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        A user's guide to animal welfare science
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1.       fee ee    ELSEVIER    TRENDS in Ecology and Evolution Vol 21 No 2 February 2006    Full text provided by www sciencedirect com    science  Q  DIRECT      A user s guide to animal welfare    science    Marian Stamp Dawkins    Department of Zoology  South Parks Road  Oxford  OX1 3PS  UK    Here    provide a guide for those new to the burgeoning  field of animal welfare science as to what this compre   hensive  relatively young discipline is all about  Drawing  on all branches of biology  including behavioural  ecology and neuroscience  the science of animal welfare  asks three big questions  Are animals conscious  How  can we assess good and bad welfare in animals  How  can we use science to improve animal welfare in  practice    also provide guidelines for an evidence   based approach to welfare issues for policy makers  and other users of animal welfare research     Introduction   Animal welfare issues increasingly force themselves  onto everyone   s agenda  scientists have to provide  environmental enrichment for their animals for welfare  reasons  1   zoos are under pressure for confining wild  animals in cages  2   farming practices are criticized and  debated  3  and people are urged to buy free range eggs  and meat on the grounds that the welfare of the animals  is better  Many welfare issues are so vast that people  avoid thinking about them  For example  2 10    broiler  chickens are killed for meat worldwide each year  4    and uncounted millions of rodents are 
2.  issues     what improves animal health and what the  animals themselves want     will help to bring about  genuine improvements to animal welfare in practice and  ensure that legislators can make evidence based  decisions  Although the most controversial issues in  animal welfare concern the way humans treat captive  and domestic animals  the ways those animals respond are  rooted in their evolutionary past and in how their wild  ancestors responded to threats to their fitness  Beha   vioural ecologists thus have a major contribution to make  to animal welfare science by connecting this evolutionary  legacy to what now matters to the animals themselves     Acknowledgements  I thank Roland Bonney  Tracey Jones  Ruth Layton and Edmund Rolls for  their comments     References  1 Benefiel  A C  et al   2005  Mandatory    enriched    housing of laboratory  animals  the need for evidence based evaluation  ILAR J  46  95 105  2 Hosey  G R   2005  How does the zoo environment affect the behaviour  of captive primates  Appl  Anim  Behav  Sci  90  107   129  Weaver  S A  and Morris  M C   2004  Science  pigs and politics  a New  Zealand perspective on the phase out of sow stalls  J  Agric  Environ   Ethics 17  51   66  4 Scientific Committee on Animal Health and Animal Welfare  2000   The Welfare of Chickens Kept for Meat Production  Broilers   European  Commission  http   europa eu int comm food fs sc scah out39_eupdf   Mason  G J  and Littin  K E   2003  The humaneness of rodent pes
3.  make the link    www sciencedirect com    of investigation might be  the    honesty    or otherwise of signals  53    the dynamics of weaning conflicts  the way animals balance anti   predator strategies with their other needs and the role of social  facilitation decision making  Just understanding how the time  budgets of captive or domestic animals differ from those of their  wild or feral counterparts provides a baseline or jumping off point for    f  s     a bs  zi on a  4   gt      iyo Mo   ASS Hee CA  Taa PRANAN NATAN ASINA       S A     mae Wy vrs Se    mre       Figure I  Pig rearing system inspired by a study of feral pig behaviour  59    Photograph reproduced with permission from FAI  http   www  faifarms co uk      assessing welfare   between the complexity of behaviour and conscious    experience  For example  chimpanzees can recognize  themselves in mirrors in the sense that they use mirrors  to respond to  or examine parts of  their bodies that they  cannot usually see  22   Although this might indicate self   awareness and a conscious knowledge of self  23 24   there  is no necessary conscious component at all  25     The complexity of social life has been suggested as the  driving force for the evolution of consciousness  on the  grounds that deception  cheating and trying to  outwit cheaters require a particularly complex type of  forward planning and perspective taking  26   However   apparently complex outcomes can result from simple  rule following  27   
