Neocortical pyramidal cells possess voltage-dependent dendritic sodium channels that promote propagation of action potentials into the dendritic tree but paradoxically may fail to originate dendritic spikes. A biophysical model was constructed to reconcile these observations with known anatomical and physiological properties. When dendritic and somatic sodium channel densities compatible with electrophysiological measurements were combined with much higher densities in the axon initial segment then, regardless of the site of stimulation, spikes initiated at the initial segment and subsequently invaded the dendrites. The lower initial segment threshold arose from high current density and electrical isolation from the soma. Failure of dendritic channels to initiate spikes was due to inactivation and source-load considerations, which were more favorable for conduction of back-propagated spikes.
Co.rre1 atcon betvveen panems 1n c a Rd d. r = 0.92 (P < 0.001). B, Time courses of calcium• signals at selected glomeruli {dlfferern experinienl) a, Odours .elicit specific s1gnals upon s tiffUJiat•Oi1 (bar). b. Hexaoollcads to successive activat•on of dif.erent regions (see Inset) 2 8 5
Recording brain activity in vivo during learning is fundamental to understanding how memories are formed. We used functional calcium imaging to track odor representations in the primary chemosensory center of the honeybee, the antennal lobe, while training animals to discriminate a rewarded odor from an unrewarded one. Our results show that associative learning transforms odor representations and decorrelates activity patterns for the rewarded versus the unrewarded odor, making them less similar. Additionally, activity for the rewarded but not for the unrewarded odor is increased. These results indicate that neural representations of the environment may be modified through associative learning.
We describe a method for optically recording neuronal activity from an intact insect brain, upon natural sensory stimulation. In this preparation, the head capsule of the honeybee, Apis mellifera, is isolated from the body while leaving the entire brain undamaged. In short, a hole is first cut into the cuticule to allow optical access to the brain and to allow the removal of tracheae and glands. Then the head is cut free and placed into a dye-loaded and cooled ringer solution in a staining chamber for 1 h. Subsequently, the head is fixed in a recording chamber, covered with a cover-slip, and imaged under the microscope with a cooled CCD camera. The whole preparation leaves the antennae dry, free to move, and functional throughout the experiment, allowing for natural odour stimulation of the olfactory system. Using calcium sensitive or potential sensitive dyes (calcium-green or RH795), we could record the processing of olfactory information at the glomerular level in the antennal lobe of the bee.
Stimulation with odours has been shown to elicit characteristic patterns of activated glomeruli in the antennal lobe (AL) of honeybees. In this study we show that these patterns are dynamic in a time window of 2-3 s after stimulus onset. We measured changes in the averaged membrane potential of all cells in the glomerular neuropil by optical imaging of the voltage-sensitive dye RH795 using a slow scan CCD camera (3 frames/s). The four substances 1-hexanol, hexanal, citral and clove-oil as well as the binary mixtures hexanol+hexanal and hexanol+citral were used as stimuli (2 s stimulus duration). We found that: (1) every odour elicited an odour-specific activity pattern, and conversely every glomerulus had a characteristic odour response profile; (2) some glomeruli had a tonic, some a phasic-tonic, and some a slow phasic response pattern; (3) the difference between the glomerular response patterns increased within 2 s of stimulus presentation, which suggests that odour representations became more characteristic over stimulus time; and (4) the responses to odorant mixtures were complex and glomerulus-dependent: some responses correspond to the sum of the compounds' responses, some to the response of one of the components.
Navigation in honeybees is discussed against the background of the types of memories employed in the navigational task. Two questions are addressed. Do bees have goal-specific expectations, and when are novel routes travelled? Expectations are deduced from (1) context stimuli as determinants for local cue memories, (2) landmark-dependent path integration, (3) sequential learning of landmarks, and (4) motivation- and context-dependent memory retrieval. Novel routes are travelled under two conditions: (1) goal-cue-based piloting and (2) integration of simultaneously activated vector memories. Our data do not support the conclusion that memory integration in bees is organised by a cognitive map. The assumption of purely separate memories that are only retrieved according to the chain of events during navigational performance also appears to be inadequate. We favour the view that multiple memories are integrated using external and internal sources of information. Such configural memories lead to both specific expectations and novel routes.
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