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Escherichia coli α-hemolysin (HlyA) is a pore-forming protein of 110 kDa belonging to the family of RTX toxins. A hydrophobic region between the amino acid residues 238 and 410 in the N-terminal half of HlyA has previously been suggested to form hydrophobic and/or amphipathic α-helices and has been shown to be important for hemolytic activity and pore formation in biological and artificial membranes. The structure of the HlyA transmembrane channel is, however, largely unknown. For further investigation of the channel structure, we deleted in HlyA different stretches of amino acids that could form amphipathic β-strands according to secondary structure predictions (residues 71–110, 158–167, 180–203, and 264–286). These deletions resulted in HlyA mutants with strongly reduced hemolytic activity. Lipid bilayer measurements demonstrated that HlyAΔ71–110 and HlyAΔ264–286 formed channels with much smaller single-channel conductance than wildtype HlyA, whereas their channel-forming activity was virtually as high as that of the wildtype toxin. HlyAΔ158–167 and HlyAΔ180–203 were unable to form defined channels in lipid bilayers. Calculations based on the single-channel data indicated that the channels generated by HlyAΔ71–110 and HlyAΔ264–286 had a smaller size (diameter about 1.4 to 1.8 nm) than wildtype HlyA channels (diameter about 2.0 to 2.6 nm), suggesting that in these mutants part of the channel-forming domain was removed. Osmotic protection experiments with erythrocytes confirmed that HlyA, HlyAΔ71–110, and HlyAΔ264–286 form defined transmembrane pores and suggested channel diameters that largely agreed with those estimated from the single-channel data. Taken together, these results suggest that the channel-forming domain of HlyA might contain β-strands, possibly in addition to α-helical structures.
Previously we reported modulation of endothelial prostacyclin and interleukin-8 production, cyclooxygenase-2 expression and vasorelaxation by oleoyl-lysophosphatidylcholine (LPC 18:1). In the present study, we examined the impact of this LPC on nitric oxide (NO) bioavailability in vascular endothelial EA.hy926 cells. Basal NO formation in these cells was decreased by LPC 18:1. This was accompanied with a partial disruption of the active endothelial nitric oxide synthase (eNOS)-dimer, leading to eNOS uncoupling and increased formation of reactive oxygen species (ROS). The LPC 18:1-induced ROS formation was attenuated by the superoxide scavenger Tiron, as well as by the pharmacological inhibitors of eNOS, NADPH oxidases, flavin-containing enzymes and superoxide dismutase (SOD). Intracellular ROS-formation was most prominent in mitochondria, less pronounced in cytosol and undetectable in endoplasmic reticulum. Importantly, Tiron completely prevented the LPC 18:1-induced decrease in NO bioavailability in EA.hy926 cells. The importance of the discovered findings for more in vivo like situations was analyzed by organ bath experiments in mouse aortic rings. LPC 18:1 attenuated the acetylcholine-induced, endothelium dependent vasorelaxation and massively decreased NO bioavailability. We conclude that LPC 18:1 induces eNOS uncoupling and unspecific superoxide production. This results in NO scavenging by ROS, a limited endothelial NO bioavailability and impaired vascular function.
Coevolution of viruses and their hosts represents a dynamic molecular battle between the immune system and viral factors that mediate immune evasion. After the abandonment of smallpox vaccination, cowpox virus infections are an emerging zoonotic health threat, especially for immunocompromised patients. Here we delineate the mechanistic basis of how cowpox viral CPXV012 interferes with MHC class I antigen processing. This type II membrane protein inhibits the coreTAP complex at the step after peptide binding and peptide-induced conformational change, in blocking ATP binding and hydrolysis. Distinct from other immune evasion mechanisms, TAP inhibition is mediated by a short ER-lumenal fragment of CPXV012, which results from a frameshift in the cowpox virus genome. Tethered to the ER membrane, this fragment mimics a high ER-lumenal peptide concentration, thus provoking a trans-inhibition of antigen translocation as supply for MHC I loading. These findings illuminate the evolution of viral immune modulators and the basis of a fine-balanced regulation of antigen processing.
