.

9 Summry Fether pecking is one of the mjor problems in commercilly kept lying hens. The current reserch considers the relevnce of colour of light in the fether pecking problem. Chickens hve welldeveloped colour vision, nd even hve the bility to see into the ultrviolet rnge (ultrviolet A). It is hypothesized tht UV-A inclusion in light results in better vision nd consequently richer environment for the hens, s hens receive more detiled informtion from their environment. Accordingly, they my become less interested in ech other fethers. The following reserch questions were ddressed in the reserch: 1) Cn UV supplementtion reduce gentle nd severe fether pecking in lying hens? 2) Cn UV supplementtion increse desired types of pecking, such s ground pecking? 3) cn UV supplementtion decrese ferfulness of the hens? 4) Are spects of hen development nd egg production ffected by UV-A supplementtion? The effect of optimized lighting conditions on the behviour nd production of lying hens ws tested experimentlly with white nd brown egg lyer breed. This ws done in three consecutive experiments. The first two experiments comprised the rering period, wheres the third experiment lso included the lying period until 5 weeks of ge. In ech experiment two dpted or optimized light sources were tested, nd compred with stndrd white light. In ll cses fluorescent tubes (PL) were used. Light intensities were either 2 or 4 lux in experiment 1 (fctor), nd ws fixed round 4 lux (stndrdized brightness) in the other experiments. The results of the three experiment indicte tht with UV-A inclusion in light, gentle fether pecking t times cn be reduced nd ground pecking my be stimulted. This is possible t reltively high light intensities (round 4 lux), which is close to the desired light intensities by lying hens for their ctive behviours. UV-A inclusion in light hs fer-reducing properties. The most importnt egg production trits re not ffected by UV-A inclusion, but uniformity in weight nd proportion of eggs in nests my be t risk nd require ttention. In experiments 1 nd 3, dpted light sources ech hd their specific (dis)dvntges. In experiment 2, the most promising light source ws E2-optimized-2, which hd the highest UV-A output. Adpted lighting in itself will not exclude or reduce injurious fether pecking, s clerly shown in experiment 3: LB hens developed high levels of injurious pecking. Other fctors, such s suitble litter, my determine whether the effects of dpted light re either beneficil or detrimentl. LB hens were reltively long kept without litter, in n environment lcking stimuli, nd this my hve led to n undesirble sitution in which improved vision my hve cused more interest in ech other fethers nd thus more fether pecking. Injurious fether pecking in LSL hens, on the other hnd, ws initilly high but ws much reduced following litter provision t reltively young ge. However, positive effects of dpted light in the longer term were only seen for gentle pecking, while severe pecking ws not ffected. In prctice, litter mngement for loose housed hens vries lot. Moreover, mny hens re kept in cges, which hve reltively brren environment with very limited litter provision. LB hens re the predominnt breed kept for commercil purposes. This specific combintion my pose risk for development of fether pecking, s we my conclude from our results. Optimized light sources re therefore not redy for mrketing t this point, without knowing more bout the interction of light with other fctors. We expect the most from rich substrtes nd feed which hve high reflective properties. Adpted light by UV inclusion then my hve its beneficil effect.

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11 Tble of contents Prefce Summry 1 Introduction Mterils nd Methods Experimentl housing nd nimls Experimentl design Experiment Experiment Experiment Observtions Pecking nd generl behviour Responses to humn presence nd novelty Plumge nd skin condition scores Body weight nd uniformity Feed consumption nd egg production Sttisticl nlysis Results Effect of optimiztion of lighting conditions on fether pecking behviour Gentle fether pecking Severe fether pecking Ground pecking Effect of optimiztion of light on generl behviour Effect of optimiztion of light on fer for humns nd generl fer Effect of optimiztion of light on plumge nd skin condition Effect of optimiztion of light on body weight nd uniformity Effect of optimiztion of light on feed intke nd egg production Discussion nd conclusions Conclusions... 23

