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<p> Alterations in neural tissues associated with environmental variables have been studied for many years. Anatomical changes in the neocortex of rats in response to exposure to complex environments were observed and replicated in a number of studies both within and across species. These changes are not dependent on the age of the animal or on the duration of exposure, and have been demonstrated in structures outside of the cortex. Due to the undisputed involvement of both the neocortex and the hippocampus in learning and memory, researchers applied a widely used model system of a synaptic mechanism for learning, long-term potentiation (LTP), to the environmental enrichment paradigm and demonstrated significant enhancements in hippocampal field potentials in enriched rats. The present study examines whether the neocortex also showed evidence of plasticity in synaptic transmission. No effects for environmental enrichment were observed on the maximum amplitude of neocortical field responses evoked from the corpus callosum. To assess the plasticity of the chronic preparation used in the study, the animals were exposed to trains of pulses previously shown to induce electrical LTP in the cortex, but revealed only a slight, although significant, depression of the evoked response amplitude. An alteration in the stimulation parameters did not result in an enhanced response. Cortical depth measures suggested that the enriched environment was indeed sufficient to produce plastic changes in anatomy, if not in the efficacy of synaptic transmission. The importance of these findings in the neocortex leads us to question the validity of the LTP model of learning and memory.</p> / Thesis / Master of Science (MSc)

Humans maintain wild animals in zoological parks for the purposes of education,conservation, research, and recreation. However, abnormal behaviors may develop in animals housed in human-made environments, if those environments do not allow them to carry out their natural behaviors (such as swimming, climbing, stalking, and predation). Captive environments in zoological parks often do not provide for natural behaviors due to spatial constraints and negative public reaction. Tigers (Panthera tigris) present a difficult case; they have large home ranges in the wild and natural predatory hunting behaviors that are difficult to provide for in captivity. As the numbers of wild tigers decline, captive breeding programs have become a major focus of the zoo community, which magnifies the importance of research on tiger husbandry. A body of research exists on small felids, but little, if any, has focused on tigers. This thesis presents an analysis of the effects of the captive environment on the behaviors of 18 captive Bengal and Siberian tigers in four zoological parks in Virginia and Pennsylvania. Certain animal characteristics (such as subspecies, and age) were also related to behavior. Several characteristics of the captive environment had statistically significant effects on stereotypic and exploratory behaviors of tigers: shade availability, the presence of a body of water, cage size, the presence of a conspecific, vegetation, environmental enrichment, and substrate type. There were significant differences in the behaviors of the two subspecies studied, but the reason for the differences are unclear. The results of this study showed clearly that tigers kept in more natural and complex enclosures performed less stereotypic pacing (unnatural behavior), and more exploratory (natural) behaviors than those housed in less natural enclosures. Reducing the stress level in captive tigers will enhance the animals' overall physical and psychological well being, which will in turn increase the success of captive breeding programs. These results suggest that captive tigers should be housed in large enclosures containing natural substrate and vegetation, water pools, ample shade, a variety of resting locations, and a variety of enrichment items. / Master of Science

Welfare for captive animals in laboratory, farm, companion and zoo settings is difficult to assess and subject to interpretation. The perceived needs of animals change over time and according to budget, fashion and policy. The assessment of the value of environmental enrichment for captive animals is hampered by the lack of consistent, quantitative, objective and methodical research. Enrichment devices and other welfare improvement strategies lack long-term assessment or implementation. The concept of cognitive enrichment, or enabling captive animals to use their cognitive abilities to solve problems and meet challenges in their environment, is in the initial stages of development. Preliminary findings indicate that cognitive enrichment seems to be an effective method of improving captive animal welfare. In the research described in this thesis, several welfare measurement techniques including changes in activity budget, stereotypic behaviour and cognitive bias were compared and used to measure the effects of cognitive enrichment upon captive pigeons and dogs in order to obtain a useable cognitive enrichment paradigm that can be extrapolated to many types of captive animals. Key findings were that cognitive enrichment improved captive group-housed pigeon welfare and individually-kennelled dog welfare, indicated by significant changes in activity budget, increases in ‘optimism’, and reduction of stereotypic behaviour in subjects. Subjects used both low-tech and high-tech cognitive enrichments as intended and showed no sign of habituation. It is hoped that these findings will be used to improve captive animal welfare, and that the cognitive enrichment and cognitive bias paradigms developed will add to this field of research.