Myotonic dystrophy type 1 (DM1) lacks non-invasive and easy to measure biomarkers, still largely relying on semi-quantitative tests for diagnostic and prognostic purposes. Muscle biopsies provide valuable data, but their use is limited by their invasiveness. microRNA (miRNAs) are small non-coding RNAs regulating gene expression that are also present in biological fluids and may serve as diseases biomarkers. Thus, we tested plasma miRNAs in the blood of 36 DM1 patients and 36 controls. First, a wide miRNA panel was profiled in a patient subset, followed by validation using all recruited subjects. We identified a signature of nine deregulated miRNAs in DM1 patients: eight miRNAs were increased (miR-133a, miR-193b, miR-191, miR-140-3p, miR-454, miR-574, miR-885-5p, miR-886-3p) and one (miR-27b) was decreased. Next, the levels of these miRNAs were used to calculate a "DM1-miRNAs score". We found that both miR-133a levels and DM1-miRNAs score discriminated DM1 from controls significantly and Receiver-Operator Characteristic curves displayed an area under the curve of 0.94 and 0.97, respectively. Interestingly, both miR-133a levels and DM1-miRNAs score displayed an inverse correlation with skeletal muscle strength and displayed higher values in more compromised patients. In conclusion, we identified a characteristic plasma miRNA signature of DM1. Although preliminary, this study indicates miRNAs as potential DM1 humoral biomarkers.
As part of a wider study of floodplain vegetation along the River Murray, we carried out a field survey in 1987–1988 involving collection of floristic and vegetation condition data from 335 sample plots (each 400 m2 in area), between Hume Dam and Lake Alexandrina (including the Edward-Wakool anabranch system). The floodplain vegetation is dominated by just two tree species, River Red Gum (Eucalyptus camaldulensis) and Black Box (Eucalyptus largiflorens), but the composition of the understorey shows much greater variation, both along the river and across the floodplain. A total of 499 plant species, subspecies and varieties were recorded from the survey plots, of which 316 (63%) were native and 183 (37%) were exotic. From analysis of the floristic data we identified 37 vegetation communities, not including the vegetation of permanent wetlands and cleared areas; 21 communities were distinguished in the River Red Gum zone, 12 communities in the Black Box zone, and 4 communities on rises within the floodplain. The main floristic division among the River Red Gum communities was between Riverine Plain/ Headwaters Zone communities of the upper Murray, and Mallee Zone communities of the lower Murray. Among the Black Box communities, the main floristic division was between inner floodplain communities and outer floodplain communities, with a further division between South Australian communities and New South Wales/Victorian communities. Major factors influencing the floristic patterns included flooding frequency/duration and soil salinity.
Eucalypt health declined steadily downstream and was poorest in the lower reaches of the river below the Darling Junction, where 60% of the trees were healthy, 18% unhealthy (at least 40% of the canopy dead) and 22% dead. By comparison, at the upper end of the river, above Tocumwal, 84% of the trees were healthy, 14% unhealthy and only 2% dead. Overall, the condition of Black Box trees (44% unhealthy or dead) was worse than the condition of River Red Gum trees (29% unhealthy or dead). Eucaypt regeneration was also poorest below the Darling Junction, with regenerants present in 77% of plots upstream of the Darling but only 35% of plots downstream. The findings of poor tree health and sparse regeneration below the Darling coincide with the most heavily regulated part of the Murray, where the reduction in flooding due to upstream storages and water extraction, mainly for irrigation, has been greatest. Black Box regeneration was much sparser overall than River Red Gum regeneration (regenerants present in 69% of River Red Gum plots but only 29% of Black Box plots). The poor condition of the Black Box trees, coupled with their poor regeneration, suggests that the long-term future of this species along the Murray, particularly below the Darling Junction, is tenuous, even though it is a dominant component of the vegetation.
