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      Low Birth Weight Impairs Acquisition of Spatial Memory Task in Pigs

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          Abstract

          In commercial pig farming, an increasing number of low birth weight (LBW) piglets are born, due to selection for large litter sizes. While LBW piglets have a higher risk of pre-weaning mortality, a considerable number of these piglets survive to slaughter age. In humans, LBW is a risk factor for long-term cognitive impairments. In pigs, studies examining the post-weaning effects of LBW on cognition have reported contradictory results. Therefore, the current study aimed to assess the effects of LBW on cognitive development in pigs using an improved study design, by (1) testing a larger sample size than previous studies, (2) assessing acute and chronic stress responses to account for a potential altered stress response in LBW pigs, and (3) testing both female and male pigs to account for potential confounding effects of sex. Learning and memory of 20 LBW pigs and 20 normal birth weight (NBW) pigs, both groups consisting of 10 females and 10 males, were compared using a spatial holeboard task. In this task, pigs had to learn and remember the locations of hidden food rewards. After a pig had successfully acquired the task, it was presented with two successive reversal phases during which it was presented with a new configuration of reward locations. The holeboard allows for simultaneous assessment of working and reference memory, as well as measures of motivation, exploration, and behavioral flexibility. Mixed model ANOVAs revealed a transiently impaired reference memory performance of LBW pigs, implying they had more difficulty learning their reward configuration in the holeboard. Also, LBW piglets showed increased pre-weaning hair cortisol concentrations compared to their NBW siblings. No other effects of LBW were found. Sex had no direct or interaction effects on any measures of holeboard performance or stress. It is possible that the enriched housing conditions applied during our study had an ameliorating effect on our pigs' cognitive development. Overall, our results suggest LBW has a negative effect on post-weaning cognitive performance in pigs. This could have welfare consequences as cognitive skills are required for pigs to learn how to correctly respond to their environment.

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          Most cited references58

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          Salivary cortisol as a biomarker in stress research.

          Salivary cortisol is frequently used as a biomarker of psychological stress. However, psychobiological mechanisms, which trigger the hypothalamus-pituitary-adrenal axis (HPAA) can only indirectly be assessed by salivary cortisol measures. The different instances that control HPAA reactivity (hippocampus, hypothalamus, pituitary, adrenals) and their respective modulators, receptors, or binding proteins, may all affect salivary cortisol measures. Thus, a linear relationship with measures of plasma ACTH and cortisol in blood or urine does not necessarily exist. This is particularly true under response conditions. The present paper addresses several psychological and biological variables, which may account for such dissociations, and aims to help researchers to rate the validity and psychobiological significance of salivary cortisol as an HPAA biomarker of stress in their experiments.
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            An overview of the tasks used to test working memory in rodents.

            In rodents, working memory is a representation of an object, stimulus, or spatial location that is typically used within a testing session, but not between sessions, to guide behaviour. In this review we consider a number of the tasks used to assess this type of memory in the rodent, and highlight some of their limitations. Although the concept of working memory as applied to rodents has its origin in the experiments of David Olton and Werner Honig in the 1970s, many earlier experiments assessed the same type of memory under the guise of delayed reaction or alternation paradigms. We revisit these early tasks, and also consider the nature of working memory used on maze tasks, operant box based tasks, and non-spatial delayed non-matching to sample paradigms.
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              A critical review of chronic stress effects on spatial learning and memory.

              C Conrad (2010)
              The purpose of this review is to evaluate the effects of chronic stress on hippocampal-dependent function, based primarily upon studies using young, adult male rodents and spatial navigation tasks. Despite this restriction, variability amongst the findings was evident and how or even whether chronic stress influenced spatial ability depended upon the type of task, the dependent variable measured and how the task was implemented, the type and duration of the stressors, housing conditions of the animals that include accessibility to food and cage mates, and duration from the end of the stress to the start of behavioral assessment. Nonetheless, patterns emerged as follows: For spatial memory, chronic stress impairs spatial reference memory and has transient effects on spatial working memory. For spatial learning, however, chronic stress effects appear to be task-specific: chronic stress impairs spatial learning on appetitively motivated tasks, such as the radial arm maze or holeboard, tasks that evoke relatively mild to low arousal components from fear. But under testing conditions that evoke moderate to strong arousal components from fear, such as during radial arm water maze testing, chronic stress appears to have minimal impairing effects or may even facilitate spatial learning. Chronic stress clearly impacts nearly every brain region and thus, how chronic stress alters hippocampal spatial ability likely depends upon the engagement of other brain structures during behavioral training and testing. Copyright 2009 Elsevier Inc. All rights reserved.
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                Author and article information

                Contributors
                Journal
                Front Vet Sci
                Front Vet Sci
                Front. Vet. Sci.
                Frontiers in Veterinary Science
                Frontiers Media S.A.
                2297-1769
                26 June 2018
                2018
                : 5
                : 142
                Affiliations
                [1] 1Behavior and Welfare Group, Department of Farm Animal Health, Faculty of Veterinary Medicine, Utrecht University , Utrecht, Netherlands
                [2] 2Brain Center Rudolf Magnus, Utrecht University , Utrecht, Netherlands
                [3] 3Graduate School of Life Sciences, Utrecht University , Utrecht, Netherlands
                [4] 4Study Programme Applied Biology, HAS University of Applied Sciences , Den Bosch, Netherlands
                Author notes

                Edited by: Christian Nawroth, Leibniz-Institut für Nutztierbiologie (FBN), Germany

                Reviewed by: Marie J. Haskell, Scotland's Rural College, United Kingdom; Nicolas Devillers, Agriculture and Agri-Food Canada (AAFC), Canada

                *Correspondence: Sanne Roelofs sanneroelofs@ 123456gmail.com

                This article was submitted to Animal Behavior and Welfare, a section of the journal Frontiers in Veterinary Science

                Article
                10.3389/fvets.2018.00142
                6028702
                6ea1f332-0b23-4d22-bacc-9d1f5d9a5371
                Copyright © 2018 Roelofs, van Bommel, Melis, van der Staay and Nordquist.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 30 March 2018
                : 08 June 2018
                Page count
                Figures: 4, Tables: 2, Equations: 0, References: 71, Pages: 14, Words: 11513
                Categories
                Veterinary Science
                Original Research

                pigs,cognition,birth weight,sex differences,spatial learning,spatial holeboard task,cortisol

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