ABSTRACT   Migration is common among populations of brown trout (Salmo trutta) and Atlantic salmon (Salmo salar). However, not all individuals in the same population migrate, a phenomenon referred to as partial migration. The aim of this thesis was to investigate if an individual’s behaviour and metabolic rate influences its decision to migrate and how such knowledge may be used when trying to produce hatchery-raised smolts with as high a proportion of migrating individuals as possible. In paper I the influence of reduced food ration on the proportion and swimming speed of migrating brown trout and Atlantic salmon smolts was investigated. Furthermore, the standard metabolic rate (SMR) of migrating and non-migrating individuals was compared. In paper II, a laboratory experiment, SMR was correlated to the behaviour of individual brown trout and Atlantic salmon. Dominant fish of both species had a higher SMR than subordinates (paper II). In addition, migrant brown trout had a higher SMR than non-migrant trout when given a normal food ration, whereas no difference in SMR between migrating and non-migrating salmon could be seen (paper I). When administered low food rations, smolts of both species migrated faster than smolts given a normal food ration, and the proportion of migrating smolts was higher for salmon given less food when the size difference for smolts from the two feeding regimes was large (paper I). Other factors that influenced migration speed were the degree of smolt development and water temperature (paper I). SMR was not correlated with aggressiveness, or with different measurements of boldness. Moreover, aggression and boldness were not correlated with each other (paper II). Trout showed a higher level of aggressiveness and acclimated more rapidly to laboratory conditions than salmon (paper II). In summary, there was no support for the existence of coping styles in migratory Atlantic salmon and brown trout. Instead, metabolic rates were related to both migratory behaviour and social status. Furthermore, an individual’s decision to migrate was influenced by ration size.

Spider monkeys (Ateles sp.) are common in zoological parks, but rare in scientific publications. Studies on tool use in primates have mostly focused on impressive tool users such as chimpanzees. Spider monkeys fulfill several criteria that are known to be associated with tool use. To be able to give an appropriate environment and enrichment for spider monkeys in captivity more knowledge is needed about their cognitive abilities. In this study we wanted to see if five male spider monkeys kept in a zoo could learn to use tools to reach a reward. Experiment 1 examined the subjects’ ability to learn to use a stick-tool to extract honey from a tube and experiment 2 their ability to learn to use a rake-tool to reach a reward. Each experiment consisted of three parts; A – monkeys got tools and treat next to each other; B – monkeys were shown how to use tool to get treat by a keeper and then got tools and treats next to each other; C – monkeys got tools and treats so they just could pull out the tool and get the treat. In both experiments at least two different spider monkeys succeeded with the zero-order manipulation task to pull out the tool and get treat in part C. Longer studies need to be conducted to be able to say if spider monkeys are able to learn a more complex tool using behavior as needed in part A and B.

