TY - THES A1 - Amaechi, Mary Chimaobi T1 - A'-movement dependencies and their reflexes in Igbo N2 - In this thesis, I examine different A-bar movement dependencies in Igbo, a Benue-Congo language spoken in southern Nigeria. Movement dependencies are found in constructions where an element is moved to the left edge of the clause to express information-structural categories such as in questions, relativization and focus. I show that these constructions in Igbo are very uniform from a syntactic point of view. The constructions are built on two basic fronting operations: relativization and focus movement, and are biclausal. I further investigate several morphophonological effects that are found in these A-bar constructions. I propose that these effects are reflexes of movement that are triggered when an element is moved overtly in relativization or focus. This proposal helps to explain the tone patterns that have previously been assumed to be a property of relative clauses. The thesis adds to the growing body of tonal reflexes of A-bar movement reported for a few African languages. The thesis also provides an insight into the complementizer domain (C-domain) of Igbo. N2 - In dieser Dissertation untersuche ich verschiedene syntaktische A-quer-Bewegungsabhängigkeiten in Igbo, einer Benue-Kongo-Sprache, die im Süden Nigerias gesprochen wird. Bewegungsabhängigkeiten finden sich in Konstruktionen, in denen ein Element an den linken Satzrand bewegt wird, um informationsstrukturelle Kategorien auszudrücken, etwa in Fragen, Relativsatzbildung und Fokussierung. Ich zeige, dass diese Konstruktionen in Igbo in Hinblick auf die Syntax sehr einheitlich sind. Die Konstruktionen bauen auf zwei grundlegenden Voranstellungsoperationen auf – Relativierung und Fokusbewegung – und sind biklausal. Außerdem untersuche ich verschiedene morphophonologische Effekte, die in diesen A-quer-Konstruktionen vorkommen. Ich schlage vor, dass die Effekte Ausdruck von Bewegung sind und dadurch ausgelöst werden, dass ein Element zum Zweck der Relativsatzbildung oder Fokussierung bewegt wird. Die Dissertation liefert einen weiteren Beleg für tonale Effekte von A-quer-Bewegung, die bereits für einige afrikanische Sprachen berichtet wurden. Die Arbeit gibt auch Aufschluss über die Komplementierer-Domäne (C-domain) in Igbo. T2 - A'-Bewegungsabhängigkeiten und ihre Reflexe im Igbo KW - A-bar movement KW - clefts KW - focus KW - relativization KW - Igbo KW - A-quer-Bewegung KW - Spaltsätze KW - Fokus KW - Relativsätze KW - Igbo Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-471524 ER - TY - THES A1 - Mavrothalassiti, Eleni T1 - A.thaliana root and shoot single-cell transcriptomes and detection of mobile transcripts Y1 - 2020 ER - TY - THES A1 - Patzwald, Christiane T1 - Actions through the lens of communicative cues BT - the influence of verbal cues and emotional cues on action processing and action selection in the second year of life N2 - The PhD thesis entitled “Actions through the lens of communicative cues. The influence of verbal cues and emotional cues on action processing and action selection in the second year of life” is based on four studies, which examined the cognitive integration of another person’s communicative cues (i.e., verbal cues, emotional cues) with behavioral cues in 18- and 24-month-olds. In the context of social learning of instrumental actions, it was investigated how the intention-related coherence of either a verbally announced action intention or an emotionally signaled action evaluation with an action demonstration influenced infants’ neuro-cognitive processing (Study I) and selection (Studies II, III, IV) of a novel object-directed action. Developmental research has shown that infants benefit from another’s behavioral cues (e.g., action effect, persistency, selectivity) to infer the underlying goal or intention, respectively, of an observed action (e.g., Cannon & Woodward, 2012; Woodward, 1998). Particularly action effects support infants in distinguishing perceptual action features (e.g., target object identity, movement trajectory, final target object state) from conceptual action features such as goals and intentions. However, less is known about infants’ ability to cognitively integrate another’s behavioral cues with additional action-related communicative cues. There is some evidence showing that in the second year of life, infants selectively imitate a novel action that is verbally (“There!”) or emotionally (positive expression) marked as aligning with the model’s action intention over an action that is verbally (“Whoops!”) or emotionally (negative expression) marked as unintentional (Carpenter, Akhtar, & Tomasello, 1998; Olineck & Poulin-Dubois, 2005, 2009; Repacholi, 2009; Repacholi, Meltzoff, Toub, & Ruba, 2016). Yet, it is currently unclear which role the specific intention-related coherence of a communicative cue with a behavioral cue plays in infants’ action processing and action selection that is, whether the communicative cue confirms, contrasts, clarifies, or is unrelated to the behavioral cue. Notably, by using both verbal cues and emotional cues, we examined not only two domains of communicative cues but also two qualitatively distinct relations between behavioral cues on the one hand and communicative cues on the other hand. More specifically, a verbal cue has the potential to communicate an action intention in the absence of an action demonstration and thus a prior-intention (Searle, 1983), whereas an emotional cue evaluates an ongoing or past action demonstration and thus signals an intention-in-action (Searle, 1983). In a first research focus, this thesis examined infants’ capacity to cognitively integrate another’s intention-related communicative cues and behavioral cues, and also focused on the role of the social cues’ coherence in infants’ action processing and action selection. In a second research focus, and to gain more elaborate insights into how the sub-processes of social learning (attention, encoding, response; cf. Bandura, 1977) are involved in this coherence-sensitive integrative processing, we employed a multi-measures approach. More specifically, we used Electroencephalography (EEG) and looking times to examine how the cues’ coherence influenced the compound of attention and encoding, and imitation (including latencies to first-touch and first-action) to address the compound of encoding and response. Based on the action-reconstruction account (Csibra, 2007), we predicted that infants use extra-motor information (i.e., communicative cues) together with behavioral cues to reconstruct another’s action intention. Accordingly, we expected infants to possess a flexibly organized internal action hierarchy, which they adapt according to the cues’ coherence that is, according to what they inferred to be the overarching action goal. More specifically, in a social-learning situation that comprised an adult model, who demonstrated an action on a novel object that offered two actions, we expected the demonstrated action to lead infants’ action hierarchy when the communicative (i.e., verbal, emotional) cue conveyed similar (confirming coherence) or no additional (un-related coherence) intention-related information relative to the behavioral cue. In terms of action selection, this action hierarchy should become evident in a selective imitation of the demonstrated action. However, when the communicative cue questioned (contrasting coherence) the behaviorally implied action goal or was the only cue conveying meaningful intention-related information (clarifying coherence), the verbally/emotionally intended action should ascend infants’ action hierarchy. Consequently, infants’ action selection should align with the verbally/emotionally intended action (goal emulation). Notably, these predictions oppose the direct-matching perspective (Rizzolatti & Craighero, 2004), according to which the observation of another’s action directly resonates with the observer’s motor repertoire, with this motor resonance enabling the identification of the underlying action goal. Importantly, the direct-matching perspective predicts a rather inflexible action hierarchy inasmuch as the process of goal identification should solely rely on the behavioral cue, irrespective of the behavioral cue’s coherence with extra-motor intention-related information, as it may be conveyed via communicative cues. As to the role of verbal cues, Study I used EEG to examine the influence of a confirming (Congruent) versus contrasting (Incongruent) coherence of a verbal action intention with the same action demonstration on 18-month-olds’ conceptual action processing (as measured via mid-latency mean negative ERP amplitude) and motor activation (as measured via central mu-frequency band power). The action was demonstrated on a novel object that offered two action alternatives from a neutral position. We expected mid-latency ERP negativity to be enhanced in Incongruent compared to Congruent, because past EEG research has demonstrated enhanced conceptual processing for stimuli that mismatched rather than matched the semantic context (Friedrich & Friederici, 2010; Kaduk et al., 2016). Regarding motor activation, Csibra (2007) posited that the identification of a clear action goal constitutes a crucial basis for motor activation to occur. We therefore predicted reduced mu power (indicating enhanced motor activation) for Congruent than Incongruent, because in Congruent, the cues’ match provides unequivocal information about the model’s action goal, whereas in Incongruent, the conflict may render the model’s action goal more unclear. Unexpectedly, in the