4.  org articles groups   conservation bes_ieem_conf     18 22 July 2006  International Fish Biology  St Johns  Canada  http   www heb pac dfo mpo gc ca congress     23 29 July 2006   11th International Behavioral Ecology Congress  Tours   France   http  Awww isbe2006 com     6 10 August 2006  Animal Behaviour Society  Salt Lake City  USA  http   www animalbehavior org ABS Program Snowbird05     www sciencedirect com       
5. 05  Two neural correlates of consciousness  Trends Cogn   Sci  9  46 52   Ramirez  J M  and Cabanac  M   2003  Pleasure  the common currency  of emotions  Ann  N  Y  Acad  Sci  1000  293   295   Sherwin  C M  and Olsson  A S   2004  Housing conditions affect self   administration of anxiolytic by laboratory mice  Anim  Welf  13  33 38  Danbury  T C  et al   2000  Self selection of the analgesic drug  carprofen by lame broiler chickens  Vet  Rec  146  307 311   Bermond  B   2001  A neuropsychological and evolutionary  approach to animal consciousness and animal suffering  Anim   Welf  10  S47 S62   Rutherford  K M D   2002  Assessing pain in animals  Anim  Welf  11   31 54   Moloney  V  et al   2002  Validation of a method for assessment of acute  pain in lambs  Appl  Anim  Behav  Sci  76  215   238   Gentle  M J   1992  Pain in birds  Anim  Welf  1  235 247   Sneddon  L U  et al   2003  Do fish have nocioceptors  Evidence for the  evolution of a vertebrate sensory system  Proc  R  Soc  Lond  B Biol   Sci  270  1115 1121   Rose  J D   2002  The neurobehavioral nature of fishes and the  question of awareness and pain  Rev  Fish  Sci  10  1 38   Chandroo  K P  et al   2004  Can fish suffer  Perspectives on sentience   pain  fear and stress  Appl  Anim  Behav  Sci  86  225   250   Sherwin  C M   2001  Can invertebrates suffer  Or how robust is the  argument from analogy  Anim  Welf  10  S103 S118   Chapman  C R  and Nakamura  Y   1999  A passion for the soul  an  introduction to 
6. 1993  Affect  cognition and awareness      affective priming with optimal and suboptimal stimulus exposures   J  Pers  Soc  Psychol  64  123   739  21 Griffin  D R  and Speck  G B   2004  New evidence for animal  consciousness  Anim  Cogn  7  5 18    Go    On    www sciencedirect com    22    23    24    25    26    27    28    29    30    31    32    33    34    35    36    37    38    39    40    41    42    43    44    45    46    47    48    49    50    51    52    53    Gallup  G G   Jr   1970  Chimpanzees  self recognition  Science 167   86 87   Platek  S M  et al   2004  Where am I  The neurobiological correlates  of self and other  Cogn  Brain Res 19  114 122    Bekoff  M  and Sherman  P W   2003  Reflections on animal selves   Trends Ecol  Evol  19  176   180   Heyes  C M   1998  Theory of Mind in non human primates  Behav   Brain Sci  21  101   114   Byrne  R W  and Corp  N   2004  Neocortex size predicts deception rate  in primates  Proc  R  Soc  Lond  B Biol  Sci  271  1693   1699   Sachs  J L   2004  The evolution of cooperation  Q  Rev  Biol  79   135   160   Chen  M K  and Hauser  M   2005  Modelling reciprocation and  cooperation in primates  evidence for a punishing strategy  J  Theor   Biol  235  5 12   Bshary  R  and Schaffer  D   2002  Choosy reef fish select cleaner fish  that provide high quality service  Anim  Behav  63  557   564  Hampton  R R   2001  Rhesus monkeys know when they remember   Proc  Natl  Acad  Sci  U  S  A  98  5359 5362   Block  N   20
7. 2004  Using behaviour to assess welfare  Anim  Welf   13  S38 S7   Patterson Kane  E H  et al   2002  Rats demand social contact  Anim   Welf  11  327 332   Mason  G J  et al   2001  Frustrations of fur farmed mink  Nature 410   35 36   Whay  H R  et al   2003  Animal based measures for the assessment of  welfare state of dairy cattle  pigs and laying hens  consensus of expert  opinion  Anim  Welf  12  205   217   Farm Animal Welfare Council  1993  Second Report on Priorities for  Research and Development in Farm Animal Welfare  Ministry of  Agriculture Fisheries and Food   Stolba  A  and Wood Gush  D G M   1989  The behaviour of pigs in a  semi natural environment  Anim  Prod  48  287   298   Mason  G J  and Latham  N R   2004  Can   t stop  won   t stop  is stereotypy  a reliable animal welfare indicator  Anim  Welf  13  S57   S69   Nicol  C J  et al   2002  Study of crib biting and gastric inflammation  and ulceration in young horses  Vet  Rec  151  658 660   Millman  S T  et al   2004  The impact of applied ethologists and the  International Society of Applied Ethologists in improving animal  welfare  Appl  Anim  Behav  Sci  86  299 311   Mellor  D J   2003  Guidelines for the humane slaughter of the fetuses  of pregnant ruminants  Surveillance 30  26   28    64    65    66    67    68    69    70    71    72    Grandin  T   2003  Transferring results of behavioural research to  industry to improve animal welfare on the farm  ranch and the  slaughter plant  Appl  Anim  Be
