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Marder, E.1
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Neuronal control of leech behavior
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A review of what is known about behaviorally relevant neuronal circuits in the leech central nervous system.
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Kristan Jr. W.B., Calabrese R., and Friesen W.O. Neuronal control of leech behavior. Prog Neurobiol 76 (2005) 279-327. A review of what is known about behaviorally relevant neuronal circuits in the leech central nervous system.
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Kristan Jr., W.B.1
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Crustacean motor pattern generator networks
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Hooper S.L., and DiCaprio R.A. Crustacean motor pattern generator networks. Neurosignals 13 (2004) 50-69
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Dynamic sensorimotor interactions in locomotion
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Rossignol S., Dubuc R., and Gossard J.-P. Dynamic sensorimotor interactions in locomotion. Physiol Rev 86 (2006) 89-154
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Büschges A. Sensory control and organization of neural networks mediating coordination of multisegmental organs for locomotion. J Neurophysiol 93 (2005) 1127-1135
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Beyond the central pattern generator: amine modulation of decision-making neural pathways descending from the brain of the medicinal leech
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Serotonin is long known to powerfully modulate the expression of swimming locomotion in the leech. This study shows that serotonergic neurons in the leech brain are excited by stimuli that initiate swimming activity and when swimming is spontaneous. Intracellular recordings from command-like neurons in leech brain revealed that serotonin application inhibited high-level locomotory control neurons and led, via a reduction in drive to segmental swim-gating neurons, to a reduction of the excitatory drive to the swim oscillator circuit. Although the overall effect of serotonin is to facilitate swimming activity, serotonin may act via brain neurons to limit swimming activity at the level of the head brain.
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Crisp K.M., and Mesce K.A. Beyond the central pattern generator: amine modulation of decision-making neural pathways descending from the brain of the medicinal leech. J Exp Biol 209 (2006) 746-756. Serotonin is long known to powerfully modulate the expression of swimming locomotion in the leech. This study shows that serotonergic neurons in the leech brain are excited by stimuli that initiate swimming activity and when swimming is spontaneous. Intracellular recordings from command-like neurons in leech brain revealed that serotonin application inhibited high-level locomotory control neurons and led, via a reduction in drive to segmental swim-gating neurons, to a reduction of the excitatory drive to the swim oscillator circuit. Although the overall effect of serotonin is to facilitate swimming activity, serotonin may act via brain neurons to limit swimming activity at the level of the head brain.
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J Exp Biol
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Crisp, K.M.1
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To swim or not to swim: regional effects of serotonin, octopamine and amine mixtures in the medicinal leech
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Crisp K.M., and Mesce K.A. To swim or not to swim: regional effects of serotonin, octopamine and amine mixtures in the medicinal leech. J Comp Physiol A 189 (2003) 461-470
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Regulation of the segmental swim-generating system by a pair of identified interneurons in the leech head ganglion
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Brodfuehrer P.D., Parker H.J., Burns A., and Berg M. Regulation of the segmental swim-generating system by a pair of identified interneurons in the leech head ganglion. J Neurophysiol 73 (1995) 983-992
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Brodfuehrer, P.D.1
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Sensitization and dishabituation of swim induction in the leech Hirudo medicinalis: role of serotonin and cyclic AMP
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Zaccardi M.L., Traina G., Cataldo E., and Brunelli M. Sensitization and dishabituation of swim induction in the leech Hirudo medicinalis: role of serotonin and cyclic AMP. Behav Brain Res 153 (2004) 317-326
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Zaccardi, M.L.1
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Serotonin delays habituation of leech swim response to touch
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Alkatout B.A., Marvin N.M., and Crisp K.M. Serotonin delays habituation of leech swim response to touch. Behav Brain Res 182 (2007) 145-149
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Alkatout, B.A.1
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11
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Modification of leech behavior following foraging for artificial blood
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The authors of this study make the interesting case that the specific mode of locomotion evoked by electrical stimulation in intact leeches is modulated by prior exposure to artificial blood. Following exposure to this chemical, electrical stimulation of the rear of the leech increased the likelihood of swimming and crawling behaviors. In semi-intact preparations, exposure to full strength artificial blood, unlike a 10% solution, reduced the likelihood that nerve stimulation would evoke swimming activity. Hence, the behavioral responses of the leech are selectively modified by prior exposure to chemicals in their environment with effects differing, in an apparently adaptive way, as concentrations are altered.