entire sample, 18-month-olds’ mid-latency ERP negativity during the observation of the same action demonstration did not differ significantly depending on whether this action was congruent or incongruent with the model’s verbal action intention. Yet, post hoc analyses revealed the presence of two subgroups of infants, each of which exhibited significantly different mid-latency ERP negativity for Congruent versus Incongruent, but in opposing directions. The subgroups differed in their productive action-related language skills, with the linguistically more advanced infants exhibiting the expected response pattern of enhanced ERP mean negativity in Incongruent than Congruent, indicating enhanced conceptual processing of an action demonstration that was contrasted rather than confirmed by the verbal action context. As expected, central mu power in the entire sample was reduced in Congruent relative to Incongruent, indicating enhanced motor activation when the action demonstration was preceded by a confirming relative to a contrasting verbal action intention. This finding may indicate the covert preparation for a preferential imitation of the congruent relative to the incongruent action (Filippi et al., 2016; Frey & Gerry, 2006). Overall, these findings are in line with the action-reconstruction account (Csibra, 2007), because they suggest a coherence-sensitive attention to and encoding of the same perceptual features of another’s behavior and thus a cognitive integration of intention-related verbal cues and behavioral cues. Yet, because the subgroup constellation in infants’ ERPs was only discovered post hoc, future research is clearly required to substantiate this finding. Also, future research should validate our interpretation that enhanced motor activation may reflect an electrophysiological marker of subsequent imitation by employing EEG and imitation in a within-subjects design. Study II built on Study I by investigating the impact of coherence of a verbal cue and a behavioral cue on 18- and 24-month-olds’ action selection in an imitation study. When infants of both age groups observed a confirming (Congruent) or unrelated (Pseudo-word: action demonstration was associated with novel verb-like cue) coherence, they selectively imitated the demonstrated action over the not demonstrated, alternative action, with no difference between these two conditions. These findings suggest that, as expected, infants’ action hierarchy was led by the demonstrated action when the verbal cue provided similar (Congruent) or no additional (Pseudo-word) intention-related information relative to a meaningful behavioral cue. These findings support the above-mentioned interpretation that enhanced motor activation during action observation may reflect a covert preparation for imitation (Study I). Interestingly, infants did not seem to benefit from the intention-highlighting effect of the verbal cue in Congruent, suggesting that the verbal cue had an unspecific (e.g., attention-guiding) effect on infants’ action selection. Contrary, when infants observed a contrasting (Incongruent) or clarifying (Failed-attempt: model failed to manipulate the object but verbally announced a certain action intention) coherence, their action selection varied with age and also varied across the course of the experiment (block 1 vs. block 2). More specifically, the 24-month-olds made stronger use of the verbal cue for their action selection in block 1 than did the 18-month-olds. However, while the 18-month-olds’ use of the verbal cue increased across blocks, particularly in Incongruent, the 24-month-olds’ use of the verbal cue decreased across blocks. Overall, these results suggest that, as expected, infants’ action hierarchy in Incongruent (both age groups) and Failed-attempt (only 24-month-olds) drew on the verbal action intention, because in both age groups, infants emulated the verbal intention about as often as they imitated the demonstrated action or even emulated the verbal action intention preferentially. Yet, these findings were confined to certain blocks. It may be argued that the younger age group had a harder time inferring and emulating the intended, yet never observed action, because this requirement is more demanding in cognitive and motor terms. These demands may explain why the 18-month-olds needed some time to take account of the verbal action intention. Contrary, it seems that the 24-month-olds, although demonstrating their principle capacity to take account of the verbal cue in block 1, lost trust in the model’s verbal cue, maybe because the verbal cue did not have predictive value for the model’s actual behavior. Supporting this interpretation, research on selective trust has demonstrated that already infants evaluate another’s reliability or competence, respectively, based on how that model handles familiar objects (behavioral reliability) or labels familiar objects (verbal reliability; for reviews, see Mills, 2013; Poulin-Dubois & Brosseau-Liard, 2016). Relatedly, imitation research has demonstrated that the interpersonal aspects of a social-learning situation gain increasing relevance for infants during the second year of life (Gellén & Buttelmann, 2019; Matheson, Moore, & Akhtar, 2013; Uzgiris, 1981). It may thus be argued that when the 24-month-olds were repeatedly faced with a verbally unreliable model, they de-evaluated the verbal cue as signaling the model’s action intention and instead relied more heavily on alternative cues such as the behavioral cue (Incongruent) or the action context (e.g., object affordances, salience; Failed-attempt). Infants’ first-action latencies were higher in Incongruent and Failed-attempt than in both Congruent and Pseudo-word, and were also higher in Failed-attempt than in Incongruent. These latency-findings thus indicate that situations involving a meaningful verbal cue that deviated from the behavioral cue are cognitively more demanding, resulting in a delayed initiation of a behavioral response. In sum, the findings of Study II suggest that both age groups were highly flexible in their integration of a verbal cue and behavioral cue. Moreover, our results do not indicate a general superiority of either cue. Instead, it seems to depend on the informational gain conveyed by the verbal cue whether it exerts a specific, intention-highlighting effect (Incongruent, Failed-attempt) or an unspecific (e.g., attention-guiding) effect (Congruent, Pseudo-word). Studies III and IV investigated the impact of another’s action-related emotional cues on 18-month-olds’ action selection. In Study III, infants observed a model, who demonstrated two actions on a novel object in direct succession, and who combined one of the two actions with a positive (happy) emotional expression and the other action with a negative (sad) emotional expression. As expected, infants imitated the positively emoted (PE) action more often than the negatively emoted (NE) action. This preference arose from an increase in infants’ readiness to perform the PE action from the baseline period (prior to the action demonstrations) to the test period (following the action demonstrations), rather than from a decrease in readiness to the perform the NE action. The positive cue thus had a stronger behavior-regulating effect than the negative cue. Notably, infants’ more general object-directed behavior in terms of first-touch latencies remained unaffected by the emotional cues’ valence, indicating that infants had linked the emotional cues specifically to the corresponding action and not the object as a whole (Repacholi, 2009). Also, infants’ looking times during the action demonstration did not differ significantly as a function of emotional valence and were characterized by a predominant attentional focus to the action/object rather than to the model’s face. Together with the findings on infants’ first-touch latencies, these results indicate a sensitivity for the notion that emotions can have very specific referents (referential specificity; Martin, Maza, McGrath, & Phelps, 2014). Together, Study III provided evidence for selective imitation based on another’s intention-related (particularly positive) emotional cues in an action-selection task, and thus indicates that infants’ action hierarchy flexibly responds to another’s emotional evaluation of observed actions. According to Repacholi (2009), we suggest that infants used the model’s emotional evaluation to re-appraise the corresponding action (effect), for instance in terms of desirability. Study IV followed up on Study III by investigating the role of the negative emotional cue for infants’ action selection in more detail. Specifically, we investigated whether a contrasting (negative) emotional cue alone would be sufficient to differentially rank the two actions along infants’ action hierarchy or whether instead infants require direct information about the model’s action intention (in the form of a confirming action-emotion pair) to align their action selection with the emotional cues. Also, we examined whether the absence of a direct behavior-regulating effect of the negative cue in Study III was due to the negative cue itself or to the concurrently available positive cue masking the negative cue’s potential effect. To this end, we split the demonstration of the two action-emotion pairs across two trials. In each trial, one action was thus demonstrated and emoted (PE, NE action), and one action was not demonstrated and un-emoted (UE action). For trial 1, we predicted that infants, who observed a PE action demonstration, would selectively imitate the PE action, whereas infants, who observed a NE action