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Grindel, Switzerland

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The main town of the municipality is the village Bräuhof. Grundlsee has five neighboring communities, the communities Grünau im Almtal and Hinterstoder are located in the Upper Austrian district of Gmunden, all others in the Styrian Salzkammergut. The name was first mentioned in as Chrungilse. The spelling of the name varied over time, until finally enforced the current spelling: The name of Grundlsee is derived from the name of the lake.

However, instead of the Krungelsee this original form changed to today's Grundlsee. This was probably done by a folk etymological effect of the Middle High German grundel, grundelinc the gudgeon. The earliest evidence of human settlement activity in Grundlseer community area form Paleolithic finds in the Salzofen cave in the Dead Mountains. Charcoal remains of a Paleolithic hearth found there were dated to around 34, years ago.

Numerous relics from the Bronze Age and the Iron Age, as well as a Bronze Age settlement were found along the natural transport line of Koppentales in the neighboring community Bad Aussee. These findings can be explained in the context of Hallstatt only 20 km away, which was due to its archaeological significance eponymous for the early Iron Age BC.

Only the river name Traun from Celtic druna , the running one indicates Celtic settlement. As witnesses of the Roman rule in Noricum late Roman traces of settlement were discovered during excavations in Bad Aussee and the Altausseer Michlhallberg Sandling -Massiv. According to the previously retrieved finds, the settlement in Altaussee may have existed from the end of the 2nd century AD until the late 4th century.

It is believed there a Roman salt mining. The epoch of migration did not leave any traces in Ausseerland. The next population group that can be proved with certainty were the Slavs. Name-related traces of the Slavic settlement can be found in the entire Ausseerland place names and field names ending in -itz, -itsch, -isch , in Grundlsee eg, Stimitz , Toplitz , Zimitz.

Around , a strong influx of baju goods began. Important traces of the first contact of the Slavs with the Bavarians are again the place names. The earliest datable Eindeutschungen of place names in the Ausseerland come from the Old High German time before The name Grundlsee apud chrungilse was first mentioned on 2 August in a document.

Duke Ottokar IV was on this day at the Grundlsee and sealed there three documents. In order to fortify the new claim to power and to protect the nearby salt mines on the Sandling massif and the mule tracks, he built the small fortress Pflindsberg in neighboring Altaussee.

Philipp von Spanheim had to retire after the peace of furnace from and the castle and the Ausseerland were incorporated into in Styria. The previous history of Ausseerland is controversial. Most likely it was part of a county in the Ennstal under the Margraves of the Carinthian Mark in the 12th to 13th centuries.

The theory that the Ausseer area previously belonged to County Traunau, can not be proven. The plant developed into an administrative center with lower jurisdiction of the independent rule Pflindsberg, which was separated from the sovereign rule Grauscharn-Pürgg.

With the Reformation in the 16th century, the population of Ausseerland had become largely Protestant. From , a recatholization commission violently enforced the Counter-Reformation. From the larger military conflicts and social upheavals of the 16th and 17th centuries peasant wars, Thirty Years War the entire Ausseerland was spared. Reasons for the lack of social tensions were a relatively secure livelihood of the population and far-reaching social concessions on the part of the authorities.

The entire Salzkammergut was a closed territory, which had dedicated itself to a mono-economy. The sole source of income was salt production, to which all economic activity was coordinated. In Altaussee the salt was mined. Grundlsees role in this mono-economy was mainly the supply of brewing pans with the necessary firewood, which required salt extraction forestry.

In addition, the workers employed in this branch were mostly small part-time farmers who, together with their family members, produced a part of the essential products themselves. Due to the austerity and the severe climatic conditions, there were only relatively few full-time farmers in the entire region. During the Napoleonic wars, French troops marched through the Ausseerland several times between and In , therefore, the Koppen Pass and the Pötschen Pass were entrenched and fortified.

But fighting did not happen. In fortified the Pötschenpass again, this time with several gun positions, a powder magazine and two barracks.

The expected battles for the pass failed, as Napoleon's troops were decisively defeated in October in the Battle of Nations near Leipzig. At the beginning of the 19th century, the Salzkammergut was discovered as a summer resort.

The nearby Bad Ischl developed into a prominent health resort and was from the imperial summer residence. Due to the presence of the nobility in the region, the Ausseerland became more and more a center of attraction for the noble society.

Soon it attracted many artists and representatives of Viennese society to Grundlsee, which had become since by the abolition of the manorial system a political community. The Ausseer landscape attracted many painters. Between and , Archduke Johann had his chamber painters work in the Ausseerland, among others Jakob Gauermann, Matthäus Loder, Thomas Ender and Jakob and Rudolf Alt, where a variety of landscape paintings with Grundlseer motifs was created.

