Aposematic species have evolved conspicuous warning signals, such as bright colors and striking patterns, to deter predators. Some edible and harmless species take advantage of this deterrent effect by mimicking their appearance. Mimicry is a great example of how natural selection produces remarkable adaptations. However, while some species evolve a very close similarity to their models to effectively avoid attacks, others are successful in doing so despite an incomplete similarity, i.e. imperfect mimicry. In some cases, it is surprising how such a crude disguise can fool predators. Why and how imperfect mimicry can persist has been much discussed and considered as a problem for the theory of natural selection. It is therefore of great interest to understand what makes it possible. Predator psychology is an important factor in the evolution of aposematism and mimicry. In the past decades it has been suggested that certain components of prey appearance are more important to predators than others during prey assessment. We developed this idea by incorporating concepts from associative learning, and presented a new approach to explain imperfect mimicry. Our general hypothesis is that prey traits have different salience to predators. Certain traits are perceived as highly salient and are thus used primarily in the discrimination and generalization of prey, while traits with low salience are overshadowed and not used in the assessment. The salience of a trait can depend on how conspicuous or discriminable it is in the particular context, and can vary due to for example previous predator experience. We tested our ideas with wild blue tits and domestic chickens as predators, and artificial and semi-natural prey stimuli. In paper I we found that the trait that was perceived as most salient (color) was the one used to discriminate and generalize between prey. Mimics of that specific trait were highly avoided, despite differences in the other traits. We also found that salience is relative and context dependent (paper II). In a context where two traits were perceived as similarly salient, mimicry of a single trait offered intermediate protection, while mimicry of both offered high protection. In another context, the traits were perceived differently salient, and mimicry of one trait was enough for high protection. In paper III we tested a proposed scenario for the initiation of mimicry evolution in the edible butterfly mimic Papilio polyxenes asterius to its noxious model Battus philenor. The results showed that a partial similarity with the model in the salient black wing color offered intermediate protection from attacks, despite a general dissimilarity. This thesis investigates the major questions of imperfect mimicry: the initial step of mimicry evolution, the persistence of imperfect mimicry, and variation in mimic-model similarity. We conclude that mimicry evolution can begin in a non-mimetic species that acquires similarity to a model species in a high-salience trait. When multiple traits have similar salience, multi-trait mimicry is needed for higher protection. Mimicry can remain imperfect if the differences are in traits with low salience, and therefore under low or no selection pressure to change. To complete the picture, we showed that predators can have a biased generalization toward a more pronounced version of a salient trait (paper IV). The evolution of aposematism could therefore be explained by gradual enhancement of salient traits. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Accepted. Paper 4: Manuscript.</p>

