Schlaf fördert die Gedächtnisbildung. Nach einem anerkannten Modell beruht der förderliche Effekt des Schlafs auf einer verdeckten Reaktivierung der initial im Hippokampus gespeicherten Gedächtnisspuren während des Tiefschlafs, die durch ihre Reaktivierung in neokortikale Hirnareale zur langfristigen Speicherung transferiert werden. Obwohl die Existenz von Gedächtnisreaktivierungen im Schlaf gut belegt ist, ist bislang ungeklärt, ob diese Reaktivierungen tatsächlich in einem kausalen Zusammenhang zu den Prozessen der Gedächtnisbildung im Schlaf stehen. Zur Klärung dieser Frage habe ich in der vorliegenden Arbeit den Effekt von experimentell induzierten Gedächtnisreaktivierungen im Schlaf auf die Gedächtnisbildung untersucht. Zur Reaktivierung von Gedächtnisinhalten im Schlaf wurde ein während der Lernphase mit den Lerninhalten assoziierter Geruch verwendet. Achtzehn gesunde Probanden lernten sowohl einen deklarativen, den Hippokampus involvierenden als auch einen prozeduralen, weniger hippokampalen Gedächtnistest am Abend unter der Präsenz einen Geruchs, der sich als Kontextstimulus mit den Lerninhalten verbinden sollte. In der darauf folgenden Nacht wurde ihnen während des Tiefschlafs entweder erneut der Geruch oder eine geruchslose Substanz präsentiert. Die Abfrage am nächsten Morgen erfolgte ohne Duftdarbietung. Nach der Nacht mit Duft war die Erinnerungsleistung in dem deklarativen Gedächtnistest signifikant erhöht, im Vergleich zu der Nacht ohne Duft. Der positive Effekt des Geruchs auf die Gedächtnisbildung trat nicht auf, wenn der Geruch während der Lernphase nicht dargeboten wurde, und zeigte sich ausschließlich im Tiefschlaf, nicht dagegen nach einer erneuten Darbietung des Dufts im rapid-eye movement (REM)-Schlaf oder Wachzustand. Die prozedurale Gedächtnisbildung im Schlaf wurde in keinem der Experimente von einer Geruchsdarbietung nach dem Lernen beeinflusst. Die geruchsinduzierten Gedächtnisreaktivierungen im Tiefschlaf hatten keine Effekte auf die mittels der Elektroenzephalographie (EEG) gemessenen Hirnaktivität im Schlaf. Dagegen konnten zusätzliche Experimente unter Verwendung der funktionellen Kernspintomographie (fMRT) zeigen, dass eine erneute Geruchsdarbietung im Tiefschlaf den linken Hippokampus aktiviert, vorausgesetzt der Duft war bereits während der Lernphase präsent. Die Befunde der verbesserten Erinnerungsleistung nach einer experimentell induzierten Reaktivierung der neu enkodierten, hippokampalen Gedächtnisinhalte liefern einen eindeutigen Beleg für den kausalen Zusammenhang zwischen hippokampalen Gedächtnisreaktivierungen im Tiefschlaf und den Prozessen der deklarativen Gedächtnisbildung. Damit bestätigen sie die Kernannahme des Modells der schlaf-abhängigen Gedächtnisbildung durch Reaktivierungen im Tiefschlaf. Zusätzlich liefern die Experimente einen neuen Ansatz zur Untersuchung und Manipulierbarkeit von Gedächtnisreaktivierungen im Schlaf, durch den möglicherweise auch im klinischen oder alltäglichen Kontext positive Ergebnisse auf die Gedächtnisbildung im Schlaf zu erzielen sind.
The forward testing effect is an indirect benefit of retrieval practice. It refers to the finding that retrieval practice of previously studied information enhances learning and retention of subsequently studied other information in episodic memory tasks. Here, two experiments were conducted that investigated whether retrieval practice influences participants’ performance in other tasks, i.e., arithmetic tasks. Participants studied three lists of words in anticipation of a final recall test. In the testing condition, participants were immediately tested on lists 1 and 2 after study of each list, whereas in the restudy condition, they restudied lists 1 and 2 after initial study. Before and after study of list 3, participants did an arithmetic task. Finally, participants were tested on list 3, list 2, and list 1. Different arithmetic tasks were used in the two experiments. Participants did a modular arithmetic task in Experiment 1a and a single-digit multiplication task in Experiment 1b. The results of both experiments showed a forward testing effect with interim testing of lists 1 and 2 enhancing list 3 recall in the list 3 recall test, but no effects of recall testing of lists 1 and 2 for participants’ performance in the arithmetic tasks. The findings are discussed with respect to cognitive load theory and current theories of the forward testing effect.
