Distilling the neural correlates of conscious somatosensory perception

The ability to consciously perceive the world profoundly defines our lives as human beings. Somehow, our brains process information in a way that allows us to become aware of the images, sounds, touches, smells, and tastes surrounding us. Yet our understanding of the neurobiological processes that generate perceptual awareness is very limited. One of the most contested questions in the neuroscientific study of conscious perception is whether awareness arises from the activity of early sensory brain regions, or instead requires later processing in widespread supramodal networks. It has been suggested that the conflicting evidence supporting these two perspectives may be the result of methodological confounds in classical experimental tasks. In order to infer participants’ perceptual awareness in these tasks, they need to report the contents of their perception. This means that the neural signals underlying the emergence of perceptual awareness often cannot be dissociated from pre- and postperceptual processes. Consequently, some of the previously observed effects may not be correlates of awareness after all but instead may have resulted from task requirements. In this thesis, I investigate this possibility in the somatosensory modality. To scrutinise the task dependence of the neural correlates of somatosensory awareness, I developed an experimental paradigm that controls for the most common experimental confounds. In a somatosensory-visual matching task, participants were required to detect electrical target stimuli at ten different intensity levels. Instead of reporting their perception directly, they compared their somatosensory percepts to simultaneously presented visual cues that signalled stimulus presence or absence and then reported a match or mismatch accordingly. As a result, target detection was decorrelated from working memory and reports, the behavioural relevance of detected and undetected stimuli was equated, the influence of attentional processes was mitigated, and perceptual uncertainty was varied in a controlled manner. Results from a functional magnetic resonance imaging (fMRI) study and an electroencephalography (EEG) study showed that, when controlled for task demands, the neural correlates of somatosensory awareness were restricted to relatively early activity (~150 ms) in secondary somatosensory regions. In contrast, late activity (>300 ms) indicative of processing in frontoparietal networks occurred irrespective of stimulus awareness, and activity in anterior insular, anterior cingulate, and supplementary motor cortex was associated with processing perceptual uncertainty and reports. These results add novel evidence to the early-local vs. late-global debate and favour the view that perceptual awareness emerges at the level of modality-specific sensory cortices.Die Fähigkeit zur bewussten Wahrnehmung bestimmt maßgeblich unser Selbstbild als Menschen. Unser Gehirn verarbeitet Informationen auf eine Weise, die es uns ermöglicht, uns der Bilder, Töne, Berührungen, Gerüche und Geschmäcker, die uns umgeben, bewusst zu werden. Unser Verständnis davon, wie neurobiologische Prozesse diese bewusste Wahrnehmung erzeugen, ist jedoch noch sehr begrenzt. Eine der umstrittensten Fragen in der neurowissenschaftlichen Erforschung des perzeptuellen Bewusstseins besteht darin, ob die bewusste Wahrnehmung aus der Aktivität früher sensorischer Hirnregionen entsteht, oder aber die spätere Prozessierung in ausgedehnten supramodalen Netzwerken erfordert. Eine mögliche Erklärung für die widersprüchlichen Ergebnisse, die diesen beiden Perspektiven zugrunde liegen, wird in methodologischen Störfaktoren vermutet, die in klassischen experimentellen Paradigmen auftreten können. Um auf die Wahrnehmung der Versuchspersonen schließen zu können, müssen diese den Inhalt ihrer Wahrnehmung berichten. Das führt dazu, dass neuronale Korrelate bewusster Wahrnehmung häufig nicht sauber von prä- und postperzeptuellen Prozessen getrennt werden können. Folglich könnten einige der zuvor beobachteten Effekte, anstatt tatsächlich bewusste Wahrnehmung widerzuspiegeln, aus den Anforderungen experimenteller Paradigmen entstanden sein. In dieser Arbeit untersuche ich diese Möglichkeit in der somatosensorischen Modalität. Um zu überprüfen, inwiefern neuronale Korrelate bewusster somatosensorischer Wahrnehmung von den Anforderungen experimenteller Aufgaben abhängen, habe ich ein Paradigma entwickelt, dass die häufigsten experimentellen Störfaktoren kontrolliert. In einer somatosensorisch-visuellen Vergleichsaufgabe mussten die Versuchspersonen elektrische Zielreize in zehn verschiedenen Intensitätsstufen detektieren. Anstatt diese jedoch direkt zu berichten, sollten sie ihre somatosensorischen Perzepte mit gleichzeitig präsentierten visuellen Symbolen vergleichen, die entweder Reizanwesenheit oder -abwesenheit signalisierten. Entsprechend wurde dann eine Übereinstimmung oder Nichtübereinstimmung berichtet. Dadurch wurde die Reizwahrnehmung von Arbeitsgedächtnis und Berichterstattung dekorreliert, die Verhaltensrelevanz detektierter und nicht detektierter Reize gleichgesetzt, der Einfluss von Aufmerksamkeitsprozessen reduziert und die mit der Detektion verbundene Unsicherheit auf kontrollierte Weise variiert. Die Ergebnisse aus einer funktionellen Magnetresonanztomographie (fMRT)-Studie und einer Elektroenzephalographie (EEG)-Studie zeigen, dass die neuronalen Korrelate bewusster somatosensorischer Wahrnehmung auf relativ frühe Aktivität (~150 ms) in sekundären somatosensorischen Regionen beschränkt sind, wenn experimentelle Störfaktoren kontrolliert werden. Im Gegensatz dazu trat späte Aktivität (>300 ms), die auf die Verarbeitung in frontoparietalen Netzwerken hindeutet, unabhängig von der Reizwahrnehmung auf, und Aktivität im anterioren insulären, anterioren cingulären und supplementär-motorischen Kortex war mit der Verarbeitung von Detektionsunsicherheit und der Berichterstattung verbunden. Diese Ergebnisse liefern neue Erkenntnisse zur Debatte um die Relevanz früher, lokaler vs. später, globaler Hirnaktivität und unterstützen die Ansicht, dass perzeptuelles Bewusstsein in modalitätsspezifischen sensorischen Kortizes entsteht

