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The infusion was started in the UA and continued until backflow from the RA occurred. Following this, the cannula in the RA was closed and leakage of the infusion fluid from other vessels was prevented by placing vessel clamps.
For gelatin infusion, the infusion system was slightly modified. All injection products were liquids and of low viscosity, however, no quantitative viscosity measurements were performed.
Preparation of the silicone, Araldite and Batson's No. All experimenters wore protective clothes and gloves. In cases of silicone, Araldite and Batson's No.
Dissection was performed after complete curing of the infusion fluid using a uniform protocol. All coloured vessels were counted and the diameters of the smallest vessels were measured with a digital calliper Mahr, 16ES, resolution 0.
Then, each compartment of the forearm and hand were dissected, with special attention paid to the degree of filling of the arterial tree, venous return, extravasations and the consistency of the injected vessels during dissection, that is colour intensity, flexibility, fragility, elasticity, strength.
Additional notes about the advantages and limitations of each technique were made. A Penetration of coloured injection fluid in the subcutaneous tissue.
B Extravasations of acrylic paint. C Latex infusion. E Araldite infusion. Acrylic paint does not cure at room temperature, which complicates dissection through leakage into the surrounding tissues when vessel walls are damaged Fig.
This dilution caused longer curing times at room temperature Fig. The silicone used in this study is a very expensive and toxic product, however, this does not apply to all silicone.
Silicone resulted in a minimal colour intensity, because the pigment did not completely dissolve.
Araldite is a strong, but fragile epoxy resin, because it lacks elasticity. Dilution using a dilutioner is an essential element in lowering the viscosity of the resin to facilitate infusion.
No macroscopic damage to the cadaver specimens and vessel walls was observed due to this thermal reaction Fig. Batson's No.
The lack of elasticity complicates dissection of smaller vessels, because it easily breaks. Its toxicity complicates infusion, as a large fumigation hood or an additional room with special ventilation and respiratory masks are required.
These problems were probably related to both the formalin itself and formalin fixation. The stiffness of soft tissue and the high vascular resistance due to formalin fixation influenced the distal filling of the arterial tree.
Furthermore, the formation of blood clots by formalin itself might have caused obstruction or distal emboli, which resulted in incomplete filling.
In the current study, acrylic paint was the only injection fluid that did not cure, so that the acrylic paint leaked into the surrounding tissues in the case of damaged vessel walls during dissection.
Latex was used by more than authors in various concentrations. This study is the first that reports the technical aspects of a specific latex, therefore it is recommended that researchers always test a specific latex in a pilot experiment, as there are so many types of latex with varying properties available.
However, the optimal mixture for vascular infusion has yet to be determined. Unfortunately, the RTV silicone could not be shipped to the Netherlands in small amounts, which rendered it too expensive for inclusion in the study.
Polymerisation is an exothermic reaction, which could theoretically cause damage to vessel walls and surrounding tissues.
This study shows that all six vascular injection products are suitable to facilitate dissection. However, the study is limited by the small number of cadaver specimens in each group, especially in the description of properties, limitations and advantages of the different injection fluids.
Further sample selection bias was found in the individual variability of the cadaver specimens, who differed in age, duration of formalin fixation or freezing, diameter of blood vessels and vascular resistance.
In addition, some equivalent substitutes of injection fluids not previously described in literature were used, which makes it difficult to compare with other studies.
This study only focused on vascular injection of the arterial system of the forearm. Therefore, special considerations should be made before using these materials in other research projects.
The selection of a suitable vascular infusion fluid should depend on the specific study purpose. Latex is preferable when flexibility and elasticity are required for dissection.
Silicone offers elasticity and strength, however, less expensive alternatives are desirable and available. When a strong polymer is required, Araldite or Batson's No.
Study conception and design: Doomernik, Kruse, Reijnen, Kooloos. Acquisition of data: Doomernik and Kruse. Analysis and interpretation of data: Doomernik, Kruse, Kooloos.
Drafting of the manuscript: Doomernik. The authors would like to thank Marjan Doom and colleagues from the Department of Morphology, Faculty of Veterinary Medicine, Ghent University, Belgium for their helpful advice regarding the study protocol.
Furthermore, the authors would like to thank B. National Center for Biotechnology Information , U. Journal List J Anat v.
J Anat. In our study, the enriched-housed pigs grew faster than the barren-housed pigs, although the effects on growth were small. All pigs of the present study were housed in a naturally ventilated stable.
Both groups had covered nests with heat lamps. The temperature in the nests did not differ between the two environments. However, the nest and the entire floor in the enriched environment were covered with straw, whereas the barren-housed pigs had only a rubber mattress in the nest and no covering on the concrete floor.
Provision of straw bedding may constitute one of the most effective measures for improving pig welfare van de Weerd et al. Hayne et al.
Straw protects pigs against heat loss. Vanheukelom et al. Both straw and peat stimulate exploratory behaviour and may reduce aggression towards the penmates Vanheukelom et al.
Pigs from barren and enriched housing did not differ in basal salivary cortisol levels, measured at the end of the study.
Belz et al. A study conducted by van der Staay et al. Higher cortisosterone levels are associated in rats with impaired cognitive performance Bodnoff et al.
However, these findings are in contrast with more recent studies that found that barren-housed pigs showed lower baseline cortisol level compared to pigs from enriched environments de Jong et al.
