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From July 2002 to March 2004 the Michelson Interferometer for Passive Atmospheric Sounding (MIPAS) aboard the European Space Agency´s Environmental Satellite (Envisat) measured nearly continuously mid infrared limb radiance spectra. These measurements are utilised to retrieve the global distribution of the chlorofluorocarbon CFC-11 by applying a new fast forward model for Envisat MIPAS and an accompanying optimal estimation retrieval processor. A detailed analysis shows that the total retrieval errors of the individual CFC-11 volume mixing ratios are typically below 10% in the altitude range 10 to 25 km and that the systematic components dominate. Contribution of a priori information to the retrieval results are less than 5 to 10% and the vertical resolution of the observations is about 3 to 4 km in the same vertical range. The data are successfully validated by comparison with several other space experiments, an air-borne in-situ instrument, measurements from ground-based networks, and independent Envisat MIPAS analyses. The retrieval results from 425 000 Envisat MIPAS limb scans are compiled to provide a new climatological data set of CFC-11. The climatology shows significantly lower CFC-11 abundances in the lower stratosphere compared with the Reference Atmospheres for MIPAS (RAMstan V3.1) climatology. Depending on the atmospheric conditions the differences between the climatologies are up to 30 to 110 ppt (45 to 150%) at 19 to 27 km altitude. Additionally, time series of CFC-11 mean abundance and variability for five latitudinal bands are presented. The observed CFC-11 distributions can be explained by the residual mean circulation and large-scale eddy-transports in the upper troposphere and lower stratosphere. The new CFC-11 data set is well suited for further scientific studies.
Global distributions of profiles of sulphur hexafluoride (SF6) have been retrieved from limb emission spectra recorded by the Michelson Interferometer for Passive Atmospheric Sounding (MIPAS) on Envisat covering the period September 2002 to March 2004. Individual SF6 profiles have a precision of 0.5 pptv below 25 km altitude and a vertical resolution of 4–6 km up to 35 km altitude. These data have been validated versus in situ observations obtained during balloon flights of a cryogenic whole-air sampler. For the tropical troposphere a trend of 0.230±0.008 pptv/yr has been derived from the MIPAS data, which is in excellent agreement with the trend from ground-based flask and in situ measurements from the National Oceanic and Atmospheric Administration Earth System Research Laboratory, Global Monitoring Division. For the data set currently available, based on at least three days of data per month, monthly 5° latitude mean values have a 1 o standard error of 1%. From the global SF6 distributions, global daily and monthly distributions of the apparent mean age of air are inferred by application of the tropical tropospheric trend derived from MIPAS data. The inferred mean ages are provided for the full globe up to 90° N/S, and have a 1 o standard error of 0.25 yr. They range between 0 (near the tropical tropopause) and 7 years (except for situations of mesospheric intrusions) and agree well with earlier observations. The seasonal variation of the mean age of stratospheric air indicates episodes of severe intrusion of mesospheric air during each Northern and Southern polar winter observed, long-lasting remnants of old, subsided polar winter air over the spring and summer poles, and a rather short period of mixing with midlatitude air and/or upward transport during fall in October/November (NH) and April/May (SH), respectively, with small latitudinal gradients, immediately before the new polar vortex starts to form. The mean age distributions further confirm that SF6 is destroyed in the mesosphere to a considerable degree. Model calculations with the Karlsruhe simulation model of the middle atmosphere (KASIMA) chemical transport model agree well with observed global distributions of the mean age only if the SF6 sink reactions in the mesosphere are included in the model.
This study presents an evaluation of a pulse height condensation particle counter (PH-CPC) and an expansion condensation particle counter (E-CPC) in terms of measuring ambient and laboratory-generated molecular and ion clusters. Ambient molecular cluster concentrations were measured with both instruments as they were deployed in conjunction with an ion spectrometer and other aerosol instruments in Hyytiälä, Finland at the SMEAR II station between 1 March and 30 June 2007. The observed cluster concentrations varied and ranged from some thousands to 100 000 cm -3. Both instruments showed similar (within a factor of ~5) concentrations. An average size of the detected clusters was approximately 1.8 nm. As the atmospheric measurement of sub 2-nm particles and molecular clusters is a challenging task, we conclude that most likely we were unable to detect the smallest clusters. Nevertheless, the reported concentrations are the best estimates to date for minimum cluster concentrations in a boreal forest environment.
We report the first atmospheric observations of the Chlorofluorocarbons (CFCs) trifluorochloroethene, 3-chloropentafluoropropene and 4,4-dichlorohexafluoro-1-butene by means of Gas Chromatography with Electron Capture and Mass Spectrometric detection (GC-ECD-MS) in air samples taken at the Taunus Observatory operated by the University of Frankfurt (Main) and the Jungfraujoch High Altitude Research Station in Switzerland. These substances belong to a class of CFCs containing a double bond and are suspected to originate from the production and thermal degradation of widely used fluoropolymers like polychlorotrifluoroethene (PCTFE). Their atmospheric lifetimes are expected to be rather short. A quantitative calibration could only be derived for trifluorochloroethene but not for the other species by now. Thus, we use a relative sensitivity method to get a first indication of the observed atmospheric abundances. Identification was possible because of an air plume containing high concentrations of these substances. We suggest that the abundances found on this occasion originated from a local source. However, we have also observed the novel CFCs in air masses representative of background conditions, though with much lower concentrations. These species and some of their degradation products are toxic and could also be relevant for stratospheric and tropospheric ozone depletion.
Assessment of ecologically relevant hydrological change in China due to water use and reservoirs
(2008)
As China’s economy booms, increasing water use has significantly affected hydro-geomorphic processes and thus the ecology of surface waters. A large variety of hydrological changes arising from human activities such as reservoir construction and management, water abstraction, water diversion and agricultural land expansion have been sustained throughout China. Using the global scale hydrological and water use model WaterGAP, natural and anthropogenically altered flow conditions are calculated, taking into account flow alterations due to human water consumption and 580 large reservoirs. The impacts resulting from water consumption and reservoirs have been analyzed separately. A modified “Indicators of Hydrologic Alteration” approach is used to describe the human pressures on aquatic ecosystems due to anthropogenic alterations in river flow regimes. The changes in long-term average river discharge, average monthly mean discharge and coefficients of variation of monthly river discharges under natural and impacted conditions are compared and analyzed. The indicators show very significant alterations of natural river flow regimes in a large part of northern China and only minor alterations in most of southern China. The detected large alterations in long-term average river discharge, the seasonality of flows and the inter-annual variability in the northern half of China are very likely to have caused significant ecological impacts.
