Prevalence and associations of Trypanosoma spp. and Sodalis glossinidius with intrinsic factors of tsetse flies

Trypanosomiasis has been identified as a neglected tropical disease in both humans and animals in many regions of sub-Saharan Africa. Whilst assessments of the biology of trypanosomes, vectors, vertebrate hosts and the environment have provided useful information about life cycles, transmission, and pathogenesis of the parasites that could be used for treatment and control, less information is available about the effects of interactions among multiple intrinsic factors on trypanosome presence in tsetse flies from different sites. It is known that multiple species of tsetse flies can transmit trypanosomes but differences in their vector competence has normally been studied in relation to individual factors in isolation, such as: intrinsic factors of the flies (e.g. age, sex); habitat characteristics; presence of endosymbionts (e.g. Wigglesworthia glossinidia, Sodalis glossinidius); feeding pattern; host communities that the flies feed on; and which species of trypanosomes are transmitted. The purpose of this study was to take a more integrated approach to investigate trypanosome prevalence in tsetse flies. In chapter 2, techniques were optimised for using the Polymerase Chain Reaction (PCR) to identify species of trypanosomes (Trypanosoma vivax, T. congolense, T. brucei, T. simiae, and T. godfreyi) present in four species of tsetse flies (Glossina austeni, G. brevipalpis, G. longipennis and G. pallidipes) from two regions of eastern Kenya (the Shimba Hills and Nguruman). Based on universal primers targeting the internal transcribed spacer 1 region (ITS-1), T. vivax was the predominant pathogenic species detected in flies, both singly and in combination with other species of trypanosomes. Using Generalised Linear Models (GLMs) and likelihood ratio tests to choose the best-fitting models, presence of T. vivax was significantly associated with an interaction between subpopulation (a combination between collection sites and species of Glossina) and sex of the flies (X2 = 7.52, df = 21, P-value = 0.0061); prevalence in females overall was higher than in males but this was not consistent across subpopulations. Similarly, T. congolense was significantly associated only with subpopulation (X2 = 18.77, df = 1, P-value = 0.0046); prevalence was higher overall in the Shimba Hills than in Nguruman but this pattern varied by species of tsetse fly. When associations were analysed in individual species of tsetse flies, there were no consistent associations between trypanosome prevalence and any single factor (site, sex, age) and different combinations of interactions were found to be significant for each. The results thus demonstrated complex interactions between vectors and trypanosome prevalence related to both the distribution and intrinsic factors of tsetse flies. The potential influence of the presence of S. glossinidius on trypanosome presence in tsetse flies was studied in chapter 3. A high number of Sodalis positive flies was found in the Shimba Hills, while there were only two positive flies from Nguruman. Presence or absence of Sodalis was significantly associated with subpopulation while trypanosome presence showed a significant association with age (X2 = 4.65, df = 14, P-value = 0.0310) and an interaction between subpopulation and sex (X2 = 18.94, df = 10, P-value = 0.0043). However, the specific associations that were significant varied across species of trypanosomes, with T. congolense and T. brucei but not T. vivax showing significant interactions involving Sodalis. Although it has previously been concluded that presence of Sodalis increases susceptibility to trypanosomes, the results presented here suggest a more complicated relationship, which may be biased by differences in the distribution and intrinsic factors of tsetse flies, as well as which trypanosome species are considered. In chapter 4 trypanosome status was studied in relation to blood meal sources, feeding status and feeding patterns of G. pallidipes (which was the predominant fly species collected for this study) as determined by sequencing the mitochondrial cytochrome B gene using DNA extracted from abdomen samples. African buffalo and African elephants were the main sources of blood meals but antelopes, warthogs, humans, giraffes and hyenas were also identified. Feeding on multiple hosts was common in flies sampled from the Shimba Hills but most flies from Nguruman had fed on single host species. Based on Multiple Correspondence Analysis (MCA), host-feeding patterns showed a correlation with site of sample collection and Sodalis status, while trypanosome status was correlated with sex and age of the flies, suggesting that recent host-feeding patterns from blood meal analysis cannot predict trypanosome status. In conclusion, the complexity of interactions found suggests that strategies of tsetse fly control should be specific to particular epidemic areas. Future studies should include laboratory experiments that use local colonies of tsetse flies, local strains of trypanosomes and local S. glossinidius under controlled environmental conditions to tease out the factors that affect vector competence and the relative influence of external environmental factors on the dynamics of these interactions.

