Imitation and Invention in Song Learning in Nightingales (Luscinia megarhynchos B., Turdidae)

Song learning in oscine birds is often defined solely as a process of song imitation; nonetheless, not all songs produced by laboratory‐tutored birds are imitations of the model songs. If song learning were strictly a process of imitation, these non‐imitated songs (inventions) would be expected to contain no learned attributes. To determine whether species‐typical song attributes can be learned in the absence of imitation, we compared the imitations and inventions of laboratory‐tutored nightingales (Luscinia megarhynchos B.) with the songs of wild nightingales and the songs of laboratory‐reared, untutored nightingales. The species‐typical song attribute measured was stereotypy. We quantified stereotypy by four variables: (1) percentage of notes shared between two renditions of the same song type (2) difference in repetition rates of the same trill in two renditions of the same song type (3) acoustic similarity of the same note in two renditions of the same song type, and (4) acoustic similarity of the same note repeated within a trill. Wild songs and imitated songs were significantly more stereotypical than the songs of untutored birds for all measures. For the percentage of notes shared (1), and the acoustic similarity of notes in two renditions of the same song type (3), invented songs did not differ from the songs of untutored birds, suggesting that imitation is necessary for the acquisition of these song characteristics. However, invented songs were significantly more stereotypical than the songs of untutored birds for measures of stereotypy related to trills (2 and 4), and neither imitated nor invented songs differed significantly from the songs of wild birds in terms of trill rate stereotypy (2). Thus, it appears that the process of learning to produce trills may differ from the process of learning non‐repetitive song components: increased stereotypy in trills occurs even when the trills themselves are not copied from song models. Strict imitation does not fully account for the acquisition of some learned song attributes.     

Conditions Affecting Song Acquisition in Nightingales (Luscinia megarhynchos L.)

Hand raised nightingales were alternatively confronted with different series of songs which permitted labeling of particular learning situations and allowed detection of specific consequences of diverse learning conditions. We found that visual contact with a tutor affected both quality and quantity of acquired patterns. Without visual contact the birds acquired only song sections consisting of repeated vocal units', which proved to be relevant for species recognition. With visual contact the birds learnt every presented song type completely (→ song type sharing between tutor and learner). Furthermore, the birds developed additional song types individually: by distinct parameter variation, or by recombination of particular tutor song units (Fig. 1; Table 1). Functional aspects of this type of song acquisition and development are discussed.     

How songbirds deal with large amounts of serial information: retrieval rules suggest a hierarchical song memory

Many songbirds develop remarkably large vocal repertoires, and this has prompted questions about how birds are able to successfully learn and use the often enormous amounts of information encoded in their various signal patterns. We have studied these questions in nightingales (Luscinia megarhynchos), a species that performs more than 200 different types of songs (strophen), or more than 1000 phonetically different elements composing the songs. In particular, we investigated whether and how both song repertoires and song performance rules of nightingales were coded by auditory stimuli presented in serial learning experiments. Evaluation of singing episodes produced by our trained birds revealed that nightingales cope well with an exposure to even long strings of master song-types. They can readily acquire information encoded within and between the different master songs, and they memorize, for example, which master song-types they have experienced in the same learning context. Imitations of such song-types form distinct sequential associations that are termed “context groups”. Additionally, nightingales develop other song-type associations that are smaller in size and termed “package groups”. Package formation results from constraints of the acquisition mechanisms which obviously lead to a segmentation of auditorily perceived master song sequences. Further experimentation validated that the song memory of nightingales is organized in a hierarchical manner and holding information about “context groups” composed of packages, “package groups” composed of songs, and songs composed of song elements. The evidence suggests that implementation of such a hierarchical organization facilitates a quick retrieval of particular songs, and thereby provides an essential prerequisite for a functionally appropriate use of large vocal repertoire is in songbirds. Received: 4 October 1997 / Accepted in revised form: 26 August 1998     

DNP-Enhanced MAS NMR: A Tool to Snapshot Conformational Ensembles of α-Synuclein in Different States