demonstration would selectively emulate the UE action. As to trial 2, we expected the complementary action-emotion pair to provide additional clarifying information as the model’s emotional evaluation of both actions, which should either lead to adaptive perseveration (if infants’ action selection in trial 1 had already drawn on the emotional cue) or adaptive change (if infants’ action selection in trial 1 signaled a disregard of the emotional cue). As to trial 1, our findings revealed that, as expected, infants imitated the PE action more often than they emulated the UE action. Like in Study III, this selectivity arose from an increase in infants’ propensity to perform the PE action from baseline to trial 1. Also like in Study III, infants performed the NE action about equally often in baseline and trial 1, which speaks against a direct behavior-regulating effect of the negative cue also when presented in isolation. However, after a NE action demonstration, infants emulated the UE action more often in trial 1 than in baseline, suggesting an indirect behavior-regulating effect of the negative cue. Yet, this indirect effect did not yield a selective emulation of the UE action, because infants performed both action alternatives about equally often in trial 1. Unexpectedly, infants’ action selection in trial 2 was unaffected by the emotional cue. Instead, infants perseverated their action selection of trial 1 in trial 2, irrespective of whether it was adaptive or non-adaptive with respect to the model’s emotional evaluation of the action. It seems that infants changed their strategy across trials, from an initial adherence to the emotional (particularly positive) cue, towards bringing about a salient action effect (Marcovich & Zelazo, 2009). In sum, Studies III and IV indicate a dynamic interplay of different action-selection strategies, depending on valence and presentation order. Apparently, at least in infancy, action reconstruction as one basis for selective action performance reaches its limits when infants can only draw on indirect intention-related information (i.e., which action should be avoided). Overall, our findings favor the action-reconstruction account (Csibra, 2007), according to which actions are flexibly organized along a hierarchy, depending on inferential processes based on extra-motor intention-related information. At the same time, the findings question the direct-matching hypothesis (Rizzolatti & Craighero, 2004), according to which the identification (and pursuit) of action goals hinges on a direct simulation of another’s behavioral cues. Based on the studies’ findings, a preliminary working model is introduced, which seeks to integrate the two theoretical accounts by conceptualizing the routes that activation induced by social cues may take to eventually influence an infant’s action selection. Our findings indicate that it is useful to strive a differentiated conceptualization of communicative cues, because they seem to operate at different places within the process of cue integration, depending on their potential to convey direct intention-related information. Moreover, we suggest that there is bidirectional exchange within each compound of adjacent sub-processes (i.e., between attention and encoding, and encoding and response), and between the compounds. Hence, our findings highlight the benefits of a multi-measures approach when studying the development of infants’ social-cognitive abilities, because it provides a more comprehensive picture how the concerted use of social cues from different domains influences infants’ processing and selection of instrumental actions. Finally, this thesis points to potential future directions to substantiate our current interpretation of the findings.. Moreover, an extension to additional kinds of coherence is suggested to get closer to infants’ everyday-world of experience. KW - infancy KW - social cognition KW - action processing KW - emotions KW - language KW - imitation KW - EEG Y1 - 2020 ER - TY - THES A1 - Fontana, Federica T1 - Antagonistic activities of Vegfr3/Flt4 and Notch1b fine-tune mechanosensitive signaling during zebrafish cardiac valvulogenesis N2 - Cardiac valves are essential for the continuous and unidirectional flow of blood throughout the body. During embryonic development, their formation is strictly connected to the mechanical forces exerted by blood flow. The endocardium that lines the interior of the heart is a specialized endothelial tissue and is highly sensitive to fluid shear stress. Endocardial cells harbor a signal transduction machinery required for the translation of these forces into biochemical signaling, which strongly impacts cardiac morphogenesis and physiology. To date, we lack a solid understanding on the mechanisms by which endocardial cells sense the dynamic mechanical stimuli and how they trigger different cellular responses. In the zebrafish embryo, endocardial cells at the atrioventricular canal respond to blood flow by rearranging from a monolayer to a double-layer, composed of a luminal cell population subjected to blood flow and an abluminal one that is not exposed to it. These early morphological changes lead to the formation of an immature valve leaflet. While previous studies mainly focused on genes that are positively regulated by shear stress, the mechanisms regulating cell behaviors and fates in cells that lack the stimulus of blood flow are largely unknown. One key discovery of my work is that the flow-sensitive Notch receptor and Krüppel-like factor (Klf) 2, one of the best characterized flow-regulated transcriptional factors, are activated by shear stress but that they function in two parallel signal transduction pathways. Each of these two pathways is essential for the rearrangement of atrioventricular cells into an immature double-layered valve leaflets. A second key discovery of my study is the finding that both Notch and Klf2 signaling negatively regulate the expression of the angiogenesis receptor Vegfr3/Flt4, which becomes restricted to abluminal endocardial cells of the valve leaflet. Within these cells, Flt4 downregulates the expressions of the cell adhesion proteins Alcam and VE-cadherin. A loss of Flt4 causes abluminal endocardial cells to ectopically express Notch, which is normally restricted to luminal cells, and impairs valve morphology. My study suggests that abluminal endocardial cells that do not experience mechanical stimuli loose Notch expression and this triggers expression of Flt4. In turn, Flt4 negatively regulates Notch on the abluminal side of the valve leaflet. These antagonistic signaling activities and fine-tuned gene regulatory mechanisms ultimately shape cardiac valve leaflets by inducing unique differences in the fates of endocardial cells. N2 - Herzklappen sind essentiell für den kontinuierlichen und gerichteten Blutfluss durch den Körper. Während der Embryonalentwicklung ist die Bildung der Herzklappen stark von vom Blutfluss generierten, mechanischen Kräften abhängig. Das Endokard, ein endotheliales Gewebe, das das Herz im Inneren auskleidet, reagiert sehr sensibel auf biomechanische Einwirkungen. Endokardzellen weisen eine Signaltransduktionsmaschinerie auf, welche die Umwandlung dieser Kräfte in biochemische und elektrische Signale ermöglicht und somit unverzichtbar für die Herzmorphogenese und -physiologie ist. Allerdings fehlt uns noch immer das Verständnis der Mechanismen, mit denen Endokardzellen dynamische, biomechanische Signale wahrnehmen und wie verschiedene zelluläre Antworten ausgelöst werden können. Im Zebrafischembryo reagieren Endokardzellen im atrioventrikulärem Kanal auf Blutfluss induzierte Schubspannung mit einer Umorganisation, wobei sich aus einer Einzelschicht an Zellen eine Doppelschicht bildet. Letztere besteht aus einer luminalen Zellpopulation, die dem Blutstrom ausgesetzt ist und einer abluminalen Population, der der Kontakt zum Blut fehlt. Diese initialen morphologischen Veränderungen führen zur Ausbildung des frühen Herzklappensegels. Bisherige Studien berichteten im Besonderen über Gene die positiv von einer veränderten Schubspannung in Endokardzellen reguliert werden. Allerdings sind die Mechanismen, die das Verhalten und die Spezifizierung von den Zellen regulieren, die nicht in Kontakt mit dem Blutfluss sind, weitgehend unbekannt. Eine meiner Schlüsselentdeckungen in dieser Arbeit ist, dass zwei der am besten charakterisierten, durch Blutfluss transkriptional regulierten Faktoren, der Notch Rezeptor und der Krüppel-like factor (Klf) 2, durch Schubspannung aktiviert werden. Dies funktioniert auf zwei parallelen mechanosensitiven Signaltransduktionswegen und beide Kaskaden sind essentiell für die Umorganisation der atrioventrikulären Zellen während der Bildung der frühen zweischichtigen Klappensegeln. Eine zweite wichtige Entdeckung meiner Studien ist, dass die Expression des angiogenen Faktors Vegfr3/Flt4, die auf abluminale Endokardzellen im frühen Klappensegel beschränk ist, von beiden Signalwegen, Notch und KLf2, negativ reguliert wird. Außerdem veringert Flt4 die Expression der Zelladhäsionsproteine Alcam und VE-cadherin in abluminalen Zellen und führt die Herzklappenmorphogenese herbei. Der Verlust von Flt4 wiederum führt zu einer ektopischen Expression von Notch in abluminalen Endokardzellen, welche sonst nur in luminalen Zellen auftritt. Daher zeigt meine Arbeit, dass abluminale Endokardzellen, die keinem mechanischem Reiz ausgesetzt sind, Notch herunterregulieren und damit die Expression von Flt4 auslösen. Flt4 wiederum blockiert dann zusätzlich den Notch Signalweg in dieser Zellpopulation. Diese antagonistischen Signalaktivitäten