Ett stort antal utrotningshotade djurarter ingår idag i bevarandeprogram världen över. Små populationer hålls då i skyddade miljöer, exempelvis i djurparker, och i många fall är målet att återintroducera djur till naturen. Dessvärre är det vanligt att det uppstår problem när djur återintroduceras vilket kan bero på beteendemässiga anpassningar som uppkommit under tiden i fångenskap. Syftet med den här studien var därför att undersöka beteendemässiga aspekter på bevarandeavel och försöka ta reda på om och hur djur påverkas beteendemässigt av att hållas i skyddade fångenskapsmiljöer. I projektet användes röda djungelhöns (Gallus gallus) som en fallstudie. En granskning av beteendevariation mellan olika populationer av röda djungelhöns i fångenskap konstaterade skillnader i antipredatorbeteende, socialt beteende och födosöksbeteende. Vid en genetisk studie av samma populationer upptäcktes dessutom ett samband mellan genetisk diversitet och beteendevariation som potentiellt kan vara intressant ur ett bevarandeperspektiv. Socialt beteende, födosöksbeteende och olika aspekter av rädsla studerades vidare i populationer med olika bakgrund som fick växa upp tillsammans i en grupp. Resultaten visade att populationerna bara skilde sig åt i rädslebeteenden vilket antyder att denna typ av beteende i större utsträckning påverkas av långvarig avel i en viss fångenskapsmiljö medan socialt beteende och födosöksbeteende istället kan bero på den omedelbara sociala eller fysiska miljön. Antipredatorbeteende studerades också i en longitudinell studie av två populationer över fyra generationer och det visade sig att populationerna blev mer lika varandra ju längre tiden gick då de hölls under likadana miljöförhållanden. Det verkar alltså som om antipredatorbeteende kan förändras av avel i en viss miljö efter bara ett fåtal generationer. Utöver detta studerades även dagliga beteendemönster i olika djurparksmiljöer samt dygnsrytm av galanden hos både vilda populationer och djurparkspopulationer inom artens naturliga utbredningsområde. Resultaten tyder på att sådana beteendemönster inte påverkas nämnvärt av att djur hålls i fångenskap. Fallstudien som presenteras här är ett av de första försöken att, ur ett bevarandeperspektiv, studera hur fångenskapsmiljöer kan påverka djurs beteende och resultaten talar för att dessa aspekter är viktiga att ta hänsyn till vid planering av bevarandeavel. / A number of endangered species are currently involved in conservation breeding programs worldwide. Conservation breeding deals with propagation of captive populations, often with the ultimate aim of releasing animals into the wild. However, an alarmingly high proportion of reintroductions have not been successful in establishing viable populations, possibly due to behavioural problems caused by genetic adaptation to captivity. The main aim of this thesis was to study behavioural aspects of conservation breeding and investigate whether, and how, maintenance of small populations in captivity cause behavioural modifications, which could affect the success of reintroductions. Throughout the project, the red junglefowl (Gallus gallus) was used as a case study, representing animals maintained in captive populations. A screening of behavioural variation revealed that captive populations differ in antipredator, social and exploratory behaviours, all of which are central components of life in the wild. A correlation was also found between genetic diversity and behavioural variation. This has not been reported before and may potentially have interesting implications for conservation breeding. When studying the behaviour of populations with different backgrounds being raised together as one group, the results suggested that fear-related behaviours may be more affected by long-term breeding in a certain captive environment than social and exploratory behaviours which seem to be more influenced by the immediate social or physical environment. A longitudinal study of antipredator behaviour in two populations across four generations revealed that the populations became more similar over time when maintained under identical conditions. This demonstrates that effects of a new environment can appear after only a few generations. Furthermore, daily behavioural routines in different captive environments as well as diurnal crowing rhythms in both wild and captive populations were studied in the species’ natural region of distribution and the results suggest that such behavioural patterns are not affected by the captive environments to any notable extent. The present case study is one of the first attempts to, from a conservation perspective, study how captive environments can affect behaviour and the results imply that these aspects are important to take into consideration in conservation breeding programs.

The main aim of this thesis was to study domestication effects on foraging behaviour in chickens and to investigate whether and how domestication and selection for high production have influenced adaptability in chickens. Two domestic strains of chickens (egg layers and meat type chickens) and their wild ancestor, the red jungle fowl (RJF) were compared in different test situations with respect to foraging behaviour and adaptability. The domestic strains showed a modified foraging strategy, where they were less inclined to explore and feed from a hidden food source, i.e. they showed less contrafreeloading (CFL, the behaviour of working for food even though identical food can be easily obtained) than RJF. The difference in CFL between RJF and the layers were not altered by food deprivation, which suggests that the lower CFL in the layers represents a genetically based difference in feeding strategy. In addition, CFL decreased with age in RJF and layers and social isolation decreased CFL in RJF. Furthermore, when foraging, RJF acquired information about the quality of different food sources, which was utilised after a change in environmental conditions. Contrary to this, layers gained less information during foraging and showed an impaired spatial learning ability compared to RJF, and in this respect, layers showed a lower degree of adaptability. Chronic stress impaired the learning capacity of both breeds but RJF seemed to be overall faster to learn to locate food in a spatial learning task. Furthermore, stressed RJF started to eat faster in the spatial learning test than non-stressed RJF, and contrary to this, stressed layers showed a more passive response by prolonging the time to start feeding compared to non-stressed layers. This indicates a more active response to stress in RJF than in layers. Similarly, when RJF and layers were exposed to food deprivation, RJF showed an active response by increasing their time spent on foraging behaviour. The general results in this thesis most likely reflect different adaptive strategies, where RJF appear to be better adapted to a stochastic environment, and the domestic strains to grow and produce egg in a more predictable environment. The findings are in accordance with the resource allocation theory, which suggests that animals selected for high production are expected to reallocate a high proportion of resources into production traits and hence fewer resources might be left to other biological processes, e.g. exploratory behaviour. Selection for high production seems to influence the ability of chickens to cope with a changing environment, which may have implications for the welfare of chickens in a production environment.