Barren housing conditions can lead to chronic stress which can result in hypocortisolism in animals Natelson et al. In rodent studies, clear differences in cognitive performances and neuronal structures between barren- and enriched-housed animals were found.
Enriched-housed rats showed an increased hippocampal neurogenesis Nilsson et al. Enrichment, i.
Enrichment can increase the diversity of behaviours, especially natural behaviour, and decrease abnormal behaviour, as well as improve how the environment is used, i.
The hidden bait as appetitive stimulus encourages natural foraging behaviour in the pigs Westlund , and stimulates some natural behaviours such as inspecting the holes by lifting the balls and covering the holes with the snout rooting.
Complex learning and memory tasks may be less suited to assess the effects of environmental enrichment in animals. The pigs in the current study had access to physical activity and enrichment due to the testing in the holeboard.
The long-lasting, extended handling and habituation and the subsequent training in the holeboard apparatus may reduce some of the negative effects associated with living in a barren environment.
Testing interrupted the daily routine, especially for the barren-housed pigs. The learning enhancement persisted during ageing.
Furthermore, it was found in rats that physical activity results in better spatial learning performance Fordyce and Farrar ; Anderson et al.
Several studies have shown that pigs are motivated to work for food Puppe et al. Our holeboard apparatus used the natural rooting movements of pigs, which is in itself rewarding for pigs Studnitz and Jensen ; Studnitz et al.
A study by Puppe et al. We used positively reinforced behaviour which offers the pig control over its environment, makes the environment predictable, teaches the pig how to use its environment optimally and may increase coping abilities Young Moreover, the test procedure may be even more rewarding for the barren-housed pigs, as barren environments have been shown to result in rebound activities in test arenas as a result of thwarted motivation for exploration in the home environment e.
Wemelsfelder et al. Recent studies, moreover, suggest a general higher reward sensitivity in barren-housed animals e. Beckmann and Bardo ; Mitchell et al.
It is possible that the effect size of barren housing versus enriched housing is reduced in our test due to an unequal effect of positive reinforcement on the two groups.
The present study provides some evidence that pigs reared in an enriched environment after weaning show a better cognitive performance in a spatial holeboard task compared to pigs reared in a barren environment.
Enriched-housed pigs showed a better reference memory performance during the acquisition phase, and a marginally better general working memory performance during the reversal phase.
In addition, enriched-housed pigs were faster in the time needed per hole visit IVI during acquisition and reversal and needed less time to complete a trial than their barren-housed littermates.
The latter finding may be due to a combination of a better spatial memory performance i. The shorter IVI suggests higher motivation of enriched-housed pigs compared to barren-housed pigs.
The holeboard task is a valid measurement instrument for spatial discrimination learning in pigs.
However, it is unclear to what extent the holeboard testing procedure itself could have provided enrichment that could partially have counteracted the effects of living in a barren environment and may lead to underestimation of the effects of a barren environment.
Therefore, it may be difficult to test effects of different environments on cognitive performance in pigs using longer-lasting, appetitively motivated complex testing procedures.
We thank Elly. Zeinstra for her support in preparing the salvia cortisol samples and Christine H. Oei for analysing the samples.
National Center for Biotechnology Information , U. Animal Cognition. Anim Cogn. Published online Oct Charlotte G. Grimberg-Henrici , Paul Vermaak , J.
Elizabeth Bolhuis , Rebecca E. Nordquist , and F. Josef van der Staay. Elizabeth Bolhuis. Rebecca E. Author information Article notes Copyright and License information Disclaimer.
Corresponding author. This article has been cited by other articles in PMC. Abstract This study investigated the effects of environmental enrichment on the cognitive performance of female conventional farm growing pigs in a spatial holeboard task.
Keywords: Reference memory, General working memory, Spatial holeboard task, Salivary cortisol, Environmental enrichment, Pig Sus scrofa.
Introduction Environmental enrichment is believed to satisfy the behavioural needs of pigs to explore and forage and to help the animals to adapt to their environment Ferguson Testing area and holeboard apparatus The barren pen, the enriched pen, a small pen which was used as waiting area before testing and the testing apparatus all were located in the research stable, connected by a corridor see Fig.
Open in a separate window. Results Spatial memory Before the formal training started, the number of different holes visited was scored during six pre-training days, with all holes baited.
Reference memory RM see Fig. Working memory WM see Fig. General working memory GWM see Fig. Inter-visit interval IVI see Fig.
Trial duration TD see Fig. Physical measurements Growth curves see Fig. Saliva basal cortisol see Fig. Discussion Our study provides some support for the hypothesis that enrichment improves cognitive performance in pigs, whereas no evidence was found for an effect of the different environments on basal salivary cortisol as index of stress.
Spatial memory During the acquisition and reversal of a spatial holeboard task, enriched- and barren-housed pigs improved their cognitive performance over trial blocks, corroborating earlier studies Arts et al.
Latencies and durations Neither group reduced their LFV during acquisition nor during the reversal. Growth curves Beattie et al.
Stress and basal saliva cortisol Pigs from barren and enriched housing did not differ in basal salivary cortisol levels, measured at the end of the study.
Testing as enrichment Enrichment, i. Conclusion The present study provides some evidence that pigs reared in an enriched environment after weaning show a better cognitive performance in a spatial holeboard task compared to pigs reared in a barren environment.
Acknowledgments We thank Elly. Compliance with ethical standards Conflict of interest The authors declare that they have no conflict of interest.
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