Rote Liste und Artenlisten der bodenlebenden wirbellosen Meerestiere

1,244 species out of nine phyla of macrozoobenthos have been evaluated for the Red Lists of invertebrate species in German North Sea and Baltic Sea waters (including the Exclusive Economic Zone). It is evident that the conditions in these marine waters are basically not better than ashore, since human impact like, first of all, demersal fishing and eutrophication as well as impact on coastal water biotopes have led to massive changes in several animal populations. 32 % of all species assessed are found in the Red Lists. However, more than half of these species are extremely rare and therefore assessed as potentially threatened. Altogether data are still insufficient for any trend analysis in more than 36 % of all evaluated 1,244 species (see high share in category „D“). Due to increases in knowledge and specific investigations on selected taxa it is expected that in a few years updated Red Lists can be provided, maybe covering the entire ecosystems of the North and Baltic Seas

Intravenous alteplase for stroke with unknown time of onset guided by advanced imaging: systematic review and meta-analysis of individual patient data

Background: Patients who have had a stroke with unknown time of onset have been previously excluded from thrombolysis. We aimed to establish whether intravenous alteplase is safe and effective in such patients when salvageable tissue has been identified with imaging biomarkers. Methods: We did a systematic review and meta-analysis of individual patient data for trials published before Sept 21, 2020. Randomised trials of intravenous alteplase versus standard of care or placebo in adults with stroke with unknown time of onset with perfusion-diffusion MRI, perfusion CT, or MRI with diffusion weighted imaging-fluid attenuated inversion recovery (DWI-FLAIR) mismatch were eligible. The primary outcome was favourable functional outcome (score of 0–1 on the modified Rankin Scale [mRS]) at 90 days indicating no disability using an unconditional mixed-effect logistic-regression model fitted to estimate the treatment effect. Secondary outcomes were mRS shift towards a better functional outcome and independent outcome (mRS 0–2) at 90 days. Safety outcomes included death, severe disability or death (mRS score 4–6), and symptomatic intracranial haemorrhage. This study is registered with PROSPERO, CRD42020166903. Findings: Of 249 identified abstracts, four trials met our eligibility criteria for inclusion: WAKE-UP, EXTEND, THAWS, and ECASS-4. The four trials provided individual patient data for 843 individuals, of whom 429 (51%) were assigned to alteplase and 414 (49%) to placebo or standard care. A favourable outcome occurred in 199 (47%) of 420 patients with alteplase and in 160 (39%) of 409 patients among controls (adjusted odds ratio [OR] 1·49 [95% CI 1·10–2·03]; p=0·011), with low heterogeneity across studies (I 2=27%). Alteplase was associated with a significant shift towards better functional outcome (adjusted common OR 1·38 [95% CI 1·05–1·80]; p=0·019), and a higher odds of independent outcome (adjusted OR 1·50 [1·06–2·12]; p=0·022). In the alteplase group, 90 (21%) patients were severely disabled or died (mRS score 4–6), compared with 102 (25%) patients in the control group (adjusted OR 0·76 [0·52–1·11]; p=0·15). 27 (6%) patients died in the alteplase group and 14 (3%) patients died among controls (adjusted OR 2·06 [1·03–4·09]; p=0·040). The prevalence of symptomatic intracranial haemorrhage was higher in the alteplase group than among controls (11 [3%] vs two [<1%], adjusted OR 5·58 [1·22–25·50]; p=0·024). Interpretation: In patients who have had a stroke with unknown time of onset with a DWI-FLAIR or perfusion mismatch, intravenous alteplase resulted in better functional outcome at 90 days than placebo or standard care. A net benefit was observed for all functional outcomes despite an increased risk of symptomatic intracranial haemorrhage. Although there were more deaths with alteplase than placebo, there were fewer cases of severe disability or death. Funding: None

Description and performance of track and primary-vertex reconstruction with the CMS tracker

A description is provided of the software algorithms developed for the CMS tracker both for reconstructing charged-particle trajectories in proton-proton interactions and for using the resulting tracks to estimate the positions of the LHC luminous region and individual primary-interaction vertices. Despite the very hostile environment at the LHC, the performance obtained with these algorithms is found to be excellent. For tbar t events under typical 2011 pileup conditions, the average track-reconstruction efficiency for promptly-produced charged particles with transverse momenta of p(T) > 0.9GeV is 94% for pseudorapidities of |η| < 0.9 and 85% for 0.9 < |η| < 2.5. The inefficiency is caused mainly by hadrons that undergo nuclear interactions in the tracker material. For isolated muons, the corresponding efficiencies are essentially 100%. For isolated muons of p(T) = 100GeV emitted at |η| < 1.4, the resolutions are approximately 2.8% in p(T), and respectively, 10μm and 30μm in the transverse and longitudinal impact parameters. The position resolution achieved for reconstructed primary vertices that correspond to interesting pp collisions is 10–12μm in each of the three spatial dimensions. The tracking and vertexing software is fast and flexible, and easily adaptable to other functions, such as fast tracking for the trigger, or dedicated tracking for electrons that takes into account bremsstrahlung