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13 1 Introduction Motives nd bckground Fether pecking is one of the mjor problems in commercilly kept lying hens. Fether pecking vries from gentle fether pecking to severe fether pecking or fether pulling (Svory, 1995). Gentle fether pecking is form of fether pecking without removl of fethers, nd cuses little or no fether dmge nd is often ignored by the recipient. Severe fether pecking on the other hnd is much more forceful, nd poses mjor welfre risk for the hens. Severe fether pecking is the pecking t nd pulling out of fethers of conspecifics. This cuses fether dmge nd loss of fethers, nd eventully leds to bld ptches. Most dmge by pecking is cused to the til, rump nd bck regions (Bilcik nd Keeling, 1999). It is lso n economicl burden for the frmer. Costs of egg production increse, s loss of fethers leds to incresed body het loss, nd consequently higher feed (energetic) requirements. Bld ptches my ttrct skin or tissue pecking, which cn esclte into wounding of the victim nd cnniblism (i.e. consumption of flesh or food). Fether pecking is both observed in cges nd in lterntive systems. In lterntive systems with lrge groups, however, the problem is more difficult to control, s fether pecking my spred by socil trnsmission (Zeltner et l, 2). In lterntive housing, losses my be up to 15% of the birds per production cycle. Bek trimming s remedil mesure will be bnned in the ner future in The Netherlnds. Gentle nd severe fether pecking re not cler-cut nd my grde into ech other (Svory, 1995). It is suggested tht in dult lying hens gentle fether pecking by incresing intensity or severity my develop into severe fether pecking (McAdie nd Keeling, 22). If this is the cse (other theories lso exist, e.g. Newberry et l., 27), gentle fether pecking erly in life my be n indictor of severe pecking in dult lying hens. Fether pecking should not be confounded with ggressive pecking, which is used to mintin the dominnce hierrchy. Aggressive pecking my led to some dmge to the hed nd neck region. There is n ongoing discussion on the custion of fether pecking. One of the most importnt single cusl fctors ssocited with fether pecking is the provision of suitble floor substrte (litter). The two most influentil theories on the custion of fether pecking re relted to this fctor: fether pecking is thought to be form of redirected behviour, developing either from ground pecking for forging nd feeding (Blokhuis, 1986) or pecking during dustbthing (Vestergrd, 1994), in the bsence of suitble substrte. However, s fether pecking is not fully eliminted by vilbility of suitble substrtes, it is generlly ccepted tht the development of fether pecking is multifctoril. Other single fctors involved re nutrition (food form, dilution), group size nd stocking density, rering conditions, nd light intensity nd colour (Rodenburg, 23). Lrge vrition in performnce of fether pecking lso exists between strins of lying hens. Breeding progrmmes to solve the fether pecking problem seem possible, but so fr the results on this fctor re not consistent, nd heritbility estimtes rnge substntilly, e.g. from.4 to.56 (vn Hierden, 23). The current reserch considers the relevnce of colour of light in the fether pecking problem. Chickens hve well-developed colour vision, nd even hve the bility to see into the ultrviolet rnge (ultrviolet A: UV-A) (Prescott nd Wthes, 1999, Prescott et l., 24; Wortel et l., 1987). To compre, the humn eye is not ble to perceive ultrviolet A rdition (Figure 1; UV-A rnges between 32 nd 4 nm wvelenght). Colour ssessment by humns is therefore n indequte nd mis-leding pproch for the relevnce of colour of light for hens. Hens hve four photorective pigments (tetrchromtic) ssocited with the cone cells, compred with three (trichromtic) in humns. These four pigments re mximlly sensitive t 415, 455, 58 nd 571 nm wvelengths, compred with 419, 531 nd 558 in humns (Prescott et l., 24). 1

14 Figure 1. Spectrl sensitivity (reltive sensitivity: y-xis) of chickens (blue line) nd humns (grey line) t different wvelengths (wvelength in nm: x-xis) UV-A light is specificlly relevnt for poultry to obtin more informtion from the environment. Reflectnce of UV-A light from fethers fcilittes recognition, ssessment of nd communiction between individuls. There is vrition between breeds in colour nd the reflectivities of the fethers. Even in flock of pprently white birds, ech my hve plumge of distinctive overll reflectivity, colour nd pttern when viewed by conspecific (Prescott nd Wthes, 1999b). Reflection by surroundings improves recognition of resources, e.g substrtes nd feed (Prescott nd Wthes, 1999b). Biologicl mterils hve similr reflective chrcteristics to fethers. Swdust or strw bedding, eggshells, nd wood ll hve minimum reflectivity t 4 nm. However, rtificil lighting used in poultry housing produce little, if ny, UV-A rdition. This mens tht current light environments impose some sensory deprivtion on lying hens, by impiring niml recognition nd forging behviour (by impired recognition of substrte nd feed). Indeed, the consequences for niml recognition were shown in study with broiler breeders: sexul selection nd mting behviour were improved with UV-A light inclusion (Jones t l, 21). The impirment of forging behviour my be of relevnce for the fether pecking problem. This impirment my led to n incresed risk for fether pecking. Interestingly, but the effect cme long with other mesures, injurious pecking behviour of turkeys ws found to be reduced with UV-A inclusion (Lewis et l., 2b). Further evidence of the beneficil effects of UV light ws provided by Mddocks et l. (21). Bsl corticosterone levels were higher, suggestive of chronic stress nd fer, in chickens kept under UVdeficient rther thn full-spectrum lighting. Ferfulness hs lso been ssocited with fether pecking, nd my be initited by ferful birds (Vestergrd et l., 1993). However, ferfulness my lso be consequence of fether pecking, induced by fether dmge nd pin. The effect of UV light on reproduction of lying hens is rther unknown. Results of Lewis et l. (2) suggest tht UV light influences behviourl rther thn reproductive responses. UV light hd no effect on the timing of the ovultory cycle, but did suppress food intke. It hs been suggested tht the more sophisticted pprtus for colour vision mens tht hens hve better vision in bright light thn dim. In commercil poultry houses, however, the intensity of light is much lower thn norml dylight. Hen houses my be lit between 5 nd 3 lux. By comprison, on sunny dy illuminnce outdoors cn rech. lux while offices re usully lit to 3 lux. Reducing light intensity is employed by frmers to restrict or to prevent rising levels of fether pecking. Often, it is stted by frmers tht higher light intensities increse the risk for fether pecking. This is, however, wrong interprettion. Other fctors my be ssocited with the occurrence of fether pecking, nd then n effective mesure is to dim light. The mechnism behind this is tht with lower light intensities bnormlities in plumge like dmge, bldness, nd blood spots become less cler. 2

