Signal transformation at the input and output of the Drosophila visual system
A key function of the nervous system is to sample data from the external world, generate internal signals, and transform them into meaningful information that can be used to trigger behaviour. In order to gain insight into the underlying mechanism for signal transformation, the visual system has been extensively studied: partly owing to the stimulus being reliably presentable, and the anatomy being well described. The Drosophila visual system is one such system, with the added advantage of genetic tractability. In this thesis, I studied the filtering property of visual neurons at two levels, biophysical and circuit levels.
The first study looks at signal transformation at the biophysical level, at the input of the visual system, in photoreceptors. Voltage-gated potassium channels counteract the depolarization caused by opening of light sensitive channels, and the heterogeneous properties of their kinetics can fine-tune the photoreceptor’s frequency response to fulfill the animal’s ecological requirements. Shaker (Kv1) and Shab (Kv2) have been identified as fast and slow inactivating components of the photoreceptor’s outward currents, however a current with intermediate kinetics (IKf) has not been molecularly identified, but had been postulated to be Shal (Kv4). I focused on characterizing this current using whole-cell patch clamp in wild type and mutants, and using antibodies for Shal. My results from whole-cell patch clamp indicated that IKf in adult R1-6 cells are not Shal, from their voltage dependence and insensitivity to a Kv4 blocker. This calls for alternative molecular basis for IKf, which is likely to be a slow inactivating component of Shaker, or a combination of its many splice variants.
The second study looks at signal transformation at the circuit level, at the output end, in the third optic neuropil, lobula. Visual projection neurons project from the lobula to the central brain, and have been proposed to carry behaviourally relevant visual features to higher brain regions. It was recently shown that optogenetic activation of individual visual projection neuron types could induce distinct behaviours such as takeoff and backward walking, linking these visual neurons to specific behavioural programs downstream. Using in vivo two-photon calcium imaging, I recorded visually evoked calcium responses from three of these cell types. Cell types that showed induced takeoff and backward walking preferentially responded to dark looming stimuli or fragmented expanding local features, suggesting their role in behaviours triggered by object approach. To explore how this visual information is transformed in the downstream circuit, we identified several candidate neurons that receive input from this cell type by anatomical overlap, and then validated their connections using optogenetic activation and calcium imaging. One downstream cell-type that projects bilaterally had very similar response properties to its upstream partner, whereas another cell-type that projects ipsilaterally seemed to filter out some information from its upstream partner. This is one of the first studies that functionally characterizes lobula visual projection neurons and their downstream partners in Drosophila, and their response properties agree with the general idea that visual information becomes increasingly selective as it is sent to higher brain regions.