und fein abgestimmte Genregulationsmechanismen sorgen für Unterschiede in der Spezifizierung der Endokardzellen und formen so schließlich die Segelklappen im Herz. KW - heart development KW - cardiac valves KW - zebrafish KW - mechanosensation KW - Herzklappe KW - Herzentwicklung KW - Mechanosensation KW - Zebrafisch Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-487517 ER - TY - THES A1 - Meijer, Niels T1 - Asian dust, monsoons and westerlies during the Eocene N2 - The East Asian monsoons characterize the modern-day Asian climate, yet their geological history and driving mechanisms remain controversial. The southeasterly summer monsoon provides moisture, whereas the northwesterly winter monsoon sweeps up dust from the arid Asian interior to form the Chinese Loess Plateau. The onset of this loess accumulation, and therefore of the monsoons, was thought to be 8 million years ago (Ma). However, in recent years these loess records have been extended further back in time to the Eocene (56-34 Ma), a period characterized by significant changes in both the regional geography and global climate. Yet the extent to which these reconfigurations drive atmospheric circulation and whether the loess-like deposits are monsoonal remains debated. In this thesis, I study the terrestrial deposits of the Xining Basin previously identified as Eocene loess, to derive the paleoenvironmental evolution of the region and identify the geological processes that have shaped the Asian climate. I review dust deposits in the geological record and conclude that these are commonly represented by a mix of both windblown and water-laid sediments, in contrast to the pure windblown material known as loess. Yet by using a combination of quartz surface morphologies, provenance characteristics and distinguishing grain-size distributions, windblown dust can be identified and quantified in a variety of settings. This has important implications for tracking aridification and dust-fluxes throughout the geological record. Past reversals of Earth’s magnetic field are recorded in the deposits of the Xining Basin and I use these together with a dated volcanic ash layer to accurately constrain the age to the Eocene period. A combination of pollen assemblages, low dust abundances and other geochemical data indicates that the early Eocene was relatively humid suggesting an intensified summer monsoon due to the warmer greenhouse climate at this time. A subsequent shift from predominantly freshwater to salt lakes reflects a long-term aridification trend possibly driven by global cooling and the continuous uplift of the Tibetan Plateau. Superimposed on this aridification are wetter intervals reflected in more abundant lake deposits which correlate with highstands of the inland proto-Paratethys Sea. This sea covered the Eurasian continent and thereby provided additional moisture to the winter-time westerlies during the middle to late Eocene. The long-term aridification culminated in an abrupt shift at 40 Ma reflected by the onset of windblown dust, an increase in steppe-desert pollen, the occurrence of high-latitude orbital cycles and northwesterly winds identified in deflated salt deposits. Together, these indicate the onset of a Siberian high atmospheric pressure system driving the East Asian winter monsoon as well as dust storms and was triggered by a major sea retreat from the Asian interior. These results therefore show that the proto-Paratethys Sea, though less well recognized than the Tibetan Plateau and global climate, has been a major driver in setting up the modern-day climate in Asia. N2 - Die ostasiatischen Monsune prägen das heutige asiatische Klima, doch ihr geologischer Ursprung und ihre Antriebsmechanismen sind nach wie vor umstritten. Der südöstliche Sommermonsun bringt Feuchtigkeit, während der nordwestliche Wintermonsun Staub aus dem trockenen asiatischen Inland aufwirbelt und das chinesische Lössplateau bildet. Der Ursprung dieses Lösses und damit des Monsuns wurde vor 8 Millionen Jahren vermutet (Ma). In den letzten Jahren sind diese Lößablagerungen jedoch weiter in das Eozän (56-34 Ma) zurückverlegt worden, einer Periode, die durch bedeutende Änderungen sowohl in der regionalen Geographie als auch im globalen Klima gekennzeichnet ist. Inwieweit diese Rekonfigurationen die atmosphärische Zirkulation antrieben und ob es sich bei den lößartigen Sedimenten um monsunartige Ablagerungen handelt, bleibt jedoch umstritten. In dieser Dissertation untersuche ich die terrestrischen Ablagerungen des Xining-Beckens, die zuvor als Löss aus dem Eozän identifiziert wurden, um die paläo-umweltbedingte Entwicklung der Region abzuleiten und die geologischen Prozesse zu identifizieren, die das asiatische Klima geprägt haben. Ich überprüfe die Staubablagerungen im geologischen Archiv und komme zu dem Schluss, dass diese durch eine Mischung aus windgetriebenen und wassergelagerten Sedimenten dargestellt werden, im Gegensatz zu dem rein windgetriebenen Material, das als Löß bekannt ist. Doch durch die Verwendung einer Kombination der oberflächenmorphologien von Quartz, Herkunftsmerkmalen und unterscheidenden Korngrößenverteilungen kann windgetriebener Staub in einer Vielzahl von Umgebungen identifiziert und quantifiziert werden. Dies hat wichtige Auswirkungen auf die Nachverfolgung der Aridifizierung und der Staubflüsse in dem gesamten geologischen Archiv. Frühere Umkehrungen des Erdmagnetfeldes werden in den Ablagerungen des Xining-Beckens aufgezeichnet und ich verwende diese zusammen mit einer datierten vulkanischen Ascheschicht, um das Alter genau auf die Eozän-Periode einzugrenzen. Eine Kombination aus Pollenansammlungen, geringen Staubhäufigkeiten und anderen geochemischen Daten deutet darauf hin, dass das frühe Eozän relativ feucht war, was auf einen verstärkten Sommermonsun aufgrund des wärmeren Treibhausklimas zu dieser Zeit hinweist. Eine anschließende Verschiebung von überwiegend Süßwasser zu Salzseen spiegelt einen langfristigen Aridifizierungstrend wider, der möglicherweise durch die globale Abkühlung und die kontinuierliche Hebung des Tibetischen Plateaus angetrieben wurde. Überlagert wird diese Aridifizierung von feuchteren Intervallen, die durch eine Zunahme in Seeablagerungen gekennzeichnet werden und mit den Hochständen des inländischen proto-Paratethys-Meeres korrelieren. Dieses Meer bedeckte den eurasischen Kontinent und versorgte dadurch die winterlichen Westwinde mit zusätzlicher Feuchtigkeit im mittleren bis späten Eozän. T2 - Asiatischer Staub, Monsune und Westwind während des Eozäns KW - Paleoclimatology KW - Asia KW - Eocene KW - Stratigraphy KW - Asien KW - Stratigrafie KW - Monsoon KW - Monsun KW - Paläoklimatologie KW - Eozän Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-488687 ER - TY - THES A1 - Weißhuhn, Peter T1 - Assessing biotope vulnerability to landscape changes T1 - Bewertung der Vulnerabilität von Biotopen gegenüber Landschaftsveränderungen N2 - Largescale patterns of global land use change are very frequently accompanied by natural habitat loss. To assess the consequences of habitat loss for the remaining natural and semi-natural biotopes, inclusion of cumulative effects at the landscape level is required. The interdisciplinary concept of vulnerability constitutes an appropriate assessment framework at the landscape level, though with few examples of its application for ecological assessments. A comprehensive biotope vulnerability analysis allows identification of areas most affected by landscape change and at the same time with the lowest chances of regeneration. To this end, a series of ecological indicators were reviewed and developed. They measured spatial attributes of individual biotopes as well as some ecological and conservation characteristics of the respective resident species community. The final vulnerability index combined seven largely independent indicators, which covered exposure, sensitivity and adaptive capacity of biotopes to landscape changes. Results for biotope vulnerability were provided at the regional level. This seems to be an appropriate extent with relevance for spatial planning and designing the distribution of nature reserves. Using the vulnerability scores calculated for the German federal state of Brandenburg, hot spots and clusters within and across the distinguished types of biotopes were analysed. Biotope types with high dependence on water availability, as well as biotopes of the open landscape containing woody plants (e.g., orchard meadows) are particularly vulnerable to landscape changes. In contrast, the majority of forest biotopes appear to be less vulnerable. Despite the appeal of such generalised statements for some biotope types, the distribution of values suggests that conservation measures for the majority of biotopes should be designed specifically for individual sites. Taken together, size, shape and spatial context of individual biotopes often had a dominant influence on the vulnerability score. The implementation of biotope vulnerability analysis at the regional level indicated that large biotope datasets can be evaluated with high level of detail using geoinformatics. Drawing on previous work in landscape spatial analysis, the reproducible approach relies on transparent calculations of quantitative and qualitative indicators. At the same time, it provides a synoptic overview and information on the individual biotopes. It is expected to be most useful for nature conservation in combination with an understanding of population, species, and community attributes known for specific sites. The biotope vulnerability analysis facilitates a foresighted assessment of different