The integrity of floodplain vegetation along the Murray has been severely compromised by weed invasion. Weeds were common throughout the survey area, but were most prevalent in the climatically wetter sections of the river, at both the downstream and upstream ends (below Mannum and above Tocumwal). The median number of exotic species per plot equalled or exceeded the number of native species in these sections of the river, whereas native species outnumbered exotic species in the other river sections. Communities of the River Red Gum zone and the rises were generally weedier than those of the Black Box zone. Exotic species strongly influenced the community classification. They were the dominant overstorey species in two communities (Salix species – willows) and outnumbered native species in the understorey of another eight communities. At the lower end of the river, below Mannum, the River Red Gums that originally fringed the river had been mostly replaced by dense thickets of the exotic Weeping Willow, Salix babylonica.
Other factors that have impacted on the floodplain vegetation at the plant community level have been river regulation and soil salinisation. Stabilization of water levels in the lower Murray by construction of a series of weirs and barrages has favoured the spread of some communities at the expense of others. The favoured communities, which are characterised by stands of Common Reed, Phragmites australis, along the water’s edge, appear to be artefacts of river regulation. Salinisation has resulted in death of eucalypts and replacement of eucalypt communities by shrub communities dominated by samphires, Tecticornia species. The samphire community characteristic of the most saline sites is one of the most species-poor communities on the Murray floodplain.
Logging along the Murray in New South Wales and Victoria has resulted in extensive replacement of old growth River Red Gum forests and woodlands by more even-aged stands of straight young trees. Following the recent conversion of many areas of State Forest along the Murray in both New South Wales and Victoria to National Park or Regional Park, and thus the cessation of logging in these areas, they should now revert gradually to mature forest and woodland.
This study is the first to describe broad scale floristic patterns in the floodplain vegetation of the Murray covering most of the length of the river. It also provides data on the vegetation condition in the 1980s, and provides a benchmark of conditions before the prolonged Millenium Drought in south-eastern Australia from 1997 to 2010. More recent surveys of vegetation condition have reported a severe decline in tree health during the drought. The results from our 1987–88 survey are important because they show that the deteriorating condition of the vegetation was already evident in the 1980s and although exacerbated by the subsequent drought, it is not just a consequence of that drought. The results are consistent with the conclusion that the primary cause of the decline has been river regulation and water extraction for irrigation. The rate of deterioration has increased rapidly since the 1980s because of the drought. There has been some improvement since the breaking of the drought, but the poor condition of the River Murray floodplain vegetation, an Australian icon, remains a major conservation and management issue. The impacts of climate change – higher temperatures and reduced rainfall – have compounded the problem and will continue to do so at an increasing rate. The results of the study support listing of the floodplain vegetation of the lower reaches of the river as a critically endangered ecological community.
The riparian rainforest on the streamside levees of the coastal floodplain of the Clarence River on the North Coast of New South Wales was cleared during the 1860s by small landholders seeking fertile land. Only three small remnants remain. Using a combination of historical species lists, corner trees from surveyors’ portion plans, habitat information and the NSW Scientific Committee’s (1999) determination for lowland rainforest on floodplain a conceptual model of the original distribution of rainforest suballiances on the levees of the Clarence River coastal floodplain is proposed.
Population size, and flowering and fruiting developmental stages in the Critically Endangered species Corunastylis sp. ‘Charmhaven’ (Family Orchidaceae, formerly included within genus Genoplesium), were investigated in the Warnervale-Charmhaven area over a three year period. Population size in 2012 was 11 plants, in 2013, 14 plants and in 2014 increased to 26 plants, with new plants appearing near the original plants. Proactive management, including mowing and erecting wire protective cages around groups of orchids was partly responsible for this increase in numbers because it prevented browsing by rabbits but only ten plants carried fruits to maturity in the 2014 season to produce seed. Despite an increase in numbers over a couple of years, a population of 26 individuals is very small and warrants maintaining the current conservation listing of Critically Endangered. The population began to flower between 15th and 29th February in 2012 and from 3rd to 14th March in 2013. However in 2014 flowering began on 11th February and extended to 19th March but it took until 17th June to reach the seed dispersal stage. 2014 involved two phases of flowering; whether climatic factors were responsible for this event is not known.