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Brodfuehrer P.D., Tapyrik L., Pietras N., Zekavat G., and Convery M. Modification of leech behavior following foraging for artificial blood. J Comp Physiol A 192 (2006) 817-825. The authors of this study make the interesting case that the specific mode of locomotion evoked by electrical stimulation in intact leeches is modulated by prior exposure to artificial blood. Following exposure to this chemical, electrical stimulation of the rear of the leech increased the likelihood of swimming and crawling behaviors. In semi-intact preparations, exposure to full strength artificial blood, unlike a 10% solution, reduced the likelihood that nerve stimulation would evoke swimming activity. Hence, the behavioral responses of the leech are selectively modified by prior exposure to chemicals in their environment with effects differing, in an apparently adaptive way, as concentrations are altered.
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J Comp Physiol A
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Brodfuehrer, P.D.1
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12
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Statistics of decision making in the leech
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Garcia-Perez E., Mazzoni A., Zoccolan D., Robinson H.P.C., and Torre V. Statistics of decision making in the leech. J Neurosci 25 (2005) 2597-2608
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Garcia-Perez, E.1
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Mazzoni A., Garcia-Perez E., Zoccolan D., Graziosi S., and Torre V. Quantitative characterization and classification of leech behavior. J Neurophysiol 93 (2005) 580-593
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5-HT Modulation of identified segmental premotor interneurons in the lamprey spinal cord
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As in many other vertebrates and invertebrates, serotonin is an important modulator of behavior in lamprey. This paper describes experiments on neuromodulation of excitatory and inhibitory spinal interneurons that generate and phase motoneuron output during lamprey swimming. The main findings are that serotonin affects both cellular properties of the interneurons (reducing slow afterhyperpolarization) and properties of their synaptic interactions with motoneurons (reducing synaptic amplitude). This study begins to explain the effects of serotonin on the expression of lamprey swimming movements.
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Biró Z., Hill R.H., and Grillner S. 5-HT Modulation of identified segmental premotor interneurons in the lamprey spinal cord. J Neurophysiol 96 (2006) 931-935. As in many other vertebrates and invertebrates, serotonin is an important modulator of behavior in lamprey. This paper describes experiments on neuromodulation of excitatory and inhibitory spinal interneurons that generate and phase motoneuron output during lamprey swimming. The main findings are that serotonin affects both cellular properties of the interneurons (reducing slow afterhyperpolarization) and properties of their synaptic interactions with motoneurons (reducing synaptic amplitude). This study begins to explain the effects of serotonin on the expression of lamprey swimming movements.
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J Neurophysiol
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Biró, Z.1
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Modulation of a spinal locomotor network by metabotropic glutamate receptors
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Serotonin is an important neuromodulator of animal behaviors, but this study demonstrates that glutamate, acting via metabotropic receptors, can play a similar role. The paper describes experiments on Xenopus tadpoles in which three types of metabotropic glutamate receptors were activated by specific agonists. Different agonists increased or decreased cycle period of fictive swimming, some altered motoneuron burst durations, some did not, and some altered the amplitude of motor bursts. These findings are interesting because, although glutamate is commonly known to mediate excitatory interactions via ionotropic receptors, the modulatory role of metabotropic glutamate receptor is less widely appreciated.
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Chapman R.J., and Sillar K.T. Modulation of a spinal locomotor network by metabotropic glutamate receptors. Eur J Neurosci 26 (2007) 2257-2268. Serotonin is an important neuromodulator of animal behaviors, but this study demonstrates that glutamate, acting via metabotropic receptors, can play a similar role. The paper describes experiments on Xenopus tadpoles in which three types of metabotropic glutamate receptors were activated by specific agonists. Different agonists increased or decreased cycle period of fictive swimming, some altered motoneuron burst durations, some did not, and some altered the amplitude of motor bursts. These findings are interesting because, although glutamate is commonly known to mediate excitatory interactions via ionotropic receptors, the modulatory role of metabotropic glutamate receptor is less widely appreciated.