Transcriptional activation involves the ordered recruitment of coactivators via direct interactions between distinct binding domains and recognition motifs. The p160/SRC/NCoA coactivator family comprises three members (NCoA-1, -2 and -3), which are organized in multiprotein coactivator complexes. We had identified the PAS-B domain of NCoA-1 as an LXXLL motif binding domain. Here we show that NCoA family members are able to interact with other full-length NCoA proteins via their PAS-B domain and they specifically interact with the CBP-interaction domain (CID/AD1) of NCoA-1. Peptide competition, binding experiments and mutagenesis of LXXLL motifs point at distinct binding motif specificities of the NCoA PAS-B domains. NMR studies of different NCoA-1-PAS-B/LXXLL peptide complexes revealed similar although not identical binding sites for the CID/AD1 and STAT6 transactivation domain LXXLL motifs. In mechanistic studies, we found that overexpression of the PAS-B domain is able to disturb the binding of NCoA-1 to CBP in cells and that a CID/AD1 peptide competes with STAT6 for NCoA-1 in vitro. Moreover, the expression of an endogenous androgen receptor target gene is affected by the overexpression of the NCoA-1 or NCoA-3 PAS-B domains. Our study discloses a new, complementary mechanism for the current model of coactivator recruitment to target gene promoters.
In a combined NMR/MD study, the temperature-dependent changes in the conformation of two members of the RNA YNMG-tetraloop motif (cUUCGg and uCACGg) have been investigated at temperatures of 298, 317 and 325 K. The two members have considerable different thermal stability and biological functions. In order to address these differences, the combined NMR/MD study was performed. The large temperature range represents a challenge for both, NMR relaxation analysis (consistent choice of effective bond length and CSA parameter) and all-atom MD simulation with explicit solvent (necessity to rescale the temperature). A convincing agreement of experiment and theory is found. Employing a principle component analysis of the MD trajectories, the conformational distribution of both hairpins at various temperatures is investigated. The ground state conformation and dynamics of the two tetraloops are indeed found to be very similar. Furthermore, both systems are initially destabilized by a loss of the stacking interactions between the first and the third nucleobase in the loop region. While the global fold is still preserved, this initiation of unfolding is already observed at 317 K for the uCACGg hairpin but at a significantly higher temperature for the cUUCGg hairpin.
In Western Europe pedunculate oak (Quercus robur L.) is the forest tree with the highest number of phytophagous insect species (Yela & Lawton 1997). One of these, the green oak leaf roller Tortrix viridana L. is an oligophagous herbivorous moth with a host range limited to the genus Quercus (Hunter 1990, Du Merle 1999). During outbreaks, T. viridana often leads to defoliation of oaks in spring. The abundance of T. viridana is subject to the population size fluctuations typical for herbivorous insects, where periods of small population sizes (latent periods) alternate with periods of high population sizes (outbreak) (e.g. Schütte 1957, Horstmann 1984). Apart from many experimental studies on population dynamics of the moth (e.g. Hunter 1990, Du Merle 1999, Ivashov & al. 2002) so far little attention has been paid to the genetic variation within the species as an important aspect of the genetics of this host-parasite interaction. Simchuk & al. (1999) found changes in the heterozygosity level of different isozyme loci during outbreaks in T. viridana and molecular markers for T. viridana have been developed for analyses of genetic variation within and among populations (Schroeder & Scholz 2005). But, investigations of genetic variation within and among populations of forest pest species are important to predict future pest outbreaks. So far the processes outbreaks based on are not entirely clarified, however it is known that migration plays a major role. Using molecular markers investigations of the genetic variation are possible and offer the opportunity to analyse distribution events. In this paper first results are presented concerning the genetic variation of the green oak leaf roller at three geographic scales: (1) among trees within a population, (2) among populations at a small spatial scale of about 150 km and (3) among populations at a broader geographic scale up to 3000 km. Furthermore results of the genetic variation of oaks at the small spatial scale are represented.
Since the late 1990s, the oak processionary moth, Thaumetopoea processionea (L.), has been occurring at high population densities in eastern Austria. Particularly, infestations in areas of human settlement have created increasing interest in this insect due to health problems caused by the urticating hairs of the larvae. New methods for biological control are desirable. Like essentially all forest Lepidoptera, T. processionea is host for entomopathogenic microsporidia. These obligatory parasitic protists have been evaluated as biocontrol agents against an other oak pest, Lymantria dispar (Weiser & Novotny, 1987; Jeffords & al., 1988). Life history traits of T. processionea make this insect an even more promising target for the use of microsporidia. The larvae are highly gregarious and stay together in nests made of larval silk for resting periods and molting. Microsporidia utilize several pathways for horizontal transmission that would be aided by these features: spores can be released after host death from cadavers as well as from living larvae via silk or feces. Additionally, many microsporidia are vertically transmitted (summarized in Maddox & al., 1998). In this project, T. processionea larvae from various regions in eastern Austria were screened for the natural occurrence of microsporidia. One isolate, Endoreticulatus sp., was further studied and mass produced in a laboratory host, L. dispar, that is easy to rear and does not pose a health hazard for people working with the insects. An inoculative release was attempted on isolated trees infested with T. processionea.
Cantharidin, which is mainly found in blister beetles (Coleoptera: Meloidae), is one of the most intensively studied natural products of insect (Dettner, 1997; McCormick & Carrel, 1987). The involvement of cantharidin in courtship behaviour has been already confirmed for certain canthariphilous insects (Eisner & al. 1996a,b; Frenzel & Dettner 1994; Frenzel & al. 1992; Schütz & Dettner, 1992; Hemp & al. 1999). The function and intrinsinc role of cantharidin in the courtship behaviour of Meloids has been never fully established. McCormick & Carrel (1987) only suggested that cantharidin might be used by female meloids when selecting a mate at close range. Pinto (1974, 1975) was, in fact, the first to consider male cuticular pores as being involved in the courtship behaviour of species from the genus Linsleya and Tegrodera (Meloidae). Based on morphology and chemical analyses of Cyaneolytta sp. (Coleoptera: Meloidae), we have hereby provided some further evidences that cantharidin may act as an infochemical in courtship behaviour of meloid beetles.
Sexual deception of male bees is one of the most remarkable mechanisms of pollination (Ackermann 1986, Proctor & al. 1996). Flowers of the orchid genus Ophrys mimic females of their pollinator species, usually bees and wasps, to attract males, which try to copulate with the flowers. During this so-called “pseudocopulation” the male removes the pollinia and transfers them to another flower to ensure pollination. Apart from visual and tactile cues, floral scent was shown to be most important for eliciting mating behaviour in males (Kullenberg 1961, Schiestl & al. 1999, Ayasse & al. 2003). Pollination in Ophrys is highly specific and usually each Ophrys species attracts only one pollinator species (Paulus & Gack 1990). The high degree of specialization provides the means of reproductive isolation between the intercrossable Ophrys-species (Ehrendorfer 1980). The complex odour-bouquets released by the flowers are species-specific and often consist of more than 100 different chemical compounds (Borg-Karlson & al. 1985, Ayasse 2006). Speciation in Ophrys-orchids may be brought about by changes in the pollinator attracting floral scent. The attraction of a new pollinator may act as a pre-zygotic isolation barrier (Stebbins 1970, Paulus & Gack 1990, Soliva & al. 2001). We investigated three sympatrically occuring Ophrys-species on Sardinia. O. chestermanii and O. normanii are endemic and are both pollinated by males of the bumblebee B. vestalis. O. tenthredinifera is pollinated by Eucera nigrilabris. There are different opinions concerning the taxonomic status of O. normanii. It has been described as an actual hybrid between O. chestermanii and O. tenthredinifera (Wood 1983). Paulus & Gack (1995) suggested that it is an own species, that either has developed from a hybrid between O. chestermanii and O. normanii or that has evolved by radiation from O. tenthredinifera. By conducting behavioural-tests with B. vestalis males, performing gas chromatographic analyses and electrophysiological studies we wanted to identify pollinator attracting scent and to clarify the taxonomic status of O. normanii.