Acacia pendula, Weeping Myall, (family Fabaceae) is the most legislatively protected plant species in the New South Wales Hunter Valley. Under the NSW Threatened Species Conservation Act 1995 it is listed as an Endangered Population (in the Hunter Valley) and as a component of two Endangered Ecological Communities (one in the Hunter, one elsewhere in NSW); it is also listed as a Critically Endangered Ecological Community (in the Hunter Valley) on the Commonwealth Environment Protection and Biodiversity Conservation Act 1999 and listed as threatened in three other eastern Australian States.
To ascertain the likely original distribution of stands of Acacia pendula in the Hunter Valley, this paper examines the writings of early Australian explorers, herbarium and database records, and the species habitat attributes across NSW. None of the journals examined, including those of botanist/explorer Allan Cunningham (who originally collected Acacia pendula from the Lachlan River in 1817), Thomas Mitchell or Ludwig Leichhardt, make note of the species for the Hunter Valley. Several explorers do, however, record Acacia pendula regularly (>100 times) across other parts of NSW, Queensland, and South Australia.
Historical herbarium and database records show a paucity of records from the Hunter prior to the year 2000, after which a 37-fold increase in observations since 1951 is apparent. For the first 128 years of botanical exploration (1823 to 1951), there are no validated collections or records of Acacia pendula from the Hunter Valley. The single exception is a specimen collected by Cunningham from 1825 (lodged at Kew, UK), purported to be from ‘Hunters River’, but which is morphologically different to other collections of Acacia pendula from that time. There is some uncertainty over the origins of this specimen.
Analysis of habitats supporting Acacia pendula in NSW outside of the Hunter show them to differ significantly in geological age, soil type, rainfall and elevation from those in the Hunter.
Collectively, these findings provide a strong circumstantial case that Acacia pendula was absent from the Hunter at the time of European settlement; this has important implications for the conservation and management of Hunter stands. Rather than being a threatened species in the Hunter Valley, it is postulated that Acacia pendula has been intentionally and/or accidentally introduced to the region, and may now be imposing a new and emerging threat to the endangered grassy woodlands and forests there. There is now an urgent need for genetic studies to clarify the origins of the current Hunter Valley stands, and to define the taxonomic limits of Acacia pendula and its close relatives.
The availability of organelle genome sequences of bryophytes provides opportunity to mine this data. Therefore in this study microsatellites in chloroplast genome sequence of Pellia endiviifolia (Accession number: NC_019628), downloaded from the National Center for Biotechnology Information (NCBI) in fasta format, were identified. The sequence was mined with the help of MISA, a Perl script, to detect microsatellites. In total, 16 perfect microsatellites were identified in 120.546 kb sequence mined. An average length of 14.94 bp was calculated for mined microsatellites with a density of 1 SSR/7.09 kb. Depending on the repeat units, the length of microsatellites ranged from 12 to 18 bp. Tetranucleotides (7, 43.75%) were the most frequent repeat type, followed by mononucleotide (3, 18.75%) repeats. Dinucleotide, trinucleotide and pentanucleotide repeats were found with equal frequency (2, 12.5%). Interestingly, hexanucleotide repeats were completely absent in chloroplast genome of Pellia endiviifolia.
Coscinodon humilis was described by Milde from mica schist in the Passeiertal NE Merano (formerly southern Tyrolia in Austria, hence cited as Austria by Greven 1995. now Alto Adige in Italy). Limpricht (1890) regarded it as “verkümmerte Form von C. cribrosus”, and although Mönkemeyer (1927) still cited it, the species got forgotten by the time. Thus the species was no more mentioned by Corley et al (1981) in the European checklist and therefore no more included by Frey et al. (1995) in the German edition of the “Moos- und Farnpflanzen Europas”. Greven (1995) re-established the species in his treatment of Grimmia (and related genera) in Europe. Therefore Frey et al. (2006) included the species, which was, however, not keyed out. Finally Hill et al. (2006) listed it again in the new European checklist as a good species.