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Eur J Neurosci
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Chapman, R.J.1
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Sensory modification of leech swimming: interactions between ventral stretch receptors and swim-related neurons
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Cang J., Yu X., and Friesen W.O. Sensory modification of leech swimming: interactions between ventral stretch receptors and swim-related neurons. J Comp Physiol A 187 (2001) 569-579
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Morphological and physiological characterization of stretch receptors in leeches
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Eight pairs of putative stretch receptors are closely associated with longitudinal muscles in the leech body wall. Through the injection of Alexa Fluor hydrazide, this study presents the morphology of terminal arborizations of 7 of the eight. In addition, morphology and physiology demonstrated that one of these is a dorsal stretch receptor associated with dorsal longitudinal muscle. At least two of these axons undergo membrane potential oscillations that are phase locked to the swimming rhythm expressed in nerve cord-body wall preparations and, at a different phase angle, also in isolated nerve cords. These two are phasically modulated by muscle tension oscillations and hence could be a source of cycle-by-cycle input to the swim oscillator network.
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Fan R.J., and Friesen W.O. Morphological and physiological characterization of stretch receptors in leeches. J Comp Neurol 494 (2006) 290-302. Eight pairs of putative stretch receptors are closely associated with longitudinal muscles in the leech body wall. Through the injection of Alexa Fluor hydrazide, this study presents the morphology of terminal arborizations of 7 of the eight. In addition, morphology and physiology demonstrated that one of these is a dorsal stretch receptor associated with dorsal longitudinal muscle. At least two of these axons undergo membrane potential oscillations that are phase locked to the swimming rhythm expressed in nerve cord-body wall preparations and, at a different phase angle, also in isolated nerve cords. These two are phasically modulated by muscle tension oscillations and hence could be a source of cycle-by-cycle input to the swim oscillator network.
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J Comp Neurol
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Fan, R.J.1
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Yu X., and Friesen W.O. Entrainment of leech swimming activity by the ventral stretch receptor. J Comp Physiol A 190 (2004) 939-949
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A dye mixture (Neurobiotin and Alexa 488) reveals extensive dye-coupling among neurons in leeches; physiology confirms the connections
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Fan R.J., Marin-Burgin A., French K.A., and Friesen W.O. A dye mixture (Neurobiotin and Alexa 488) reveals extensive dye-coupling among neurons in leeches; physiology confirms the connections. J Comp Physiol 191 (2005) 1157-1171
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Fan, R.J.1
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Mapping motor neuron activity to overt behavior in the leech. I. Passive biomechanical properties of the body wall
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Wilson R.J.A., Skierezynski B.A., Meyer J.K., Skalak R., and Kristan Jr. W.B. Mapping motor neuron activity to overt behavior in the leech. I. Passive biomechanical properties of the body wall. J Comp Physiol A 178 (1996) 637-654
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22
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Muscle function in animal movement: Passive mechanical properties of leech muscle
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This paper advances our understanding of muscle function during animal movements by describing passive muscle tensions caused by step-stretch experiments and by realistic rhythmic changes in length, such as occur during leech swimming locomotion. Although tensions generated by step changes in length are complex, requiring models that comprise three nonlinear springs, and two nonlinear dashpots, tensions generated by rhythmic stretching are described well by a single nonlinear spring, without any dynamics. Hence, this paper supports the surprising conclusion that passive muscle tensions during rhythmic locomotion may not require the use of Hill-type models for muscle dynamics.
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Tian J., Iwasaki T., and Friesen W.O. Muscle function in animal movement: Passive mechanical properties of leech muscle. J Comp Physiol A 193 (2007) 1205-1219. This paper advances our understanding of muscle function during animal movements by describing passive muscle tensions caused by step-stretch experiments and by realistic rhythmic changes in length, such as occur during leech swimming locomotion. Although tensions generated by step changes in length are complex, requiring models that comprise three nonlinear springs, and two nonlinear dashpots, tensions generated by rhythmic stretching are described well by a single nonlinear spring, without any dynamics. Hence, this paper supports the surprising conclusion that passive muscle tensions during rhythmic locomotion may not require the use of Hill-type models for muscle dynamics.