Our own language: Scots verse translation and the second-generation Scottish renaissance

This thesis examines the role of Scots language verse translation in the second-generation or post-war Scottish Renaissance. The translation of European poetry into Scots was of central importance to the first-generation Scottish Renaissance of the nineteen twenties and thirties. As Margery Palmer McCulloch has shown, the wider cultural climate of Anglo-American modernism was key to MacDiarmid’s conception of the interwar Scottish Renaissance. What was the effect on second-generation poet-translators as the modernist moment passed? Are the many translations undertaken by the younger poets who emerged in the course of the nineteen forties and fifties a faithful reflection of this cultural inheritance? To what extent are they indicative of a new set of priorities and international influences? The five principal translators discussed in this thesis are Douglas Young (1913-1973), Sydney Goodsir Smith (1915-1975), Robert Garioch (1909-1981), Tom Scott (1918-1995) and William J. Tait (1918-1992). Each is the subject of a chapter, in many cases providing the first or most extensive treatment of particular translations. While the pioneering work of John Corbett, Bill Findlay and J. Derrick McClure, among other scholars, has drawn attention to the long history of literary translation into Scots, this thesis is the first extended critical work to take the verse translations of the post-MacDiarmid makars as its subject. The nature and extent of MacDiarmid’s influence is considered throughout, as are the wider discourses around language and translation in twentieth-century Scottish poetry. Critical engagement with a number of key insights from theoretical translation studies helps to situate these writers’ work in its global context. This thesis also explores the ways in which the specific context of Scots translation allows scholars to complicate or expand upon theories of translation developed in other cultural situations (notably Lawrence Venuti’s writing on domestication and foreignisation). The five writers upon whom this thesis concentrates were all highly individual, occasionally idiosyncratic personalities. Young’s polyglot ingenuity finds a foil in Garioch’s sharp, humane wit. Goodsir Smith’s romantic ironising meets its match in Scott’s radical certainty of cause. Tait’s use of the Shetlandic tongue sets him apart. Nonetheless, despite the great variety of style, form and tone shown by each of these translators, this thesis demonstrates that there are meaningful links to be made between them and that they form a unified, coherent group in the wider landscape of twentieth-century Scottish poetry. On the linguistic level, each engaged to some extent in the composition of a ‘synthetic’ or ‘plastic’ language deriving partly from literary sources, partly from the spoken language around them. On a more fundamental level, each was committed to enriching this language through translation, within which a number of key areas of interest emerge. One of the most important of these key areas is Gaelic – especially the poetry of Sorley MacLean, which Young, Garioch and Goodsir Smith all translated into Scots. This is to some extent an act of solidarity on the part of these Scots poets, acknowledging a shared history of marginalisation as well as expressing shared hopes for the future. The same is true of Goodsir Smith’s translations from a number of Eastern European poets (and Edwin Morgan’s own versions, slightly later in the century). The translation of verse drama by poets is another key theme sustained throughout the thesis, with Garioch and Young attempting to fill what they perceived as a gap in the Scots tradition through translation from other languages (another aspect of these writers’ legacy continued by Morgan). Beyond this, all of the writers discussed in this thesis translated extensively from European poetries from Ancient Greece to twentieth-century France. Their reasons for doing so were various, but a certain cosmopolitan idealism figures highly among them. So too does a desire to see Scotland interact with other European nations, thus escaping the potentially narrowing influence of post-war British culture. This thesis addresses the legacy of these writers’ translations, which, it argues, continue to exercise a perceptible influence on the course of poetry in Scotland. This work constitutes a significant contribution to a much-needed wider critical re-assessment of this pivotal period in modern Scottish writing, offering a fresh perspective on the formal and linguistic merits of these poets’ verse translations. Drawing upon frequently obscure book, pamphlet and periodical sources, as well as unpublished manuscripts in the National Library of Scotland and the Shetland Archives, this thesis breaks new ground in its investigation of the role of Scots verse translation in the second-generation Scottish Renaissance.