The orchid genus Epipactis is represented by 25 species in Europe (Richards 1982). Epipactis helleborine (L.) Crantz is the most common and widely distributed species of the genus (Wiefelspütz 1970), and is a prime example for wasp-flowers, because it is mainly pollinated by social wasps (Hymenoptera: Vespidae), like Vespula vulgaris and V. germanica (Müller 1873). Darwin (1888) already noticed that E. helleborine is almost exclusively ignored by bees and bumblebees, an observation that was confirmed in recent investigations (Keppert 2001). The flowers of E. helleborine show morphological, physiological and phenological adaptations to the visit and the pollination by Vespidae (Keppert 2001). They possess a reddish-brown or dirty purple coloration of the inflorescence (Keppert 2001), have relatively small, mostly bulbous blossoms with a broad entrance and bulbous widened, nectar-rich juice holders (Müller 1873, 1881; Schremmer 1962). Although there is much reported about wasp-pollinated flowers there is little known about the signals that are responsible for the attraction of wasps. Wiefelspütz (1970) proclaimed the statement that only the visual stimulus is responsible for the wasp attraction. Recently studies, however, assumed that odour is involved in the wasp attraction (Keppert 2001). Hölzler (2003) showed that the main attraction of the wasp-flower Epipactis for pollinators is its olfactory stimulus. It remains an unanswered question why E. helleborine flowers almost exclusively attracts social wasps, as opposed to bees and bumblebees. In this study we analysed the role of floral volatiles which are responsible for the specific attraction of social wasps. We supposed a mimicry-system in E. helleborine for the specific attraction of pollinators for the following reasons. So-called “green leaf volatiles” (GLVs) are emitted by plants while herbivorous insects, for example caterpillars, feed on them. GLVs thereby attract predators or parasitoids of the herbivorous insects (Dicke & Sabelis 1988; Turlings & al. 1990, 1995; Dicke & Vet 1999). Among the GLVs so far identified in former studies there are aldehydes, compounds that were also found in flower extracts of E. helleborine (Hölzler 2003). Therefore, we postulated that E. helleborine flowers produce GLVs in order to attract prey hunting social wasps for pollination. We performed bioassays and analysed flower odour gained to headspace-sampling using gas chromatography (GC), mass spectrometry (GC-MS) and gas chromatography coupled with electrophysiological analysis (GC-EAD) to investigate the hypothesis that E. helleborine flowers mimic “green leaf volatiles” (GLVs) to attract their pollinators.
Solitary bees are important pollinators of flowers. Besides nectar they collect pollen at flowers mainly to provide their larvae with food. Many bee species collect pollen only on a few closely related plant species (oligolecty) (Müller & al. 1997). Little is known about the visual and olfactorial signals they use for host-plant finding (Wcislo & Cane 1996). However, bees can olfactorily distinguish between different pollen species (von Frisch 1923), and a species-specific chemistry of pollen odour is known for some plant species (Bergström & al. 1995, Dobson & al. 1999). Further, it was shown that naïve oligolectic bees recognize their host-plant on the basis pollen volatiles (Dobson & Bergström 2000) and that flower-experienced bees could use pollen odours to assess pollen availability (Dobson & al. 1999). Besides scent, also visual cues are of relevance for host-plant finding, and bees orientate especially spectral contrasts. Biotests with dummy flowers revealed that colour contrast and not intensity and dominant wavelengths are influencing innate behavioural responses (Lunau 1990). Further it was shown that naïve bumblebees were most motivated to land on a flower when visual stimuli from the antheres are combined with olfactorial stimuli from the pollen (1992). We choose Osmia adunca P., which is highly specialized on Echium L., as a model to investigate the importance of floral cues for an oligolectic bee. Because bees learn to associate odours with reward more rapidly than visual cues (Menzel 1985), we hypothesize that scent plays a major role in attraction flower-experienced O. adunca females. We used gas chromatography to compare the scent of three Echium species with the scent of a closely related Anchusa species, and a spectrometer to compare the colour of the three Echium species. Additionally we conducted a biotest to determine the importance of visual and olfactorial signals of Echium for host-plant finding of experienced O. adunca females.
In agroecological research it has been appreciated only fairly recently that plant-insect interactions and other ecological processes depend on scales much larger than a single habitat (Wiens et al. 1997). Crop-pest interactions have mainly been studied on single pest species by focusing either on the impact of field parameters or on landscape structure but only rarely included both factors (Östman et al. 2001). Here we investigated how the abundances of three major insect pest species in oilseed rape (OSR) responded to field parameters and landscape characteristics at various spatial scales. Pest species considered in the current study include (i) ceutorhynchid stem weevils that lay eggs in leaf petioles or midribs of OSR plants while the larvae tunnel in the stems; (ii) pollen beetles that feed on pollen and destroy flower buds and (iii) brassica pod midge that lay eggs into OSR pods where the hatched larvae consume the seeds as well as tissue of the pod walls and cause the pods to split prematurely (Alford et al. 2003). Studying these different groups of pests is especially important because they attack different parts of the crop, use different habitats as overwintering sites and also differ in their mobility; with the exception of pollen beetles these pest species have never been studied in a landscape context. The specific objectives of this study were to determine (i) whether the major OSR pest species differ in their relation to field and landscape characteristics and (ii) at which spatial scales landscape variables are effective.
Agricultural intensification is a major threat to biological diversity worldwide. Land management activities enhancing landscape diversity are therefore regarded as a key strategy to halt species loss in cultural landscapes. Diverse and abundant communities of predatory arthropods, e.g. spiders (Araneae), have a high potential to suppress pest populations (Symondson et al. 2002) and could therefore contribute to allow reductions of pesticide use. Crop fields alone are usually not able to sustain diverse and individual-rich populations of predatory arthropods, because agricultural management results in disturbances and habitat deteriorations (harvest, soil cultivation, pesticide application) that kill or drive away large parts of the populations. Therefore semi-natural and perennial habitats in agricultural landscapes are considered to be of great importance for beneficial arthropods. On the one hand they offer refuge habitats in times when arable fields are hostile, e.g. fields with bare grounds during winter (Schmidt & Tscharntke 2005). On the other hand, viable populations of predatory arthropods in semi-natural habitats can serve as sources for (re-) colonisation of arable fields (Schmidt & Tscharntke 2005). Because of these exchanges between crop and non-crop areas it is important to include the surrounding landscape when investigating field-scale processes. We investigated the relations between spider assemblages in arable fields and the surrounding landscape in 29 fields of winter oilseed rape (OSR) in an agricultural landscape in eastern Austria. The objectives of this study were to estimate (1) how much spider assemblages in oilseed rape fields are influenced by the surrounding landscape, (2) the relative influence of landscape variables compared to field-scale variables and (3) at which spatial scales landscape variables are effective.