Myxobacteria are on order of Gram-negative, soil dwelling bacteria that feature an impressive number of properties: they can glide on solid surfaces by using two different motility motors, subsist by preying on other microorganisms, are often producers of multiple natural products, and upon adverse environmental conditions, they are able to form multicellular structures called “fruiting bodies”. The process, in which these macroscopically visible structures arise from independent single cells, has been the predominant subject of myxobacterial research for many decades. More precisely, researchers have strived for the discovery of genes, proteins and small molecules that act as signals, receivers or modulators of this complex process. In this regard, the species Myxococcus xanthus has evolved into the model organism due to its relatively simple and reliable handling in a laboratory environment. The research underlying this thesis focused on the identification and biosynthesis of lipids that may act as intercellular signaling molecules during the course of fruiting body formation of the myxobacterium Myxococcus xanthus as part of the “E-signal” system. In general, lipids containing branched-chain fatty acids with an uneven number of carbon atoms were found to be important players in this particular process. Nevertheless, their exact roles remain largely unknown as of this day. The first publication that is part of this thesis deals with an aspect that even strengthened the importance of role of iso-branched compounds in myxobacteria: myxobacterial metabolism is able to transform precursors of iso-lipids to isoprenoids. It addresses the question whether isoprenoids in general are important for fruiting body formation. Phenotypic analysis of mutants impaired in the biosynthesis of the central isoprenoid precursor 3-hydroxymethylglutaryl-Coenzyme A (3-HMG-CoA) from acetate and/or branched chain keto acids and their genetic and metabolic complementation clearly showed that isoprenoids are essential for fruiting body formation and confirmed that leucine derived isovalerate is an important source for isoprenoid precursors in myxobacteria. The second, and by far and away most tedious and sophisticated study, addressed the question as to how myxobacteria form fatty acid derived iso-branched ether lipids and to what extent they are important for fruiting body formation and sporulation. In a previous study, those unusual lipids were identified as specific biomarkers for myxobacterial development. No biochemical pathways to ether lipids specific for prokaryotes were known by then. In this study, a putative candidate gene that may be in involved in ether lipid biosynthesis was investigated. A combination of gene disruption and complementation experiments, phenotypic analysis and monitoring of ether lipid formation by means of GC-MS demonstrated its involvement in myxobacterial ether lipid biosynthesis and the importance of these lipids for the developmental process. Heterologous expression and biochemical testing of this gene together with in-silico sequence analysis and docking experiments confirmed the functions of its predicted domains. The discussion section provides an additional suggestion on how the ether bond formation is performed. Furthermore and most importantly, iso-branched ether lipids were found to be essential for sporulation but not for fruiting body formation. In summary, one or several molecules derived from an iso-branched alkylglycerol seem to play a role during sporulation in M. xanthus and a multidomain enzyme unique for myxobacteria is involved in their biosynthesis. The last manuscript addresses the complexity of lipid metabolism in myxobacteria. Prior to this work, there was limited knowledge about the exact composition of the myxobacterial lipidome and no method was available to monitor putative changes in the myxobacterial lipidome down to the single molecular species for studying lipid biosynthesis or regulation. An ultra-performance liquid chromatography coupled with mass spectrometry based method with electrospray ionization (UPLC-ESI-MS) utilizing standard equipment and a water/acetonitrile/isopropanol based eluent system proved to be geared for the construction of lipid profiles for wild type and mutant cells of M. xanthus and to show their differences. Fragmentation spectra based structure elucidation of lipid molecular species resulted in the identification of 99 molecular species comprising glycerophosphoethanolamines, glycerophosphoglycerols, glycerolipids, ceramides and ceramide phosphoinositols. The latter have never been described for any prokaryotes before. Three dimensional plots were created from the relative intensity differences of the single molecular ion species between the different samples to provide an efficient and versatile visualization of the data and enable the researcher to quickly detect differences.
