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- Department Psychologie (6)
- Institut für Jüdische Studien und Religionswissenschaft (6)
Although the basic structure of biological membranes is provided by the lipid bilayer, most of the specific functions are carried out by membrane proteins (MPs) such as channels, ion-pumps and receptors. Additionally, it is known, that mutations in MPs are directly or indirectly involved in many diseases. Thus, structure determination of MPs is of major interest not only in structural biology but also in pharmacology, especially for drug development. Advances in structural biology of membrane proteins (MPs) have been strongly supported by the success of three leading techniques: X-ray crystallography, electron microscopy and solution NMR spectroscopy. However, X-ray crystallography and electron microscopy, require highly diffracting 3D or 2D crystals, respectively. Today, structure determination of non-crystalline solid protein preparations has been made possible through rapid progress of solid-state MAS NMR methodology for biological systems. Castellani et. al. solved and refined the first structure of a microcrystalline protein using only solid-state MAS NMR spectroscopy. These successful application open up perspectives to access systems that are difficult to crystallise or that form large heterogeneous complexes and insoluble aggregates, for example ligands bound to a MP-receptor, protein fibrils and heterogeneous proteins aggregates. Solid-state MAS NMR spectroscopy is in principle well suited to study MP at atomic resolution. In this thesis, different types of MP preparations were tested for their suitability to be studied by solid-state MAS NMR. Proteoliposomes, poorly diffracting 2D crystals and a PEG precipitate of the outer membrane protein G (OmpG) were prepared as a model system for large MPs. Results from this work, combined with data found in the literature, show that highly diffracting crystalline material is not a prerequirement for structural analysis of MPs by solid-state MAS NMR. Instead, it is possible to use non-diffracting 3D crystals, MP precipitates, poorly diffracting 2D crystals and proteoliposomes. For the latter two types of preparations, the MP is reconstituted into a lipid bilayer, which thus allows the structural investigation in a quasi-native environment. In addition, to prepare a MP sample for solid-state MAS NMR it is possible to use screening methods, that are well established for 3D and 2D crystallisation of MPs. Hopefully, these findings will open a fourth method for structural investigation of MP. The prerequisite for structural studies by NMR in general, and the most time consuming step, is always the assignment of resonances to specific nuclei within the protein. Since the last few years an ever-increasing number of assignments from solid-state MAS NMR of uniformly carbon and nitrogen labelled samples is being reported, mostly for small proteins of up to around 150 amino acids in length. However, the complexity of the spectra increases with increasing molecular weight of the protein. Thus the conventional assignment strategies developed for small proteins do not yield a sufficiently high degree of assignment for the large MP OmpG (281 amino acids). Therefore, a new assignment strategy to find starting points for large MPs was devised. The assignment procedure is based on a sample with [2,3-13C, 15N]-labelled Tyr and Phe and uniformly labelled alanine and glycine. This labelling pattern reduces the spectral overlap as well as the number of assignment possibilities. In order to extend the assignment, four other specifically labelled OmpG samples were used. The assignment procedure starts with the identification of the spin systems of each labelled amino acid using 2D 13C-13C and 3D NCACX correlation experiments. In a second step, 2D and 3D NCOCX type experiments are used for the sequential assignment of the observed resonances to specific nuclei in the OmpG amino acid sequence. Additionally, it was shown in this work, that biosynthetically site directed labelled samples, which are normally used to observe long-range correlations, were helpful to confirm the assignment. Another approach to find assignment starting points in large protein systems, is the use of spectroscopic filtering techniques. A filtering block that selects methyl resonances was used to find further assignment starting points for OmpG. Combining all these techniques, it was possible to assign nearly 50 % of the observed signals to the OmpG sequence. Using this information, a prediction of the secondary structure elements of OmpG was possible. Most of the calculated motifs were in good aggreement with the crystal structures of OmpG. The approaches presented here should be applicable to a wide variety of MPs and MP-complexes and should thus open a new avenue for the structural biology of MPs.