The lower wood-feeding Australian termite Mastotermes darwiniensis Froggatt (Fig. 1) is the only living member of the family Mastotermitidae. The complex symbiotic hindgut flora consists of protozoa (formerly named Archaezoa; Cleveland & Grimstone 1964; Brugerolle & al. 1994; Berchtold & König 1995; Fröhlich & König 1999a, b), bacteria (Berchtold & König 1996; Berchtold & al. 1999), archaea (Fröhlich & König 1999a, b) and yeasts (Prillinger & al. 1996; Schäfer & al. 1996). The digestive system of Mastotermes darwiniensis consists of the foregut with the crop and the gizzard, the midgut, and the hindgut (Noirot & Noirot-Timothée 1969; 1995). The hindgut consists of five segments (P1 – P5): the proctodeal segment, the enteric valve, the paunch, the colon and the rectum. The paunch is the main microbial fermentation chamber, but the colon also contains microorganisms. The paunch is subdivided into a dilated thin-walled region (P3a) and a thick walled more tubular region (P3b) (Fig. 1c). In the case of Mastotermes darwiniensis oxygen diffusion gradients could be detected up to 100 μm below the epithelium (Berchtold & al., 1999).
In our present-day landscape in Central Europe major parts of the xylobiontic especially of the saproxylic beetle fauna belong to the group of endangered species assemblages (Speight 1989, Geiser 1994). Oaks, in Central Europe mainly Quercus robur and Q. petraea, are well known for their large number of associated insect species and harbour the highest beetle diversity, especially for dead wood inhabiting species, of all broadleaved tree species in this region (e.g. Palm 1959). A characteristic species associated with oaks in its life-cycle is the endangered Great Capricorn Cerambyx cerdo. C. cerdo is one of the protected species explicitly named in the Habitats Directive of the European Union with the goal of maintaining existing populations and establishing long-term survival (Council of the European Communities 1992). The last remaining colonised areas of this longhorn beetle in Central Europe are well known for the enormous number of very rare xylobiontic beetle species. Thus, we are interested in the following research questions: 1) Are there typical species associated with C. cerdo? 2) If so, what kind of relationship do these associated species have to C. cerdo from a nature conservation point of view?
A high-precision pressure probe is described which allows non-invasive online-monitoring of the water relations of intact leaves. Real-time recording of the leaf water status occurred by data transfer to an Internet server. The leaf patch clamp pressure probe measures the attenuated pressure, Pp, of a leaf patch in response to a constant clamp pressure, Pclamp. Pp is sensed by a miniaturized silicone pressure sensor integrated into the device. The magnitude of Pp is dictated by the transfer function of the leaf, Tf, which is a function of leaf patch volume and ultimately of cell turgor pressure, Pc, as shown theoretically. The power function Tf=f(Pc) theoretically derived was experimentally confirmed by concomitant Pp and Pc measurements on intact leaflets of the liana Tetrastigma voinierianum under greenhouse conditions. Simultaneous Pp recordings on leaflets up to 10 m height above ground demonstrated that changes in Tf induced by Pc changes due to changes of microclimate and/or of the irrigation regime were sensitively reflected in corresponding changes of Pp. Analysis of the data show that transpirational water loss during the morning hours was associated with a transient rise in turgor pressure gradients within the leaflets. Subsequent recovery of turgescence during the afternoon was much faster than the preceding transpiration-induced water loss if the plants were well irrigated. Our data show the enormous potential of the leaf patch clamp pressure probe for leaf water studies including unravelling of the hydraulic communication between neighbouring leaves and over long distances within tall plants (trees).
Chloroplast function depends on the translocation of cytosolically synthesized precursor proteins into the organelle. The recognition and transfer of most precursor proteins across the outer membrane depend on a membrane inserted complex. Two receptor components of this complex, Toc34 and Toc159, are GTPases, which can be phosphorylated by kinases present in the hosting membrane. However, the physiological function of phosphorylation is not yet understood in detail. It is demonstrated that both receptors are phosphorylated within their G-domains. In vitro, the phosphorylation of Toc34 disrupts both homo- and heterodimerization of the G-domains as determined using a phospho-mimicking mutant. In endogenous membranes this mutation or phosphorylation of the wild-type receptor disturbs the association of Toc34, but not of Toc159 with the translocation pore. Therefore, phosphorylation serves as an inhibitor for the association of Toc34 with other components of the complex and phosphorylation can now be discussed as a mechanism to exchange different isoforms of Toc34 within this ensemble.
Gene trapping is used to introduce insertional mutations into genes of mouse embryonic stem cells (ESCs). It is performed with gene trap vectors that simultaneously mutate and report the expression of the endogenous gene at the site of insertion and provide a DNA tag for rapid identification of the disrupted gene. Gene traps have been employed worldwide to assemble libraries of mouse ESC lines harboring mutations in single genes, which can be used to make mutant mice. However, most of the employed gene trap vectors require gene expression for reporting a gene trap event and therefore genes that are poorly expressed may be under-represented in the existing libraries. To address this problem, we have developed a novel class of gene trap vectors that can induce gene expression at insertion sites, thereby bypassing the problem of intrinsic poor expression. We show here that the insertion of the osteopontin enhancer into several conventional gene trap vectors significantly increases the gene trapping efficiency in high-throughput screens and facilitates the recovery of poorly expressed genes.
Comparative studies suggest that at least some bird species have evolved mental skills similar to those found in humans and apes. This is indicated by feats such as tool use, episodic-like memory, and the ability to use one´s own experience in predicting the behavior of conspecifics. It is, however, not yet clear whether these skills are accompanied by an understanding of the self. In apes, self-directed behavior in response to a mirror has been taken as evidence of self-recognition. We investigated mirror-induced behavior in the magpie, a songbird species from the crow family. As in apes, some individuals behaved in front of the mirror as if they were testing behavioral contingencies. When provided with a mark, magpies showed spontaneous mark-directed behavior. Our findings provide the first evidence of mirror self-recognition in a non-mammalian species. They suggest that essential components of human self-recognition have evolved independently in different vertebrate classes with a separate evolutionary history.