15 Accordingly, hens re less invited to peck t the fethers nd skin of other hens. A similr effect is seen with red light, strtegy lso often pplied in prctice when fether pecking occurs. Strtegies of dim or red light do t first sight not negtively ffect egg production. Lying hens re much more tolernt to low light intensities thn erlier stocks (Renem et l., 21). Specificlly for red light, it must be emhsized tht lthough numbers of eggs lid re not negtively ffected, eggs my become smller sized (Er et l., 27; Pyrzk et l., 1987). Other dvntges of dimming light re reduction in energy costs nd higher food conversion efficiency. The ltter is very probbly due to reduced hen ctivity. Indeed, it is known tht hen ctivity is positively ssocited with light intensity. Hens do show preference for lower light intensities for performing certin behviours such s resting nd perching, but will move to brighter res for more ctive behviours such s feeding, drinking nd forging (Dvis et l., 1999). When given choice, hens prefer to employ ctive behviour, such s eting, in 2 lux (Prescott nd Wthes, 22). Mrtin (1989) showed tht ctivity in hens reduced t 5 lux or less. It my therefore be expected tht reduction in intensity will influence the wvelengths emitted by light source nd tht t dim levels chickens re unble to employ their full repertoire of visul cpbilities with potentilly importnt consequences for their behviour. Also, young birds were more ferful t lux thn t 55-8 lux (Hughes nd Blck, 1974). Very low intensities (<5 lux) even my cuse eye bnormlities s the functionl development of vision my be ffected, especilly when these conditions occur during rering. At these very low light intensities, hens re lso restricted in moving round the house, nd to jump between horizontl perches (Tylor et l., 23). Lst but not lest, dim conditions lso negtively ffect stff working conditions. A minimum intensity of 2 lux is recommended on grounds of niml welfre, stff working conditions nd esthetics. However, the cited results bove on fer nd ctivity threshold, might indicte tht the minimum lux should be higher, i.e. between 5 nd lux. It must be emphsized tht for poultry with different spectrl sensitivities to the humn, the lux unit is incorrect. For exmple, for typicl fluorescent nd incndescent luminires illuminted to the sme lux level nd consequently isoluminnt for humns, hens perceive the incndescent bulb s 2% brighter thn the fluorescent tube (Prescott et l., 24). Aim of the reserch The concern is tht current light environments in hen houses impose some sensory deprivtion on the hen. The effect of light colour nd light intensity on behviour nd especilly on the onset of injurious behviour like fether pecking, merits further investigtion. It is hypothesized tht UV-A inclusion in light results in better vision nd consequently richer environment for the hens, s hens receive more detiled informtion from their environment. Accordingly, they my become less interested in ech other fethers. To test the potentil of dpted lighting for lying hen behviour, Philips Lighting sked the Animl Sciences Group of Wgeningen University nd Reserch Centre (Wgeningen UR) to study this experimentlly in smll scle trils. In time-spn of three yers, 3 experiments were performed. Promising dpted light sources were provided by Philips Lighting. The following reserch questions were ddressed in the reserch: 1) Cn UV supplementtion reduce gentle nd severe fether pecking in lying hens? 2) Cn UV supplementtion increse desired types of pecking, such s ground pecking? 3) Cn UV supplementtion decrese ferfulness of the hens? 4) Are spects of hen development nd egg production ffected by UV-A supplementtion? Underlying questions were: ) Do white nd brown egg lyer breeds rect differently to dpted light environments? b) Is there n effect of light intensity in the rnge 2 to 4 lux? c) Are generl time budgets of other behviours ffected by dpted light environments, e.g those relted to resting, feeding or comfort? 3

