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  • 1
    Electronic Resource
    Electronic Resource
    Springer
    Biological cybernetics 40 (1981), S. 171-179 
    ISSN: 1432-0770
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Computer Science , Physics
    Notes: Abstract The Culex circadian pacemaker's response to phase-resetting light signals was studied in the first 3 cycles of darkness following a 12h light exposure. (1) In both cycles 1 and 2 there is a clear change from “type 1” to “type 0” phase-resetting as the resetting signal is prologed (Fig. 2). (2) Mosquitoes in cycle 1 are about half as sensitive to phase-resetting as those in cycles 2 or 3 (the criterion being the minimum pulse duration required to produce type 0 phase-resetting) (Fig. 2). (3) Each cycle appears to have a corkscrew-shaped phaseresetting surface and a phase singularity (Figs. 4, 5, and 7). The hypothesis that the Culex pacemaker reaches a stable limit cycle within the first cycle leads to an economical explanation of the results.
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  • 2
    Electronic Resource
    Electronic Resource
    Springer
    Biological cybernetics 40 (1981), S. 181-194 
    ISSN: 1432-0770
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Computer Science , Physics
    Notes: Abstract The stability of the Culex circadian oscillation in darkness was tested by perturbing it with bright light pulses. (1) In the absence of perturbation the overt circadian rhythm persists in darkness for at least 2 weeks (Fig. 5). (2) The circadian pacemaker, as perceived by the response to a standard phaseresetting signal, also follows a regular cycle in the absence of perturbation (Fig. 4). (3) Almost all light pulse perturbations quickly shift the phase of both overt rhythm and pacemaker, and the system resumes normal oscillation within 24 h (Figs. 6, 7, 8, and 9). (4) By contrast, perturbations driving the pacemaker near its phase singularity cause unpredictable phase shifts and/or disrupt individuals' overt rhythms for many cucles (Figs. 12, 13, and 14). Such pulses also sharply reduce the perceived amplitude of the pacemaker's oscillation (Figs. 10, 11, and 15). Most mosquitoes resume normal oscillation within a week but in some cases abnormalities persist for at least 2 weeks. The results re-emphasize the analogy between the circadian pacemaker and a simple limit cycle oscillator. The lasting anomalous behaviour of some mosquitoes after some pulses can either be explained by invoking a weakly unstable singularity, or simply treated as a minor, and perhaps revealing, deviation from prediction.
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  • 3
    Electronic Resource
    Electronic Resource
    [s.l.] : Nature Publishing Group
    Nature 303 (1983), S. 240-242 
    ISSN: 1476-4687
    Source: Nature Archives 1869 - 2009
    Topics: Biology , Chemistry and Pharmacology , Medicine , Natural Sciences in General , Physics
    Notes: [Auszug] Many aspects of the CNS of the leech, Hirudo medicinalis, are well understood3, but very little is known about how it processes visual information. The leech's visual system includes five pairs of eyes, numbered 1-5 in a front-to-rear sequence on the surface of the head (Fig. la), and numerous ...
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  • 4
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 154 (1984), S. 781-788 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The two C cells and the single S cell in each segmental ganglion of the leech nerve cord are strongly electrically coupled, as are the S cells of adjacent ganglia. Impulses arising in any of these neurons propagate rapidly, via the fast conducting system formed by the axons of the S cells, to all other neurons in the network. 2. The dye Lucifer Yellow crosses the electrical junctions among the C and S cells. Exploiting this property of the dye, I traced the fast conducting system rostrally into the subesophageal ganglion, and caudally into the tail-brain. 3. The dye revealed 9 previously unidentified neurons in the subesophageal ganglion: 2 neurons resembling C cells in each subganglion, and a single asymmetrical neuron, designated S0, in the most anterior subganglion. 4. Physiologically, the S0 cell resembles the S cells of the segmental ganglia, and it may be homologous to them despite its unusual structure. 5. Strong electrical coupling (coefficient 40%) between the S0 cell and the S1 cell (the S cell in the first segmental ganglion) ensures that an impulse in one neuron is always matched by an impulse in the other. 6. Input from tactile and photic receptors on the head excites the S0 and S1 cells, eliciting impulses from initiation sites in the processes of the S1 cell, within the subesophageal ganglion. 7. Lucifer Yellow revealed 21 previously unidentified neurons in the tail-brain; these evidently consist of an S cell and 2 C cells in each subganglion. 8. A recently-proposed model for the origin of the unpaired neurons of the leech nervous system explains the asymmetry of the S0 cell.
