Mount Carmel in the Commune: Promoting the Holy Land in Central Italy in the 13th and 14th Centuries

The Carmelite friars were the last of the major mendicant orders to be established in Italy. Originally an eremitical order, they arrived from the Holy Land in the 1240s, decades after other mendicant orders, such as the Franciscans and Dominicans, had constructed churches and cultivated patrons in the burgeoning urban centers of central Italy. In a religious market already saturated with friars, the Carmelites distinguished themselves by promoting their Holy Land provenance, eremitical values, and by developing an institutional history claiming to be descendants of the Old Testament prophet Elijah. By the end of the 13th century the order had constructed thriving churches and convents and leveraged itself into a prominent position in the religious community. My dissertation analyzes these early Carmelite churches and convents, as well as the friars’ interactions with patrons, civic governments, and the urban space they occupied. Through three primary case studies – the churches and convents of Pisa, Siena and Florence – I examine the Carmelites’ approach to art, architecture, and urban space as the order transformed its mission from one of solitary prayer to one of active ministry.

My central questions are these: To what degree did the Carmelites’ Holy Land provenance inform the art and architecture they created for their central Italian churches? And to what degree was their visual culture instead a reflection of the mendicant norms of the time?

I have sought to analyze the Carmelites at the institutional level, to determine how the order viewed itself and how it wanted its legacy to develop. I then seek to determine how and if the institutional model was utilized in the artistic and architectural production of the individual convents. The understanding of Carmelite art as a promotional tool for the identity of the order is not a new one, however my work is the first to consider deeply the order’s architectural aspirations. I also consider the order’s relationships with its de facto founding saint, the prophet Elijah, and its patron, the Virgin Mary, in a more comprehensive manner that situates the resultant visual culture into the contemporary theological and historical contexts.

Physical and Genetic Analysis of the CUP1 Tandem Array in the Yeast Saccharomyces cerevisiae

The genomes of many strains of baker’s yeast, Saccharomyces cerevisiae, contain multiple repeats of the copper-binding protein Cup1. Cup1 is a member of the metallothionein family, and is found in a tandem array on chromosome VIII. In this thesis, I describe studies that characterized these tandem arrays and their mechanism of formation across diverse strains of yeast. I show that CUP1 arrays are an illuminating model system for observing recombination in eukaryotes, and describe insights derived from these observations.

In our first study, we analyzed 101 natural isolates of S. cerevisiae in order to examine the diversity of CUP1-containing repeats across different strains. We identified five distinct classes of repeats that contain CUP1. We also showed that some strains have only a single copy of CUP1. By comparing the sequences of all the strains, we were able to elucidate the mechanism of formation of the CUP1 tandem arrays, which involved unequal non-homologous recombination events starting from a strain that had only a single CUP1 gene. Our observation of CUP1 repeat formation allows more general insights about the formation of tandem repeats from single-copy genes in eukaryotes, which is one of the most important mechanisms by which organisms evolve.

In our second study, we delved deeper into our mechanistic investigations by measuring the relative rates of inter-homolog and intra-/inter-sister chromatid recombination in CUP1 tandem arrays. We used a diploid strain that is heterozygous both for insertion of a selectable marker (URA3) inside the tandem array, and also for markers at either end of the array. The intra-/inter-sister chromatid recombination rate turned out to be more than ten-fold greater than the inter-homolog rate. Moreover, we found that loss of the proteins Rad51 and Rad52, which are required for most inter-homolog recombination, did not greatly reduce recombination in the CUP1 tandem repeats. Additionally, we investigated the effects of elevated copper levels on the rate of each type of recombination at the CUP1 locus. Both types of recombination are increased at high concentrations of copper (as is known to be the case for CUP1 transcription). Furthermore, the inter-homolog recombination rate at the CUP1 locus is higher than the average over the genome during mitosis, but is lower than the average during meiosis.

The research described in Chapter 2 is published in 2014.