Ribosome biogenesis in eukaryotes requires the participation of a large number of ribosome assembly factors. The highly conserved eukaryotic nucleolar protein Nep1 has an essential but unknown function in 18S rRNA processing and ribosome biogenesis. In Saccharomyces cerevisiae the malfunction of a temperature-sensitive Nep1 protein (nep1-1ts) was suppressed by the addition of S-adenosylmethionine (SAM). This suggests the participation of Nep1 in a methyltransferase reaction during ribosome biogenesis. In addition, yeast Nep1 binds to a 6-nt RNA-binding motif also found in 18S rRNA and facilitates the incorporation of ribosomal protein Rps19 during the formation of pre-ribosomes. Here, we present the X-ray structure of the Nep1 homolog from the archaebacterium Methanocaldococcus jannaschii in its free form (2.2 Å resolution) and bound to the S-adenosylmethionine analog S-adenosylhomocysteine (SAH, 2.15 Å resolution) and the antibiotic and general methyltransferase inhibitor sinefungin (2.25 Å resolution). The structure reveals a fold which is very similar to the conserved core fold of the SPOUT-class methyltransferases but contains a novel extension of this common core fold. SAH and sinefungin bind to Nep1 at a preformed binding site that is topologically equivalent to the cofactor-binding site in other SPOUT-class methyltransferases. Therefore, our structures together with previous genetic data suggest that Nep1 is a genuine rRNA methyltransferase.
Bency Eichorn learns in kollel and, on the side, has been researching about various segulos. For his wedding he authored a book, Simchas Zion, discussing the segulah of keeping the afikomom from year-to-year. The post below is a small part of a much larger project on this segulah and has been adapted for the blog.
Market uptake of pegylated interferons for the treatment of hepatitis C in Europe : meeting abstract
(2008)
Introduction and Objectives Hepatitis C virus (HCV) infection is a leading cause of chronic liver disease with life threatening sequelae such as end-stage liver cirrhosis and liver cancer. It is estimated that the infection annually causes about 86,000 deaths, 1.2 million disability adjusted life years (DALYs), and ¼ of the liver transplants in the WHO European region. Presently, only antiviral drugs can prevent the progression to severe liver disease. Pegylated interferons combined with ribavirin are considered as current state-of-the-art treatment. Objective of this investigation was to assess the market uptake of these drugs across Europe in order to find out whether there is unequal access to optimised therapy. Material and Methods We used IMS launch and sales data (April 2000 to December 2005) for peginterferons and ribavirin for 21 countries of the WHO European region. Market uptake was investigated by comparing the development of country-specific sales rates. For market access analysis, we converted sales figures into numbers of treated patients and related those to country-specific hepatitis C prevalence. To convert sales figures into patient figures, the amount of active pharmaceutical ingredients (API) sold was divided by average total patient doses (ATPD), derived by a probability tree-based calculation algorithm accounting for genotype distribution, early stopping rules, body weight, unscheduled treatment stops and dose reductions Ntotal=APIPegIFNalpha-2a/ATPDPegIFNalpha-2a+APIPegIFN&alpha-2b/ATPDPegIFNalpha-2b For more concise result presentation the 21 included countries were aggregated into four categories: 1. EU founding members (1957): Belgium, France, Germany, Italy and Netherlands; 2. Countries joining EU before 2000: Austria (1995), Denmark (1973), Finland (1995), Greece (1981), Republic of Ireland (1973), Spain (1986), Sweden and UK (1973) 3. Countries joining EU after 2000: Czech Republic (2004), Hungary (2004), Poland (2004) and Romania (2007); 4. EU non-member states: Norway, Russia, Switzerland and Turkey. Results Market launch and market uptake of the investigated drugs differed considerably across countries. The earliest, most rapid and highest increases in sales rates were observed in the EU founding member states, followed by countries that joined the EU before 2000, countries that joined the EU after 2000, and EU non-member states. Most new EU member states showed a noticeable increase in sales after joining the EU. Market access analysis yielded that until end of 2005, about 308 000 patients were treated with peginterferon in the 21 countries. Treatment rates differed across Europe. The number of patients ever treated with peginterferon per 100 prevalent cases ranged from 16 in France to less than one in Romania, Poland, Greece and Russia. Discussion Peginterferon market uptake and prevalence adjusted treatment rates were found to vary considerably across 21 countries in the WHO European region suggesting unequal access to optimised therapy. Poor market access was especially common in low-resource countries. Besides budget restrictions, national surveillance and prevention policy should be considered as explanations for market access variation. Although our results allowed for the ranking of countries in order of market access, no final conclusions on over- or undertreatment can be drawn, because the number of patients who really require antiviral treatment is unknown. Further research based on pan-European decision models is recommended to determine the fraction of not yet successfully treated but treatable patients among those ever diagnosed with HCV. ...
Many new gene copies emerged by gene duplication in hominoids, but little is known with respect to their functional evolution. Glutamate dehydrogenase (GLUD) is an enzyme central to the glutamate and energy metabolism of the cell. In addition to the single, GLUD-encoding gene present in all mammals (GLUD1), humans and apes acquired a second GLUD gene (GLUD2) through retroduplication of GLUD1, which codes for an enzyme with unique, potentially brain-adapted properties. Here we show that whereas the GLUD1 parental protein localizes to mitochondria and the cytoplasm, GLUD2 is specifically targeted to mitochondria. Using evolutionary analysis and resurrected ancestral protein variants, we demonstrate that the enhanced mitochondrial targeting specificity of GLUD2 is due to a single positively selected glutamic acid-to-lysine substitution, which was fixed in the N-terminal mitochondrial targeting sequence (MTS) of GLUD2 soon after the duplication event in the hominoid ancestor ~18–25 million years ago. This MTS substitution arose in parallel with two crucial adaptive amino acid changes in the enzyme and likely contributed to the functional adaptation of GLUD2 to the glutamate metabolism of the hominoid brain and other tissues. We suggest that rapid, selectively driven subcellular adaptation, as exemplified by GLUD2, represents a common route underlying the emergence of new gene functions.
C2-symmetric bisamidines : chiral Brønsted bases catalysing the Diels-Alder reaction of anthrones
(2008)
C2-symmetric bisamidines 8 have been tested as chiral Brønsted bases in the Diels- Alder reaction of anthrones and N-substituted maleimides. High yields of cycloadducts and significant asymmetric inductions up to 76% ee are accessible. The proposed mechanism involves proton transfer between anthrone and bisamidine, association of the resulting ions and finally a cycloaddition step stereoselectively controlled by the chiral ion pair.
Oscillatory activity in human electro- or magnetoencephalogram has been related to cortical stimulus representations and their modulation by cognitive processes. Whereas previous work has focused on gamma-band activity (GBA) during attention or maintenance of representations, there is little evidence for GBA reflecting individual stimulus representations. The present study aimed at identifying stimulus-specific GBA components during auditory spatial short-term memory. A total of 28 adults were assigned to 1 of 2 groups who were presented with only right- or left-lateralized sounds, respectively. In each group, 2 sample stimuli were used which differed in their lateralization angles (15° or 45°) with respect to the midsagittal plane. Statistical probability mapping served to identify spectral amplitude differences between 15° versus 45° stimuli. Distinct GBA components were found for each sample stimulus in different sensors over parieto-occipital cortex contralateral to the side of stimulation peaking during the middle 200–300 ms of the delay phase. The differentiation between "preferred" and "nonpreferred" stimuli during the final 100 ms of the delay phase correlated with task performance. These findings suggest that the observed GBA components reflect the activity of distinct networks tuned to spatial sound features which contribute to the maintenance of task-relevant information in short-term memory.