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  • 5
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 155 (1984), S. 543-557 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The lateral visual (LV) cells of the medicinal leech,Hirudo medicinalis, are excited by photoreceptors having axons in the ipsilateral DC and DD nerves. Activating the photoreceptors photically or electrically yields at least three kinds of event in the LV cell (Fig. 1B). 2. There are very small events, which appear to be electrical coupling potentials from the photoreceptors (Fig. 6); there are large events, which appear to be action potentials arising in the main neurite and propagating out the axon (Figs. 2, 5); and finally, there are unusual medium-sized events. 3. These medium-sized events appear to be ‘stacks’ of several small spike-like components (Fig. 7A). These small spikes have different thresholds and latencies, suggesting that they arise at different initiation sites (Fig. 7B). 4. Small spikes are not chemical excitatory postsynaptic potentials (EPSPs), since they persist in high Mg saline (Fig. 7). They are not antidromic impulses arising distally in the axon, since they do not collide with orthodromic action potentials (Fig. 8). Nor, given the initiation kinetics of small spikes, are they coupling potentials from neurons having axons in the optic nerves (Figs. 11, 12). 5. Moderate hyperpolarization increases the amplitude of small spikes, but strong hyperpolarization blocks them (Fig. 9). Very strong hyperpolarizing pulses elicit small spikes by anode break excitation (Fig. 10). 6. These results suggest that small spikes may be active responses that arise locally in the dendrites of the LV cell but fail to propagate actively into the main neurite and axon (Fig. 16). 7. Alternatively, small spikes may be coupling potentials from unidentified interneurons that are strongly electrically coupled to the LV cell. In order to account for the results, however, there would have to be at least 24 such interneurons. The failure to find any coupling neurons, despite extensive searching, makes this alternative possibility increasingly unlikely. 8. The small spikes activated by axons in the DC nerve appear to be distinct from those activated by axons in the DD nerve (Figs. 13, 14). 9. Within the main neurite the small spikes behave like conventional EPSPs: they sum temporally and spatially, and they interact with inhibitory synaptic input to set the firing level of the LV cell (Figs. 12A, 15). 10. Thus, there appear to be two stages of integration in the activation of the LV cell: at the first stage, EPSPs from the photoreceptors sum to elicit small spikes; at the second stage, the small spikes sum to generate action potentials.
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  • 6
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 156 (1985), S. 707-717 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The anterior visual (AV) cell is a bilaterally paired visual interneuron in the supraesophageal ganglion of the medicinal leech (Fig. 1). 2. The input resistance of the AV cell increases with depolarization from rest and decreases with hyperpolarization (Fig. 2); overall, the current-voltage relationship is sigmoid (Fig. 4A). These features persist when chemical synaptic transmission is blocked by high Mg saline (Figs. 4B, 5). 3. Because of this current-voltage relationship, for certain applied currents the AV cell has two steady-state membrane potentials. Under those conditions current pulses switch the AV cell from one steady state to the other (Fig. 5C). 4. The spontaneous spike-like depolarizations in the AV soma (Fig. 2) appear to be synaptic potentials rather than failed impulses, since injected current neither blocks nor stimulates them, nor does TEA affect them (Fig. 3). The current-voltage relationship of the AV cell explains the complicated dependence of these ‘large EPSPs’ on membrane potential (Fig. 6). 5. When any ipsilateral eye is illuminated an AV cell shows a graded depolarization that persists in high Mg saline (Figs. 8, 9D, 10). Moreover, the AV cell appears to be lucifer yellow dye-coupled to the photoreceptors of the ipsilateral eyes (Fig. 1). 6. When any contralateral eye is illuminated an AV cell receives a volley of large EPSPs (Fig. 10). This input does not persist in high Mg saline and is therefore probably polysynaptic. 7. Although there is no synaptic connection between the AV cells, the large EPSPs in an AV cell do match synaptic potentials in several other identified neurons in the supraesophageal ganglion (Fig. 12), including the other AV cell (Fig. 11C). 8. In addition, large EPSPs in an AV cell match EPSPs in the contralateral LVa cell (Fig. 13B) and IPSPs in the ipsilateral LVa cell (Fig. 13D). Depolarization of the LVa cell elicits large EPSPs in the contralateral AV cell (Fig. 13C). 9. These results suggest that the AV cells are second order visual neurons that are specialized to respond positively to slight increases in light intensity, and that the AV cells are tightly integrated with the other known visual interneurons.
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  • 7
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 138 (1980), S. 201-211 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary TheCulex flight activity circadian oscillator, free-running in darkness, was perturbed by single bright light pulses. The phase and duration of the pulse were varied systematically (Winfree's “singularity trap” procedure) revealing the oscillator's phase-resetting surface. The corkscrew shape of this surface, predicted by theCulex model and verified here, suggests that the circadian dynamic system has a limit cycle.
