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Linda Keeling

Linda Keeling
Linda Keeling is Professor of Animal Welfare and her research is related to the welfare of farm, companion and sport animals. She is an ethologist, interested in asking fundamental questions related to social behaviour, motivation and cognition in animals, but also applied questions related to the effects of housing and management on animals, and the development of animal welfare assessment methodologies.


Linda Keeling's research is grouped into three main areas. These are investigating emotional states in animals (especially postive states), the effects of early experience on later behaviour, and the role of animal welfare in sustainable development. Her broad experience within animal welfare science means that she is active in multidisciplinary projects and also in science-based animal welfare policy development. 

She is a member of the EU Platform on Animal Welfare, member of the Royal Swedish Academy of Forestry and Agriculture and former chairperson of the Biology Committee at the Swedish Academy of Sciences. She has been a member of the Animal Health and Animal Welfare panel at the European Food Safety Authority and is currently in their working group on broilers. She is Honorary Doctor at the University of Linköping in Sweden.


Linda Keeling is involved in many studies, but the large research projects that she is currently responsible for are;

The use of positive animal welfare indicators in the dairy industry: Science and policy 

Collaborators: Björn Forkman, Jacob Bull, Ann Nyman, Diana de Oliveira

The quality of life of an individual is a combination of positive and negative experiences. While most of the research on animal welfare has focused on the negative aspects of animal welfare, there is a move towards acknowledging the importance of the positive aspects and hence a growing interest in assessing positive emotions in animals. The aim of this project is to find practical and validated indicators for positive animal welfare in dairy cattle to give a more complete picture of the their welfare as well highlight the benefits of going beyond the ‘bottom line’ of complying with legislation and towards a more sustainable and competitive dairy industry.

Identifying areas of synergy and conflict between different sustainability goals in animal production: A quantitative approach using existing databases

Collaborators: Anders Glimsär, Eddie Bokkers, Jan Hultgren

Despite the considerable effort put into defining goals for sustainable animal production and, more recently, tools to assess them, there is little work on how the different goals interact. The Swedish Board of Agriculture, the Swedish Environmental Protection Agency and the National Food Agency collect data to check compliance with animal welfare, biodiversity, on-farm emissions and food safety. We will use their unique databases to investigate the complex network of interactions between these different areas of sustainability. The overall aim is to identify areas of potential synergy and conflict in sustainable animal production, and to initiate constructive discussion on these areas with the authorities. Representatives of the three authorities form the reference group for this project.

Preparing for the challenges and opportunities of life: effects of early experience on adaptive plasticity in laying hens

Collaborators: Ruth Newberry and Inma Estevez

Farm animals are often reared in one type of environment but housed as adults in another. When moved, a lack of skills needed to adapt rapidly to the new environment can compromise welfare. We hypothesize that exposure to a diverse and stimulating early environment will prepare chicks to better cope with challenges and exploit opportunities later in life as adult laying hens. Besides identifying how best to prepare chicks to cope with future stressors (so minimizing reductions in welfare), this study investigates how developmental plasticity can enhance positive aspects of welfare, such as exploration skills that help hens quickly find food and use nest boxes and other new resources when moved to the laying house.


Selected publications

Linda Keeling has published 146 original peer reviewed papers. H-index 44 (according to Web of Science).

Publications over the past 7 years.

Keeling L.J., Marier E.A., Olmos Antillón G., Blokhuis H.J., Staaf Larsson B. and Stuardo L., (2022) A global study to identify a potential basis for policy options when integrating animal welfare into the UN Sustainable Development Goals. Front. Anim. Sci. 3:974687. doi: 10.3389/fanim.2022.974687

Nazar, F.N., Skånberg, L., McCrea, K., Keeling, L.J., (2022) Increasing Environmental Complexity by Providing Different Types of Litter and Perches during Early Rearing Boosts Coping Abilities in Domestic Fowl Chicks. Animals 12, 1969.

Hultgren, J., Hiron, M., Glimskär, A., Bokkers, E.A.M., Keeling, L.J., (2022). Environmental Quality and Compliance with Animal Welfare Legislation at Swedish Cattle and Sheep Farms. Sustainability 14, 1095. doi:10.3390/su14031095

Tamminen, L-M., Keeling L.J., Svensson A., Briot L. and Emanuelson U., (2021) Unraveling the Complexity to Observe Associations Between Welfare Indicators and Hair Cortisol Concentration in Dairy Calves. Front. Anim. Sci. 2:793558. doi: 10.3389/fanim.2021.793558

Verbeek, E., Keeling, L., Landberg, R., Lindberg, J.E., Dicksved, J., (2021). The gut microbiota and microbial metabolites are associated with tail biting in pigs. Scientific Reports 11. doi:10.1038/s41598-021-99741-8