8. For example     cheats    do not have to  be detected by an elaborate working out of    what would I  do in his place      but lose out to simple strategies such as  tit for tat and its variants  28  in which animals  remember what happened in past encounters and act  accordingly  Even simpler    win stay lost shift    strategies  result in animals moving away from places where social  rewards are not forthcoming  29  so that non cooperators  automatically lose out on long term benefits with no  consciousness on either side    Recently  particular attention has been focused on the  possibility that animals monitor the state of their own  memories  Monkeys perform better on some tasks when  allowed to choose for themselves whether to take the test  or an easier but less rewarding option than when the  experimenter decides for them  suggesting that the  monkeys knew how much they knew  380   Although we  should not forget that computers can be programmed to    TRENDS in Ecology and Evolution Vol 21 No 2 February 2006 79    behave differently depending on how uncertain they are  about an outcome  experiments of this sort have opened up  new possibilities for asking animals about what they  think they know even though we cannot use words to  ask them  9 11   Studies of animal cognition continue to  amaze us with what animals can do and deliver a major  assault  if not a complete solution  to the problem of  animal consciousness     Pleasure and suffering   The second category of 
9. It would not have to be  conscious  The gap between doing and feeling is still there   however small it might seem     Anatomy and brain function  The most direct way of deciding whether animals are  conscious would be to compare their physiological equip   ment and brain functions with our own  85   Pain in  mammals  36 37  and  toa lesser extent  birds  38  has been  well studied and  more recently  the discovery of nociocep   tors in fish  39  has led to the view that they also feel pain  and can suffer  40 41   On the basis of physiological and  behavioural responsiveness  even invertebrates cannot be  dismissed  42   But definite answers to the questions of  whether they consciously experience pain as we do  does  their pain hurt   and suffering  does it matter to them    remain frustratingly elusive  10 43   particularly as our  own pain can be modulated by input from different brain  pathways  44   Far from resolving the question of animal  consciousness  comparative brain anatomy has lead  McPhail  16  to conclude that only humans are conscious   and Baars  45  to come to the different conclusion that at  least all vertebrates are    Even the promise of being able to use non invasive  brain imaging techniques such as PET and fMRI  to    www sciencedirect com    compare human and animal brains  23  will be a long time  bearing fruit  It has been more difficult than anticipated to  find the neural basis of consciousness in humans  12 31   Consciousness  as an evolved fe
10. ature  is properly part of  biology  but  it is difficult to study     How can good and bad welfare be recognized    The second major question that animal welfare science deals  with is how to measure and assess animal welfare  7 46    Poor physical health  caused by disease  injury or deformity   is relatively straightforward to recognize and can often be  quantified  for example  by scoring how well an animal is  walking or the size of lesions on its body  Other  less obvious  measures of decreased health  such as depressed immune  function  47   resource dependent changes in the sex ratio of  offspring and reduced food intake  48   have recently been  imported from other disciplines  These can often give an  indication that all is not well with the animal before clinical  symptoms become obvious  To go further and investigate the  mental health state of an animal  wecan use both physiology  and behaviour    The physiological measures of welfare that have been  used so far have been autonomic responses  such as  increased heart rate and raised levels of hormones such as  corticosteroids     stress hormones     7 49   Although these  are objective measures  they can be difficult to interpret in  welfare terms because many of these changes are part of  the adaptive way in which the animal responds to its  environment  and because apparently pleasurable activi   ties  such as sex and hunting prey  can lead to similar  changes to those that are apparently unpleasant  such as  e