Master of Science / Department of Psychology / Mary E. Cain / Previous research has shown that both the environment and psychostimulant use influence dopamine levels via the mesolimbic dopamine pathway. C-fos expression has also been observed following exposure to novel environments and psychostimulants. The present study looked to determine the effects of acute amphetamine exposure on locomotor activity and c-fos expression in the basolateral and central nucleus of the amygdala, for rats raised in either an enriched condition (EC), impoverished condition (IC), or social condition (SC). Rats were reared in either the EC, IC, or SC for 30 days, after which they received an acute amphetamine injection (1.0 mg/kg) and locomotor activity was measured. Following the locomotor test rats were perfused and immunohistochemistry was used to measure c-fos levels in the basolateral and central nucleus of the amygdala. Results showed that EC amphetamine rats had significantly greater locomotor activity compared to EC saline rats. There were no significant group or treatment differences in c-fos expression in the ACe. In the BLA SC amphetamine rats had significantly greater c-fos expression than EC amphetamine rats. Overall, the current study revealed that environmental enrichment and amphetamine do significantly alter locomotor activity and c-fos expression in the BLA.

O desenvolvimento humano pode ser influenciado pelo ambiente. Estímulos recebidos ao longo da vida determinam seu progresso e sucesso. Estímulos positivos levam ao desenvolvimento de habilidades, melhorando funções cognitivas e da memória, enquanto estímulos negativos podem predispor a patologias como o estresse. Eventos estressantes durante a infância aumentam a predisposição para o desenvolvimento de transtornos psiquiátricos ao longo da vida. A separação materna, modelo animal de estresse pós-natal, promove diversas alterações comportamentais e encefálicas. Animais submetidos à separação materna apresentam comportamentos que mimetizam doenças psiquiátricas humanas. Por outro lado, diversos trabalhos sugerem que o enriquecimento ambiental pode ter efeito benéfico na reversão ou atenuação de modificações comportamentais e encefálicas promovidas por modelos animais de depressão, esquizofrenia, ansiedade e hiperatividade. Esses aspectos motivaram-nos a estudar se as alterações causadas por estresse podem ser revertidas ou atenuadas pelo enriquecimento do ambiente. Há evidências que sugerem um importante envolvimento de células gliais e de transportadores de glutamato presentes nessas células em modelos animais de transtornos psiquiátricos. Sendo assim, investigamos a expressão de mRNA e proteínas de dois transportadores de glutamato gliais e um neuronal, do receptor de glutamato AMPA, de marcadores gliais GFAP, S100?, glutamina sintase (GS) e do marcador de neurônios maduros NeuN na camada molecular e granular do giro denteado do hipocampo de ratos de 60 dias. Observamos que a separação materna diminui a expressão das proteínas GLAST, GLT-1, GS e NeuN, reduz a expressão dos genes Gria1 (AMPA) e S100?, e aumenta a expressão da proteína EAAC1 no giro denteado. Nossos dados sugerem uma reversão das alterações causadas pela separação materna em relação ao gene Gria1/AMPA e às proteínas GLAST, GLT-1 e EAAC1 após o enriquecimento ambiental. Portanto, o enriquecimento ambiental pode reverter as modificações causadas pela separação materna nas vias glutamatérgicas. Esses efeitos benéficos podem ser investigados para auxiliar no tratamento de transtornos psiquiátricos relacionados à separação materna. / Human development can be influenced by the environment. Stimuli received throughout life determine its progress and success. Positive stimuli lead to development of skills, improving cognitive and memory functions, while negative stimuli may predispose to pathologies such as stress. Stressful events during childhood increase the predisposition to psychiatric disorders throughout life. Maternal separation, an animal model of postnatal stress, promotes several behavioral and encephalic changes. Animals submitted to maternal separation stage behaviors associated with psychiatric diseases in humans. On the other hand, some researches have suggested that environmental enrichment may have some beneficial effects on the reversal or attenuation of behavioral and encephalic modifications promoted by animal models of depression, schizophrenia, anxiety and hyperactivity, which motivates us to study if these changes, stirred by this kind of stress, can be reversed or mitigated by environmental enrichment. There are evidences suggesting the involvement of glial cells and glutamate transporters existent in these cells in psychiatric disorders and animal models of these disorders. Therefore, we investigated mRNA and protein expression of two glial and one neuronal glutamate transporters, AMPA glutamate receptor, glial markers GFAP, S100?, glutamine synthase (GS), and the NeuN neuronal marker in the molecular and granular layer of the hippocampal gyrus in sixty-days-old rats. We observed that maternal separation decreases expression of GLAST, GLT-1, GS and NeuN proteins, reduces Gria1 (AMPA) and S100? gene expression, and increases EAAC1 protein expression in the dentate gyrus. After environmental enrichment, our data suggests a reversal of the maternal separation changes in the Gria1/AMPA gene and the GLAST, GLT-1 and EAAC1 proteins. Therefore, environmental enrichment may reverse the maternal separation changes in the glutamatergic pathways. These beneficial effects may be investigated to aid in the treatment of psychiatric disorders related to maternal separation.