16 2 Mterils nd Methods The effect of optimized lighting conditions on the behviour nd production of lying hens ws tested experimentlly with white nd brown egg lyer breed. This ws done in three consecutive experiments. All procedures involving niml hndling nd testing were pproved by the Animl Experimentl Committee of the Animl Sciences Group of Wgeningen University nd Reserch Centre. 2.1 Experimentl housing nd nimls The two breeds used in the experiments were Lohmn Brown (LB) hens nd Lohmn Selected Leghorns (LSL). These two breeds were obtined from commercil htchery (Verbeek, The Netherlnds), nd re known for their difference in behviourl trits nd ctivity (LSL more ctive). Accordingly they were lso expected to differ in propensity for fether pecking. The hens used for the experiments were not bek trimmed. Experiments 1 nd 2 were performed t the experimentl poultry fcility Spelderholt in Lelystd, The Netherlnds. Experiment 3 ws done in climte controlled fcility, lso in Lelystd, The Netherlnds. Figure 2. Experimentl poultry fcility Spelderholt (left) nd the climte controlled fcility (right), Lelystd, The Netherlnds In ech experiment, birds rrived s dy-old chicks. They were llocted t rndom to floor pens, s described in more detil under 2.2. Experimentl design. The pens were built of wire nd hens could not see their flock mtes in other pens by dditionl wooden prtitions between the pens. The pens used in experiments 1 nd 2 mesured.95 by.75 meter (height:.8 m) nd were supplied with wood shvings s substrte on the floor from dy to week 6 of ge. As floors were solid, the mteril on the floor grdully becme mixture of wood shvings, (dry) mnure, nd fethers. The pens in experiment 3 were vrible in size nd mesured mximlly 3. by 1. meter (vrible height: mximum 1.5 meter). The hens were housed on fully sltted floors t the strt of the experiment. Provision of substrte on prtly solid floor (1/3 of floor surfce) ws done t lter ge (see lso experiment 3). The pens were provided with perches (15 cm/hen) nd lying nests (5 hens per nest) when the hens were round 17 weeks of ge. The lying nests were plced outside ech pen. The helth sttus of the hens ws monitored dily. The vccintion scheme pplied in the rering period ws comprble to tht in commercil hen frming. The hens hd free ccess to feed nd wter. Feed ws provided in feeding troughs (length of.75 meter) nd wter ws vilble through nipple drinkers. From to 3 weeks of ge, the hens received stndrd commercil strter diet. During the further rering period (until the ge of 17 weeks), two different stndrd commercil rering (grower) diets were provided. From 17 weeks onwrds (experiment 3), hens received stndrd commercil lyer diet. The environmentl temperture ws lowered from 33 C on dy one to 2 C t 6 weeks of ge onwrds. On dys 1 nd 2 of ge the chicks received 24 hours of light. Between 2 nd 7 dys of ge, light ws on for 16 hours per dy. From 7 dys to 6 weeks of ge onwrds the light regime grdully decresed by one hour per week to 9 hour light period. Until 17 weeks of ge, this light regime 4

17 styed the sme. From this ge onwrds (experiment 3), the light period ws grdully extended by one hour per week to 15 hour light period t 23 weeks of ge. Light intensities vried ccording to experimentl design nd light source. All mterils concerning the light instlltions nd sources were supplied by Philips Lighting BV. In ll cses fluorescent tubes (PL) were used. 2.2 Experimentl design Experiment 1 In experiment 1, totl of 24 LSL nd 24 LB hens were used. The experiment ws done in the rering period only, i.e with young hens from birth to the ge of 17 weeks. The hens were housed in two identicl comprtments. Ech comprtment ws divided in 6 subcomprtments (rows) nd consisted of 4 pens. In ech pen, hens were housed in groups of (48 pens in totl). Within ech subcomprtment, one light source ws tested, together with the two breeds. Mingling of light between subcomprtments ws prevented by prtitioning by fences on which blck plstic ws ttched. Groups of hens were llotted to 1 of the 12 tretments ccording to 3 (light source) x 2 (light intensity) x 2 (breed) fctoril rrngement, with 4 replictes per tretment: Light source (code in itlics): 1. E1-Stndrd: white light (Mster PL-S 83/4P; 7 W). 2. E1-optimized-1: Combintion white light (mster PL-S 84/4P; 9W) nd Blcklight Blue (PL-S 9W/8 2P). 3. E1-optimized-2: dpted light source developed by Philips (detils unknown with experimenters). Light intensity (the 3 light sources were illuminted to the sme lux level) : 1. 2 lux (often recommended for commercil poultry frming) 2. 4 lux (more close to behviourl needs) Breed: 1. Lohmn Selected Leghorn (LSL) 2. Lohmn Brown (LB) Figure 3. Experimentl set-up Experiment 1: overview (left), E1-optimized-2 light (right) Experiment 2 Experiment 2 ws lso done in the rering period only. A totl of 24 LSL nd 24 LB hens were housed in the sme two comprtments s for experiment 1. Now ech comprtment ws divided into 12 subcomprtments. Ech subcomprtment consisted of 2 pens, in which one light source nd the two breeds of lying hens were tested. Different lux levels were used to rech the sme brightness of the light sources s perceived by the hens, with 4 lux for stndrd lighting s reference. As for experiment 1, hens were housed in groups of (48 pens in totl). Compred to experiment 1, dditionl fences with blck plstic were plced to prevent mingling of light between subcomprtments. Groups of hens were originlly ssigned to 1 of 12 tretments, ccording to 6 (light source) x 2 (breed) fctoril rrngement. However, it ppered during the experiment tht light emission of some 5