The forward testing effect refers to the finding that retrieval practice of previously studied information enhances learning and retention of subsequently studied other information. While most of the previous research on the forward testing effect examined group differences, the present study took an individual differences approach to investigate this effect. Experiment 1 examined whether the forward effect has test-retest reliability between two experimental sessions. Experiment 2 investigated whether the effect is related to participants’ working memory capacity. In both experiments (and each session of Experiment 1), participants studied three lists of items in anticipation of a final cumulative recall test. In the testing condition, participants were tested immediately on lists 1 and 2, whereas in the restudy condition, they restudied lists 1 and 2. In both conditions, participants were tested immediately on list 3. On the group level, the results of both experiments demonstrated a forward testing effect, with interim testing of lists 1 and 2 enhancing immediate recall of list 3. On the individual level, the results of Experiment 1 showed that the forward effect on list 3 recall has moderate test-retest reliability between two experimental sessions. In addition, the results of Experiment 2 showed that the forward effect on list 3 recall does not depend on participants’ working memory capacity. These findings suggest that the forward testing effect is reliable at the individual level and affects learners at a wide range of working memory capacities alike. The theoretical and practical implications of the findings are discussed.
The last decades of stress research have yielded substantial advancements highlighting the importance of the phenomenon for basic psychological functions as well as physical health and well-being. Progress in stress research heavily relies on the availability of suitable and well validated laboratory stressors. Appropriate laboratory stressors need to be able to reliably provoke a response in the relevant parameters and be applicable in different research settings or experimental designs. This thesis focuses on the Cold Pressor Test (CPT) as a stress induction technique. Three published experiments are presented that show how the advantages of the CPT can be used to test stress effects on memory processes and how some of its disadvantages can be met by a simple modification that retains its feasibility and validity. The first experiment applies the CPT in a substantial sample to investigate the consolidation effects of post-learning sympathetic arousal. Stressed participants with high increases in heart rate during the CPT showed enhanced memory performance one day after learning compared to both the warm water control group and low heart rate responders. This finding suggests that beta-adrenergic activation elicited shortly after learning enhances memory consolidation and that the CPT induced heart rate response is a predictor for this effect. Moreover, the CPT proved to be an appropriate stressor to test hypothesis about endogenous adrenergic effects on memory processes. The second experiment addresses known practical limitations of the standard dominant hand CPT protocol. A bilateral feet CPT modification is presented, the elicited neuroendocrine stress response assessed and validated against the standard CPT in a within-subjects design. The bilateral feet CPT elicited a substantial neuroendocrine stress response. Moreover, with the exception of blood pressure responses, all stress parameters were enhanced compared to the standard CPT. This shows that the bilateral feet CPT is a valid alternative to the standard CPT. The third experiment further validates the bilateral feet CPT and its corresponding control procedure by employing it in a typical application scenario. Specifically, the bilateral feet CPT was used to modulate retrieval of event files in a distractor-response binding paradigm that required lateralized bimanual responses. Again, the bilateral feet CPT induced significant increases in heart rate, blood pressure and cortisol, no such increases could be observed in the warm water control condition. Moreover, stressed participants showed diminished retrieval compared to controls. These results provide further evidence for the feasibility and validity of the bilateral feet CPT and its warm water control procedure. Together the experiments presented here highlight the usefulness of the CPT as a tool in psychophysiological stress research. It is especially well suited to test hypothesis concerning stress effects on memory processes and its applicability can be further increased by the bilateral feet modification.
The brain is the central coordinator of the human stress reaction. At the same time, peripheral endocrine and neural stress signals act on the brain modulating brain function. Here, three experimental studies are presented demonstrating this dual role of the brain in stress. Study I shows that centrally acting insulin, an important regulator of energy homeostasis, attenuates the stress related cortisol secretion. Studies II and III show that specific components of the stress reaction modulate learning and memory retrieval, two important aspects of higher-order brain function.