In the present paper, we concentrate on (selected) Bantu and Nilotic bare-passive strategies and lay out the basis for a typology of transitive passive constructions in these languages. We argue that bare-passives constitute an optimal strategy to change prominence relations between arguments, in languages that strongly hold to the default mapping between the highest thematic role available and the grammatical subject (i.e. Spec,TP). The Nilotic and Bantu languages discussed here differ in their way of satisfying this default mapping. In particular, impersonal bare-passives satisfy it by resorting to an agentive place-holder (an indefinite subject marker) and realizing the logical agent as a lower thematic/semantic role (e.g. instrument or locative). Left-dislocation and so called 'subjectobject' reversal bare-passives realize the default matching between agent and subject in a more straightforward way, but locate the patient in a higher argument position within the inflectional domain (Spec,TopP). As argued in Hamlaoui and Makasso (2013) and Hamlaoui (2013), and in line with Noonan (1977), the present languages display a clauseinternal split between subjecthood (being the grammatical subject in Spec,TP) and topicality (being the subject of the predication, in an inflectional-domain internal Spec,TopP).
Inversion constructions in Bantu have been discussed from a variety of perspectives over the last decades. Well-known construction types include locative inversion and subject-object reversal, while more recently semantic locative inversion and instrument inversion have been described. Theoretical studies of Bantu inversion constructions have focused on different aspects of the construction, including the licensing and grammatical function, information structure properties and the formal characteristics of pre- and postverbal NPs. With respect to the status of preverbal NPs in inversion constructions, different analyses have probed the status of the NP as subject or topic, or, more recently, as the subject of a Predication relation. The present paper summarises and compares different analyses of the preverbal domain in inversion constructions and brings out empirical and conceptual similarities and differences. In addition, different analyses are related to comparative studies of Bantu inversion constructions, so as to probe how attested variation across Bantu relates to findings of different formal accounts. The paper aims to summarise current research on the preverbal domain in inversion constructions and to indicate directions for future work.
Nsong is a western Bantu language spoken in the neighbourhood of Kikwit (5°2'28"S 18°48'58"E, Kwilu District, Bandundu Province, DRC) and encoded as B85d in the New Updated Guthrie List (Maho 2009). To this B80 or Tiene-Yanzi group also belongs Mbuun, encoded as B87 by Guthrie (1971: 39) and spoken in the wider vicinity of Idiofa (4°57'35"S 19°35'40", Kwilu District, Bandundu Province, Democratic Republic of the Congo). Both languages are closely related. They share a high percentage of fundamental and other vocabulary as well as several rather atypical phonological innovations (Bostoen & Koni Muluwa 2014; Koni Muluwa 2014; Koni Muluwa & Bostoen 2012). Preliminary elicitation-based research on Mbuun has pointed out that the pre-verbal domain plays a crucial role in the marking of argument focus in Mbuun (Bostoen & Mundeke 2011, 2012). In this paper, we assess whether this is also the case in Nsong on the basis of a text corpus which the first author has been collecting, transcribing and annotating in 2013 and 2014 as part of an endangered language documentation project funded by the DoBeS program of the Volkswagen Foundation through a 3-year grant (2012-2015). More information on the project can be found on http://www.kwilubantu.ugent.be/. This Nsong text corpus exclusively consists of oral discourse and currently counts 48.022 tokens and 11.973 types. The team’s 2013 fieldwork aimed at documenting Nsong speech events in as many different cultural settings as possible. As a result, the corpus comprises different text genres, such as political speeches, historical traditions, folk music, tales, proverbs, hunting language, ceremonial language used during circumcision and twin rites, and popular biological knowledge. In line with previous research on Mbuun, we concentrate here on mono-clausal argument focus constructions, even if preliminary research has pointed out that bi-clausal focus structures are more common in the Nsong corpus.