11. ced welfare  7  and those that are positively  beneficial for the animal  60  can equally be established  using the    two question    approach  Some stereotypies  actually seem to benefit the health of the animal  Repeated  biting of wooden doors or food troughs shown by some  stabled horses  61   for example  is associated with reduced  gastric ulcers  possibly because the    abnormal    behaviour  stimulates the production and swallowing of salive which  in turn protects the stomach from excess acid  Others  such  as bar biting in restricted sows  where the animal rubs its  mouth until it bleeds  indicate poor welfare on the most  basic health grounds  7   As emphasized earlier  there is no  one single measure of welfare     How can science be used to improve animal welfare in  practice    Having measured good and bad welfare  it is then  necessary to translate what has been found into    Box 2  Improving animal welfare in practice    Genuine improvements in the welfare of animals are most likely to  occur if legislators and other users of animal welfare science follow  certain key guidelines     i  Suggested improvements in animal welfare  such as environ   mental enrichments  giving animals more space  should be based on  answers to two questions  will they genuinely improve animal  health  Will it give the animals something they want     ii  Many different sorts of measurements     health  behaviour and  physiology     might have to be made to answer these as there is n
12. cial farms in cooperation with farmers  and other    end users    will greatly increase the applica   bility of the results of welfare research compared to  relying on smaller scale studies carried out in a laboratory   This involvement can take the form of epidemiological  studies of what is currently happening on farms  65  and  even participation in farm level experiments  For  example  11 major broiler chicken producers in the UK  and Denmark recently agreed to manipulate the stocking  density     crowding     as measured by kgm     of whole  chicken houses in a coordinated experiment that involved   gt 2 7 million birds  66   This gave policy makers a better  preview of the probable effect of enacting legislation to  change stocking density than any small scale laboratory  could have done and indicated that limiting stocking  density would have much less effect on chicken welfare  than previously supposed  Working directly with farmers  has the further advantage that other factors  such as food  safety  can be studied simultaneously so that policy  decisions can be based not only on what is best for animal  welfare  but also in the wider context of human health   effects on the environment and what the public wants to  see happening  67     Second  critically evaluating already published data  through the evidence based approach that has revolutio   nized clinical medicine  68  and is beginning to be used in  veterinary medicine  69  and conservation  70 71  greatly  inc
13. e hardest problem in biology  12 13   We have no idea  how the populations of nerve cells that comprise our  brains give rise to conscious thoughts and emotions or  even the most basic feelings of pain and hunger   sentience  in ourselves  This ignorance of how our own  consciousness is caused makes it difficult to know what to  look for as evidence for it in other species  particularly  those  such as birds  whose brains are anatomically  different from ours  14     The situation is further complicated by recent dis   coveries from psychology that many complex actions in  humans  e g  driving a car or playing a musical instru   ment  can be carried out quite unconsciously  11 12 15      www sciencedirect com 0169 5347     see front matter    2005 Elsevier Ltd  All rights reserved  doi 10 1016 j tree 2005 10 017    78 TRENDS in Ecology and Evolution Vol 21 No 2 February 2006    Box 1  The behavioural ecology of animal welfare  a personal view     Adopting an evolutionary approach could help us to understand what  might improve animal welfare  for the following reasons     i  The domesticated and captive species whose welfare  concerns so many people carry with them an evolutionary legacy  of responses that  in the wild  helped them to survive and  reproduce  e g  fleeing from predators  seeking water or shelter    Figure  l      ii  These mechanisms are of two types  direct threats to fitness   which can be assessed as reduced health and loss of reproductive  success  and ant