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J Comp Physiol A
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Tian, J.1
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An elastic rod model for anguilliform swimming
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This paper describes a very thorough modeling study of the mechanics underlying anguilliform swimming, in which the animal is viewed as an elliptical elastic rod. Time-dependent curvature changes are imposed onto the rod to generate a traveling wave that, through fluid interactions, simulates animal swimming movements. The study includes both mathematical analyses of forces on the continuous rod and numerical simulations of a rod comprising up to 80 discrete segments. The investigation includes the effects of stiffness (analogous to passive and active muscle tensions), body shape, and even motor neuron patterns on swim execution. This study is an important theoretical contribution to an essential step in the formulation of complete models of swimming locomotion. These will comprise central oscillators, sensory feedback, body and fluid mechanics, initiation and termination control mechanisms, and neuromodulation.
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McMillen T., and Holmes P. An elastic rod model for anguilliform swimming. J Math Biol 53 (2006) 843-886. This paper describes a very thorough modeling study of the mechanics underlying anguilliform swimming, in which the animal is viewed as an elliptical elastic rod. Time-dependent curvature changes are imposed onto the rod to generate a traveling wave that, through fluid interactions, simulates animal swimming movements. The study includes both mathematical analyses of forces on the continuous rod and numerical simulations of a rod comprising up to 80 discrete segments. The investigation includes the effects of stiffness (analogous to passive and active muscle tensions), body shape, and even motor neuron patterns on swim execution. This study is an important theoretical contribution to an essential step in the formulation of complete models of swimming locomotion. These will comprise central oscillators, sensory feedback, body and fluid mechanics, initiation and termination control mechanisms, and neuromodulation.
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J Math Biol
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McMillen, T.1
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Using a systems approach, this paper presents a mathematical model of intermediate complexity to analyze the segmental and intersegmental circuits that generate the swimming rhythm of the leech. Using electrophysiological synaptic and impulse conduction data from physiological experiments, the time dependent and static properties are modeled for individual ganglia. The segmental models are then linked by identified intersegmental interactions to form a coordinated system. This novel approach, with an intermediate level of biophysical realism, generates quantitative predictions of period, phase, and amplitude, and also reveals the relationships between model parameters and system properties.
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Zheng M., Friesen W.O., and Iwasaki T. Systems-level modeling of neuronal circuits for leech swimming. J Comput Neurosci 22 (2007) 21-38. Using a systems approach, this paper presents a mathematical model of intermediate complexity to analyze the segmental and intersegmental circuits that generate the swimming rhythm of the leech. Using electrophysiological synaptic and impulse conduction data from physiological experiments, the time dependent and static properties are modeled for individual ganglia. The segmental models are then linked by identified intersegmental interactions to form a coordinated system. This novel approach, with an intermediate level of biophysical realism, generates quantitative predictions of period, phase, and amplitude, and also reveals the relationships between model parameters and system properties.
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J Comput Neurosci
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Esch T., Mesce K.A., and Kristan Jr. W.B. Evidence for sequential decision making in the medicinal leech. J Neurosci 22 (2002) 11045-11054
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Crisp K.M., and Mesce K.A. A cephalic projection neuron involved in locomotion is dye coupled to the dopaminergic neural network in the medicinal leech. J Exp Biol 207 (2004) 4535-4542
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Stimulating a single nerve in an isolated nervous system in exactly the same way again and again produced either swimming or crawling in the whole nerve cord. The neurons that predicted swimming or crawling earliest also biased the choice when they were depolarized or hyperpolarized. The activity of these neurons individually did not predict the outcome; only as a group of six or more were they reliable predictors. This study strongly suggests that the choice between these two behaviors is a subtle one, made by the concerted activity of a neuronal population.
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Briggman K.L., Abarbanel H.D.I., and Kristan Jr. W.B. Optical imaging of neuronal populations during decision-making. Science 307 (2005) 896-901. Stimulating a single nerve in an isolated nervous system in exactly the same way again and again produced either swimming or crawling in the whole nerve cord. The neurons that predicted swimming or crawling earliest also biased the choice when they were depolarized or hyperpolarized. The activity of these neurons individually did not predict the outcome; only as a group of six or more were they reliable predictors. This study strongly suggests that the choice between these two behaviors is a subtle one, made by the concerted activity of a neuronal population.