Intrinsically disordered proteins dynamically sample a wide conformational space and therefore do not adopt a stable and defined three-dimensional conformation. The structural heterogeneity is related to their proper functioning in physiological processes. Knowledge of the conformational ensemble is crucial for a complete comprehension of this kind of proteins. We here present an approach that utilizes dynamic nuclear polarization-enhanced solid-state NMR spectroscopy of sparsely isotope-labeled proteins in frozen solution to take snapshots of the complete structural ensembles by exploiting the inhomogeneously broadened line-shapes. We investigated the intrinsically disordered protein α-synuclein (α-syn), which plays a key role in the etiology of Parkinson’s disease, in three different physiologically relevant states. For the free monomer in frozen solution we could see that the so-called “random coil conformation” consists of α-helical and β-sheet-like conformations, and that secondary chemical shifts of neighboring amino acids tend to be correlated, indicative of frequent formation of secondary structure elements. Based on these results, we could estimate the number of disordered regions in fibrillar α-syn as well as in α-syn bound to membranes in different protein-to-lipid ratios. Our approach thus provides quantitative information on the propensity to sample transient secondary structures in different functional states. Molecular dynamics simulations rationalize the results.     

Genetic Diversity in the Major Capsid L1 Protein of HPV-16 and HPV-18 in the Netherlands

ObjectivesIntratypic molecular variants of human papillomavirus (HPV) type-16 and -18 exist. In the Netherlands, a bivalent vaccine, composed of recombinant L1 proteins from HPV-16 and -18, is used to prevent cervical cancer since 2009. Long-term vaccination could lead to changes in HPV-16 and -18 virus population, thereby hampering vaccination strategies. We determined the genetic diversity of the L1 gene in HPV-16 and -18 viral strains circulating in the Netherlands at the start of vaccination in order to understand the baseline genetic diversity in the Dutch population.MethodsDNA sequences of the L1 gene were determined in HPV-16 (n = 241) and HPV-18 (n = 108) positive anogenital samples collected in 2009 and 2011 among Dutch 16- to 24-year old female and male attendees of the sexually transmitted infection (STI) clinics. Phylogenetic analysis was performed and sequences were compared to reference sequences HPV-16 ({"type":"entrez-nucleotide","attrs":{"text":"AF536179","term_id":"33330936"}}AF536179) and HPV-18 ({"type":"entrez-nucleotide","attrs":{"text":"X05015","term_id":"60975"}}X05015) using BioNumerics 7.1.ResultsFor HPV-16, ninety-five single nucleotide polymorphism (SNPs) were identified, twenty–seven (28%) were non-synonymous variations. For HPV-18, seventy-one SNPs were identified, twenty-nine (41%) were non-synonymous. The majority of the non-silent variations were located in sequences encoding alpha helix, beta sheet or surface loops, in particular in the immunodominant FG loop, and may influence the protein secondary structure and immune recognition.ConclusionsThis study provides unique pre-vaccination/baseline data on the genetic L1 diversity of HPV-16 and -18 viruses circulating in the Netherlands among adolescents and young adults.     

Two Channels of Charge Generation in Perylene Monoimide Solid‐State Dye‐Sensitized Solar Cells

The mechanism of charge generation in solid‐state dye‐sensitized solar cells using triarylamine‐substituted perylene monoimide dyes is studied by vis‐NIR broadband pump‐probe transient absorption spectroscopy. The experiments demonstrate that photoinduced electron injection into the TiO₂ can only occur in regions where Li⁺, from the commonly used Li‐TFSI additive salt, is present on the TiO₂ surface. Incomplete surface coverage by Li⁺ means that some dye excitons cannot inject their electron into the TiO₂. However it is observed in the solar cell structure that some of the dye excitons that cannot directly inject an electron still contribute to free charge generation by the previously hypothesized reductive quenching mechanism (hole transfer to the solid‐state hole transporter followed by electron injection from the dye anion into the TiO₂). The contribution of reductive quenching to the quantum efficiency of charge generation is significant, raising it from 68% to over 80%. Optimization of this reductive quenching pathway could be exploited to maintain high quantum efficiency in dyes with greater NIR absorption to achieve overall enhancements in device performance. It is demonstrated that broadband NIR transient spectroscopy is necessary to obtain population kinetics in these systems, as strong Stark effects distort the population kinetics in the visible region.