One out of several distinguishing features of multiple sclerosis (MS) is the epidemiological variation of geographic distribution. Population-based studies on the prevalence and incidence of MS in Sweden have previously been performed only in Göteborg. Another feature of MS is the clinical variation between individuals. To a large extent data on the clinical characteristics of MS come from studies on cases frequenting MS clinics and therefore, may be biased. Also rare are population-based studies of the consequences of MS-related incapacity on socio­economic factors. As for MS aetiology, both environment and genes are involved. Human herpesviruses are often the main suspected environmental aetiological agents. Our aim was to estimate the prevalence of MS in Västerbotten County for 1 January 1990, the incidence during a 10-year period 1988-97, and the prevalence 31 December 1997; and also to present detailed clinical data including onset symptoms and the disability distribution for the latter two MS populations. Furthermore, we wanted to estimate the prevalence of sick leave, professional assistance, and housing; and also, to study the risk factors for sick leave. In order to investigate the association between MS and human herpesviruses, samples were identified in two regional population-based serumbank registers. This linkage identified samples collected from before MS-onset in 73 MS cases and after MS onset in 161 cases The prevalence and incidence populations were identified through multiple sources. Diagnostic ascertainment, the reliability of clinical data, and additional information were assured from a questionnaire with follow-up interview and neurological examination. The onset adjusted crude prevalence of MS was 125/100,000 (95% CI: 112-140) in January 1990, and 154/100,000 (95% Cl: 139-170) in December 1997. The increase was mainly attributable to a higher incidence than mortality. The crude incidence rate 1988-97 was 5.2/100,000 (95% CI: 4.4-6.2). The disability distribution in the 1997 prevalence population in Västerbotten was compared to the disability distribution in a Canadian MS population, which has been used for publications on the natural history of MS. One difference from the Canadian studies appears to be the better recognition of cases with more benign disease. Nevertheless almost half of prevalent MS cases aged 18-64 years were fully sick-listed, and one-fourth of all prevalent cases received professional assistance. High disability level was the strongest predictor for sick leave. All MS cases showed signs of past Epstein-Barr virus (EBV) infection. High activity to EBV (EBNA-1 but not VCA) and human herpesvirus 6 (HHV-6) significantly (borderline significance for HHV-6) increased the risk to develop MS. These estimates show that Västerbotten County is a high risk area for MS. Both incidence and prevalence were significantly higher when compared to estimates from Göteborg. The comparison with the Canadian MS population shows that MS might be a slightly more benign disease than previously recognized. Still, the consequences of MS regarding socio-economic aspects are considerable. We suggest that EBV is a prerequisite for the development of MS. Individuals that will develop MS exhibit an altered immune response against the EBV virus characterised by high activities to EBNA-1 in the absence of high VCA activities, this being most pronounced in the five-year period preceding MS onset. A pathogenetic role is suggested for EBV and remains possible also for HHV-6. / digitalisering@umu

An appreciation of the role of mutualism is essential when studying ecology and evolution in most ecosystems. Information covering aspects of mutualistic interactions can serve as a complement to the somewhat one-sided perspective from the 1950’s and 60’s that is used when teaching biology. In this thesis I applied an in-depth approach in which variation in the interspecific interaction between Aphis fabae aphids and Lasius niger ants was studied both in the field and in the laboratory. An emphasis was put on studies spanning several consecutive aphid generations. This approach revealed important differences between ant tended aphids and those without ants. In the lab, I found an initial decrease in aphid adult size and reproductive investment in the first generations after the start of ant tending, which was followed by a recovery to the pre-tending situation after about four generations. Another laboratory experiment showed an increase in alate (winged aphid) production from exposure to aphid alarm pheromones, and an even stronger decrease in alate production from ant attendance, suggesting that ants have gained the upper hand in an evolutionary conflict over aphid dispersal. Results from a field experiment further emphasized the possibility of negative effects of ants on aphids, showing that ant-tended aphid colonies experienced a higher rate of parasitoid attacks, produced fewer alates and embryos in adult aphids. The thesis highlights the scope for variation in the net effect of the interaction for aphids, and argues that, depending on the environmental circumstances, the interaction may sometimes and perhaps even often not really be a case of mutualism. / <p>At the time of the doctoral defense, the following paper was unpublished and had a status as follows: Paper 4: Manuscript.</p>

Egg-laying hens coming from commercial hatchery go through hatchery procedures considered as stressful and engaging prolonged stress response in adult chickens. The aim of our study was to evaluate the impact of commercial hatching procedure on the affective state of chicks, on their short- and long-term memory and on their need for social reinstatement. To assess the affective state of the chicks we used a cognitive bias protocol integrating the ecological response of a chick to the picture of another chick, to an owl and to an ambiguous cue mixing features of both the chickand the owl pictures. Short-term memory was evaluated by using a delayed matching-to-sample experiment (with 10, 30,60 and 120 s delays), with conspecifics as sample stimuli. We assessed long-term memory with an arena containing multiple doors leading to conspecifics, in which a chick had to remember which door was open after a delay of one hour or three hours. Finally, we observed the need for social reinstatement through a sociality test arena allowing a chick to be more or less close to conspecifics. We found that chicks coming from commercial hatchery were in a depressive affective state compare to control group. Those chicks also showed higher need for social reinstatement and loss weight. No differences were found regarding short- and long-time working memory between the two groups, but the methods used during these experiments will be discussed. Studying how commercial procedures impact the cognition and more specifically the emotions and state of mind of chickens, is a necessary step forward into the understanding of farm animals’ welfare.