land uses, aiding in identifying options to slow habitat loss to sustainable levels. It can also be incorporated into planning of restoration measures, guiding efforts to remedy ecological damage. Restoration of any specific site could yield synergies with the conservation objectives of other sites, through enhancing the habitat network or buffering against future landscape change. Biotope vulnerability analysis could be developed in line with other important ecological concepts, such as resilience and adaptability, further extending the broad thematic scope of the vulnerability concept. Vulnerability can increasingly serve as a common framework for the interdisciplinary research necessary to solve major societal challenges. N2 - Weltweit wurden in den vergangenen Jahrzehnten massive Veränderungen in der Landnutzung vorgenommen. Diese gingen meistens mit dem Verlust natürlicher Habitate einher. Um die Folgen auf die verbliebenen naturnahen Biotope zu bewerten, sind Analyseinstrumente notwendig, die neben einzelnen Flächen auch kumulative Effekte auf der Landschaftsebene einbeziehen. Das interdisziplinäre Konzept der Vulnerabilität kann einem solchen Analyseinstrument den Rahmen bieten. Bisher wurde es kaum für die Bewertung ökologischer Systeme herangezogen. Dabei kann eine flächendeckende Biotop-Vulnerabilitätsanalyse jene Gebiete identifizieren, die vom Landschaftswandel am stärksten betroffen sind und die gleichzeitig die geringsten Erholungsaussichten aufweisen. Dazu wurde eine Reihe ökologischer Indikatoren gesichtet und entwickelt, um die drei Vulnerabilitätsgrößen Exposition, Sensitivität und Anpassungskapazität abzudecken. Dabei wurden die Lagebeziehungen einzelner Biotope sowie die Eigenschaften der jeweils ansässigen Artengemeinschaft ausgewertet. Der errechnete Index kombiniert sieben voneinander weitestgehend unabhängige Indikatoren und stellt eine Übersicht der Biotop-Vulnerabilität dar. Liegt eine flächendeckende Vulnerabilitätsbewertung vor, können räumliche Häufungen von hohen Werten sowie die Verteilung der besonders hohen und besonders niedrigen Werte über die Biotoptypen hinweg analysiert werden. Dies erscheint besonders sinnvoll für Flächengrößen mit Relevanz für die Raumplanung und die Verteilung der Naturschutzflächen. Es stellte sich heraus, dass in der Planungsregion Brandenburg vor allem die Biotope mit hoher Abhängigkeit von der Wasserverfügbarkeit, sowie die Gehölze enthaltenden Offenlandbiotope (z.B. Streuobstwiesen) besonders vulnerabel gegenüber Landschaftsveränderungen sind. Im Gegensatz dazu erscheint die Mehrheit der Waldbiotope weniger verwundbar zu sein. Trotz der Möglichkeit zur Ableitung solcher verallgemeinerten Aussagen für einige Biotoptypen legt die Werteverteilung nahe, Naturschutzmaßnahmen mehrheitlich spezifisch für einzelne Flächen zu entwerfen. Größe, Form und räumlicher Kontext einzelner Biotopflächen üben zusammengenommen häufig einen dominanten Einfluss auf die Vulnerabilität gegenüber Landschaftsveränderungen aus. Die Demonstration der Biotop-Vulnerabilitätsanalyse auf regionaler Ebene zeigt, dass mit Methoden der Geoinformatik auch große Biotop-Datensätze detailliert ausgewertet werden können. Die damit erzielte naturschutzfachliche Analyse basiert auf transparent berechneten qualitativen und quantitativen Indikatoren und ist damit vollständig nachvollziehbar. Sie bietet gleichzeitig einen großräumigen Überblick sowie Informationen zu den einzelnen Biotopflächen. Mit der Vulnerabilitätsanalyse von Biotopen wird die vorausschauende Abwägung zwischen verschiedenen Landnutzungen erleichtert. Dies kann dazu beitragen, dass der Habitatverlust in Zukunft auf ein nachhaltigeres Maß gebremst wird. Auch kann eine solche Biotopbewertung in die Planung von Renaturierungsmaßnahmen einfließen, um ökologische Schäden zunächst dort zu beheben, wo große Synergieeffekte im Biotopverbund und mit anderen Naturschutzzielen zu erwarten sind. Die Biotop-Vulnerabilitätsanalyse konnte im Einklang mit wichtigen ökologischen Konzepten wie Resilienz und Anpassungsfähigkeit entwickelt werden. Sie erweitert damit den ohnehin breiten thematischen Anwendungsbereich des Vulnerabilitätskonzepts. Somit kann Vulnerabilität zunehmend als Brückenkonzept dienen und der zur Lösung der großen gesellschaftlichen Herausforderungen notwendigen interdisziplinären Forschung einen gemeinsamen Rahmen geben. KW - vulnerability KW - biotope KW - habitat KW - index KW - landscape KW - spatial ecology KW - landscape ecology KW - Vulnerabilität KW - Biotop KW - Habitat KW - Index KW - Landschaft KW - raumbezogene Ökologie KW - Landschaftsökologie Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-442777 ER - TY - THES A1 - Balt, Miriam T1 - Assessment of early numeracy development BT - contributions to designing a progression-based instrument to monitor learning N2 - Early numeracy is one of the strongest predictors for later success in school mathematics (e.g., Duncan et al., 2007). The main goal of first grade mathematics teachers should therefore be to provide learning opportunities that enable all students to develop sound early numeracy skills. Developmental models, or learning progressions, can describe how early numerical understanding typically develops. Assessments that are aligned to empirically validated learning progressions can support teachers to understand their students learning better and target instruction accordingly. To date, there have been no progression-based instruments made available for German teachers to monitor their students’ progress in the domain of early numeracy. This dissertation contributes to the design of such an instrument. The first study analysed the suitability of early numeracy assessments currently used in German primary schools at school entry to identify students’ individual starting points for subsequent progress monitoring. The second study described the development of progression-based items and investigated the items in regards to main test quality criteria, such as reliability, validity, and test fairness, to find a suitable item pool to build targeted tests. The third study described the construction of the progress monitoring measure, referred to as the learning progress assessment (LPA). The study investigated the extent to which the LPA was able to monitor students’ individual learning progress in early numeracy over time. The results of the first study indicated that current school entry assessments were not able to provide meaningful information about the students’ initial learning status. Thus, the MARKO-D test (Ricken, Fritz, & Balzer, 2013) was used to determine the students’ initial numerical understanding in the other two studies, because it has been shown to be an effective measure of conceptual numerical understanding (Fritz, Ehlert, & Leutner, 2018). Both studies provided promising evidence for the quality of the LPA and its ability to detect changes in numerical understanding over the course of first grade. The studies of this dissertation can be considered an important step in the process of designing an empirically validated instrument that supports teachers to monitor their students’ early numeracy development and to adjust their teaching accordingly to enhance school achievement. N2 - Frühes mengen- und zahlenbezogenes Wissen (early numeracy) ist einer der stärksten Prädiktoren für spätere Lernerfolge in der Schulmathematik. Hauptziel der Mathematiklehrkräfte der ersten Klassen sollte es daher sein, Lernmöglichkeiten anzubieten, die es allen Schüler*innen erlauben, fundierte mengen- und zahlenbezogene Fähigkeiten zu erwerben. Entwicklungsmodelle (learning progressions) beschreiben, wie sich frühes mengen- und zahlenbezogenes Verständnis typischerweise entwickelt. Diagnostische Tests (assessments), die sich an empirisch validierten Entwicklungsmodellen orientieren, können Lehrkräfte dabei unterstützen, die Leistungen ihrer Schüler*innen besser einzuschätzen und den Unterricht entsprechend darauf anzupassen. Bislang gibt es keine entwicklungsbasierten Instrumente, mit denen deutsche Lehrkräfte die Lernfortschritte ihrer Schüler*innen im Bereich des frühen mengen- und zahlenbezogenen Wissens erfassen können. Diese Dissertation trägt zur Gestaltung eines solchen Instruments bei. Die erste Studie untersucht, inwiefern sich derzeit an deutschen Grundschulen eingesetzte Instrumente zur mathematischen Schuleingangs-diagnostik dafür eignen, das individuelle Ausgangsniveau der Schüler*innen für eine anschließende Lernverlaufsdiagnostik zu bestimmen. In der zweiten Studie wird die Konstruktion von entwicklungsorientierten Items beschrieben. Es wurde untersucht, inwiefern die Items die Testgütekriterien Reliabilität, Validität und Testfairness erfüllen, um einen Item-Pool aufzubauen, der für adaptives Testen eingesetzt werden kann. Die dritte Studie beschreibt die Konstruktion einer Lernverlaufsdiagnostik, learning progress assessment genannt (LPA) und untersucht, inwieweit das LPA die individuellen Lernfortschritte der Schüler*innen hinsichtlich früher arithmetischer Konzepte im Verlauf der ersten Klasse erfassen kann. Die Ergebnisse der ersten Studie zeigten, dass die derzeit an den Grundschulen eingesetzten Verfahren zur Schuleingangsdiagnostik keine aussagekräftigen Informationen über die Erfassung von Lernausgangslagen zulassen. Daher wurde in den beiden anderen Studien der MARKO-D verwendet, um das arithmetische Wissen der Schüler*innen zum Schulanfang zu erfassen. Beide Studien liefern belastbare Evidenz für die Qualität des LPA und dessen Fähigkeit, Veränderung hinsichtlich arithmetischen Wissens im Laufe der ersten Klasse zu messen. Die in dieser