14. evidence for animal consciousness  centres more explicitly on basic sensations of pain   pleasure and suffering  also called sentience or     phenomenal consciousness     31   One approach is to ask  humans about their conscious experiences of pain or  pleasure and then to use the similarity of behaviour of  humans and other animals  such as rats  when both are   say  deprived of water  to infer that the thirsty rat  similar  the thirsty human  has the same conscious experience of  pleasure in the taste of water  32     Impressive in this context is the phenomenon of self   medication  i e  animals learning to give themselves drugs  that are known to give relief from pain or anxiety in  humans   33   Broiler chickens given the choice between  two coloured foods  one of which contained Carprofen  a  non steroidal anti inflammatory drug that relieves pain in  humans  will learn to choose the colour with the analgesic   but only if they themselves are lame and have obvious leg  weaknesses  Healthy birds have no such preference  34    For many people  this is enough to show that birds with  bad legs experience pain in the same way that we do  but  strictly speaking  a non conscious explanation is possible   A vehicle could be designed to fill itself with one of several  kinds of lubricant  based on which one had  in the past   been shown to result in the lowest fuel consumption  It  could be said to find one lubricant more    rewarding     more  likely to be ingested in future   
15. g that access to water to  swim in is something that is important to them    Faced with a diversity of different welfare    measures      some of which seem to contradict each other  animals do    80 TRENDS in Ecology and Evolution Vol 21 No 2 February 2006    not always choose what is best for their long term health   for example   animal welfare scientists have taken one of  two complementary approaches  One strategy is to take  the sum of as many measures as possible  behavioural   health  physiological  and to use a check list of different  ones  57   and to    triangulate    on what is good welfare  The  other is to focus on just the two questions     are the animals  healthy    and    do they have what they want      54   For  example  the fifth of the    Five Freedoms    that the Farm  Animal Welfare Council  58  argued were essential to good  welfare is the    Freedom to perform most natural patterns  of behaviour     But is natural behaviour essential for  welfare  After all  being chased by a predator is natural  for most wild animals but it is not obviously good for  welfare  By asking whether the health of an animal is  improved by being allowed to behave naturally and  whether the animal shows evidence of wanting to do the  behaviour  we can distinguish between natural behaviour  that does improve welfare and that which might not  59    Similarly  the distinction between stereotypies  repeated  fixed behaviours with no apparent function  that might  indicate redu
16. hav  Sci  81  215   228   Nicol  C J  et al   2003  Matched concurrent case control study of risk  factors for feather pecking in hens on free range commercial farms in  the UK  Br  Poult  Sci  44  515   523   Dawkins  M S  et al   2004  Chicken welfare is influenced more by  housing conditions than by stocking density  Nature 427  342 344  Glass  C A  et al   2005  Measuring the value to the public of pig  welfare imporovements  a contingent valuation approach  Anim  Welf   14  61 69   Abalos  E   2005  The tools and techniques of evidence based  medicine  Best Pract  Res  Clin  Obstet  Gynaecol  19  15   26   Aragon  C L  and Budsberg  S C   2005  Applications of evidence   based medicine  Cranial cruciate ligament injury repair in the dog   Vet  Surg  34  93 98   Sutherland  W J  et al   2004  The need for evidence based  conservation  Trends Ecol  Evol  19  305 308   Cote  I M  et al   2005  Measuring coral reef decline through meta   analysis  Philos  Trans  R  Soc  Lond  B Biol  Sci  360  385   395  Bashaw  M J  et al   2001  A survey assessment of variables relating to  stereotypy in captive giraffe and okapi  Appl  Anim  Behav  Sci  73   235 247    Forthcoming Conferences    Are you organizing a conference  workshop or meeting that would be of interest to TREE readers  If so  please e mail the details to  us at TREE  elsevier com and we will feature it in our Forthcoming Conference filler     6 11 August 2006  91st ESA Annual Meeting  Memphis  USA  http   www esa org 
17. icipated threats to fitness  where the animal behaves     as if    trying to avoid some future danger  6 7   For example  a perfectly  healthy animal with no predator in sight might be highly motivated to  seek cover because natural selection has favoured those of its  ancestors that had an emotional state of fear in open spaces and  immediately sought cover     iii  Good welfare consists of good physical health  no direct threats  to fitness  and the animal having what it wants  no anticipated threats  to fitness that are stimulating the animal to escape or seek something  it needs      iv  The ethical issues in animal welfare arise from the fact that the  direct and indirect threats to animal fitness are caused by human beings     v  Paradoxically  