Keeling, L.J., Winckler, C., Hintze, S. and Forkman, B. (2021) Towards a Positive Welfare Protocol for Cattle: A Critical Review of Indicators and Suggestion of How We Might Proceed. Front. Anim. Sci. 2:753080. doi: 10.3389/fanim.2021.753080

Verbeek, E., Dicksved, J., and Keeling, L., (2021) Supplementation of Lactobacillus early in life alters attention bias to threat in piglets. Scientific Reports, 11:1: Article Number: 10130. DOI: 10.1038/s41598-021-89560-2

Olmos Antillón, G., Tunón, H., de Oliveira, D., Jones, M., Wallenbeck, A., Swanson, J., Blokhuis, H. and Keeling, L., (2021) Animal Welfare and the United Nations’ Sustainable Development Goals - Broadening Students’ Perspectives. Sustainability 2021, 13, 3328.

Skånberg, L., Bramgaard Kjærsgaard Nielsen, C.  and Keeling, L. ,(2021). Litter and perch type matter already from the start: exploring preferences and perch balance in laying hen chicks. Poultry Science. 100 :2, 431-440.

Sirovnik, J., Voelkl, B., Keeling, L.J., Wurbel, H. and Toscano, M.J. (2021). Breakdown of the ideal free distribution under conditions of severe and low competition. Behav Ecol Sociobiol 75, 31.

Nicol, C., Bejder, L., Green, L., Johnson, C., Keeling, L., Noren, D., Vander Hoop, J. and Simmonds, M. (2020). Anthropogenic Threats to Wild Cetacean Welfare and a Tool to Inform Policy in This Area. Frontiers in Veterinary Science 7:57. doi: 10.3389/fvets.2020.00057

Tamminen, L-M, Hranac, C.R., Dicksved, J., Eriksson, E.,  Emanuelson, U. and Keeling L.J. (2020) Socially engaged calves are more likely to be colonised by VTEC O157:H7 than individuals showing signs of poor welfare . Scientific Reports 10:6320

Trindade P.H.E., Hartmann, E., Keeling, L.J., Andersen, P.H., Ferraz, G.C., Paranhos da Costa, M.J.R. (2020) Effect of work on body language of ranch horses in Brazil. PLoS ONE 15(1): e0228130.

Tamminen, L-M., Söderlund, R., Wilkinson, D. A., Torsein, M., Eriksson, E.  Churakov, M., Dicksved, J., Keeling, L.J., and Emanuelson, U. (2020). Risk factors and dynamics of verotoxigenic Escherichia coli O157:H7 on cattle farms: An observational study combining information from questionnaires, spatial data and molecular analyses. Preventive Veterinary Medicine 170 (2019) 104726

Keeling, L.J., Tunon, H., Olmos Antillón, G., Berg C.,  Jones, M., Stuardo, L., Swanson, J.,  Wallenbeck, A., Winckler, C., Blokhuis, H.(2019). Animal Welfare and the United Nations Sustainable Development Goals. Frontiers in Veterinary Science, section Animal Behavior and Welfare, 10 October 2019.

Zupan, M., Rehn, T. de Oliveira, D., Malovrh, S., Keeling, L.J. (2019). Individual play patterns stimulated by a familiar object are groupdriven. Scientific Reports. 9:6092.

de Oliveira, D., Keeling, L.J., Paranhos da Costa., M.J. R. (2019). Individual variation over time in piglet´s reactions to early handling and its association to weight gain. Applied Animal Behaviour Science 215 (2019) 7–12.

Campderrich, I., Nazar, FN., Wichman, A., Marin, RH., Estevez, I., Keeling, LJ. (2019) Environmental complexity: A buffer against stress in the domestic chick. PLoS ONE, 4(1) Article Number: e0210270, DOI:10.1371/journal.pone.0210270

Comin, A., Jeremiasson, A., Kratzer, G. and Keeling, L. (2019) Revealing the structure of the associations between housing system, facilities, management and welfare of commercial laying hens using Additive Bayesian Networks. Preventive Veterinary Medicine, 164: 23-32. DOI:10.1016/j.prevetmed.2019.01.004

Tamminen, L-M, Söderlund, R.,  Wilkinson D.A., Torsein, M., Eriksson, E., Churakov, M., Dicksved, J., Keeling, L.J. Emanuelson, U. (2019) Risk factors and dynamics of verotoxigenic Escherichia coli O157:H7 on cattle farms: An observational study combining information from questionnaires, spatial data and molecular analyses. Preventive Veterinary Medicine 170 , 104726,  DOI:10.1016/j.prevetmed.2019.104726

Buller, H., Blokhuis, H., Jensen, P. and Keeling, L. (2018) Towards farm animal welfare and sustainability. Animals 8 (81), doi:10.3390/ ani8060081 

de Oliveira, D and Keeling, L.J. (2018) Routine activities and emotion in the life of dairy cows: Integrating body language into an affective state framework. PLoS ONE 13 (5), 10.1371/journal.pone.0195674 