Another painter working in Grundlsee is Johann Matthias Ranftl, after whom the Ranftlmühle, built in , is named on the Stimitzbach. The Kronprinz-Rudolf-Bahn was opened in and the Ausseerland was finally infrastructurally well-developed for tourism. In tourist steamboat traffic was started in Grundlsee with the wooden steamboat Erzherzog Johann.

Among the artists and intellectuals who spent their summer vacation in Grundlsee, or settled there permanently, the industrialist and folklorist Konrad Mautner, the neurologist and founder of psychoanalysis Sigmund Freud and the conductor Herbert von Karajan are particularly noteworthy.

After the annexation of Austria to the German Reich in , the entire Ausseerland was incorporated into the administrative unit Oberdonau Upper Austria. The autonomy of the communities Bad Aussee, Grundlsee and Altaussee was dissolved and a mayor was established in Bad Aussee.

The municipal offices Grundlsee and Altaussees were henceforth field offices of Bad Aussee. The Ausseerland subsequently attracted numerous Nazi personalities. For example, three National Socialist Gauleiters regularly spent their holidays in the neighboring town of Altaussee: In , they had begun to set up a depot for art goods in the nearby salt mine Altaussee.

In August of the same year began the storage of art treasures from Austrian churches, monasteries and museums to protect them from bombing. Its association with the second leg-bearing segment in tardigrades suggests that the second trunk ganglion is a homologue of the arthropod tritocerebrum, whereas the first ganglion corresponds to the deutocerebrum.

We therefore conclude that the tardigrade brain consists of a single segmental region corresponding to the arthropod protocerebrum and, accordingly, that the tardigrade head is a non-composite, one-segmented structure. The composition of the arthropod head has been the subject of much controversy in the past and still remains one of the most contentious issues in the fields of comparative morphology and developmental biology today.

The typical arthropod head is a composite structure consisting of several segments, which might have been added successively in the course of arthropod evolution [3] — [5]. The homology of the cephalic segments has been clarified recently in arthropods and onychophorans [6] , [7] , but the contradictory data from tardigrades [8] — [10] currently do not allow an adequate comparison with these two animal groups.

The main problem in deciphering the organisation of the nervous system in tardigrades is due to their tendency to contract their bodies while being fixed, and their minute size, as for example the entire brain of the species studied herein would fit into the soma of a single insect neuron Fig. Our understanding of the segmental composition of the tardigrade head depends on the unresolved issue of whether the tardigrade brain is a non-segmental structure or composed of one, two, or more segments [8] — [15].

The results of two recent immunocytochemical studies arrive at opposing conclusions, as they suggest that the tardigrade brain is either unsegmented and comparable to the ring-shaped brain of cycloneuralians [9] , or tripartite consisting of proto-, deuto- and tritocerebrum and, therefore, similar to the brain of arthropods [10].

While both studies mainly focused on the distribution of commissures and neuropils in the tardigrade brain, this approach generally does not seem to be sufficient for resolving the number of segmental brain regions since a single segment can be composed of multiple neuropils [4]. Thus, it is essential to provide other lines of evidence in order to clarify the segmental composition of the tardigrade brain.

Light micrographs; both images are to scale. Note that the entire anterior end of the tardigrade, including the brain and consisting of hundreds of cells [79] , would fit into a single neuronal cell body of the insect. A Cobalt-filled dorsal unpaired median neuron DUM neuron from the thoracic ganglion of the locust Locusta migratoria.

B Specimen of Macrobiotus cf. The position of the stomatogastric ganglia has been used successfully in the past to align the segmental regions of the brain and the head segments in arthropods [5] , [16]. In most major arthropod groups, the stomatogastric ganglia, which innervate the ectodermal foregut and the endodermal midgut structures, are typically associated with the third body segment [5] , [16] — [19].

Thus, clarifying the organisation and position of the stomatogastric nervous system in tardigrades might also help to align the anterior body segments between tardigrades and arthropods. Additional nerves and neurites supplying the buccal apparatus, including the stylet and the anterior sensory structures, have been described in detail using confocal microscopy [9] , [10] , but the data on the innervation pattern of the midgut are still lacking in tardigrades.

To close this gap and to clarify the number of head segments in tardigrades, we applied a variety of cytochemical and immunolabelling markers in conjunction with confocal laser-scanning microscopy to visualise the nervous tissues of adult specimens of the limno-terrestrial tardigrade Macrobiotus cf. For these experiments, we optimised our asphyxiation technique to obtain specimens with a fully extended body, which maximised the resolution of neuroanatomical structures in these minute animals Figure 1B.