Odor discrimination ability was tested in four female South African fur seals (Arctocephalus pusillus pusillus) using a food-rewarded two-choice instrumental conditioning paradigm. The seals’ ability to distinguish between members of homologous series of aliphatic ketones (2-butanone to 2-heptanone) and alcohols (1-butanol to 1-heptanol) was assessed. The results showed that three out of four seals successfully discriminated between all of their stimulus combinations in both classes of odorants. One seal succeeded to reach the discrimination criterion with all 2-ketones but failed with all 1-alcohols. No significant correlation between odor discrimination performance and structural similarity of the odorants in terms of differences in carbon chain length was found in either of the two chemical classes. Furthermore, it was found that the 2-ketones were significantly better discriminated than the 1-alcohols. The fact that both classes of odorants are known to be present in the natural environment of seals provides a possible explanation as to why most of the seals were able to successfully discriminate between them. The results of the present study support the notion that the sense of smell may play an important role in behavioral contexts such as social communication, foraging and reproductive behavior of fur seals.

I investigated the understanding of human communicative motives in domestic dogs. Dogs use human communicative cues, like the pointing gesture when searching for hidden food, but it is uncertain how dogs interpret human communication. 32 dogs were presented with two communicative contexts in an object choice task experimental design. In a cooperative context the experimenter informed the subject where food was hidden by pointing and giving a verbal indication. In a competitive context the experimenter held out her arm towards the correct location in a stop gesture and firmly said no. To be successful in the competitive context the subject had to understand the experimenters communicative motive and make an inference from the prohibition (i.e. she would only prohibit it if there was something good there). The average correct choices were compared between the conditions. The dogs successfully followed the cooperative communication. They showed a trend towards choosing the baited cup in the competitive condition. A second study tested if the stop gesture affected the dogs’ choice, since it is not known how dogs interpret gestures. The pointing cue was now presented with the prohibiting command and the stop gesture was presented with the cooperative verbal cue. The dogs used the cooperative communication but did not understand the competitive context. A difference between the contexts was found. The dogs did not differentiate between the gestures. In conclusion dogs do not make inferences from competitive communication or prohibition but are specialized in utilizing cooperative communication

Animals raised in captivity often fail to express appropriate anti-predator behaviour when reintroduced into the wild. The European bison (Bison bonasus) is a species that was close to extinction in the early 20th century but was saved in the last moment by intense captive breeding and subsequent reintroduction into the wild. In this study, seven groups of European bison living in different locations in Sweden were studied to investigate whether there was any difference in the anti-predator behaviour depending on the type of enclosure they were kept in. Olfactory and auditory stimuli from moose, as a control, and from two predators, wolf and bear, and visual stimulus (silhouette of a wolf) were presented to the animals and their response to them and behaviour following presentation were analysed. The results showed that European bison kept in barren enclosures responded stronger to auditory stimuli than those that were kept in naturalistic enclosures. The results further showed that the animals had a stronger response to the visual stimulus than to the auditory stimuli. The animals changed their behaviour after stimuli presentations compared to a pre-test baseline. They moved, stood still and ate for a significantly longer period of time and they rested for a shorter period of time after being presented olfactory, auditory and visual stimuli than during pre-test baseline.