Dissertation vorgestellten Studien können als wichtiger Schritt zur Entwicklung eines empirisch validierten Instruments betrachtet werden, das Lehrkräfte dabei unterstützt, die Entwicklung frühen mengen- und zahlenbezogenen Wissens zu erfassen und ihren Unterricht entsprechend anzupassen und damit den Lernerfolg der Schüler*innen zu fördern. KW - assessment KW - learning progression KW - early numeracy KW - primary school KW - mathematics KW - Diagnostik KW - Lernverlauf KW - numerische Basisfähigkeiten KW - Grundschule KW - Mathematik Y1 - 2020 ER - TY - THES A1 - Schinköth, Michaela T1 - Automatic affective reactions to exercise-related stimuli BT - towards a better understanding of exercise motivation BT - für ein besseres Verständnis der Sportmotivation N2 - Even though the majority of individuals know that exercising is healthy, a high percentage struggle to achieve the recommended amount of exercise. The (social-cognitive) theories that are commonly applied to explain exercise motivation refer to the assumption that people base their decisions mainly on rational reasoning. However, behavior is not only bound to reflection. In recent years, the role of automaticity and affect for exercise motivation has been increasingly discussed. In this dissertation, central assumptions of the affective–reflective theory of physical inactivity and exercise (ART; Brand & Ekkekakis, 2018), an exercise-specific dual-process theory that emphasizes the role of a momentary automatic affective reaction for exercise-decisions, were examined. The central aim of this dissertation was to investigate exercisers and non-exercisers automatic affective reactions to exercise-related stimuli (i.e., type-1 process). In particular, the two components of the ART’s type-1 process, that are, automatic associations with exercise and the automatic affective valuation to exercise, were under study. In the first publication (Schinkoeth & Antoniewicz, 2017), research on automatic (evaluative) associations with exercise was summarized and evaluated in a systematic review. The results indicated that automatic associations with exercise appeared to be relevant predictors for exercise behavior and other exercise-related variables, providing evidence for a central assumption of the ART’s type-1 process. Furthermore, indirect methods seem to be suitable to assess automatic associations. The aim of the second publication (Schinkoeth, Weymar, & Brand, 2019) was to approach the somato-affective core of the automatic valuation of exercise using analysis of reactivity in vagal HRV while viewing exercise-related pictures. Results revealed that differences in exercise volume could be regressed on HRV reactivity. In light of the ART, these findings were interpreted as evidence of an inter-individual affective reaction elicited at the thought of exercise and triggered by exercise-stimuli. In the third publication (Schinkoeth & Brand, 2019, subm.), it was sought to disentangle and relate to each other the ART’s type-1 process components—automatic associations and the affective valuation of exercise. Automatic associations to exercise were assessed with a recoding-free variant of an implicit association test (IAT). Analysis of HRV reactivity was applied to approach a somatic component of the affective valuation, and facial reactions in a facial expression (FE) task served as indicators of the automatic affective reaction’s valence. Exercise behavior was assessed via self-report. The measurement of the affective valuation’s valence with the FE task did not work well in this study. HRV reactivity was predicted by the IAT score and did also statistically predict exercise behavior. These results thus confirm and expand upon the results of publication two and provide empirical evidence for the type-1 process, as defined in the ART. This dissertation advances the field of exercise psychology concerning the influence of automaticity and affect on exercise motivation. Moreover, both methodical implications and theoretical extensions for the ART can be derived from the results. N2 - Obwohl die meisten Menschen wissen, dass Sport gesund ist, hat ein hoher Prozentsatz Mühe, die empfohlenen Bewegungsumfänge zu erreichen. Sozial-kognitive Theorien, die üblicherweise zur Erklärung von Sportmotivation angewendet werden, stützen sich auf die Annahme, dass Menschen ihre Entscheidungen hauptsächlich auf Grund rationaler Überlegungen treffen. Unser Verhalten ist jedoch nicht immer rational. In den letzten Jahren ist die Rolle von Automatizität und Affekt für die Sportmotivation daher zunehmend diskutiert worden. In dieser Dissertation wurden zentrale Annahmen der affective–reflective theory zur Erklärung von körperlicher Inaktivität und Sporttreiben (ART; Brand & Ekkekakis, 2018), einer sportspezifischen Zwei-Prozesstheorie, die die Rolle einer momentanen automatischen affektiven Reaktion für Sportentscheidungen betont, überprüft. Das zentrale Ziel dieser Dissertation war die Untersuchung automatisch affektiver Reaktionen von Sportlern und Nicht-Sportlern auf sportbezogene Stimuli (i.e., Typ-1-Prozess). Insbesondere wurden die beiden in der ART beschrieben Komponenten dieser automatisch affektiven Reaktion, d.h. die automatischen Assoziationen zu Sport und die automatische affektive Valuation von Sport, untersucht. In der ersten Publikation (Schinkoeth & Antoniewicz, 2017) wurde die Forschung zu automatischen Assoziationen zu Sport in einem systematischen Review zusammengefasst und evaluiert. Die Ergebnisse deuteten darauf hin, dass automatische Assoziationen relevante Prädiktoren für das Sportverhalten und andere sportbezogene Variablen zu sein scheinen, was den Beweis für eine zentrale Annahme des Typ-1-Prozesses der ART lieferte. Darüber hinaus scheinen indirekte Methoden geeignet zu sein um automatische Assoziationen zu messen. Das Ziel der zweiten Publikation (Schinkoeth, Weymar, & Brand, 2019) war es, sich dem somato-affektiven Kern der automatischen Valuation von Sport mittels Analyse der Reaktivität der vagalen HRV bei Betrachtung von Sportbildern zu nähern. Die Ergebnisse zeigten, dass die HRV-Reaktivität Unterschiede im Sportumfang hervorsagen konnte. Im Licht der ART wurden diese Befunde als Hinweis auf eine interindividuelle affektive Reaktion interpretiert, die beim bloßen Gedanken an Sport hervorgerufen und durch die Sportbilder ausgelöst wurde. In der dritten Publikation (Schinkoeth & Brand, 2019, subm.) wurde versucht, die Typ-1-Prozesskomponenten der ART - automatische Assoziationen und die affektive Valuation von Sport - zu trennen und in Beziehung zueinander zu setzen. Automatische Assoziationen zu Sport wurden mit einer rekodierungsfreien Variante eines impliziten Assoziationstests (IAT) gemessen. Die Analyse der HRV-Reaktivität wurde genutzt, um sich einer somatischen Komponente der affektiven Valuation zu nähern, und Gesichtsreaktionen in einer Mimikaufgabe (GR) dienten als Indikatoren für die Valenz der automatischen affektiven Reaktion. Das Bewegungsverhalten wurde mittels Selbstbericht bewertet. Die Messung der Valenz der affektiven Valuation mit der GR-Aufgabe funktionierte in dieser Studie nicht. Es konnte aber gezeigt werden, dass die HRV-Reaktivität durch den IAT-Score hervorgesagt werden konnte und wiederum das selbstberichtete Sportverhalten statistisch vorhersagen konnte. Diese Ergebnisse bestätigen und erweitern somit die Ergebnisse der Publikation zwei und liefern empirische Evidenz für den Typ-1-Prozess, wie er in der ART definiert ist. Die Ergbenisse dieser Dissertation tragen dazu bei die Forschung Rund um den Einfluss von Automatizität und Affekt auf die Sportmotivation entscheidend voran zu treiben. Darüber hinaus lassen sich aus den Ergebnissen sowohl methodische Implikationen als auch theoretische Erweiterungen für die ART ableiten. T2 - Automatisch affektive Reaktionen auf Sportstimuli KW - exercise KW - motivation KW - affect KW - automatic KW - dual-process KW - Sport KW - Motivation KW - Affekt KW - Automatizität KW - Zwei-Prozess Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-471115 ER - TY - THES A1 - AL-Rawi, Shadha T1 - Biochemical studies to determine the role of Early Starvation 1 (ESV1) protein and its homologue Like-Early Starvation 1 (LESV) during starch degradation N2 - Depending on the biochemical and biotechnical approach, the aim of this work was to understand the mechanism of protein-glucan interactions in regulation and control of starch degradation. Although starch degradation starts with the phosphorylation process, the mechanisms by which this process is controlling and adjusting starch degradation are not yet fully understood. Phosphorylation is a major process performed by the two dikinases enzymes α-glucan, water dikinase (GWD) and phosphoglucan water dikinase (PWD). GWD and PWD enzymes phosphorylate the starch granule surface; thereby stimulate starch degradation by hydrolytic enzymes. Despite these important roles for GWD and PWD, so far the biochemical processes by which these enzymes are able to regulate and adjust the rate of phosphate incorporation into starch during the degradation process haven‘t been understood. Recently, some proteins were found associated with the starch granule. Two of these proteins are named Early Starvation Protein 1 (ESV1) and its homologue Like-Early Starvation Protein 1 (LESV). It was supposed that both are involved in the control of starch degradation, but their function has not been clearly known until now. To understand how ESV1 and LESV-glucan interactions are regulated and affect the starch breakdown, it was analyzed the influence of ESV1 and LESV proteins on the phosphorylating