it is the evolved mechanisms for coping with  anticipated threats to fitness  such as the fear that leads an animal to  avoid a dangerous place  that causes more concern about welfare than  the direct threats themselves  Humane death  for example  is regarded  as less of a welfare problem than causing pain  fear or frustration to a  healthy animal   6      vi  Behavioural ecologists have a major role to play in under   standing the mechanisms by which different species respond to  threats to their fitness and thus in defining what constitutes    welfare      health and what the animal wants  for different species  Fruitful lines    Some human patients with certain sorts of brain damage  can successfully reach out and touch object
18. killed  most  commonly with anticoagulant poisons  because they  are pests  5     Here I provide a guide to the burgeoning science  of animal welfare  now one of the most comprehensive of  all the biological sciences  a discipline that embraces  behavioural ecology  evolution  neuroscience  animal  behaviour  genetics  cognitive science and even conscious   ness studies  The guide is intended for    users    of animal  welfare science  not so much those already working in the  field  but the many other people for whom the results of  welfare research increasingly matter  such as philoso   phers interested in animal emotions  politicians and  legislators confronted with public demands for improve   ments in the treatment of animals and behavioural  ecologists interested in the ways animals make decisions  and respond to real and perceived threats to their fitness   6   Indeed  I hope to encourage more readers of TREE to  take up some of the challenges raised by studying the  welfare of animals  Box 1   The views expressed are  my own     Corresponding author  Dawkins  M S   marian dawkins zoo ox ac uk    Available online 10 November 2005    What is animal welfare science    Animal welfare is no more     and no less     difficult to define  than is human welfare  6 7   When we talk about good  welfare for humans  we mean that someone is in good  health and that their emotions are generally positive  that  they are    fit and feeling good     8   Poor human welfare  comes not on
19. ly from ill health  injury and disease but also  from states such as stress  frustration  boredom  lone   liness or grief  Many of these mental symptoms also  appear as physical symptoms  but not always  so that  for  humans  we make a distinction between physical well   being and mental well being  So it is with animals  6 8    Good animal welfare also starts with physical health   which is why animal welfare sciences has its roots in  veterinary medicine  although it does not stop there  Good  welfare implies that animals also have positive emotions   such as pleasure and contentment  rather than negative  ones  such as fear or frustration  which we humans label     suffering     7 8   Animal welfare science has met this  challenge by asking three big questions   i  Are animals  conscious   ii  How can we assess good and bad welfare in  animals   iii  How can science be used to improve animal  welfare in practice     Are animals conscious    Although many people believe  without the need for  science  that other animals consciously experience pain  and suffering in ways that are similar to the ways in which  humans do  an exciting development in animal welfare  science is that it is now beginning to embrace advances in  other disciplines that tackle the issues of consciousness  itself  9 11  and so can potentially appeal to those more  sceptical people for whom the issue of animal conscious   ness is not obvious    From a scientific point of view  human consciousness is  th
20. memphis     13 19 August 2006   24th International Ornithological Congress  Hamburg   Germany   http   www i o c org    22 26 August 2006   1st European Congress of Conservation Biology  Eger   Hungary   http   www eccb2006 org     4 6 September 2006   ECBB  3rd Joint European Conference on Behavioural  Biology  Belfast  Northern Ireland  http   asab nottingham ac uk meetings asab php    5 7 September 2006   British Ecological Society Annual Meeting  University of  Oxford  UK   http   www britishecologicalsociety org     10 14 September 2006  American Society for Fish Biology  Lake Placid  USA  http   www fisheries org html index shtml     3 7 October 2006  American Ornithologist Union  Veracruz  Mexico  http   www naoc2006 org     18 21 October 2006  Society of Vertebrate Palaeontology  http   www vertpaleo org    23 24 March 2006   Trade offs in conservation  deciding what to save   Zoological Society of London  UK  http   www zoo cam ac uk ioz meetings htm    4 9 June 2006  ASLO Summer meeting  Victoria  Canada  http   aslo org meetings victoria2006     23 27June 2006   Annual meeting of the Society for the Study of Evolution   State University of New York at Stony Brook  USA  http   Isvl la asu edu evolution symp06 html    24 29 June 2006  Society for Conservation Biology  San Jose  USA  http   conbio org 2006     11 12 July 2006   BES IEEM Ecological Impact Assessments  Science and  Best Practice  Bath Spa University College  Bath  UK  http   www britishecologicalsociety
