Of mice, birds, and men: the mouse ultrasonic song system has some features similar to humans and song-learning birds.

Humans and song-learning birds communicate acoustically using learned vocalizations. The characteristic features of this social communication behavior include vocal control by forebrain motor areas, a direct cortical projection to brainstem vocal motor neurons, and dependence on auditory feedback to develop and maintain learned vocalizations. These features have so far not been found in closely related primate and avian species that do not learn vocalizations. Male mice produce courtship ultrasonic vocalizations with acoustic features similar to songs of song-learning birds. However, it is assumed that mice lack a forebrain system for vocal modification and that their ultrasonic vocalizations are innate. Here we investigated the mouse song system and discovered that it includes a motor cortex region active during singing, that projects directly to brainstem vocal motor neurons and is necessary for keeping song more stereotyped and on pitch. We also discovered that male mice depend on auditory feedback to maintain some ultrasonic song features, and that sub-strains with differences in their songs can match each other's pitch when cross-housed under competitive social conditions. We conclude that male mice have some limited vocal modification abilities with at least some neuroanatomical features thought to be unique to humans and song-learning birds. To explain our findings, we propose a continuum hypothesis of vocal learning.

Specialized motor-driven dusp1 expression in the song systems of multiple lineages of vocal learning birds.

Mechanisms for the evolution of convergent behavioral traits are largely unknown. Vocal learning is one such trait that evolved multiple times and is necessary in humans for the acquisition of spoken language. Among birds, vocal learning is evolved in songbirds, parrots, and hummingbirds. Each time similar forebrain song nuclei specialized for vocal learning and production have evolved. This finding led to the hypothesis that the behavioral and neuroanatomical convergences for vocal learning could be associated with molecular convergence. We previously found that the neural activity-induced gene dual specificity phosphatase 1 (dusp1) was up-regulated in non-vocal circuits, specifically in sensory-input neurons of the thalamus and telencephalon; however, dusp1 was not up-regulated in higher order sensory neurons or motor circuits. Here we show that song motor nuclei are an exception to this pattern. The song nuclei of species from all known vocal learning avian lineages showed motor-driven up-regulation of dusp1 expression induced by singing. There was no detectable motor-driven dusp1 expression throughout the rest of the forebrain after non-vocal motor performance. This pattern contrasts with expression of the commonly studied activity-induced gene egr1, which shows motor-driven expression in song nuclei induced by singing, but also motor-driven expression in adjacent brain regions after non-vocal motor behaviors. In the vocal non-learning avian species, we found no detectable vocalizing-driven dusp1 expression in the forebrain. These findings suggest that independent evolutions of neural systems for vocal learning were accompanied by selection for specialized motor-driven expression of the dusp1 gene in those circuits. This specialized expression of dusp1 could potentially lead to differential regulation of dusp1-modulated molecular cascades in vocal learning circuits.

CHASING THE SINGERS: THE TRANSITION OF LONG-SONG (URTYN DUU) IN POST-SOCIALIST MONGOLIA

Long-song (Urtyn duu) is a prominent Mongolian traditional folk song genre that survived throughout the socialist period (1921-1990) and throughout the political transformation of Mongolia from socialism to democratic capitalism after the Soviet Union was dismantled and terminated its aid to Mongolia in 1990. This dissertation, based on research conducted from 2006 to 2010, presents and investigates the traces of singers' stories and memories of their lives, songs, and singing, through the lens of the discourse on change and continuity in, and as, folk tradition. During the socialist period, this genre was first considered backward, and was then subtly transformed into an urban national style, with the formation of a boundary between professionalism and amateurism among long-song singers and with selective performance of certain songs and styles. This boundary was associated with politics and ideology and might be thought to have ended when the society entered its post-socialist period. However, the long-song genre continued to play a political role, with different kinds of political meaning one the one hand and only slight musical modification on the other. It was now used to present a more nostalgic and authentic new Mongolian identity in the post-socialist free market. Through my investigation, I argue that the historical transition of Mongolia encompassed not merely political or economic shifts, but also a deeper transformation that resulted in new cultural forms. Long-song provides a good case study of the complicated process of this cultural change.

Male mice song syntax depends on social contexts and influences female preferences.

In 2005, Holy and Guo advanced the idea that male mice produce ultrasonic vocalizations (USV) with some features similar to courtship songs of songbirds. Since then, studies showed that male mice emit USV songs in different contexts (sexual and other) and possess a multisyllabic repertoire. Debate still exists for and against plasticity in their vocalizations. But the use of a multisyllabic repertoire can increase potential flexibility and information, in how elements are organized and recombined, namely syntax. In many bird species, modulating song syntax has ethological relevance for sexual behavior and mate preferences. In this study we exposed adult male mice to different social contexts and developed a new approach of analyzing their USVs based on songbird syntax analysis. We found that male mice modify their syntax, including specific sequences, length of sequence, repertoire composition, and spectral features, according to stimulus and social context. Males emit longer and simpler syllables and sequences when singing to females, but more complex syllables and sequences in response to fresh female urine. Playback experiments show that the females prefer the complex songs over the simpler ones. We propose the complex songs are to lure females in, whereas the directed simpler sequences are used for direct courtship. These results suggest that although mice have a much more limited ability of song modification, they could still be used as animal models for understanding some vocal communication features that songbirds are used for.