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Science
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Imaging dedicated and multifunctional neural circuits generating distinct behaviors
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This study used voltage-sensitive dyes to record from 317 of the 400 neurons in a leech segmental ganglion while the swimming and crawling central pattern generators were active. Of these 317, 188 oscillated in phase with crawling and 90 oscillated in phase with swimming. Of the 90 swim-phased neurons, 84 of them were also phased-locked to crawling. This implies that many of the neurons in the swim CPG are also used in the crawl CPG. This suggests a very high degree of shared neurons (multifunctionality) in the two pattern generators.
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Briggman K.L., and Kristan Jr. W.B. Imaging dedicated and multifunctional neural circuits generating distinct behaviors. J Neurosci 26 (2006) 10925-10933. This study used voltage-sensitive dyes to record from 317 of the 400 neurons in a leech segmental ganglion while the swimming and crawling central pattern generators were active. Of these 317, 188 oscillated in phase with crawling and 90 oscillated in phase with swimming. Of the 90 swim-phased neurons, 84 of them were also phased-locked to crawling. This implies that many of the neurons in the swim CPG are also used in the crawl CPG. This suggests a very high degree of shared neurons (multifunctionality) in the two pattern generators.
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Kristan Jr. W.B., and Gillette R. Decision-making in small neuronal networks. In: North G., and Greenspan R. (Eds). Invertebrate Neurobiology (2007), Cold Spring Harbor Laboratory Press 533-554
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(2007)
Invertebrate Neurobiology
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Kristan Jr., W.B.1
Gillette, R.2
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38
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33646095767
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Persistent responses to brief stimuli: feedback excitation
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The authors investigated the long-held, but weakly supported view that sustained locomotory movements induced by even very brief stimulation arise from reciprocal excitatory interactions in neuron circuits. Using whole-cell patch recording techniques they demonstrated that such mutually excitatory interactions exist in the caudal hindbrain and rostral spinal cord of Xenopus tadpoles. Electrophysiological and modeling studies were used to show that these interactions could be the source of sustained excitatory drive to maintain swimming activity. The paper provides the best evidence to date that excitatory synaptic feedback can lead to prolonged excitation in locomotory networks.
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Li W.-C., Soffe S.R., Wolf E., and Roberts A. Persistent responses to brief stimuli: feedback excitation. J Neurosci 26 (2006) 4026-4035. The authors investigated the long-held, but weakly supported view that sustained locomotory movements induced by even very brief stimulation arise from reciprocal excitatory interactions in neuron circuits. Using whole-cell patch recording techniques they demonstrated that such mutually excitatory interactions exist in the caudal hindbrain and rostral spinal cord of Xenopus tadpoles. Electrophysiological and modeling studies were used to show that these interactions could be the source of sustained excitatory drive to maintain swimming activity. The paper provides the best evidence to date that excitatory synaptic feedback can lead to prolonged excitation in locomotory networks.
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(2006)
J Neurosci
, vol.26
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Li, W.-C.1
Soffe, S.R.2
Wolf, E.3
Roberts, A.4
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39
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0036824323
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Cellular bases of a vertebrate locomotor system-steering, intersegmental and segmental co-ordination and sensory control
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Grillner S., and Wallén P. Cellular bases of a vertebrate locomotor system-steering, intersegmental and segmental co-ordination and sensory control. Brain Res Rev 40 (2002) 92-106
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Brain Res Rev
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Grillner, S.1
Wallén, P.2
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40
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33644611212
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Development of swimming in the medicinal leech; the gradual acquisition of a behavior
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French K.A., Chang J., Reynolds S., Gonzalez R., Kristan III W.B., and Kristan Jr. W.B. Development of swimming in the medicinal leech; the gradual acquisition of a behavior. J Comp Physiol 191 (2005) 813-821
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(2005)
J Comp Physiol
, vol.191
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French, K.A.1
Chang, J.2
Reynolds, S.3
Gonzalez, R.4
Kristan III, W.B.5
Kristan Jr., W.B.6
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