Food waste is a major issue from various perspectives. During the process from harvest to the end consumer, almost one-third of food produced is wasted. It is not just the wasted food during the process that is concerning, there are issues in sustainability related to food waste that need to be considered. Moreover, there has been little attention to the issue of food waste in the downstream part of the supply chain and how specific behaviours affect the issue of wasting food.This thesis explores the food waste of fresh fruits and vegetables from a consumer´s perspective. To specify, the purpose of the study is to investigate the drivers of the disposal pattern of fresh fruit and vegetables, with an emphasis on the behaviour of "Generation Y" (born 1980 – 1995). Therefore, a revised model of the Theory of Planned Behaviour is applied. Also, to understand the behaviour of "Generation Y" regarding disposal, it is expected to identify impacts on the supply chain.A deductive approach is applied to this thesis. The qualitative study was conducted with open-ended survey questions to supplement the results with the answers of the respondents. The empirical data is collected from consumers within the “Generation Y” who usually purchase their fresh fruits and vegetables for their respective households. The data was analysed using the coding analysis which involves summarization and categorization of data.The results of the research reveal that external attitudinal factors such as price and marketing perception, storage habits, and quality consciousness and internal attitudinal factors such as sustainable environmental awareness, health consciousness, and subjective norms influence the respondents’ disposal behaviour. Moreover, the sustainable attitude of “Generation Y’’ leads to most of the consumers’ waste reduction, and highly influences the supply chain.

Horses are herbivores and can graze for up to 20 hours per day. It is well known that animals are motivated to perform natural behaviours and horses have, except from a physiological need to graze, a strong motivation to perform their natural feeding behaviour. This review aimed to give insight into how horses are kept, how the way they are kept affects their behaviour and welfare, if there are reasons why some horses cannot be kept on pasture, and if alternative feeding strategies can improve horse welfare by allowing horsesto perform their natural feeding behaviour. Most horses are kept in stables and are put on feeding regimes that do not resemble their natural feeding behaviour. This can lead to development of stereotypies, and when prevented from carrying out their feeding behaviour, oral stereotypies such as crib-biting can result. Horses kept free range display very few, if any, stereotypies and is therefore the best way to keep horses from a welfare approach. However, obesity and grass-related illnesses can restrict horses from being kept on pasture, thus, alternative feeding methods for stabled horses is imperative. Use of edible bedding, presenting several types of forage, providing ad libitumforage, or using slow feeders are alternative methods that could be implemented by horseowners and caretakers to improve horse welfare. Horses can however respond differently to these methods and a method that suits one horse might not be appropriate for another.

Dolphins use stereotyped, individually distinctive, frequency modulated whistles, referred to as signature whistles, in order to broadcast their identity. In this study, we trained six dolphins at Kolmården Zoo, Sweden, to be called over, either upon hearing their own signature whistle (SW) or upon hearing a biologically irrelevant ”trivial” sound (TS), with the aim to prove the Whistle caller concept. The Whistle caller concept is based on the fact that dolphins occasionally use other dolphins’ signature whistles in order to address specific group members and convene.  Our hypotheses were that (1) dolphins call-over trained using their SW would learn the behaviour faster than dolphins trained using TSs, and (2) dolphins trained with their SW would be able to discriminate between different SWs better than dolphins trained with a TS would be at discriminating between different TSs.  Three out of three dolphins were successfully call-over trained using their SW, and two out of three dolphins using their assigned TS. When discriminating between different sounds, two of the dolphins trained using their SW performed significantly better than one of the dolphins trained using a TS. However, there were large intra-group differences in the results, indicating that we cannot eliminate the possibility that these results stem from individual differences in these dolphins’ ability to learn new behaviours overall, rather than an understanding of the sounds they heard. We suggest that future studies focus on (1) male-female differences in discrimination success when applying the Whistle caller concept, (2) how the characteristics of the trivial sounds affect discrimination success, and (3) the option of calling more than one animal at a time by sending out several SWs in succession.