For comparison, we analysed selected elements of the nervous system innervating the foregut and midgut structures in the onychophoran Euperipatoides rowelli. Our findings provide a clear framework for aligning the anterior body segments in tardigrades and arthropods. Specimens of Macrobiotus cf. No specific permits were required for the collection of tardigrades since the location is not privately-owned or protected in any way and no endangered or protected species were involved.

Incubations with antibodies were carried out either overnight or for several days. The following primary antibodies were used:. This antibody has been used successfully to label neuron fibres in various metazoans, including cnidarians [27] , acoelans [28] , annelids [29] and chaetognaths [30].

In addition, we used two antisera that recognise different neuromodulators that are common in invertebrates. Its specificity has been established by the manufacturer and it has been used to label FMRFamide containing neurons in a wide variety of invertebrates, including cnidarians [34] , acoelomorphs [35] , rotifers [36] , molluscs [37] , annelids [38] and numerous insects e.

Its specificity has been established and it has been shown to recognise serotonergic neurons for example in molluscs [40] , insects [41] and rats [42]. After several rinses in PBS, some specimens and sections were incubated for one hour in a solution containing phalloidin-rhodamine Invitrogen to stain f-actin as described previously [32].

Scanning electron microscopy was carried out as described previously [24]. Confocal images of entire specimens of Macrobiotus cf. The brain and the four trunk ganglia are linked by paired, somata-free connectives. An additional pair of outer connectives joins the first trunk ganglion with the brain Figure 3A—D.

See also Figures S1 , S2 for the colour-blind. RFamide-like immunoreactivity red and DNA labelling green. Maximum projections of confocal z-series. Anterior is up in all images except for B in which anterior is left. Arrowheads in A—C point to varicose swellings in the core of each trunk ganglion.

Arrows in B and D indicate two anterior, extra-cerebral RFamide-like immunoreactive cell bodies. A Specimen in ventral view showing four trunk ganglia linked by somata-free connectives. B Specimen in lateral view revealing the dorsal position of the brain dotted line. C Anterior end in ventral view.

D Anterior end in dorsal view with details of central brain neuropil. Dotted line indicates the shape of the brain with its lobes. A, B Specimens in ventral and ventrolateral view, respectively. C, D Depth-coded projections of confocal z-series of anterior ends of an extended in C and a contracted specimen in D. Note the complex fibre network within the central brain neuropil, which appears as a unitary structure with the nerve ring in the contracted specimen.

Anti-tubulin immunolabelling reveals numerous contralateral projections and commissure-like structures within each ganglion and an additional extra-ganglionic commissure in front of the second, third and fourth trunk ganglia Figure S3.

We also observed vivid RFamide-like immunoreactive staining of two varicose swellings that lie in close apposition to each other in the core of each trunk ganglion arrowheads in Figure 2A—C and Figures S1 , S2. The shape of these structures differs between the ganglia of the same individual but is similar in the same ganglion of different specimens. Since they lack a nucleus they are clearly not somata. Their small size maximum diameter 2. Since we observed no other clearly RFamide-like labelled neuropil structures in the trunk ganglia, these varicosities could possibly represent individual synaptic terminals in close association.

DNA labelling and analyses of confocal z-series reveal that the brain of Macrobiotus cf. Since some of these nuclei are likely to be from non-neuronal cell types, such as glia cells, the tardigrade brain might contain fewer than neurons.

In shape, the brain appears lobate in dorsal view, as it consists of one pair of outer lobes, one pair of inner lobes and an unpaired median lobe. There is no obvious grouping of somata into clusters within the lobes.

Anti-RFamide and anti-serotonin immunolabelling reveals somata of individually identifiable neurons of different size, which are arranged in a bilaterally symmetric pattern Figure 4A—D. Specimens in dorsal view; anterior is up. Note the bilaterally symmetric arrangement of individually identifiable neuronal somata. A RFamide-like immunoreactivity; maximum projection of confocal z-series.

Inset shows details of two extra-cerebral RFamide-like immunoreactive somata situated in front of the brain asterisks. B Volume rendering of the same dataset as in A. The brain contours are illustrated in blue autofluorescence , RFamide-like immunoreactive structures in yellow.

Inset shows details of a ring-like structure surrounding the mouth. C Serotonin-like immunoreactivity; maximum projection of confocal z-series. Note the strong signal in the mouth cone. D Volume rendering of the same dataset as in C. Serotonin-like immunoreactive neural structures are illustrated in green, the mouth cone in light-blue, autofluorescent calcified stylet structures in magenta.