Skånberg, L., Gauffin, O., Norling, Y., Lindsjö, J. and Keeling, L.J. (2018) Cage size affects comfort, safety and the experienced security of working dogs in cars. Applied Animal Behaviour Science 205, 132-40. DOI: 

Zidar, J, Campderrich, I., Jansson, E., Wichman, E., Winberg, S., Keeling, L. and Løvlie. H. (2018) Environmental complexity buffers against stress-induced negative judgement bias in female chickens. Scientific Reports 8:5404 | DOI:10.1038/s41598-018-23545-6

Alpigiani, I., Cortinas Abrahantes, J.C., Michel V, Huneau-Salaün A., Chemaly, M., Keeling, L.J.  Gervelmeyer A., Baccia, C., Brindania, F., Bonardia, S. and Berthe, F. (2017). Associations between animal welfare indicators and Campylobacter spp. in broiler chickens under commercial settings: A case study. Prev. Vet. Med. 147:186-193.

Eriksson, M., Keeling, L.J., Rehn, T. (2017) Cats and owners interact more with each other after a longer duration of separation. PLoS ONE 12 (10):e0185599

Hitchens, P., Hultgren, J., Frössling, J., Emanuelson, U., & Keeling, L.J.  (2017) Circus and zoo animal welfare in Sweden: An epidemiological analysis of data from regulatory inspections by the official competent authorities. Animal Welfare 26;373.382.

Hitchens, P., Hultgren, J., Frössling, J., Emanuelson, U., & Keeling, L. (2017). An epidemiological analysis of equine welfare data from regulatory inspections by the official competent authorities. Animal, 11(7), 1237-1248. doi:10.1017/S1751731116002512

Lind, A-K., Hydbring-Sandberg, E., Forkman, B. and Keeling, L. (2017). Assessing stress in dogs during a visit to the veterinary clinic: Correlations between dog behavior in standardized tests and assessments by veterinary staff and owners. Journal of Vet. Behav - Clinical applications and research, 17: 24-31

Keeling, L., Newberry, R.C. and Estevez.I (2017) Flock size during rearing affects pullet behavioural synchrony and spatial clustering. Applied Animal Behaviour Science 194; 36-41.

Rehn, T. and Keeling, L.J. (2017) Links between the owner’s adult attachment style and support-seeking behaviour of their dog. Frontiers in Psychology. 8:2059 doi 10.3389/psyg.2017.02059 

Brunberg, E. I., Rodenburg. B. T., Rydhmer, L., Kjaer, J.B., Jensen, P and Keeling, L.J. (2016) Omnivores Going Astray: A Review  and New Synthesis of Abnormal Behavior in Pigs and Laying Hens. Front. Vet. Sci. 3:57.  doi: 10.3389/fvets.2016.00057

I Alpigiani, C Bacci, L J Keeling, M D Salman, F Brindani, S Pongolini,  PL Hitchens  and S Bonardi (2016) The associations between animal-based welfare measures and the presence of indicators of food safety in finishing pigs. Animal Welfare 25:  355-363.

Keeling, L. J. Boe, K. E. Christensen, J. W. Hyyppa, S. Jansson, H. Jorgensen, G. H. M. Ladewig, J. Mejdell, C. M., Rundgren, M. Sarkijarvi, S. Sondergaard, E. and Hartmann, E. (2016) Injury incidence, reactivity and ease of handling of horses kept in groups: a matched case control study in four Nordic countries. Applied Animal Behaviour Science 185; 59-65.

Lindahl, C.; Pinzke, S.; Herlin, A.; Keeling, L. J. (2016) Human-animal interactions and safety during dairy cattle handling-Comparing moving cows to milking and hoof trimming. Journal of Dairy Science, 99 (3): 2131-2141

Hitchens, P.L., Hultgren, J., Frössling, J., Emanuelson U., and Keeling, L.J., (2016) Prevalence and risk factors for overweight horses at premises in Sweden assessed using official animal welfare control data. Acta Veterinaria Scandinavica 58:31-35.

Rehn, T., Keeling, L.J. (2016). Measuring dog-owner relationships: Crossing boundaries between animal behaviour and human psychology. Applied Animal Behaviour Science 183: 1-9.

Zupan, M., Buskas, J., Altimiras, J., Keeling, L.J. (2016) Assessing positive emotional states in dogs using heart rate and heart rate variability. Physiology & Behavior 155: 102-111

Zupan, M., Rehn, T., De Olivera, D. and Keeling, L.J. (2016) Promoting positive states: the effects of early human handling on play and exploratory behaviour in pigs. Animal 10:135-141