By studying how captive animals use outdoor space, we can determine which areas fulfill their biological and behavioral needs to improve enclosure design for animal welfare and visitor experience. The aim of the study was to determine the differential use of outdoor space in a mixed exhibit of captive ring-tailed lemurs (Lemur catta) and red ruffed lemurs (Varecia rubra), and to assess both the effect of adding environmental enrichment and possible visitor and meteorological effects on the lemurs’ use of space and behavior. For one month, five-minute interval scan sampling was used to record lemur location and behavior, as well as the meteorological conditions and presence of zoo visitors in the lemurs’ outdoor enclosure. That baseline was followed by four alternating two-week periods of food or structural enrichment, both with and without visitors. Enrichment increased the lemurs’ use of the enriched sectors, decreased resting, and increased locomotion. Structural enrichment increased their exploratory behaviors and stimulated arboreal locomotion. Both species preferred sunny and warm conditions for sunbathing and resting, while moving more around the enclosure in cooler and cloudier weather. The visitor effect on the lemurs’ use of space was weaker than the effects of enrichment and meteorological conditions. The availability of areas to hide and possible habituation to humans may have played a role in the reduced visitor effect. The results of the study showed inter-specific differences in the lemurs’ use of space and behavior and a complex combined effect of enrichment, visitor presence and meteorological conditions.

An optimal method for stunning animals before slaughter should result in instantaneous and irreversible insensibility. Today, there are various stunning and slaughter practices used around the world for farmed fish. With aquaculture being a growing food sector, the welfare of the animals used has become increasingly important in the consciousness of consumers, researchers and regulatory bodies. With growing research into the subject matter, an overview to summarize and examine how these practices impact on the welfare of the fish, and how well they conform to animal welfare legislation, is warranted to minimize the suffering of farmed fish. Stunning practices used in aquaculture include methods such as electrical and percussive stunning, carbon dioxide and asphyxiation. Each with its own level of effectiveness in terms of how fast the method results in loss of consciousness, whether or not the effect is reversible and how the welfare of the fish is affected as determined by behavioural and physiological stress responses. It was concluded that there is no unambiguous answer as to which stunning method is optimal in regard to animal welfare in modern day aquaculture. The optimal method for a given facility is influenced by factors like practicalities relating to each individual method as well as legislation on EU and national level. Suggestions were made for future research.

Incidental bycatch in gillnets is a substantial threat to small cetaceans. Using Acoustic Deterrent Devices, “pingers”, have successfully reduced bycatch of harbour porpoises in gillnets. However, seals can use pingers as “dinner-bells” to easier find gillnets in order to raid and destroy them, further aggravating the existing conflicts between seals and coastal fisheries. Therefore, in the present study, the efficiency of two alleged “seal-safe” pingers, an experimental Banana pinger “SSB” and a Future Oceans F70 pinger “FO”, in deterring harbour porpoises from the vicinity of gillnets and thereby reducing bycatch in commercial gillnet fisheries, was tested. This was done by deploying click detectors, “C-PODs”, recording Detection Positive Minutes per hour, at each end of gillnets, provided with the two pinger types or no pingers at all. Bycatch instances were recorded into logbooks by participating fishermen and verified using video footage from on-board video cameras. Results showed that video monitoring was a reliable method for verifying the number of bycatches of porpoises and seals, but not seabirds, recorded in the fishermen’s logbooks. The experimental SSB pingers and the FO pingers significantly reduced porpoise presence, measured as Detection Positive Minutes per hour in the vicinity of the nets, compared to gillnets without pingers. However, the sample size was too small to yield a significant result regarding the bycatch reducing efficiency and dinner bell effect of the experimental pingers. Nevertheless, bycatch trends suggest that pingers did in fact reduce porpoise bycatch. Although both successful, FO pingers were slightly more efficient in deterring porpoises than SSB pingers. The SSB pinger sounds had bigger directionality variations than the FO pinger, which may have affected its deterrent effects. Therefore, additional trials are needed to further investigate this aspect.