18 light sources did not fully meet the set requirements. Other chrcteristics, on the bsis of UV-A output, were therefore used for sttisticl nlysis. The fctoril setup therefore chnged into 3 (light source) x 2 rrngement. Light source (code in itlics): 1. E2-Stndrd: white light (Mster PL-S 83/4P; 7 W); UV-A output: to 7.5 (unit unknown with experimenters); 8 replictes. 2. E2-optimized-1: dpted light source developed by Philips; UV-A output 7.5 to 13 (detils nd unit unknown with experimenters); 34 replictes 3. E2-optimized--2: dpted light source developed by Philips; UV-A output 13 to 2 (detils nd unit unknown with experimenters); 6 replictes Breed: 1. Lohmn Selected Leghorn (LSL) 2. Lohmn Brown (LB) Figure 4. Experimentl set-up Experiment 2: E2-stndrd light (left), dpted light source (right) Experiment 3 A totl of 27 LSL nd 27 LB hens ws studied until the ge of 5 weeks. They were housed in 3 identicl climte-controlled rooms. Ech room consisted of 6 pens. These pens were stepwise enlrged in size ccording to ge of the birds: 1 m2 between nd 4 weeks, 1.25 m2 between 4 nd 7 weeks, 2 m2 between 7 nd 17 weeks, nd 3 m2 from 17 weeks of ge onwrds. Accordingly, the number of feed troughs lso incresed, from 1 between nd 7 weeks, 2 between 7-17 weeks, to 3 from 17 weeks onwrds. To stimulte fether pecking behviour, substrte ws not provided t the strt of the experiment nd floors were fully sltted. Moreover, group size ws enhnced compred to experiments 1 nd 2, to 3 hens per pen. However, due to n outbrek of severe fether pecking nd cnniblism in LSL hens round 4 to 5 weeks of ge, litter ws provided t 6 weeks of ge (mortlity t the end of rering: 8.5%). At tht point, fether pecking mong LB ws much less, nd litter ws provided t 14 weeks of ge (mortlity t the end of rering: 4.4%). The groups of hens were llotted to 1 of 6 tretments, ccording to 3 (light source) x 2 (breed) fctoril design. In ech room, ll combintions were present, nd ech tretment hd 3 replictes. Brightness of the light sources ws stndrdized, with 4 lux for stndrd lighting s reference: Light source (code in itlics): 1. E3-Stndrd: white light (Mster PL-S 83/4P; 7 W). 2. E3-optimized-1: dpted light source developed by Philips (code: green; detils unknown with experimenters) 3. E3-optimized-2: dpted light source developed by Philips (code: red; detils unknown with experimenters). Breed: 1. Lohmn Selected Leghorn (LSL) 2. Lohmn Brown (LB) 6

19 Figure 5. Experimentl set-up Experiment 3: E2-stndrd light (left), E3-optimized-1 light (right) 2.3 Observtions Pecking nd generl behviour To evlute the effect of lighting conditions, pecking behviour nd generl behviour were observed every 4 weeks in the rering period, i.e. t 4 (or 5), 8, 12 nd 16 weeks of ge. Additionlly, in the lying period observtions were done t 24, 32, 4 nd 48 weeks of ge (experiment 3). In n observtion week, the behviour of the hens ws observed in the home pen. Observtions were lwys performed between 8:3.m. nd 5: p.m., by either two (experiment 3) or 3 observers (experiment 1 nd 2). Observtions were crried out on two dys per test week, usully on Tuesdy nd Thursdy. On one dy, the behviour of the hens in ll pens ws noted 8 times (in totl 16 times per pen in one observtion week). The occurrence of pecking behviour (see Tble 1; ethogrm) ws determined through one-minute scn smplings, following dpttion periods of one minute. Pecking behviour ws either scored s seen or not seen, independent of the number of nimls engged in the behviour. Subsequently, t the end of the one-minute observtions, generl time budgets of vrious other behviours ws noted, by counting the number of hens performing certin behviour (see Tble 1; ethogrm). 7

20 Tble 1. The ethogrm used for observtion of behviour in the home pen Lbel Description Pecking behviours Gentle fether pecking Severe fether pecking Aggressive pecking Ground or litter pecking Other behviours Wlking Scrtching Eting nd drinking Resting Preening Dustbthing Mild or gentle pecking t fethers of other hens, generlly performed in multiple bouts, no removl of fethers. No rection of recipient. Severe or forceful pecking on or vigorous pulling the fethers of other hens. This my be single pecks or pulls. Cn result in wounding the other bird. Often with rection of recipient. Forceful pecks, directed t hed or neck region. Pecking t the floor or litter. Wlking, running, jumping or flying. Ground scrtching, hed usully in lower position thn the rump. Eting from food trough or drinking from nipple drinkers/cups. Sitting or stnding inctive (no movement of the legs), perching. Comfort behviour: utopecking, nibbling, stroking, combing, hed rubbing Responses to humn presence nd novelty Comfort behviour: lying down on the floor or in substrte nd trying to work dust or other loose mteril into the fethers. As fether pecking is suggested to be ssocited with ferfulness, simple behviourl tests were done to ssess fer for humns nd generl fer. Fer for humns ws ssessed by humn pproch test. This test ws performed three times in experiment 1 (t 5, 8 nd 16 weeks of ge) nd once in experiments 2 (t 5 weeks of ge) nd 3 (t 15 weeks of ge). From distnce of 1.5 meter, n experimenter wlked stright forwrd to pen, en then stood still t distnce of.5 meter from the front of the pen. Ech pen ws virtully divided in two equl sections: bck nd front section. The experimenter then noted the ltency time for five hens to pproch the experimenter by entering the front section of the pen. The test lsted mximlly 2 minutes. Generl fer ws ssessed in experiments 2 nd 3 by introducing unfmilir or novel objects into the pens (experiment 2: t 15 weeks; experiment 3: t 15 nd 44 weeks). The novel objects were, respectively, bottle of shmpoo in experiment 2, nd piece of white pper (15 weeks) nd coloured stick (44 weeks) in experiment 3. Both objects were plced in the front section of the pens. In experiment 2, the ltency time for the first hen to enter the front section of the pen ws determined. In experiment 3, it ws determined when the piece of pper ws first pecked t. Agin, mximum time of the test ws set on 2 minutes. For the test with the coloured stick, during 2 minutes the number of hens within henlength of the stick ws noted every 3 seconds (expressed in % of number of hens in pen; cummultive score). 8