Microsaccades are very small, involuntary flicks in eye position that occur on average once or twice per second during attempted visual fixation. Microsaccades give rise to EMG eye muscle spikes that can distort the spectrum of the scalp EEG and mimic increases in gamma band power. Here we demonstrate that microsaccades are also accompanied by genuine and sizeable cortical activity, manifested in the EEG. In three experiments, high-resolution eye movements were corecorded with the EEG: during sustained fixation of checkerboard and face stimuli and in a standard visual oddball task that required the counting of target stimuli. Results show that microsaccades as small as 0.15° generate a field potential over occipital cortex and midcentral scalp sites 100 –140 ms after movement onset, which resembles the visual lambda response evoked by larger voluntary saccades. This challenges the standard assumption of human brain imaging studies that saccade-related brain activity is precluded by fixation, even when fully complied with. Instead, additional cortical potentials from microsaccades were present in 86% of the oddball task trials and of similar amplitude as the visual response to stimulus onset. Furthermore, microsaccade probability varied systematically according to the proportion of target stimuli in the oddball task, causing modulations of late stimulus-locked event-related potential (ERP) components. Microsaccades present an unrecognized source of visual brain signal that is of interest for vision research and may have influenced the data of many ERP and neuroimaging studies.
Human information processing depends critically on continuous predictions about upcoming events, but the temporal convergence of expectancy-based top-down and input-driven bottom-up streams is poorly understood. We show that, during reading, event-related potentials differ between exposure to highly predictable and unpredictable words no later than 90 ms after visual input. This result suggests an extremely rapid comparison of expected and incoming visual information and gives an upper temporal bound for theories of top-down and bottom-up interactions in object recognition.
We examined individual differences in masked repetition priming by re-analyzing item-level response-time (RT) data from three experiments. Using a linear mixed model (LMM) with subjects and items specified as crossed random factors, the originally reported priming and word-frequency effects were recovered. In the same LMM, we estimated parameters describing the distributions of these effects across subjects. Subjects’ frequency and priming effects correlated positively with each other and negatively with mean RT. These correlation estimates, however, emerged only with a reciprocal transformation of RT (i.e., -1/RT), justified on the basis of distributional analyses. Different correlations, some with opposite sign, were obtained (1) for untransformed or logarithmic RTs or (2) when correlations were computed using within-subject analyses. We discuss the relevance of the new results for accounts of masked priming, implications of applying RT transformations, and the use of LMMs as a tool for the joint analysis of experimental effects and associated individual differences.
Throughout its empirical research history eye movement research has always been aware of the differences in reading behavior induced by individual differences and task demands. This work introduces a novel comprehensive concept of reading strategy, comprising individual differences in reading style and reading skill as well as reader goals. In a series of sentence reading experiments recording eye movements, the influence of reading strategies on reader- and word-level effects assuming distributed processing has been investigated. Results provide evidence for strategic, top-down influences on eye movement control that extend our understanding of eye guidance in reading.
It has recently been demonstrated that the presentation of a rare target in a visual oddball paradigm induces a prolonged inhibition of microsaccades. In the field of electrophysiology, the amplitude of the P300 component in event-related potentials (ERP) has been shown to be sensitive to the stimulus category (target vs. non target) of the eliciting stimulus, its overall probability, and the preceding stimulus sequence. In the present study we further specify the functional underpinnings of the prolonged microsaccadic inhibition in the visual oddball task, showing that the stimulus category, the frequency of a stimulus and the preceding stimulus sequence influence microsaccade rate. Furthermore, by co-recording ERPs and eye-movements, we were able to demonstrate that, despite being largely sensitive to the same experimental manipulation, the amplitude of P300 and the microsaccadic inhibition predict each other very weakly, and thus constitute two independent measures of the brain’s response to rare targets in the visual oddball paradigm.
Covert shifts of attention are usually reflected in RT differences between responses to valid and invalid cues in the Posner spatial attention task. Such inferences about covert shifts of attention do not control for microsaccades in the cue target interval. We analyzed the effects of microsaccade orientation on RTs in four conditions, crossing peripheral visual and auditory cues with peripheral visual and auditory discrimination targets. Reaction time was generally faster on trials without microsaccades in the cue-target interval. If microsaccades occurred, the target-location congruency of the last microsaccade in the cuetarget interval interacted in a complex way with cue validity. For valid visual cues, irrespective of whether the discrimination target was visual or auditory, target-congruent microsaccades delayed RT. For invalid cues, target-incongruent microsaccades facilitated RTs for visual target discrimination, but delayed RT for auditory target discrimination. No reliable effects on RT were associated with auditory cues or with the first microsaccade in the cue-target interval. We discuss theoretical implications on the relation about spatial attention and oculomotor processes.