21. o  single measure of welfare     iii  Answers should  as far as possible  be derived in situ     in the  place where there is concern for animals such as on commercial  farms and in zoos  rather than from small scale studies which might  not reflect the real world  This means using large scale experiments   an epidemiological approach or systematic meta analysis of  published data     iv  Giving animals    the benefit of the doubt    and enacting  legislation without such evidence might give short term satisfaction  to campaigners but could make genuine improvements more difficult  to achieve by distracting resources and attention     v  Improvements in animal welfare should be studied in the  context of human health  food safety and environmental protection     www sciencedirect com    improvements in the treatment of real animals     those  on commercial farms and in zoos  for example  Animal  welfare science has already resulted in changes to the laws  governing the treatment of animals  62 63  but there  remains much research that does not get taken up in  practice  64   Although there are many possible reasons for  this  including commercial pressures against change  and  a lack of funding to develop a research idea into a practical  solution  the way in which research is conceived and  conducted might have also been partly responsible  Two  recent developments might help to change that  Box 2     First  it is increasingly recognized that carrying out  research on commer
22. pain for consciousness researchers  Conscious  Cogn   8  391 422   Rainville  P   2002  Brain mechanisms of pain affect and pain  modulation  Curr  Opin  Neurobiol  12  195   200   Baars  B J   2001  There are no known differences in brain  mechanisms of consciousness between humans and other mammals   Anim  Welf  10  S31   S40   Appleby  M C  and Hughes  B O  eds   1997  Animal Welfare  CAB  International   Irwin  M   1999  The immune correlates of depression  Adv  Exp  Med   Biol  461  1   24   Dallman  M F   2001  Stress and sickness decrease food intake and  body weight  How does this happen  When does this adaptive response  progress to pain and suffering   In Coping with Challenge  Welfare in  Animals including Humans  Broom  D M   ed    pp  301 316  Dahlem  University Press   Korte  S M   2001  Corticosteroids in relation to fear  anxiety and  psychopathology  Neurosci  Biobehav  Rev  25  117   142   Rushen  J   1991  Problems associated with the interpretation of  physiological data in the assessment of animal welfare  Appl  Anim   Behav  Sci  28  381 386   Toates  F   1995  Stress  Conceptual and Biological Aspects  John  Wiley  amp  Sons   Oatley  K  and Jenkins  J M   1996  Understanding Emotions   Blackwells   Weary  D M  and Fraser  D   1995  Calling by domestic piglets  reliable  signals of need  Anim  Behav  50  1047   1055    54    55    56    57    58    59    60    61    62    63    TRENDS in Ecology and Evolution Vol 21 No 2 February 2006    Dawkins  M S   
23. rease the value of research findings  Although it might  seem obvious to say that animal welfare decisions should  be evidence based  this is by no means always the case in  practice  For example  a recent European Union report on  the welfare of broiler chickens  4  based its conclusions for  legislation on  gt  500 publications  but made no attempt to  rank the publications on their research quality  magnitude  of effect  sample size or whether the studies were  conducted on commercial farms or in small groups  By  contrast  controversial issues such as whether    enrich   ments    really improve welfare  2  or what can be done to  reduce stereotypies in zoo animals  59 72  are now being  tackled more usefully by objective evaluation of large  numbers of already published research papers     Conclusions   Animal welfare science thinks big  It asks big questions  about animal consciousness  animal health and animal  emotions and it tackles big issues that affect millions of  people and billions of animals  To do so  it draws on a  variety of new techniques and approaches so that it is now  one of the most comprehensive of all the biological    TRENDS in Ecology and Evolution Vol 21 No 2 February 2006 81    sciences  Animal consciousness is central to the study  of animal welfare but is still  tantalizingly  the     hard problem    and needs to be respected as such   There is no single measure of animal welfare  no  convenient equivalent of a litmus test  but focusing on  two