21 2.3.3 Plumge nd skin condition scores Plumge nd skin condition were evluted to ssess the extent of problems ssocited with fether pecking. Dmge to the fethers nd skin of hens ws ssessed by method slightly dpted from Bilcik nd Keeling (1999). This method ssesses fether nd skin condition on body prts. The scle ws (intct fethers) to 5 (completely bld) for fether condition, nd 5 (intct skin; no injury) to 9 (severe wounds) for skin condition. Furthermore, n overll score ws given for dirtiness: (clen) to 3 (dirty). The condition scores were obtined t 6, 9, 13 nd 17 weeks of ge in experiments 1 nd 2, nd t 8 (only LSL hens) nd 17 nd 5 weeks of ge in experiment 3. All hens were scored Body weight nd uniformity Development of the hens ws determined by informtion on body gin nd uniformity. The hens were weighed per pen t the strt of ll experiments nd t 5 weeks of ge in experiment 3. At 17 weeks of ge, ll hens were weighed individully. Informtion on individul weights ws used to determine uniformity, ie the percentge of hens tht ws within % (+/- %) of verge weight. The higher uniformity, the more hens re in the sme stge of development, which is beneficil for efficient pplying feeding nd lighting schedules in the lying period Feed consumption nd egg production In experiment 3, feed consumption ws recorded every 4 weeks, nd egg production weekly, from 2 weeks of ge onwrds. Feed consumption ws determined by weighing feed nd feed troughs every 4 weeks, on fixed dy of the week. Eggs were collected ech dy, nd floor eggs were kept seprtely. Once week, lso on fixed week dy, the number of eggs ws counted to determine rte of ly nd weight of the eggs. The proportion of second grde eggs nd eggs with open breks ws determined. The informtion on egg weight nd feed intke ws lso used to ssess feed conversion, i.e. kg feed/kg egg. 2.4 Sttisticl nlysis Anlyses were performed with Genstt 11.1 (28, VSN Interntionl Ltd, GenStt Procedure Librry Relese PL19.1). Significnce level ws set t.5. Split-plot design were used for experiments 1 nd 2, nd completely rndomized block design ws used for experiment 3. In experiments 1 nd 3, continuous dt, e.g. responses to novel object, body weight, feed intke, egg weight, were nlyzed with nlysis of vrince (ANOVA). In experiment 2, continuous dt were nlyzed by regression nlysis (liner mixed model REML, fits vrince-components model by residul (or restricted) mximum likelihood). For determining differences between light sources nd interctive effects the procedures PAIRTEST (performs t-tests for pirwise differences) en PPAIR (displys results of t-tests for pirwise differences) were used. Percentges (close to ), e.g for behviours, mortlity, floor eggs, second grde eggs, were nlyzed with n IRREML procedure, which fits devince-components model by residul (or restricted) mximum likelihood). For determining differences between light sources nd interctive effects the procedures RPAIR en PPAIR were used. RPAIR gives t-tests for ll pirwise differences of mens from regression or GLM nd PPAIR displys results of t-tests for pirwise differences. Plumge nd skin condition scores were nlyzed by the threshold model of McCullgh or by logistic regression when the exterior scores were uneqully divided. 9

22 3 Results Behviourl ctivity time budgets nd other chrcteristics often vried substntilly between the two breeds. Therefore, it ws decided to present the results seprtely for LSL nd LB hens. In experiment 1, results for the two light intensities only occsionlly differed significntly. Therefore, results for light intensity re not presented seprtely in tbles or figures, but where pplicble differences re mentioned. 3.1 Effect of optimiztion of lighting conditions on fether pecking behviour Gentle fether pecking Rering period Results for gentle fether pecking in the rering period re shown in Figure 6. Levels of gentle fether pecking with optimized lighting re presented in comprison with the results under stndrd lighting conditions (-line). In the first experiment, t 5 weeks of ge, the occurrence of gentle fether pecking ws decresed with E1-optimized-1, compred to E1-stndrd lighting (significnt min effect for the fctor light source: p<.1). In ddition, but in the presence of both optimized light sources, gentle fether pecking ws reduced t 8 weeks of ge. This effect ws only observed in Lohmn Brown hens (significnt interction between the fctors breed nd light source: p<.1). A decrese in gentle fether pecking, specificlly for 8 weeks old LSL hens, ws observed in experiment 2 with E2-optimized-2 lighting, when compred to E2-stndrd nd E2-optimized-1 conditions (significnt interction between the fctors breed nd light source: P<.5). In experiment 3, high gentle fether pecking levels were observed. At 5 weeks of ge, gentle fether pecking levels seemed lower in LSL hens with E3-optimized-1. The interction between the fctors light source nd breed ws, however, not significnt. Lying period Figure 7 shows levels of gentle fether pecking during the lying period of experiment 3. At 24 weeks of ge, gentle fether pecking ws decresed in LSL hens nd incresed in LB hens with E3- optimized-1 light reltive to the other two light sources. The interctive effect of light source nd breed lmost reched significnce (p=.1). At 4 weeks, gentle fether pecking tended to be lower with the two optimized light sources, but only in LSL hens (interction between the fctors breed nd light source: p=.1). Gentle pecking seemed to be decresed with the two dpted light sources t 48 weeks in LB hens, but effects were not ner significnce.