Only in primates is permanent male-female association the most widespread social structure of all. The continuous presence of resident males in the social group can have significant impacts on female fitness, both in forms of costs and benefits. In this study I investigate particular short-term benefits and costs of resident males to females in a population of ursine colobus (Colobus vellerosus). I hypothesise that for females permanent association with males result in certain benefits and certain costs, exceeding those provided or imposed by other females. The results indicate that female derive greater benefits from males than from females during intergroup encounters and in the form of vigilance since males were the main participants in intergroup encounter and were more vigilant than females. I could not confirm any type of behaviour employed by resident males that is costly to females. However, the rarity and subtleness of some costly male behaviours imply that more data is needed before making a conclusion on their absence or occurrence in this population and I purpose that herding behaviour could occur at my study site. Moreover, multi-male groups (MM-groups) showed higher rates of vigilance than single-male groups (SM-groups) and had a tendency to experiencing fewer intergroup encounters than SM-groups. I interpret the former as a result of the demanding social conditions in the MM-groups. The latter indicate that females may benefit from MM-group living through a decrease in intergroup encounters.

For many years, primates have been considered to be animals with a poorly developed sense of smell. However, in recent years several studies have shown that at least some primate species have a high olfactory sensitivity for a variety of odorants. The present study used a two-choice instrumental conditioning paradigm to test the olfactory sensitivity for six aromatic aldehydes in four spider monkeys (Ateles geoffroyi). With helional, cyclamal,canthoxal and lilial all animals discriminated concentrations below 1 ppm from the odorless solvent, with single individuals even scoring better. With 3-phenyl-propionic aldehyde all animals detected concentrations below 2 ppb, and with bourgeonal even below 0.3 ppb. The detection thresholds of the odorants changed systematically with molecular structure. Addition of a dioxo or methoxy group to the benzene ring led to an increase in threshold values,while the absence of a methyl group close to the aldehyde moiety was linked to a low threshold value for the odorant. The study shows that spider monkeys have a well developed olfactory sensitivity for aromatic aldehydes.

Den biologiska studien har visat att värphöns genom domesticeringen förändrat och anpassat sina beteenden till att leva med människor. De har utvecklat en minskad rädsla för människor och en ökad stresstålighet. I denna studie har höns återhämtning efter en stressupplevelse studerats. För att se hur domesticeringen har påverkat återhämtningen jämfördes beteenden mellan raserna White Leghorn och red junglefowl. Frågeställningarna som användes för att undersöka detta var: Hur lång tid tar det innan höns återhämtar sina naturliga beteenden, ej stressrelaterade, efter en stressupplevelse? Finns det några skillnader i återhämtningen beroende av kön eller mellan raserna White Leghorn och red junglefowl? Resultatet visade inte på en specifik tid för återhämtning men på att en tydlig återhämtning förekom. Återhämtningen visades i beteendena Relaxed Behaviour, Comfort Behaviour, Perch, Vocalisation och Stand Alert genom att uppvisandet av beteendena skiljde sig signifikant med tiden. Även signifikanta skillnader visades mellan raserna, både i beteenden som visade på återhämtning och i beteenden som inte gjorde det. En slutsats om vilken ras som återhämtade sig snabbast var svår att dra. Detta på grund av att beteendeskillnaderna visade på olika tendenser gällande återhämtning. Skillnaderna i återhämtningen mellan könen, visades tendenser på att hanarna återhämtade sig snabbare. Domesticering har även i denna studie studerats utifrån ett lärarperspektiv. Undersökningen visade att domesticering kan ingå i biologiundervisningen i områdena evolution och genetik i år 7-9. / The biological study has shown that behaviour of laying hens has changed through domestication and the hens have adapted to live with humans. They have developed less fear of humans and more resistance to stress. In this study hens’ recovery after a stressful experience has been studied. A comparison between the breeds White Leghorn and red junglefowl has been made to see how the domestication has affected the recovery. The questions of this study are therefore: How long does it take for hens’ to recover their natural behaviour, not stressrelated, after a stressful experience? Are there any differences in recovery between the different breeds White Leghorn and red junglefowl? The results did not show a specific time for recovery, but a distinct recovery was noticed in Relaxed Behaviour, Comfort Behaviour, Perch, Vocalisation and Stand Alert. This beacause of the behaviours differed significantly with time. Also significant differences existed between the breeds, both in behaviours that showed recovery and in behavious that did not. A conclusion of what breed that recovered fastest was difficult to make. This because of the differences of behaviour showed different tendencies in recovery. The differences in recovery between the sexes showed tendencies that the males recovered fastest. The domestication has also been studied from a teacherperspective. The study showed that domestication can be a part of biologyeducation in the fields of evolution and genetics in the grades 7-9.