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  • 8
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 156 (1985), S. 719-727 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The H cell is an unpaired neuron originating in subganglion 3 of the subesophageal ganglion. Its processes form an ‘H’ with long axis on the midline of the nervous system (Fig. 1). The soma is off-center, originating from the junction of the main neurite (the crossbar of the ‘H’) and one or the other axon (upright of the ‘H’). 2. Illuminating any eye excites the H cell, producing a volley of impulses riding on a compound synaptic potential. The synaptic potential is graded with light intensity and it persists in 40 mmol/l Mg saline (Fig. 3). 3. Touching the skin inhibits the H cell by a polysynaptic pathway that includes the mechanosensory T cells (Fig. 4). 4. The H cell has two impulse initiation sites, one in each axon. Impulses in the axon ipsilateral to the soma cause large spikes in the soma; those in the other axon cause small spikes (Fig. 2). 5. Large spikes match synaptic potentials in the AV (anterior visual) cell ipsilateral to the soma of the H cell (Figs. 5, 6); small spikes match synaptic potentials in the contralateral AV cell. 6. This synaptic potential has a large inhibitory component and a small excitatory component (Fig. 7). The inhibitory component is caused by an increase in Cl conductance (Figs. 7, 8); the excitatory component may be electrical (Fig. 9). 7. The connection between the H cell and the AV cell may be polysynaptic since it is blocked by high Ca/Mg saline (Fig. 9).
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  • 9
    Electronic Resource
    Electronic Resource
    Springer
    Journal of comparative physiology 156 (1985), S. 697-706 
    ISSN: 1432-1351
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Summary 1. The medicinal leech has five pairs of simple eyes (Fig. 1A). The photoreceptors of some eyes are known to excite one of the bilateral pair of lateral visual (LVa) cells in the first segmentai ganglion (G1). A new pair of neurons, the LVb cells, has been found in subganglion 4 (SubEG4) of the subesophageal ganglion. Since G1 and SubEG4 develop from adjacent ganglionic primordia, the LVa and LVb cells may be serial homologues. 2. The somata and dendrites of the LVa and LVb cells lie in corresponding positions (Fig. 3), and their axons run side-by-side and terminate together in the supraesophageal ganglion. 3. Both neurons have bulbous contralateral branches (Figs. 2, 3), and fibrous ipsilateral dendrites that match the two zones where photoreceptors of the ipsilateral eyes 3–5 have synaptic terminals (Fig. 1C). Moreover, both neurons are Lucifer Yellow dye-coupled to the photoreceptors of those same eyes (Fig. 2B). 4. When the ipsilateral eyes 3–5 are illuminated, the LVa and LVb cells both show a characteristic complex response that includes a compound EPSP produced by electrical coupling potentials from the photoreceptors (Figs. 5, 6), small spike-like events, and conventional-looking action potentials (Fig. 4). 5. The compound EPSP is thought to elicit local spikes in ipsilateral dendritic branches, which then spread electrotonically into the main neurite and soma as small spikes. In turn, the summation of these small spikes in the neurite evidently elicits propagating action potentials. 6. LV cells produce extended plateau-like depolarizations (Fig. 7) in weak or continuously varying light, or in response to current injection. These slow potentials appear to be generated by a noninactivating Na conductance and a TEA-sensitive K conductance (Fig. 8). Since slow potentials are accentuated when extracellular Ca is replaced by Ba (Fig. 9), Ca appears to participate in the response, presumably via a Ca-dependent K channel. 7. Although the LVa and LVb cells may be homologous, they are not identical (Fig. 5). For example, unlike the LVa cells, the LVb cells are not inhibited by input from the contralateral eyes 3–5. Moreover, eyes 1 and 2 weakly inhibit the LVa cells, but weakly excite the LVb cells.
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  • 10
    Electronic Resource
    Electronic Resource
    [s.l.] : Nature Publishing Group
    Nature 280 (1979), S. 677-679 
    ISSN: 1476-4687
    Source: Nature Archives 1869 - 2009
    Topics: Biology , Chemistry and Pharmacology , Medicine , Natural Sciences in General , Physics
    Notes: [Auszug] Fig. 1 a, Outline of the two-dimensional limit cycle model. The two variables are left unspecified, the oscillator is merely assumed to be attracted in darkness to a closed cycle, here represented by the bold circle. This cycle, the limit cycle, is assumed to be stable. If the oscillator is ...
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