23 Exp 1 Gentle pecking LSL Exp 1 Gentle pecking LB lighting (-line) (%) ** E1-stndrd E1-optimized-1 E1-optimized-2-15 (26) (16) (14) (13) -2 (23) (25) (9) (7) lighting (-line) (%) ** b b E1-stndrd E1-optimized-1 E1-optimized-2 Exp 2 Gentle pecking LSL Exp 2 Gentle pecking LB lighting (-line) (%) ** b E2-stndrd E2-optimized-1 E2-optimized-2-15 (19) (16) (14) (27) (16) (9) (11) (9) lighting (-line) (%) ** E2-stndrd E2-optimized-1 E2-optimized-2 Exp 3 Gentle pecking LSL Exp 3 Gentle pecking LB 3 3 lighting (-line) (%) E3-stndrd E3-optimized-1 E3-optimized-2 (71) (85) (6) (33) (15) (56) (38) (15) lighting (-line) (%) E3-stndrd E3-optimized-1 E3-optimized-2 Figure 6. Gentle fether pecking in experiments 1 to 3: rering period. The vlues between brckets represent bsolute percentges with stndrd lighting. **When letters re presented, there is (significnt) interction between the fctors breed nd light source; different letters present significnt within breed-differences. 11

24 Exp 3 Gentle pecking LSL Exp 3 Gentle pecking LB 3 3 lighting (-line) (%) (29) (19) (21) () E3-stndrd E3-optimized-1 E3-optimized-2 lighting (-line) (%) (15) (27) (33) (44) E3-stndrd E3-optimized-1 E3-optimized-2 Figure 7. Gentle fether pecking in experiment 3: lying period. The vlues between brckets represent bsolute percentges with stndrd lighting Severe fether pecking Rering period In experiments 1 nd 2, levels of severe fether pecking were rther low for ll tretment groups, nd no differences were observed between tretment groups (Figure 8). Severe fether pecking in experiment 3, however, reched high levels t rther young ge, especilly observed in LSL hens under stndrd lighting t 5 weeks of ge (but no significnt effect of light source). To control the outbrek of fether pecking mong LSL hens, litter ws provided t 6 weeks of ge. At lter ge, fether pecking lso incresed in Lohmn Brown hens, nd ccordingly litter ws provided t 14 weeks of ge. As cn be seen in Figure 8, optimized lighting conditions rther incresed thn suppressed severe fether pecking in Lohmn Brown hens in the bsence of litter, t 8 nd 12 week of ge (but no significnt or nerly significnt interctions). Lying period In ccordnce with the erlier observtions of the hens in experiment 3 t 8 nd 12 weeks of ge, both optimized light sources pprently enhnced severe fether pecking in LB hens (t 4 nd 48 weeks of ge), but not in LSL hens (Figure 9). Differences between tretment groups did, however, not (nerly) rech significnce, due to high vrition between replictes. 12

25 Exp 1 Severe pecking LSL Exp 1 Severe pecking LB lighting (-line) (%) b -6 b (26) (12) (16) (7) (14) (1) (13) () -8 (23) (2) (25) () (9) (1) (7) () E1-stndrd E1-optimized-1 E1-optimized-2 lighting (-line) (%) ** E1-stndrd E1-optimized-1 E1-optimized-2 Exp 2 Severe pecking LSL Exp 2 Severe pecking LB lighting (-line) (%) (2) () (2) () E2-stndrd E2-optimized-1 E2-optimized-2 lighting (-line) (%) (2) () (2) (2) E2-stndrd E2-optimized-1 E2-optimized-2 Exp 3 Severe pecking LSL Exp 3 Severe pecking LB lighting (-line) (%) (63) (29) (31) () E3-stndrd E3-optimized-1 E3-optimized-2 lighting (-line) (%) () () (48) () E3-stndrd E3-optimized-1 E3-optimized-2 Figure 8. Severe fether pecking in experiments 1 to 3: rering period. The vlues between brckets represent bsolute percentges with stndrd lighting. 13

