der dynamische Bereich. Eine solche divisive Kennlinienänderung kann die mathema-tische Operation der Division für das Nervensystem bereitstellen. Obwohl Division sehr wichtig für die neuronale Informationsverarbeitung ist, wurden bisher sehr wenig Mechanismen gefunden , die für einen divisiven Effekt (auch ‘gain control’ genannt) verantwortlich sein können. Deshalb wurde durch verschieden Kreuzadaptationsver-suche und durch pharmakologische Eingriffe nach dem verantwortlichen Mechanis-mus gesucht. Es konnte gezeigt werden, dass präsynaptisch Hemmung für die divisive Operation verantwortlich ist. Durch Strominjektionen ausgelöste intrinsische Adapta-tion im AN2 selbst hatte dagegen einen rein subtraktiven Effekt.
D5 Selektive Informationskodierung in der Hörbahn der Grille
Das Kapitel 6 beschäftig sich nochmals mit dem AN2 der Grille. Auch hier werden die Ergebnisse elektrophysiologischer Untersuchungen beschrieben. Der Fokus bei diesen Experimenten lag dabei allerdings auf Adaptation auf einer längeren Zeitskala, die mit Zeitkonstanten im Bereich von 1-2 Sekunden beschrieben werden kann.
Die Experimente in Kapitel 6 stellen einerseits diese langsam verlaufende Adap-tation dar und hatten andererseits das Ziel, die Frage zu beantworten, wozu diese
‘langsame’ Adaptation dient. Dabei wurden zwei alternative Hypothesen getestet:
(1) die Adaptation verändert die Kennlinie so, dass möglichst die gesamte akustis-che Umgebung des Tieres abgebildet werden kann (Infomax-Theorie), (2) die Kennlin-ien werden durch Adaptation so verändert, dass nur das lauteste Signal von mehreren gleichzeitig vorhandenen dargestellt wird, wobei der ‘Hintergrund’ unterdrückt wird (selektive Kodierung).
Um diese beiden Hypothesen zu testen, wurde eine Umgebung mit mehreren gle-ichzeitig vorhandenen, akustischen Signalen simuliert. Die Hörbahn wurde jeweils für längere Zeit entsprechenden Stimuli ausgesetzt, um im Anschluss die durch Adap-tation veränderten Kennlinien zu messen. Die beiden Hypothesen liefern dabei un-terschiedliche Voraussagen. Weder die Vorhersagen für die Infomax-Theorie, noch diejenigen, die sich aus der selektiven Kodierung ergab, trafen genau zu. Eine an-schließende Analyse der Transinformation (‘mutual information’) zeigte aber, dass Ad-aptation dazu führte, selektiv die Information über den Hintergrund zu unterdrücken, während die Information für das jeweils lauteste ‘Signal’ konstant blieb. Diese Anal-yse lieferte also Hinweise dafür, dass langsame Adaptation im AN2 der Grille selektive Kodierung des lautesten Signals ermöglicht.
D6 Schlussbetrachtung
In Kapitel 7 werden die Ergebnisse der einzelnen Studien zusammengefasst und un-ter dem Gesichtspunkt allgemeiner Konstruktionsprinzipien für Adaptation in sen-sorischen Bahnen betrachtet. Drei größere Themenkomplexe werden dabei in funk-tioneller Hinsicht diskutiert: (1) das Verhältnis des zeitlichen Verlaufes der Adaptation zu der Zeitskala, auf der sich relevante Aspekte des sensorischen Eingangs verändern, (2) wie sich widersprüchliche Ansprüche bei der Prozessierung von Signalen auf die Konstruktion der Adaptation auswirken, und (3) wo Adaptationsmechanismen inner-halb der sensorischen Bahn platziert sein sollten, unter besonderer Berücksichtigung von Divergenz, Konvergenz und von Nichtlinearitäten in der Sinnesbahn.
Zusammenfassend lässt sich sagen, dass sowohl die Adaptationsmechanismen, als auch deren genaue Platzierung innerhalb der sensorischen Bahn wesentlich für Sin-nesleistungen sind. Ein umfassendes Verständnis neuronaler Adaptation ist nur unter Berücksichtigung dieser Konstruktionsprinzipien, sowie der für das Tier jeweils rele-vanten Signale möglich.
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