Manter animais em cativeiro implica no dever ético de lhes proporcionar saúde física e psicológica. Procedimentos conhecidos como Enriquecimento Ambiental buscam elevar o bem-estar de animais cativos. Neste trabalho, três artefatos foram testados para avaliar sua eficácia como enriquecimento ambiental para macacos-prego (Cebus apella). Dois destes artefatos (Brinquedo e Caixa de forrageamento) já haviam sido testados anteriormente por Boinski et al. (1999). O terceiro artefato (Ferramenta) foi testado pela primeira vez e buscava proporcionar aos animais a possibilidade de executar um comportamento típico da espécie: a quebra de cocos. Usamos medidas comportamentais e de corticosterona fecal para medir os efeitos dos artefatos. Foram encontradas algumas diferenças significativas entre as condições experimentais (controles e com artefatos), e entre a freqüência de interação com os artefatos, porém os resultados não nos possibilitam afirmar se algum dos artefatos seria um enriquecimento ambiental eficaz para macacos-prego cativos. Acreditamos que experimentos ainda devem ser feitos para elucidar os efeito de fatores como: o ambiente externo ao recinto, os procedimentos de manutenção direcionados aos animais e a metodologia de aplicação dos artefatos, sobre o bem-estar de animais cativos. / Keeping animals in captivity implies in an ethical duty of offering conditions that foster their physical and psychological health. Procedures known as Environmental Enrichment seek to enhance animals’ life quality. Here we tested three stimuli efficiency as environmental enrichment for captive capuchin monkeys (Cebus apella). Two which had been previously tested (Toy and Box) by Boinski et al. (1999). The third one (Tool) was tested for the first time and aimed giving the animals the opportunity to perform a species-typical behavior: cracking open nuts. The stimuli had their effects measured by behavioral and fecal corticosterone sampling. Some statistical significant differences were found between experimental conditions (control and stimuli) and between the frequencies of interaction with the stimuli. The data, however, did not point to any of the tested stimuli as effective environmental enrichment for capuchin monkeys. We believe, therefore, that more research should be conducted in order to clarify the effects of factors such as the environment outside the cages, the maintenance procedures, and the stimulus presentation procedures, on the well-being of captive animals.

O enriquecimento ambiental (EA) apresenta efeitos positivos sobre a dependência ao etanol e é possível que atue como um estressor leve, ajudando o animal a lidar melhor com situações adversas. Camundongos foram separados em três grupos: controle (CO), EA e estresse (ES). Foram estudados os efeitos do EA e ES no comportamento tipo ansiedade, memória episódica, preferência condicionada a lugar (CPP) com etanol, corticosterona e quantidade de receptor de oxitocina (OT). O EA diminuiu a latência de entrada no braço aberto, sugerindo efeito ansiolítico, ao contrário do ES. A memória não diferiu entre os grupos e o EA mostrou maior preferência condicionada ao etanol, comparado com CO. A corticosterona não mostrou diferença. A autorradiografia apontou diminuição da ligação de OT nos núcleos olfatórios e córtex frontal dos animais EA e aumento da ligação de OT na amígdala do ES em relação ao CO, sugerindo maior comportamento pró-social e maior ansiedade, respectivamente. Concluímos que o ambiente é capaz de afetar circuitos cerebrais envolvidos no estresse e dependência. / Environmental enrichment (EE) presents positive effects upon alcohol addiction and its possible that EE acts as a mild stressor, which would help animal cope better with future stressful situations. Mice were separated in three groups: control (CO), EE and stress (ST). We studied the effects of EE and ST on anxiety-like behavior, episodic memory, conditioned place preference (CPP) to ethanol, corticosterone and quantity of oxytocin receptor. EE decreased the latency to enter the open arm, suggesting an anxiolytic effect, unlike ST. Memory did not differ between groups and EE demonstrated a higher preference to ethanol compared to CO. Corticosterone levels didnt differ among groups. Autoradiography showed decreased binding of OT in the olfactory nuclei and frontal cortex of EE animals and enhanced binding of OT in the amygdala of ST animals compared to CO, suggesting higher pro-social behavior and higher anxiety-like behavior, respectively. We conclude that the environment is capable of affecting important brain circuitries involved in stress and drug addiction.