Cytotoxic T-lymphocytes play an important role in the protection against viral infections, which they detect through the recognition of virus-derived peptides, presented in the context of MHC class I molecules at the surface of the infected cell. The transporter associated with antigen processing (TAP) plays an essential role in MHC class I–restricted antigen presentation, as TAP imports peptides into the ER, where peptide loading of MHC class I molecules takes place. In this study, the UL49.5 proteins of the varicelloviruses bovine herpesvirus 1 (BHV-1), pseudorabies virus (PRV), and equine herpesvirus 1 and 4 (EHV-1 and EHV-4) are characterized as members of a novel class of viral immune evasion proteins. These UL49.5 proteins interfere with MHC class I antigen presentation by blocking the supply of antigenic peptides through inhibition of TAP. BHV-1, PRV, and EHV-1 recombinant viruses lacking UL49.5 no longer interfere with peptide transport. Combined with the observation that the individually expressed UL49.5 proteins block TAP as well, these data indicate that UL49.5 is the viral factor that is both necessary and sufficient to abolish TAP function during productive infection by these viruses. The mechanisms through which the UL49.5 proteins of BHV-1, PRV, EHV-1, and EHV-4 block TAP exhibit surprising diversity. BHV-1 UL49.5 targets TAP for proteasomal degradation, whereas EHV-1 and EHV-4 UL49.5 interfere with the binding of ATP to TAP. In contrast, TAP stability and ATP recruitment are not affected by PRV UL49.5, although it has the capacity to arrest the peptide transporter in a translocation-incompetent state, a property shared with the BHV-1 and EHV-1 UL49.5. Taken together, these results classify the UL49.5 gene products of BHV-1, PRV, EHV-1, and EHV-4 as members of a novel family of viral immune evasion proteins, inhibiting TAP through a variety of mechanisms.
The degradation of the poly(A) tail is crucial for posttranscriptional gene regulation and for quality control of mRNA. Poly(A)-specific ribonuclease (PARN) is one of the major mammalian 3’ specific exo-ribonucleases involved in the degradation of the mRNA poly(A) tail, and it is also involved in the regulation of translation in early embryonic development. The interaction between PARN and the m7GpppG cap of mRNA plays a key role in stimulating the rate of deadenylation. Here we report the solution structures of the cap-binding domain of mouse PARN with and without the m7GpppG cap analog. The structure of the cap-binding domain adopts the RNA recognition motif (RRM) with a characteristic a-helical extension at its C-terminus, which covers the b-sheet surface (hereafter referred to as PARN RRM). In the complex structure of PARN RRM with the cap analog, the base of the N7-methyl guanosine (m7G) of the cap analog stacks with the solvent-exposed aromatic side chain of the distinctive tryptophan residue 468, located at the C-terminal end of the second b-strand. These unique structural features in PARN RRM reveal a novel cap-binding mode, which is distinct from the nucleotide recognition mode of the canonical RRM domains.
We performed a bioinformatical analysis of protein export elements (PEXEL) in the putative proteome of the malaria parasite Plasmodium falciparum. A protein family-specific conservation of physicochemical residue profiles was found for PEXEL-flanking sequence regions. We demonstrate that the family members can be clustered based on the flanking regions only and display characteristic hydrophobicity patterns. This raises the possibility that the flanking regions may contain additional information for a family-specific role of PEXEL. We further show that signal peptide cleavage results in a positional alignment of PEXEL from both proteins with, and without, a signal peptide.
While the adaptor SKAP-55 mediates LFA-1 adhesion on T-cells, it is not known whether the adaptor regulates other aspects of signaling. SKAP-55 could potentially act as a node to coordinate the modulation of adhesion with downstream signaling. In this regard, the GTPase p21ras and the extracellular signal-regulated kinase (ERK) pathway play central roles in T-cell function. In this study, we report that SKAP-55 has opposing effects on adhesion and the activation of the p21ras -ERK pathway in T-cells. SKAP-55 deficient primary T-cells showed a defect in LFA-1 adhesion concurrent with the hyper-activation of the ERK pathway relative to wild-type cells. RNAi knock down (KD) of SKAP-55 in T-cell lines also showed an increase in p21ras activation, while over-expression of SKAP-55 inhibited activation of ERK and its transcriptional target ELK. Three observations implicated the p21ras activating exchange factor RasGRP1 in the process. Firstly, SKAP-55 bound to RasGRP1 via its C-terminus, while secondly, the loss of binding abrogated SKAP-55 inhibition of ERK and ELK activation. Thirdly, SKAP-55−/− primary T-cells showed an increased presence of RasGRP1 in the trans-Golgi network (TGN) following TCR activation, the site where p21ras becomes activated. Our findings indicate that SKAP-55 has a dual role in regulating p21ras-ERK pathway via RasGRP1, as a possible mechanism to restrict activation during T-cell adhesion.
Diabrotica virgifera virgifera LeConte, in its original North American habitat also known as western corn rootworm beetle, actively continues its expansion to new territories and uses Homo sapiens as its prime vector. It took only 15 years to spread to and occupy the southeastern and central parts of Europe, so far with the exception of Denmark where it has not been documented as of 2007. Economic thresholds have been reached and surpassed only in Southeast European countries like Slovakia, Hungary, Serbia, Eastern Croatia, Romania and Northern Italy. But both, the area affected and the severity of symptoms are increasing. Model calculations by a number of authors (Baufeld & Enzian, 2005 a and b; Hongmei Li & al. 2006, CLIMEX model) indicate a definitive propensity of D. v. virgifera to expand its currently occupied territory to regions with moderate temperatures and Zea mays cultivation. East Africa and Eastern Asia are included in the list of potential candidates for future inadvertent introduction. In most discussions it is tacitly and erroneously assumed that Z. mays is the only or the only important host of D. v. virgifera. Our recent observations in Eastern Slovenia on the oil pumpkin Cucurbita pepo indicate, however, that this simplifying assumption is notlonger strictly valid. It has to be modified in light of new evidence. Here, we report a few field experiments conducted in August of 2006 clarifying the host status of C. pepo in a European country.