24. s in front of  them but then say that they are not conscious of seeing  them at all  16   They are thus simultaneously blind   consciously  and sighted  unconsciously guided reaching    For much of what we do  there appear to be multiple  routes to the same behaviour  only some of which involve  consciousness  15   But if the same action  e g  breathing  or stretching a limb  can occur in humans through either a  conscious or an unconscious pathway  the argument that if  the behaviour of another animal is similar to that of a  human  that animal too must be conscious  17  is  weakened  An animal could be doing the same behaviour  but using its evolutionarily older unconscious circuits  18    Even human emotions can be unconscious  15 19   For  example  people can be subliminally influenced in how  they interpret a stimulus by a brief  50 ms  flash of a  happy or sad face immediately beforehand  even when  they have no conscious awareness of having seen any  faces  20     Greater understanding of the human brain has thus  made deciding which non human animals are conscious  more difficult than ever  However  new research findings  have opened up some intriguing ways forward     Animal cognition   Griffin  21  argued that consciousness in humans evolved  to enable adaptively flexible behaviour and that  therefore   the best way to find evidence for consciousness in animals  would be to look for adaptively flexible behaviour in other  species  The problem  as we have seen  is to
25. scaping a predator  50 51   Even in humans  it is difficult  to judge whether someone is feeling angry  fearful or just  plain excited just from knowing what their autonomic  responses are  52   We will never understand the  physiology of animal emotions just by looking at  autonomic responses  We need to understand what is  going on in their brains  23     Behaviour has the advantage that it can be studied  non invasively and can give a direct insight into the view  of the situation from the perspective of the animal   For example  the squeals of piglets give an    honest     indication of how hungry they are  53   The apparently  simple question     Does the animal have what it wants     is  the key both to whether it is being treated in ways it  dislikes  e g  pain it wants to avoid  and to whether it is  deprived  it wants something it does not have   54     We now have several different ways of    asking    animals  what they want and whether they find the situations they  are in pleasurable or distressing   For example  rats have  been shown to press a bar more for the reward of being  allowed access to a cage containing other rats than they  will for an empty cage  indicating that being with social  companions is something they want  55   Caged mink will  push doors to gain access to a water bath and will continue  to do so even when the doors are loaded with weights  but  they will not make this effort for an empty cage or a cage  full of novel objects  56   suggestin
26. t  control  Anim  Welf  12  1 37  6 Dawkins  M S   1998  Evolution and animal welfare  Q  Rev  Biol  73   305 328  7 Broom  D M  ed   2001  Coping with Challenge  Welfare in Animals  including Humans  Dahlem University Press  8 Webster  A J F  et al   2004  Welfare assessment  indices from clinical  observation  Anim  Welf  13  S93 S98  9 Mendl  M  and Paul  E S   2004  Consciousness  emotion and animal  welfare  insights from cognitive science  Anim  Welf  13 Suppl     S17 S25  10 Bateson  P   2004  Do animals suffer like us  Vet  J  168  110   111  11 Paul  E S  et al   2005  Measuring emotional responses in animals  the  utility of a cognitive approach  Neuro  Biobehav  Rev  29  469 491  12 Blackmore  S   2003  Consciousness  An Introduction  Hodder  amp   Stoughton  13 Koch  C   2004  The Quest for Consciousness  A Neuroscientific  Approach  Roberts  amp  Co  14 Burish  M J  et al   2004  Brain architecture and social complexity in  modern and ancient birds  Brain Behav  Evol  63  107   124  15 Rolls  E T   2005  Emotion Explained  Oxford University Press  16 Weiskrantz  L   2003  Roots of visual awareness  Prog  Brain Res  144   229 241  17 de Waal  F   2005  Animals and us  suspicious minds  New Scientist  2502  48  18 McPhail  E M   1998  The Evolution of Consciousness  Oxford  University Press  19 Berridge  K C  and Winkielman  P   2003  What is an unconscious  emotion   The case for unconscious    liking      Cogn  Emotion 17   181 211  20 Murphy  S  and Zajonc  R   
    
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