Music, emotion, and autobiographical memory: they're playing your song.

Very long-term memory for popular music was investigated. Older and younger adults listened to 20-sec excerpts of popular songs drawn from across the 20th century. The subjects gave emotionality and preference ratings and tried to name the title, artist, and year of popularity for each excerpt. They also performed a cued memory test for the lyrics. The older adults' emotionality ratings were highest for songs from their youth; they remembered more about these songs, as well. However, the stimuli failed to cue many autobiographical memories of specific events. Further analyses revealed that the older adults were less likely than the younger adults to retrieve multiple attributes of a song together (i.e., title and artist) and that there was a significant positive correlation between emotion and memory, especially for the older adults. These results have implications for research on long-term memory, as well as on the relationship between emotion and memory.

BOULANGER, SATIE, AND DEBUSSY: THEIR SPHERES OF INFLUENCE ON FRENCH MELODIE AND AMERICAN ART SONG

This dissertation project explored the spheres of influence on art song by Nadia Boulanger, Erik Satie, and Claude Debussy within Boulangeries, Les Six, and Les Apaches. After World War I, American composers flocked to Paris to study with Boulanger. Boulanger gave her students the confidence to explore their native talents instead of mimicking foreign models. Works by Aaron Copland, Virgil Thomson, Theodore Chanler, John Duke, and Richard Hundley were included in the first dissertation recital on January 31, 2010: The Legacy of Nadia Boulanger: Her Influence on American Song Composers. Satie established a new modern French musical style, and was a catalyst for the formation of Les Six. Ned Rorem came to Paris, and had a close association with Les Six. Works by Satie, and three members of Les Six, Francis Poulenc, Arthur Honegger, Darius Milhaud; and Rorem were featured in the second recital on September 1, 2010: Satie, Selected Members of Les Six, and Rorem in Paris. Debussy was one of the most significant French composers in the late nineteenth century, predating Boulanger and Satie. Young composers exploring new directions were inspired by Debussy, forming the group Les Apaches. The final recital, April 7, 2011, featured works by Debussy and two members of Les Apaches, Maurice Ravel and Manuel de Falla: Debussy: A Catalyst for Les Apaches, Ravel and Falla. Falla‘s less well-known repertoire was presented. This dissertation showed the influence of these three major figures and that they embraced innovation in their own time, along with their followers. Recordings of these three performances may be obtained from the Michelle Smith Performing Arts Library in Clarice Smith Performing Arts Center at the University of Maryland, College Park.

THE ART SONG OF SOUTH AMERICA: AN EXPLORATION THROUGH PERFORMANCE

The repertoire included in this dissertation was presented over the course of three recitals, The Songs of Argentina, The Songs of Brazil, Chile and Venezuela, and The Songs of Perú and Colombia. Each recital was supplemented by written program notes and English translations of the Spanish, Portuguese and Quechua texts. The selections presented in this study was chosen in an effort to pair the works of internationally renowned composers like Argentine composers Alberto Ginastera and Carlos Guastavino, and Brazilian composer Heitor Villa-Lobos, with those of lesser-known composers, including Venezuelan composer Juan Bautista Plaza, Peruvian composers Edgar Valcárcel, Theodoro Valcárcel, and Rosa Mercedes Ayarza de Morales, and Colombian composer Jaime Léon. Each composer represents a milestone in the development of art song composition in South America. All three recitals were recorded and are available on compact discs in the Digital Repository at the University of Maryland (DRUM). This dissertation was completed in May, 2011.