The codling moth, Cydia pomonella (Lep., Tortricidae), is a significant pest of orchard crops such as apple and pear in Southern Germany, and can cause severe economic damage to apple crops. Due to resistance to conventional pesticides and the growing market for organic fruit, Cydia pomonella Granulovirus (CpGV) has been used to control C. pomonella in Germany for over 10 years. Recently, populations exhibiting resistance to CpGV have been reported. In this study, we have used amplified fragment length polymorphism (AFLP) markers to estimate genetic variations between eight different C. pomonella populations, which were obtained from different locations exhibiting varying levels of resistance to CpGV. Three different AFLP primer combinations generated a total of 194 AFLP fragments, ranging from 57.84 to 424.11 bp, with an average of 59.23 amplified fragments per primer combination. The total number of segregating fragments ranged from 181 to 115 and resulted in a high loci polymorphism of 100% in most cases, except for two populations, where it was found to be 88.1% and 93.3%. An analysis of genetic variation based on the obtained AFLP markers resulted in high gene diversity (Hj) values, ranging between 0.2884 to 0.3508. Hj values also indicated a loss in gene diversity within a population over time. The Wright Fixation Index (FST) values indicated a low to moderate genetic differentiation in the populations. The cluster analysis (UPGMA), based on genetic distance values, showed that the majority of C. pomonella populations from different locations were clearly distributed into distinct groups and showed a large genetic variability.
The bee fauna of Taiwan was studied intensively in the first half of last century and was based in large parts on the extensive material collected by Hans Sauter between 1902 and 1914. Subsequent studies on bees of Taiwan have only been sporadic. Within a cooperation between the above mentioned institutions the bee fauna was reinvestigated. It was shown how insufficiently the bee fauna of Taiwan had been investigated so far, in particular, the higher mountain regions. Now about 150 species of bees, belonging to 32 different genera, are known from Taiwan, ten of which have been described or recognized as new for science by the recent cooperation.
Taxonomic, systematic, and biogeography knowledge on the Palaearctic species of Pristaulacus Kieffer 1900 is summarized. Twenty-one valid species are recognized. The most important morphological characters taken into consideration are: shape, cuticular sculpture, and pubescence of head; index length/width of antennomeres; shape, sculpture and cuticular processes of mesosoma, especially of pronotum and mesonotum; number and shape of teeth on claw; shape and sculpture of metasoma; ovipositor length compared with wing and antenna length; and colour pattern (e.g., the dark spots on fore wing, and the colour of hind tarsus). Several characters of the genital capsule of the male were proved to be very useful for species identification, e.g., the shape of the paramere, volsella, cuspis, and digitus. Based on analysis of twenty-five morphological characters, eight species groups are recognized. The critical revision of the chorological data, including many new records, introduced relevant changes of the geographical distribution pattern of most species. Twelve species are restricted to the western part of the Palaearctic Region and eight species are restricted to its eastern part; only one species, P. gibbator, has a wider distribution, including both western and eastern parts of the Palaearctics.
Rising atmospheric CO2 is regarded as the main driver of global warming (Crowley, 2000). While temperature changes directly affect plants and animals (Root et al., 2003; Parmesan, 2006), the effects of CO2 on herbivores are mediated through changes in nutrient quality. Elevated concentrations of atmospheric CO2 are likely to increase photosynthetic activity and thus provide more C-based compounds which may alter plant chemical profiles and plant–herbivore–natural enemy interactions. There are several scenarios how insects will react when confronted with a different food quality. A nutrient poor diet, induced by nitrogen dilution, may result in compensatory feeding with either no adverse effects on insect performance or with negative effects on insect growth due to low digestibility of plant structural compounds (e.g. lignin) or toxic effects of secondary metabolites (e.g. tannins). Here we present data from on-tree feeding trials with larvae of the generalist herbivore Lymantria dispar and one of its natural enemies, the hymenopteran endoparasitoid Glyptapanteles liparidis, studied in 2005. The experiments were conducted at the Swiss free-air CO2 enrichment (FACE) site near Basel, in an approximately 80-100-yr-old, mixed-species forest. The data link changes in foliar chemistry of three tree species (Quercus petraea, Fagus sylvatica, Carpinus betulus) exposed to 540 ppm CO2 with herbivore and parasitoid performance.
The impact of European integration on the German system of pharmaceutical product authorization
(2008)
The European Union has evolved since 1965 into an influential political player in the regulation of pharmaceutical safety standards. The objective of establishing a single European market for pharmaceuticals makes it necessary for member-states to adopt uniform safety standards and marketing authorization procedures. This article investigates the impact of the European integration process on the German marketing authorization system for pharmaceuticals. The analysis shows that the main focal points and objectives of European regulation of pharmaceutical safety have shifted since 1965. The initial phase saw the introduction of uniform European safety standards as a result of which Germany was obliged to undertake “catch-up” modernization. From the mid-1970s, these standards were extended and specified in greater detail. Since the mid-1990s, a process of reorientation has been under way. The formation of the European Agency for the Evaluation of Medicinal Products (EMEA) and the growing importance of the European authorization procedure, combined with intensified global competition on pharmaceutical markets, are exerting indirect pressure for EU member-states to adjust their medicines policies. Consequently, over the past few years Germany has been engaged in a competition-oriented reorganization of its pharmaceutical product authorization system the outcome of which will be to give higher priority to economic interests.
We study the responses of residential property and equity prices, inflation and economic activity to monetary policy shocks in 17 countries, using data spanning 1986-2006, using single-country VARs and panel VARs in which we distinguish between groups of countries depending on their financial systems. The effect of monetary policy on property prices is about three times as large as its impact on GDP. Using monetary policy to guard against financial instability by offsetting asset-price movements thus has sizable effects on economic activity. While the financial structure influences the impact of policy on asset prices, its importance appears limited.
This paper explores the role of trade integration—or openness—for monetary policy transmission in a medium-scale New Keynesian model. Allowing for strategic complementarities in price-setting, we highlight a new dimension of the exchange rate channel by which monetary policy directly impacts domestic inflation. Although the strength of this effect increases with economic openness, it also requires that import prices respond to exchange rate changes. In this case domestic producers find it optimal to adjust their prices to exchange rate changes which alter the domestic currency price of their foreign competitors. We pin down key parameters of the model by matching impulse responses obtained from a vector autoregression on U.S. time series relative to an aggregate of industrialized countries. While we find evidence for strong complementarities, exchange rate pass-through is limited. Openness has therefore little bearing on monetary transmission in the estimated model.
Bayesian learning provides the core concept of processing noisy information. In standard Bayesian frameworks, assessing the price impact of information requires perfect knowledge of news’ precision. In practice, however, precision is rarely dis- closed. Therefore, we extend standard Bayesian learning, suggesting traders infer news’ precision from magnitudes of surprises and from external sources. We show that interactions of the different precision signals may result in highly nonlinear price responses. Empirical tests based on intra-day T-bond futures price reactions to employment releases confirm the model’s predictions and show that the effects are statistically and economically significant.
The popular Nelson-Siegel (1987) yield curve is routinely fit to cross sections of intra-country bond yields, and Diebold and Li (2006) have recently proposed a dynamized version. In this paper we extend Diebold-Li to a global context, modeling a potentially large set of country yield curves in a framework that allows for both global and country-specific factors. In an empirical analysis of term structures of government bond yields for the Germany, Japan, the U.K. and the U.S., we find that global yield factors do indeed exist and are economically important, generally explaining significant fractions of country yield curve dynamics, with interesting differences across countries.