O objetivo deste trabalho foi avaliar o efeito de duas dietas líquidas , associadas ao uso de enriquecimento ambiental (EA) em bezerros alojados individualmente. O delineamento experimental foi um fatorial 2 x 2, sendo avaliadas duas dietas líquidas (leite ou sucedâneo lácteo) e ausência ou presença de EA (bico de mamadeira e coçador). Foram utilizados 24 bezerros recém-nascidos da raça Holandês, distribuidos em 4 tratamentos: Leite, Leite + EA, Sucedâneo, Sucedâneo + EA. O sucedâneo lácteo continha em sua composição 22,33 % PB, 15,96% EE e 1,07% de FDN, na matéria seca, e foi diluído para uma concentração de 14% de sólidos. O leite apresentava 3,37% PB, 3,38% EE na matéria seca e 12,25% de sólidos. Os animais foram alojados em abrigos individuais e tinham livre acesso a água e ao concentrado. Foi utilizado um programa alimentar step up/step down, com desaleitamento realizado aos 56 dias de vida. Não houve efeito da associação das diferentes dietas líquidas com EA sobre o desempenho dos animais (P>0,05), mas houve tendência (P<0,082) de redução do consumo de concentrado para animais com acesso ao EA. O desempenho animal foi sigificativamente afetado pelo fator idade (P<0,05). A associação de dieta líquida com EA não afetou os parâmetros metabólicos, mas animais aleitados com leite apresentaram maiores concentrações de B-Hidroxibutirato e ureia plasmática (P>0,05). Da mesma forma, a combinação de dietas com EA não afetou o escore fecal, mas animais aleitados com sucedâneo apresentaram fezes mais fluidas (P>0,05). Não houve alteração no repertório comportamental dos animais em função de diferentes dietas líquidas associadas ao EA (P>0,05). Durante o período de avaliação de duas horas após o aleitamento da manhã, os animais passaram a maior parte do tempo em ócio (60%), seja em pé ou deitado. Os outros 40% do tempo foram gastos com atividades relacionadas a alimentação, como pastejando (8,5%), consumindo concentrado (3,8%); atividades fisiológicas, como dormindo (5,45%), ruminando (1,3%), defecando (0,6%), urinando (0,1%); ou ainda atividades comportamentais, como explorando o ambiente (6,3%) e se lambendo (6,7%). Os animais utilizaram o EA por aproximadamente 1,4% do tempo somente e vocalizaram durante 1,7% ou realizaram mamada não nutritiva por 1,9%. Houve tendência (P<0,0627) para redução na vocalização e função da disponibilidade de EA. O fornecimento de diferentes dietas líquidas associadas ou não ao EA não afetaram o desempenho, metabolismo ou comportamento animal. No entanto, não se descarta a possibilidade de ganhos no bem-estar dos animais com o uso de enriquecimento ambiental uma vez que, os animais vocalizam menos durante o processo de desaleitamento. / The objective of this study was to evaluate the effect of two liquid diets, associated to the use of environmental enrichment (EE) in individually housed calves. For the evaluation a 2 x 2 factorial experimental design, with two liquid diets (milk or milk replacer) and absence or presence of EE (bottle nipple and scraper) evaluated. Twenty-four newborn Holstein calves were distributed in to four treatments: Milk, Milk + EE, Milk replacer, Milk replacer + EE. The milk replacer contained 22.33% CP, 15.96% EE and 1.07% NDF, in the dry matter, and was diluted to a concentration of 14% solids. The milk presented 3.37% CP, 3.38 EE in the dry matter and 12, 25% solids. The animals were housed in individual shelters and had free access to water and concentrate. A step up/step down diet program was used, with weaning performed at 56 days of age. There was no effect of the association of different liquid diets with EA on the performance of the animals (P> 0.05), but there was a tendency (P <0.082) to reduce the concentrate intake for animals with access to EE. The animal performance parameters were significantly affected by the age factor (P<0.05). The association of liquid diet with EE did not affect metabolic parameters, but milk-fed animals had higher concentrations of B-Hydroxybutyrate and plasma urea-N (P>0.05). Likewise, the combination of liquid diets with EE did not affect fecal score, but animals fed with milk replacer had more fluid faeces (P>0.05). There was no change in the behavioral repertoire of the animals as a function of liquid diets associated with EA (P> 0.05). During the two-hour evaluation period after liquid diet feeding, the animals spent most of their time in leisure (60%), either standing or laying down. The other 40% of the time was spent on feeding related activities such as grazing (8.5%), consuming concentrate (3.8%); physiological activities, such as sleeping (5.45%), ruminating (1.3%), defecating (0.6%), urinating (0.1%); or behavioral activities, such as exploring the environment (6.3%) and self-grooming (6.7%). The animals used the EE for approximately 1.4% of the time only, and vocalized for 1.7% or performed non-nutritive feeding for 1.9%. There was a trend (P <0.0627) for reduction in vocalization and function of EE availability. Feeding different liquid diets associated or not with EA did not affect performance, metabolic or animal behavior. However, the possibility of improved animal welfare with the use of environmental enrichment is not ruled out since the animals vocalize less during the process of weaning.