PIANO AS A PARTNER IN TWENTIETH­ CENTURY ART SONG REPERTOIRE: A RECORDING PROJECT

The piano's role in art song repertoire has evolved from a modest one during its formative years to the versatility, complexity and creativity found in the twentieth-century. The art song repertoire of the twentieth century is vast and has secured the reputation for being the most diverse, innovative, illustrative, atmospheric and colorful in all of art song literature. Within this time period, there are compositions that reach back to the romantic works of nineteenth century, others which combine old and new traditions, and finally those which adopt new means and new ends. In choosing the material for this project, I have focused on compositions with uniquely challenging and unusual piano accompaniments in order to achieve a balance between well- known and rarely performed works, as well as those pieces that combine various languages and styles. Selections range from Claude Debussy, Richard Strauss, Sergey Rachmaninoff, Ralph Vaughan Williams, Roger Quilter, Francis Poulenc, Fernando Obradors, and Joaquin Rodrigo to composers such as Samuel Barber, Marc Blitzstein, Dominick Argento, William Bolcom, and John Duke, including arrangements of traditional spirituals by Harry T. Burleigh and Florence B. Price, all of which helped to establish the American Art Song. My objective is to trace the development of the twentieth-century art song from the late Romantic Period through nationalistitrends to works which show the influences of jazz and folk elements. The two CD's for this dissertation recording project are available on compact discs which can be found in the Digital Repository at the University of Maryland (DRUM). The performers were Daniel Armstrong, baritone, Giles Herman, baritone, Thomas Glenn, tenor, Valerie Yinzant, soprano, Aaron Odom, tenor, Jennifer Royal, soprano, Kenneth Harmon, tenor, Karen Sorenson, soprano and Maxim Ivanov, baritone.

Core and Shell Song Systems Unique to the Parrot Brain.

The ability to imitate complex sounds is rare, and among birds has been found only in parrots, songbirds, and hummingbirds. Parrots exhibit the most advanced vocal mimicry among non-human animals. A few studies have noted differences in connectivity, brain position and shape in the vocal learning systems of parrots relative to songbirds and hummingbirds. However, only one parrot species, the budgerigar, has been examined and no differences in the presence of song system structures were found with other avian vocal learners. Motivated by questions of whether there are important differences in the vocal systems of parrots relative to other vocal learners, we used specialized constitutive gene expression, singing-driven gene expression, and neural connectivity tracing experiments to further characterize the song system of budgerigars and/or other parrots. We found that the parrot brain uniquely contains a song system within a song system. The parrot "core" song system is similar to the song systems of songbirds and hummingbirds, whereas the "shell" song system is unique to parrots. The core with only rudimentary shell regions were found in the New Zealand kea, representing one of the only living species at a basal divergence with all other parrots, implying that parrots evolved vocal learning systems at least 29 million years ago. Relative size differences in the core and shell regions occur among species, which we suggest could be related to species differences in vocal and cognitive abilities.

SELECTED TWENTIETH-CENTURY BRITISH & AMERICAN ENGLISH SONG REPERTOIRE WRITTEN FOR SOPRANO

English language song (both British and American) is influenced by a variety of cultures, races, and musical forms and has produced a broad range of song repertoire. Like songs in all countries and throughout history, these songs can be classified into three categories: imitative songs, experimental songs, and songs of individuality. Music experimentation, necessary and welcome as it is, can hardly command broad international attention. Thus, the songs of this dissertation performance project are chosen from the first and third categories: imitative songs and individual songs in the composer's own unique style. This project concentrates its exploration on twentieth-century solo songs written in English. Although twentieth-century British & American composers also produced solos and chamber music in other languages, this dissertation focuses upon their English repertoire. This performance project consists of three programs: one British repertoire and two American. The first program titled An Evening of British Song examines twentieth-century British song written by Roger Quilter, Peter Warlock, William Walton, Benjamin Britten, Thomas F. Dunhill, Ivor Gurney, and Frank Bridge. It was presented on December 12, 2001, in Homer Ulrich Recital Hall with the collaborative pianist Meriel Owen. The second program titled An Evening of American Song I comprises music written by Dominick Argento, Samuel Barber, Ned Rorem, Leonard Bernstein, and Lee Hoiby. It was presented on October 23, 2002, in Joseph & Alma Gildenhorn Recital Hall with the collaborative pianist R. Timothy McReynolds. The third program titled An Evening of American Song II written by John Duke, John Corigliano, Charles Ives, Richard Hundley, Lori Laitman, Frederick Loewe, George Gershwin, and Jerome Kern was presented on December 18, 2003, again in Joseph & Alma Gildenhorn Recital Hall with the collaborative pianist R. Timothy McReynolds and the flutist Jessica Dunnavant. Each of these three dissertation recitals occurred at the University of Maryland in College Park and was recorded. These CD recordings are held by the Michelle Smith Performing Arts Library at the University of Maryland.

Flexible diel vertical migration behaviour of zooplankton in the Irish Sea

The diel vertical migration (DVM) of the whole plankton community was investigated in the central and coastal Irish Sea. Generally, more than 60% of the plankton did not perform significant DVM. A correlation analysis of the weighted mean depths of different organisms and their potential predators suggested relationships between two groups, Oithona spp., copepod nauplii and fish larvae, and between Pseudocalanus elongatus, Calanus spp. and chaetognaths. The organisms showing significant DVM were chaetognaths (Sagitta spp.), Calanus spp. and P. elongatus. Calanus spp. showed clear ontogenic variations in DVM, and along with P. elongatus demonstrated great flexibility both in the amplitude and direction of migration. P. elongatus did not migrate in the coastal area and Calanus spp. showed a clear reverse migration. The direction of migration appeared to be related to the vertical position of the chaetognaths in the water column during the day.