Arrowheads in C and D point to two bilateral neurites, which join the dorsal portion of the ring nerve see also Movie S2. In addition, two extra-cerebral RFamide-like immunoreactive cell bodies are found dorsally behind the brain Figure 4B and Movie S1.

Each of these two somata sends out a neurite, which enters the brain between the inner and median brain lobes Movie S1. Additional two RFamide-like and ten serotonin-like immunoreactive somata are located anterior to the brain Figure 4A—D.

The two RFamide-like immunoreactive cell bodies cluster together in the dorsomedian head region inset in Figure 4A. The anterior cell body gives rise to two lateroposterior neurites and to an unpaired anterior neurite, which forms a ring-like structure around the mouth inset in Figure 4B , whereas the weakly labelled posterior cell body sends out a single posterior neurite towards the brain Movie S1.

The ten extra-cerebral, serotonin-like immunoreactive cell bodies are not associated with the brain but are spatially distributed around the mouth cone. One lies dorsomedially, one ventromedially, two are closely associated with the lateral portions of the buccal tube and three bilateral pairs lie further posteriorly Figure 4C, D , Figure S4 and Movie S2.

The ten extra-cerebral, serotonin-like immunoreactive somata send single neurites to the buccal tube. The lateral portions of the central neuropil receive fibres from the anterior-most and outer connectives, which link the brain to the first trunk ganglion Figure 3C, D. In addition, single fibres and bundles of neurites extend from the paired anterolateral and posterolateral sensory fields to the central brain neuropil Figure 3A—D.

In front of the central neuropil, there is a prominent ring nerve surrounding the buccal tube Figure 3A—C and Figure S5. This nerve receives fibres from the brain as well as from the two connectives linking the brain to the first trunk ganglion Figures 3A—C and Movie S3. The ring nerve gives rise to numerous neurites projecting anteriorly to the peribuccal lamellae surrounding the mouth opening Figure 3A—C , 5A—C , Figure S5 and Movie S3.

Moreover, it receives fibres from bundles of neurites innervating the stylet apparatus Figure 5B, C. These bundles arise ventrally from the central brain neuropil and project posteriorly to innervate the musculature of the stylet apparatus Figures 3C , 5B, C , Figure S6 and Movie S3.

The bundles are linked by three ventral commissures, whereas anteriorly they send off several longitudinal projections that join the ring nerve Figure 5C. A Scanning electron micrograph showing the peribuccal lamellae surrounding the mouth opening.

B Anterior end of a specimen labelled with phalloidin-rhodamine to reveal the stylet and pharyngeal musculature. Projection of a confocal z-series; anterior is up. Projection of confocal z-series; anterior is up. For the sake of clarity, only elements of the stomodeal complex were selected from each optical section for the projection. Dotted lines demarcate the corresponding body region in the specimen labelled with phalloidin-rhodamine.

Projections of confocal z-series. A Anterior half of a specimen in dorsal view. Maximum projection of a confocal z-series. Note serotonin-like immunoreactivity in the stomatogastric ganglion. Arrowheads point to three posterior neurites. B Detail of the stomatogastric ganglion. Neuronal cell bodies are marked by asterisks. C Detail of a lateral posterior neurite arrowhead , which has an input into the second trunk ganglion arrow.

Projection of a confocal z-series. Although our data show that the ring nerve and the central brain neuropil are clearly distinguishable in extended specimens Figure 3C , they appear as a fused, unitary structure in contracted specimens Figure 3D.

Similarly, the anterior neurites innervating the peribuccal lamellae surrounding the mouth are clearly visible in extended specimens but obscured after contraction Figure 3C, D and Figure S7.

The extent of distortion of neural structures situated in the head is particularly evident in the position of the anterolateral and posterolateral sensory fields, which are widely separated in fully extended specimens, but in close proximity in contracted animals. Notably, the central neuropil and all cell bodies of RFamide-like and serotonin-like immunoreactive neurons of the brain in Macrobiotus cf.

The fibres of the anterior connectives from the first trunk ganglion circumvent the buccal tube via the ring nerve and connect with the dorsal brain, without condensing to form any additional neuropil structure or traversing any further conglomeration of nuclei Figures 2B, C , 4A, B , Figures S1 , S2 and Movie S1. Corresponding findings were obtained for 21 critically inspected preparations immunolabelled with an anti-FMRFamide antibody.

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The municipality is an S-shaped linear village located in the Thierstein district. Moreover, Hejnol and Schnabel [43] found no anlage of this ganglion in the embryo of the tardigrade Thulinia stephaniae using 4D microscopy.

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