26 Exp 3 Severe pecking LSL Exp 3 Severe pecking LB 5 5 lighting (-line) (%) (15) () () (6) E3-stndrd E3-optimized-1 E3-optimized-2 lighting (-line) (%) (15) (13) (8) (31) E3-stndrd E3-optimized-1 E3-optimized-2 Figure 9. Severe fether pecking in experiment 3: lying period. The vlues between brckets represent bsolute percentges with stndrd lighting Ground pecking Rering period Figure shows the results for ground pecking behviour. In experiment 1, more ground pecking ws observed with E1-optimized-2 t 5 weeks of ge (83%), compred to the E1-stndrd nd E1- optimized-1 light sources (respectively, 73 nd 74%; significnt min effect for the fctor light source: p<.5). Experiment 2 showed significnt effects of light source on ground pecking t severl ges. At 8 weeks of ge, ground pecking levels were generlly higher with E2-optimized-2 (75%; 66 nd 67%, for E2- stndrd lighting nd E2-optimized-1, respectively; significnt min effect for the fctor light source: p<.5). lighting, ground pecking ws either more frequently (LB hens, 12 weeks) or less frequently observed (LSL hens, 16 weeks) under E2-optimized-2 lighting conditions (significnt interctions between the fctors breed nd light source: p<.5) In experiment 3, ground pecking ws stimulted by provision of litter from 6 weeks of ge onwrds in LSL, nd from 14 weeks onwrds in LB hens. At 12 weeks of ge, LSL hens ppered to hve more pecks directed to the litter with both optimized light sources, but the increses were not significnt. Results for LB hens re not shown in Figure, s there ws only one observtion period, ie t 16 weeks of ge. Ground pecking levels did not differ for LB hens t this ge: 9, 85 nd 88%, for E3- stndrd, E3-optimized-1, nd E3-optimized-2, respectively. Lying period At 32 weeks of ge, but not t the other ges nd only observed with LB hens, ground pecking ws incresed with E3-optimized-1 (Figure 11; significnt interction between the fctors breed nd light source: p=.5). 14

27 Exp 1 Ground pecking LSL Exp 1 Ground pecking LB 2 2 lighting (-line) (%) (26) (79) (16) (77) (14) (45) (13) (37) E1-stndrd E1-optimized-1 E1-optimized-2 lighting (-line) (%) (67) (59) (31) (3) E1-stndrd E1-optimized-1 E1-optimized-2 Exp 2 Ground pecking LSL Exp 2 Ground pecking LB lighting (-line) (%) ** ** b b (86) (81) (78) (61) E2-stndrd E2-optimized-1 E2-optimized-2 lighting (-line) (%) ** (61) (52) (45) (14) b ** E2-stndrd E2-optimized-1 E2-optimized-2 Exp 3 Ground pecking LSL 2 lighting (-line) (%) () () () (81) (48) () (88) E3-stndrd E3-optimized-1 E3-optimized-2 Figure. Ground pecking in experiments 1 to 3: rering period. The vlues between brckets represent bsolute percentges with stndrd lighting. **When letters re presented, there is (significnt) interction between the fctors breed nd light source; different letters present significnt within breed-differences. 15

28 Exp 3 Ground pecking LSL Exp 3 Ground pecking LB 2 2 lighting (-line) (%) ** -15 () (94) (96) (92) E3-stndrd E3-optimized-1 E3-optimized-2 lighting (-line) (%) ** b b -15 (88) (85) (92) (83) E3-stndrd E3-optimized-1 E3-optimized-2 Figure 11. Ground pecking in experiment 3: lying period. The vlues between brckets represent bsolute percentges with stndrd lighting. **When letters re presented, there is (significnt) interction between the fctors breed nd light source; different letters present significnt within breed-differences. 3.2 Effect of optimiztion of light on generl behviour Very often, time spent in vrious other behviours thn those relted to pecking, did not differ between tretment groups in the 3 experiments. Rering period For resting nd comfort behviours, but never for wlking, scrtching nd eting/drinking behviours, some significnt differences were found between tretment groups t specific ges. These re mentioned here. In experiment 1, with E1-optimized-2, t 12 weeks of ge more resting behviour ws observed t 2 lux (56%) compred to 4 lux (46%) (significnt interction between fctors light source nd light intensity: p<.5). In experiment 2, comfort behviour ws incresed with E2-optimized-2 t two ges. At 8 weeks of ge, more dustbthing ws observed (1.4%), compred to the other two light sources (both.7%) (significnt min effect for the fctor light source: p<.1). At 12 weeks of ge, hens housed with E2- optimized-2 showed more preening (13%), nd the min effect for light source tended to be signicnt (p=.1) (stndrd: 8%; E2-optimized-1: 6%). In experiment 3, t 5 weeks of ge, hens showed more comfort (preening) behviour with both optimized light sources (E3-optimized-1: 4.4% ; E3-optimized-2: 4.6%) compred to E3-stndrd lighting (2.8%) (significnt min effect for the fctor light source: p<.5). A similr effect (min effect light source: p<.5) ws observed t 16 weeks of ge, but then n increse in preening ws only seen in the presence of E3-optimized-1. Lying period Generl behviour ws not ffected by light source. 3.3 Effect of optimiztion of light on fer for humns nd generl fer Results re shown in Tble 2 for both rering (experiments 1 to 3) nd lying periods (experiment 3). At 5 weeks of ge in experiment 1, LB hens took longer to enter the front prt of the cge in humn presence with E1-stndrd lighting compred to E1-optimized-1 lighting. Introduction of novel object (bottle) in experiment 2 showed tht it lsted longer for LSL hens to pproch the object with E2-stndrd lighting compred to E2-optimized-2. Findings in experiment 3 were comprble with those in experiment 2: hens with stndrd lighting generlly tended to hve longer ltency to peck t the novel object (piece of pper), compred to E3-optimized-2. At 44 weeks, the percentge of hens within henlength of coloured stick did not differ between tretments. 16

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3.

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3. The nlysis of vrince (ANOVA) Although the t-test is one of the most commonly used sttisticl hypothesis tests, it hs limittions. The mjor limittion is tht the t-test cn be used to compre the mens of only

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