enzyme GWD and PWD and hydrolysing enzymes ISA, BAM, and AMY. However, the analysis determined the location of LESV and ESV1 in the chloroplast stroma of Arabidopsis. Mass spectrometry data predicted ESV1and LESV proteins as a product of the At1g42430 and At3g55760 genes with a predicted mass of ~50 kDa and ~66 kDa, respectively. The ChloroP program predicted that ESV1 lacks the chloroplast transit peptide, but it predicted the first 56 amino acids N-terminal region as a chloroplast transit peptide for LESV. Usually, the transit peptide is processed during transport of the proteins into plastids. Given that this processing is critical, two forms of each ESV1 and LESV were generated and purified, a full-length form and a truncated form that lacks the transit peptide, namely, (ESV1and tESV1) and (LESV and tLESV), respectively. Both protein forms were included in the analysis assays, but only slight differences in glucan binding and protein action between ESV1 and tESV1 were observed, while no differences in the glucan binding and effect on the GWD and PWD action were observed between LESV and tLESV. The results revealed that the presence of the N-terminal is not massively altering the action of ESV1 or LESV. Therefore, it was only used the ESV1 and tLESV forms data to explain the function of both proteins. However, the analysis of the results revealed that LESV and ESV1 proteins bind strongly at the starch granule surface. Furthermore, not all of both proteins were released after their incubation with starches after washing the granules with 2% [w/v] SDS indicates to their binding to the deeper layers of the granule surface. Supporting of this finding comes after the binding of both proteins to starches after removing the free glucans chains from the surface by the action of ISA and BAM. Although both proteins are capable of binding to the starch structure, only LESV showed binding to amylose, while in ESV1, binding was not observed. The alteration of glucan structures at the starch granule surface is essential for the incorporation of phosphate into starch granule while the phosphorylation of starch by GWD and PWD increased after removing the free glucan chains by ISA. Furthermore, PWD showed the possibility of starch phosphorylation without prephosphorylation by GWD. Biochemical studies on protein-glucan interactions between LESV or ESV1 with different types of starch showed a potentially important mechanism of regulating and adjusting the phosphorylation process while the binding of LESV and ESV1 leads to altering the glucan structures of starches, hence, render the effect of the action of dikinases enzymes (GWD and PWD) more able to control the rate of starch degradation. Despite the presence of ESV1 which revealed an antagonistic effect on the PWD action as the PWD action was decreased without prephosphorylation by GWD and increased after prephosphorylation by GWD (Chapter 4), PWD showed a significant reduction in its action with or without prephosphorylation by GWD in the presence of ESV1 whether separately or together with LESV (Chapter 5). However, the presence of LESV and ESV1 together revealed the same effect compared to the effect of each one alone on the phosphorylation process, therefore it is difficult to distinguish the specific function between them. However, non-interactions were detected between LESV and ESV1 or between each of them with GWD and PWD or between GWD and PWD indicating the independent work for these proteins. It was also observed that the alteration of the starch structure by LESV and ESV1 plays a role in adjusting starch degradation rates not only by affecting the dikinases but also by affecting some of the hydrolysing enzymes since it was found that the presence of LESV and ESV1leads to the reduction of the action of BAM, but does not abolish it. N2 - Ziel dieser Arbeit war es, den Mechanismus der Protein-Glucan-Wechselwirkungen bei der Regulation und Kontrolle des Stärkeabbaus zu verstehen. Der Stärkeabbau beginnt mit dem Phosphorylierungsprozess, der von den beiden Dikinasen, der a-Glucan, Wasserdikinase (GWD) und der Phosphoglucanwasserdikinase (PWD) durchgeführt wird. Kürzlich wurden einige Proteine gefunden, die mit dem Stärkegranulum assoziiert sind. Zwei dieser Proteine heißen Early Starvation 1 (ESV1) und das Homolog Like-Early Starvation (LESV), Es wurde vorgeschlagen, dass beide an der Kontrolle des Stärkeabbaus beteiligt sind, aber ihre Funktion ist bisher nicht bekannt. Um zu verstehen, wie ESV1- und LESV-Glucan-Wechselwirkungen reguliert werden und den Stärkeabbau beeinflussen, wurde der Einfluss der beiden Proteine auf die Phosphorylierungsenzyme GWD und PWD, sowie die Hydrolasen isoamylase, betaamylase, und alpha-amylase ntersucht. Dabei ergab die Analyse, dass LESV und ESV1 nicht nur stark an der Oberfläche, sondern auch in den tieferen Schichten der Stärkegranula binden. Obwohl beide Proteine in der Lage sind, an die Stärkestruktur zu binden, zeigte nur LESV eine Bindung an Amylose, während für ESV1 keine Bindung beobachtet werden konnte. Die Veränderung der Glucanstrukturen an der Oberfläche der Stärkekörner ist für den Einbau von Phosphat wesentlich, so nahm beispielsweise die Phosphorylierung der Stärke durch GWD und PWD nach Entfernung der freien Glucanketten mittels ISA zu. Darüber hinaus konnte ebenso gezeigt werden, dass PWD auch ohne eine Präphosphorylierung durch GWD die Glucosyleinheiten innerhalb der Stärke phosphorylieren kann. Die Bindung von LESV und ESV1 führt zu einer Veränderung der Glucanstrukturen von Stärken, wodurch die Aktivität der Dikinasen (GWD und PWD) und somit die Geschwindigkeit des Stärkeabbaus wahrscheinlich besser gesteuert werden kann. Es wurden keine Wechselwirkungen zwischen LESV und ESV1 oder zwischen jedem von ihnen mit GWD und PWD oder zwischen GWD und PWD festgestellt, was auf die unabhängige Arbeit von diesen Proteinen hinweist. Es wurde auch beobachtet, dass die Modifikation der Stärkestruktur durch LESV und ESV1 eine Rolle bei der Anpassung der Stärkeabbauraten spielt, nicht nur durch Beeinflussung der Dikinasen, sondern auch durch die Beeinflussung einiger hydrolysierender Enzyme wie BAM. Den so zeigte die Amylase eine eindeutige Reduktion ihrer katalytischen Wirkung in Präsenz von LESV und ESV1. Daraus resumierend kann davon ausgegangen werden, dass die beiden Proteine ESV1 und LESV für die Feinregulation des Stärkeabbaus von höchster Relevanz sind. T2 - Biochemische Studien zur Bestimmung der Rolle des ESV1-Proteins (Early Starvation 1) und seines Homologen Like-Early Starvation 1 (LESV) während des Stärkeabbaus KW - Early starvation protein KW - Like-Early starvation protein KW - Glucan water dikinase KW - Phosphoglucan water dikinase KW - Phosphorylation process KW - Starch metabolism KW - Early Starvation 1 KW - Glucan-Wasser-Dikinase KW - Like-Early Starvation 1 KW - Phosphoglucan-Wasser-Dikinase KW - Phosphorylierungsprozess KW - Stärkestoffwechsel Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-483956 ER - TY - THES A1 - Friese, André T1 - Biogeochemistry of ferruginous sediments of Lake Towuti, Sulawesi, Indonesia N2 - Ferruginous conditions were a prominent feature of the oceans throughout the Precambrian Eons and thus throughout much of Earth’s history. Organic matter mineralization and diagenesis within the ferruginous sediments that deposited from Earth’s early oceans likely played a key role in global biogeochemical cycling. Knowledge of organic matter mineralization in ferruginous sediments, however, remains almost entirely conceptual, as modern analogue environments are extremely rare and largely unstudied, to date. Lake Towuti on the island of Sulawesi, Indonesia is such an analogue environment and the purpose of this PhD project was to investigate the rates and pathways of organic matter mineralization in its ferruginous sediments. Lake Towuti is the largest tectonic lake in Southeast Asia and is hosted in the mafic and ultramafic rocks of the East Sulawesi Ophiolite. It has a maximum water depth of 203 m and is weakly thermally stratified. A well-oygenated surface layer extends to 70 m depth, while waters below 130 m are persistently anoxic. Intensive weathering of the ultramafic catchment feeds the lake with large amounts of iron(oxy)hydroxides while the runoff contains only little sulfate, leading to sulfate-poor (< 20 µM) lake water and anoxic ferruginous conditions below 130 m. Such conditions are analogous to the ferruginous water columns that persisted throughout much of the Archean and Proterozoic eons. Short (< 35 cm) sediment cores were collected from different water depths corresponding to different bottom water redox conditions. Also, a drilling campaign of the International Continental Scientific Drilling Program (ICDP) retrieved a 114 m long sediment core dedicated for geomicrobiological investigations from a water depth of 153 m, well below the depth of oxygen penetration at the time of sampling. Samples collected from these sediment cores form the fundament of this thesis and were used to perform a suite of biogeochemical and microbiological analyses. Geomirobiological investigations depend on uncontaminated samples. However, exploration of subsurface environments relies on drilling, which requires the use of a drilling fluid. Drilling fluid infiltration during drilling can not be avoided. Thus, in order to trace contamination of the sediment core and to identify uncontaminated samples for further analyses a simple and inexpensive technique for assessing contamination during drilling operations was developed and applied during the ICDP drilling campaign. This approach uses an aqeous