The present paper aims at describing different pre-verbal focus strategies in Kisikongo (H16a), spoken in the vicinity of Mbanza Kongo, northern Angola. This western Bantu language is part of the Kikongo Language Cluster (KLC), stretching from southern Gabon to northern Angola, including Cabinda and parts of Congo-Brazzaville and Congo-Kinshasa. Kikongo exhibits a clause-internal pre-verbal argument focus position, which has rarely been reported in Bantu languages, except in Mbuun (B87) (Bostoen and Mundeke 2012) and Nsong (B85d) (Koni Muluwa and Bostoen, this volume), both spoken in the neighboring Kwilu region of the DRC. The more extensively studied eastern and southern Bantu languages generally have a post-verbal argument focus position (cf. Watters 1979, Morimoto 2000, Creissels 2004, Güldemann 2007, Buell 2009, van der Wal 2009, among others). In addition to this mono-clausal argument focus strategy, Kisikongo also relies on different bi-clausal constructions to focus arguments, i.e. cleft-constructions.
Nen and Nyokon are unique among the Bantu languages in allowing full nominal objects between the tense/aspect marker and the verb. Despite the fact that the two languages are neighbours and related they make different use of this positional option. In Nen the position is the default one for objects and the post-verbal position renders an object discrete and suitable for quantified objects and for contrast. In Nyokon the position before the verb is functionally equivalent to the one after the verb. The difference is related to the fact that Nyokon allows the preverbal object only in certain tenses whereas in Nen it is not restricted. But contrasted objects in Nyokon too appear after the verb. There is a construction in which both positions are filled with a constituent. This construction is modelled on a secondary predication construction.
Locative inversion in Cuwabo
(2014)
This paper proposes a detailed description of locative inversion (LI) constructions in Cuwabo, in terms of morphosyntactic properties and thematic restrictions. Of particular interest are the use of disjoint verb forms in LI, and the co-existence of formal and semantic LI, which challenges the widespread belief that the two constructions cannot be found in the same language.
This paper deals with left and right dislocation in Embɔsi, a Bantu language (C25) spoken in Congo-Brazzaville. The prosody of dislocation has gathered considerable attention, as it is particularly informative for the theories of the syntax-prosody mapping of Intonation Phrases (a.o. Selkirk, 2009, 2011; Downing, 2011). Concentrating on selected Bantu languages, Downing (2011) identifies two main phrasing patterns. She primarily distinguishes languages in which only right dislocated phrases display a lack of prosodic integration ("asymmetric" languages), from languages in which both left and right dislocations phrase separately ("symmetric" languages). Hiatus avoidance processes, boundary tones and register expansion/reduction indicate that Embɔsi displays a somewhat more intricate phrasing pattern. In this language, both left and right dislocated items sit outside of the Intonation Phrase formed by the core-clause, but only the latter form their own Intonation Phrase. We also discuss the prosody of multiple dislocations (i.e. with two dislocated arguments), which have not so far received all the attention they deserve. What we observe in Embɔsi is that either the two dislocated items phrase together and are not integrated to the core Intonation Phrase, or only the outermost dislocated element phrases separately.
Introduction
(2014)
Bantu languages have been at the heart of the research on the interaction between syntax, prosody and information structure. In these predominantly SVO languages, considerable attention has been devoted to postverbal phenomena. By addressing issues related to Subjects, Topics and Object-Verb word orders, the goal of the present papers is to deepen our understanding of the interaction of different grammatical components (syntax, phonology, semantics/pragmatics) both in individual languages and across the Bantu family. Each paper makes a valuable contribution to ongoing discussions on the preverbal domain.
It has long been observed that subjects cross-linguistically have topic properties: they are typically definite, referential and/or generic (Givón 1976). Bantu languages are said to illustrate this generalization: preverbal position for NPs is equated with both subject and topic status and postverbal position with focus (and non-subject). However, there is a growing body of work showing that preverbal subjects are not necessarily syntactically or semantically equivalent to topics. For example, Zerbian’s (2006) careful study of preverbal position in Northern Sotho shows that preverbal subjects meet few of the semantic tests for aboutness topics. The study of restrictions on preverbal subjects in Durban Zulu presented in this paper builds on Zerbian (2006) and Halpert (2012). In particular, we investigate the interpretational properties of preverbal indefinite subjects. These subjects show us that preverbal subjects carry a presupposition of existence. We explore an analysis connecting the "strong reading" of preverbal subjects with how high the verb moves in Zulu (following Tsai’s 2001 work on Mandarin).