O enriquecimento ambiental (EA) é capaz de alterar a percepção a estímulos nociceptivos, bem como de aumentar a resposta analgésica induzida por opioides. Considerando que a dor neuropática é um grave problema de saúde pública e o tratamento para esta condição ainda é insatisfatório e acarreta efeitos adversos severos, os objetivos deste trabalho foram avaliar a interferência do bem-estar animal na sensibilidade dolorosa de ratos frente a diferentes estímulos nociceptivos e investigar possíveis mecanismos envolvidos neste efeito. Os animais foram submetidos à avaliação da ansiedade e da sensibilidade dolorosa, em modelo de neuropática, frente a estímulos nociceptivos mecânicos e térmicos. Foi verificado que um protocolo de EA elaborado e iniciado desde o nascimento foi capaz de reverter totalmente a dor neuropática de animais submetidos à constrição crônica do nervo isquiático (CCI). Este efeito foi completamente abolido quando os animais enriquecidos foram tratados com naloxona, um antagonista opioide não seletivo. Análises de Western Blot não mostraram diferenças na expressão de receptores opioides em regiões relacionadas ao processamento e controle da dor, porém os níveis circulantes de beta-endorfina e met-encefalina aumentaram na presença de dor crônica nos animais enriquecidos. Os níveis séricos de corticosterona também se apresentaram aumentados nos animais com EA, independentemente da neuropatia, mas o tratamento com mifepristona, um antagonista de receptores de glicocorticoides, não alterou a analgesia dos animais operados. Ainda, o EA também reduziu a imunorreatividade para serotonina na medula espinal de animais com CCI. Além do efeito analgésico, o EA também reduziu o marcador de lesão neuronal ATF-3 no gânglio da raiz dorsal e, no local da constrição, reduziu a degeneração neuronal característica do modelo, induzindo ainda, a presença predominantemente de macrófagos do tipo M2. Este trabalho reforça a importância do bem-estar na prevenção do desenvolvimento da dor neuropática e mostra uma abordagem não farmacológica que pode aumentar a resiliência de animais contra estímulos nocivos. / Environmental enrichment (EE) is capable of altering the perception of nociceptive stimuli, as well as increasing the analgesic response induced by opioids. Considering that neuropathic pain is a serious public health problem and the treatment for this condition is still unsatisfactory and induces severe side effects, the aims of this study were to evaluate the interference of animal welfare in the sensitivity to different nociceptive stimuli and to investigate possible mechanisms involved in this effect. Animals were submitted to the evaluation of anxiety and pain sensitivity in a model of neuropathic pain, against mechanical and thermal nociceptive stimuli. It was seen that an elaborated EE starting from birth was able to totally reverse the neuropathic pain of animals submitted to chronic constriction injury of the sciatic nerve (CCI). This effect was completely abolished when enriched animals were treated with naloxone, a nonselective opioid antagonist. Western blot analysis did not show differences in opioid receptor expression in regions related to pain processing and control, however, circulating levels of beta-endorphin and met-enkephalin were increased in the presence of chronic pain in enriched animals. Serum corticosterone levels were also increased in animals with EE regardless of neuropathy, but treatment with mifepristone, a nonselective glucocorticoid receptor antagonist, did not alter the analgesia of operated animals. Moreover, EE reduced serotonin immunoreactivity in the spinal cord of CCI animals. In addition to analgesic effect, EE also reduced the neuronal injury marker ATF-3 at the dorsal root ganglia and, at the site of constriction, decreased the neuronal degeneration characteristic of the model, inducing the presence of M2 macrophages subtype predominantly. This work reinforces the importance of well-being in preventing the development of neuropathic pain and shows a non-pharmacological approach that may increase animal resilience against noxious stimuli.