fluorescent pigment dispersion commonly used in the paint industry as a particulate tracer. It has the same physical properties as conventionally used particulate tracers. However, the price is nearly four orders of magnitude lower solving the main problem of particulate tracer approaches. The approach requires only a minimum of equipment and allows for a rapid contamination assessment potentially even directly on site, while the senstitivity is in the range of already established approaches. Contaminated samples in the drill core were identified and not included for further geomicrobiological investigations. Biogeochemical analyses of short sediment cores showed that Lake Towutis sediments are strongly depleted in electron acceptors commonly used in microbial organic matter mineralization (i.e. oxygen, nitrate, sulfate). Still, the sediments harbor high microbial cell densities, which are a function of redox conditions of Lake Towuti’s bottom water. In shallow water depths bottom water oxygenation leads to a higher input of labile organic matter and electron acceptors like sulfate and iron, which promotes a higher microbial abundance. Microbial analyses showed that a versatile microbial community with a potential to perform metabolisms related to iron and sulfate reduction, fermentation as well as methanogenesis inhabits Lake Towuti’s surface sediments. Biogeochemical investigations of the upper 12 m of the 114 m sediment core showed that Lake Towuti’s sediment is extremely rich in iron with total concentrations up to 2500 µmol cm-3 (20 wt. %), which makes it the natural sedimentary environment with the highest total iron concentrations studied to date. In the complete or near absence of oxygen, nitrate and sulfate, organic matter mineralization in ferruginous sediments would be expected to proceed anaerobically via the energetically most favorable terminal electron acceptors available - in this case ferric iron. Astonishingly, however, methanogenesis is the dominant (>85 %) organic matter mineralization process in Lake Towuti’s sediment. Reactive ferric iron known to be available for microbial iron reduction is highly abundant throughout the upper 12 m and thus remained stable for at least 60.000 years. The produced methane is not oxidized anaerobically and diffuses out of the sediment into the water column. The proclivity towards methanogenesis, in these very iron-rich modern sediments, implies that methanogenesis may have played a more important role in organic matter mineralization thoughout the Precambrian than previously thought and thus could have been a key contributor to Earth’s early climate dynamics. Over the whole sequence of the 114 m long sediment core siderites were identified and characterized using high-resolution microscopic and spectroscopic imaging together with microchemical and geochemical analyses. The data show early diagenetic growth of siderite crystals as a response to sedimentary organic matter mineralization. Microchemical zoning was identified in all siderite crystals. Siderite thus likely forms during diagenesis through growth on primary existing phases and the mineralogical and chemical features of these siderites are a function of changes in redox conditions of the pore water and sediment over time. Identification of microchemical zoning in ancient siderites deposited in the Precambrian may thus also be used to infer siderite growth histories in ancient sedimentary rocks including sedimentary iron formations. N2 - Während des Präkambriums und damit während des Großteils der Erdgeschichte, zeichneten sich die Ozeane durch ihren hohen Eisengehalt aus. Sowohl die Remineralisierung von organischem Material, als auch die Diagenese in den Sedimenten, die in den frühen Ozeanen der Erde abgelagert wurden, hatte höchstwahrscheinlich bedeutende Auswirkungen auf die globalen biogeochemischen Stoffkreisläufe. Unser Verständnis des Abbaus von organischem Material in eisenhaltigen Sedimenten ist jedoch sehr begrenzt, da moderne Analogsysteme extrem selten sind und bis heute nicht erforscht wurden. Der Towutisee auf der Insel Sulawesi in Indonesien ist ein solches modernes Analogsystem und Ziel dieser Doktorarbeit war es, die Raten und Pfade des Abbaus von organischem Material in den modernen eisenhaltigen Sedimenten des Towutisees zu erforschen. Der Towutisee ist der größte tektonische See in Südostasien und ist von mafischen und ultramafischen Gesteinen des Ost-Sulawesi-Ophioliten umgeben. Er hat eine maximale Wassertiefe von 203 m und ist schwach thermisch stratifiziert. Bis zu einer Tiefe von 70 m herrschen oxische Bedingungen, während die Wassersäule unterhalb von 130 m permanent anoxisch ist. Intensive Verwitterungsprozesse des ultramafischen Einzugsgebietes führen zu einem hohen Eintrag von Eisen(oxy)hydroxiden, während der Oberflächenabfluss nur wenig Sulfat enthält. Die Konzentrationen von Sulfat in der Wassersäule sind daher außergewöhnlich gering (< 20µM). Diese physikochemischen Verhältnisse sind analog zu denen der Ozeane des Archaikums und des Proterozoikums. Kurze (< 35 cm) Sedimentkerne wurden von verschiedenen Wassertiefen und unterschiedlichen Redox-Bedingungen des Bodenwassers entnommen. Darüber hinaus, wurde, im Zuge einer Bohrkampagne des International Continental Scientific Drilling Programs (ICDP) am Towutisee, ein 114 m langer Sedimentkern aus einer Wassertiefe von 153m, also deutlich unterhalb des Sauerstoffgradienten, erbohrt. Dieser war ausschließlich für geomikrobiologische Probenahmen und Untersuchungen vorgesehen. Die Proben, die aus diesen Sedimentkernen entnommen wurden, bilden das Fundament dieser Doktorarbeit und wurden für biogeochemische und mikrobiologische Untersuchungen verwendet. Unkontaminierte Proben sind für geomikrobiologische Untersuchungen unabdingbar. Das Erforschen von Gebieten unterhalb der Oberfläche ist jedoch auf Bohrungen angewiesen, welche wiederum den Einsatz einer Bohrspülung erfordern. Leider ist es unvermeidlich, dass diese im Zuge des Bohrprozesses in den erbohrten Sedimentkern eindringen. Die einzige Möglichkeit unkontaminierte Proben zu gewinnen ist es daher, den Grad der Kontamination des Bohrkerns nachzuverfolgen und unkontaminierte Proben für weitere Analysen zu identifizieren. Dazu wurde im Zuge dieser Doktorarbeit eine einfache und kostengünstige Methode zur Kontaminationskontrolle während Bohroperationen entwickelt und während der ICDP Bohrkampagne auf dem Towutisee angewandt. Als Tracer kam eine Farbe zum Einsatz, deren physikalische Eigenschaften denen von partikulären Tracern ähnelt. Der Preis dieser Farbe ist im Vergleich zu bisher verwendeten partikulären Tracern, jedoch vier Größenordnungen geringer und löst damit das Hauptproblem dieser Tracer. Die Methode benötigt nur ein Mindestmaß an Equipment und ermöglicht eine schnelle Identifizierung von Kontaminationen, möglicherweise sogar vor Ort. Die Sensitivität der Methode ist im Bereich von etablierten Kontaminationskontrollen. Kontaminierte Proben des erbohrten Sedimentkerns wurden mit dieser Methode identifiziert und nicht für weitere geomikrobiologische Untersuchungen verwendet. Biogeochemische Analysen der Kurzkerne zeigen, dass die Sedimente des Towutisees sehr arm an Elektronenakzeptoren sind, die für den mikrobiellen Abbau von organischem Material verwendet werden (d.h. Sauerstoff, Nitrat und Sulfat). Nichtsdestotrotz zeichnen sich die Sedimente des Towutisees durch hohe Zellzahlen aus, die von den Redox-Bedingungen des Bodenwassers abhängig sind. In niedrigen Wassertiefen führt oxygeniertes Bodenwasser zu einem erhöhten Eintrag von labilem organischen Material sowie Elektronenakzeptoren wie Eisen und Sulfat, wodurch hohe Zellzahlen resultieren. Mikrobiologische Analysen zeigen, dass die Sedimente des Towutisees durch eine vielseitige, mikrobielle Gemeinschaft bevölkert werden, die in der Lage ist, Stoffwechsel, wie Eisenreduktion, Sulfatreduktion, Fermentation sowie Methanogenese auszuführen. Biogeochemische Untersuchungen der oberen 12 m des 114 m langen Sedimentkerns zeigen, dass die Sedimente des Towutisees mit 2500 µM cm-3 extrem hohe Eisengehalte (20 Gew. %) aufweisen und damit das eisenreichste natürliche sedimentäre System sind, welches bisher erforscht wurde. Nach unserem bisherigen Verständnis über biogeochemische Stoffkreisläufe sollte, in Abwesenheit von Sauerstoff, Nitrat oder Sulfat, organisches Material über den energetisch günstigsten verfügbaren Elektronenakzeptoren abgebaut werden – in dem Fall Eisen (III). Erstaunlicherweise jedoch, ist Methanogenese der dominante (> 85 %) Remineralisierungsprozess in den Sedimenten des Towutisees. Mikrobiell theoretisch verfügbares reaktives Eisen (III) hingegen bleibt stabil über die oberen 12 m des Sedimentkerns und damit über mehr als 60.000 Jahre. Produziertes Methan wird nicht anaerob oxidiert und diffundiert aus dem Sediment in die Wassersäule. Die Dominanz von Methanogenese in diesen eisenreichen Sedimenten impliziert, dass dieser Prozess im Präkambrium vermutlich eine viel bedeutendere Rolle in der Remineralisierung von organischem Material eingenommen hat, als bisher angenommen. Methan, als bedeutendes Treibhausgas, war demnach möglicherweise ein wichtiger Regulator des Klimas in der frühen Erdgeschichte. T2 - Biogeochemie eisenreicher Sedimente des Lake Towuti, Sulawesi, Indonesien KW - Geomicrobiology KW - Biogeochemistry KW - Organic matter mineralization KW - Early Earth KW - Contamination Control KW - Biogeochemie KW - Kontaminationskontrolle KW - Frühe Erdgeschichte KW - Geomikrobiologie KW - Mikrobieller Abbau von organischen Material Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-475355 ER -