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    Tibet and the British Raj, 1904-47: The influence of the Indian political department officers.

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    Following Colonel Younghusband's Mission to Lhasa in 1903-04, officers selected by the Indian Political Department were stationed in Tibet under the command of the Political Officer Sikkim. This study examines aspects of the character, role and influence of these officers, whom I collectively term the 'Tibet cadre', and demonstrates that the cadre maintained a distinct collective identity and ethos, which was reflected in their approach to Anglo-Tibetan policies, and in the image of Tibet which resulted from the Anglo-Tibetan encounter. British India's northern frontier was the location for powerful imperial mythologies, such as the "Great Game", which were a part of cadre identity. Conditions on the frontier were believed to suit a particular type of individual, and officers of that type, capable of upholding British prestige while gaining an empathy with Tibet and Tibetans, were favoured for cadre service. A similar type of character was sought among the local intermediaries, the most successful of whom were given cadre postings. As frontiersmen following the traditions of Younghusband, their 'founding father', the cadre promoted 'forward' policies, designed to counter the perceived Russian threat to British India by extending British influence over the Himalayas. But Whitehall refused to support these policies to avoid damaging relations with China and other powers who regarded Tibet as part of China. The increased control exerted by central government over the imperial periphery in this period meant that, although the Tibet cadre did succeed in their primary aim of establishing British representation in Lhasa, they were unable to exert a dominant influence on policy-making either in Whitehall or in Lhasa. The cadre largely controlled the flow of information from Tibet, and they contributed a great deal to the construction of an image of Tibet, particularly through the books they wrote. But although individual officers such as Sir Charles Bell developed a deep understanding of Tibet, this did not fully emerge in the final image, which had passed through layers of censorship designed to ensure that the image served British interests

    Pleiotropy of FRIGIDA enhances the potential for multivariate adaptation.

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    An evolutionary response to selection requires genetic variation; however, even if it exists, then the genetic details of the variation can constrain adaptation. In the simplest case, unlinked loci and uncorrelated phenotypes respond directly to multivariate selection and permit unrestricted paths to adaptive peaks. By contrast, 'antagonistic' pleiotropic loci may constrain adaptation by affecting variation of many traits and limiting the direction of trait correlations to vectors that are not favoured by selection. However, certain pleiotropic configurations may improve the conditions for adaptive evolution. Here, we present evidence that the Arabidopsis thaliana gene FRI (FRIGIDA) exhibits 'adaptive' pleiotropy, producing trait correlations along an axis that results in two adaptive strategies. Derived, low expression FRI alleles confer a 'drought escape' strategy owing to fast growth, low water use efficiency and early flowering. By contrast, a dehydration avoidance strategy is conferred by the ancestral phenotype of late flowering, slow growth and efficient water use during photosynthesis. The dehydration avoidant phenotype was recovered when genotypes with null FRI alleles were transformed with functional alleles. Our findings indicate that the well-documented effects of FRI on phenology result from differences in physiology, not only a simple developmental switch
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