Measuring financial asset return and volatilty spillovers, with application to global equity markets
(2008)
We provide a simple and intuitive measure of interdependence of asset returns and/or volatilities. In particular, we formulate and examine precise and separate measures of return spillovers and volatility spillovers. Our framework facilitates study of both non-crisis and crisis episodes, including trends and bursts in spillovers, and both turn out to be empirically important. In particular, in an analysis of nineteen global equity markets from the early 1990s to the present, we find striking evidence of divergent behavior in the dynamics of return spillovers vs. volatility spillovers: Return spillovers display a gently increasing trend but no bursts, whereas volatility spillovers display no trend but clear bursts.
Research with Keynesian-style models has emphasized the importance of the output gap for policies aimed at controlling inflation while declaring monetary aggregates largely irrelevant. Critics, however, have argued that these models need to be modified to account for observed money growth and inflation trends, and that monetary trends may serve as a useful cross-check for monetary policy. We identify an important source of monetary trends in form of persistent central bank misperceptions regarding potential output. Simulations with historical output gap estimates indicate that such misperceptions may induce persistent errors in monetary policy and sustained trends in money growth and inflation. If interest rate prescriptions derived from Keynesian-style models are augmented with a cross-check against money-based estimates of trend inflation, inflation control is improved substantially.
Religious conversion has become a dangerous social and individual problem. In Latin America, a traditional Catholic area, Protestant sects are successfully con-verting more and more Catholics into their own communities. Therefore the Pope demands a strict control of these activities. In India e.g., the Catholic hierarchy is critizising the Indian governments which have forbidden conversion on non-spiritual reasons. Hindu organizations have started even very successfully to re-convert Indian Christians particularly of Dalit and tribal background. Buddhists are very successful in indirect and even direct conversion of many Westerners. Wah-habit missionaries spread their Neo-Islam in the Muslim societies and get more and more even non-Muslim converts. We should add the forcible and sometimes ex-tremely cruel conversions the atheistic states had executed since the last century. ...
In the course of the ME period, HAVE began to encroach on territory previously held by BE. According to Rydén and Brorström (1987); Kytö (1997), this occurred especially in iterative and durational contexts, in the perfect infinitive and modal constructions. In Early Modern English (henceforth EModE), BE was increasingly restricted to the most common intransitives come and go, before disappearing entirely in the 18th and 19th centuries. This development raises a number of questions, both historical and theoretical. First, why did HAVE start spreading at the expense of BE in the first place? Second, why was the change conditioned by the factors mentioned by Rydén and Brorström (1987) and Kytö (1997)? Third, why did the change take on the order of 800 years to go to completion? Fourth, what implications does the change have for general theories of auxiliary selection? In this paper we’ll try to answer the first question by focusing on one the earliest clearly identifiable advance of HAVE onto BE territory – its first appearance with the verb come, which for a number of reasons is an ideal verb to focus on. First, come is by far the most common intransitive verb, so we get large enough numbers for statistical analysis. Second, clauses containing the past participle of come with a form of BE are unambiguous perfects: they cannot be passives, and they did not continue into modern English with a stative reading like he is gone. Third, and perhaps most importantly, come selected BE categorically in the early stages of English, so the first examples we find with HAVE are clear evidence for innovation. We will present evidence from a corpus study showing that the first spread of HAVE was due to a ban on auxiliary BE in certain types of counterfactual perfects, and will propose an account for that ban in terms of Iatridou’s (2000) Exclusion theory of counterfactuals.
Verbs, nouns and affixation
(2008)
What explains the rich patterns of deverbal nominalization? Why do some nouns have argument structure, while others do not? We seek a solution in which properties of deverbal nouns are composed from properties of verbs, properties of nouns, and properties of the morphemes that relate them. The theory of each plus the theory of howthey combine, should give the explanation. In exploring this, we investigate properties of two theories of nominalization. In one, the verb-like properties of deverbal nouns result from verbal syntactic structure (a “structural model”). See, for example, van Hout & Roeper 1998, Fu, Roeper and Borer 1993, 2001, to appear, Alexiadou 2001, to appear). According to the structural hypothesis, some nouns contain VPs and/or verbal functional layers. In the other theory, the verbal properties of deverbal nouns result from the event structure and argument structure of the DPs that they head. By “event structure” we mean a representation of the elements and structure of a linguistic event, not a representation of the world. We refer to this view as the “event model”. According to the event model hypothesis, all derived nouns are represented with the same syntactic structure, the difference lying in argument structure – which in turn is critically related to event structure, in the way sketched in Grimshaw (1990), Siloni (1997) among others. In pursuing these lines of analysis, and at least to some extent disentangling their properties, we reach the conclusion that, with respect to a core set of phenomena, the two theories are remarkably similar – specifically, they achieve success with the same problems, and must resort to the same stipulations to address the remaining issues that we discuss (although the stipulations are couched in different forms).
Class features as probes
(2008)
In this article, we adress (i) the form and (ii) the function on inflection class features in minimalist grammar. The empirical evidence comes from noun inflection systems involving fusional markers in German, Greek, and Russian. As for (i), we argue (based on instances of transparadigmatic syncretism) that class features are not privative; rather, class information must be decomposed into more abstract, binary features. Concerning (ii), we propose that class features qualify as the very device that brings about fusional infection: They are uninterpretable in syntax and actas probes on stems, with matching inflection markers as goels, and thus trigger morphological Agree operations that merge stem and inflection marker before syntax is reached.
Background: Polymorphisms within the insulin gene can influence insulin expression in the pancreas and especially in the thymus, where self-antigens are processed, shaping the T cell repertoire into selftolerance, a process that protects from ß-cell autoimmunity.
Methods: We investigated the role of the -2221Msp(C/T) and -23HphI(A/T) polymorphisms within the insulin gene in patients with a monoglandular autoimmune endocrine disease [patients with isolated type 1 diabetes (T1D, n = 317), Addison´s disease (AD, n = 107) or Hashimoto´s thyroiditis (HT, n = 61)], those with a polyglandular autoimmune syndrome type II (combination of T1D and/or AD with HT or GD, n = 62) as well as in healthy controls (HC, n = 275).
Results: T1D patients carried significantly more often the homozygous genotype "CC" -2221Msp(C/T) and "AA" -23HphI(A/T) polymorphisms than the HC (78.5% vs. 66.2%, p = 0.0027 and 75.4% vs. 52.4%, p = 3.7 × 10-8, respectively). The distribution of insulin gene polymorphisms did not show significant differences between patients with AD, HT, or APS-II and HC.
Conclusion: We demonstrate that the allele "C" of the -2221Msp(C/T) and "A" -23HphI(A/T) insulin gene polymorphisms confer susceptibility to T1D but not to isolated AD, HT or as a part of the APS-II.