“…However, in discrete-trial chemosensory blocking and overshadowing experiments, we have observed opposite effects: Chemosensory stimuli potentiate conditioning to light (Farley & Jin, 1997;Farley, Reasoner, & Janssen, 1997). Nonetheless, accepting Rogers and Matzel's (1995) results at face value, we have previously discussed in detail their possible interpretation and significance (Farley et al, 1997, pp. 336-338).…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
confidence: 93%
“…On the contrary, the animal has learned two, mutually antagonistic responses. In contrast, control animals that received nonreinforced scallop exposures during the first phase of training, followed by compound light, scallop, and rotation pairings [designated as groups (AϪ)A:P and (IOϩ/AϪ)A:P from Experiments 1 and 2 of Rogers & Matzel (1995)] showed foot contraction to light due to the unopposed light-shaking association.…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
confidence: 99%
“…Rogers and Matzel (1995) used chemosensory cues as contextual stimuli and training methods essentially identical to those in Rogers et al (1996). No direct measures of conditioning to the chemosensory cue that signaled shaking were reported in this study (e.g., changes in feeding latencies, preference for the reinforced chemosensory contextual cue in choice tests, etc.).…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
We critically review chemosensory conditioning studies with molluscs and find that, in many studies, the influence of nonassociative processes complicates, obscures, and renders ambiguous the unique contribution of associative learning. These nonassociative processes include sensory adaptation, habituation, sensitization, and changes in feeding motivation. They arise from both the food extracts that have often been used as conditioned stimuli and the aversive stimuli that have been used as unconditioned stimuli.
“…However, in discrete-trial chemosensory blocking and overshadowing experiments, we have observed opposite effects: Chemosensory stimuli potentiate conditioning to light (Farley & Jin, 1997;Farley, Reasoner, & Janssen, 1997). Nonetheless, accepting Rogers and Matzel's (1995) results at face value, we have previously discussed in detail their possible interpretation and significance (Farley et al, 1997, pp. 336-338).…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
confidence: 93%
“…On the contrary, the animal has learned two, mutually antagonistic responses. In contrast, control animals that received nonreinforced scallop exposures during the first phase of training, followed by compound light, scallop, and rotation pairings [designated as groups (AϪ)A:P and (IOϩ/AϪ)A:P from Experiments 1 and 2 of Rogers & Matzel (1995)] showed foot contraction to light due to the unopposed light-shaking association.…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
confidence: 99%
“…Rogers and Matzel (1995) used chemosensory cues as contextual stimuli and training methods essentially identical to those in Rogers et al (1996). No direct measures of conditioning to the chemosensory cue that signaled shaking were reported in this study (e.g., changes in feeding latencies, preference for the reinforced chemosensory contextual cue in choice tests, etc.).…”
Section: Additional Reports Of Contextual Conditioning In Hermissendamentioning
We critically review chemosensory conditioning studies with molluscs and find that, in many studies, the influence of nonassociative processes complicates, obscures, and renders ambiguous the unique contribution of associative learning. These nonassociative processes include sensory adaptation, habituation, sensitization, and changes in feeding motivation. They arise from both the food extracts that have often been used as conditioned stimuli and the aversive stimuli that have been used as unconditioned stimuli.
“…Studies have also shown that a chemosensory CS, when paired with rotation, suppresses bite-strike responses normally elicited by the chemosensory CS prior to conditioning (Farley et al 1990a). Rogers and Matzel (1996) reported that an excitatory context produced by presenting unsignaled USs (rotation) in a context of chemosensory stimuli blocked later conditioned foot-shortening produced by a light CS paired with rotation US within that context. More recently, it was reported that explicitly unpaired presentations of the CS and US produced conditioned inhibition expressed by increased phototactic behavior (Britton and Farley 1999).…”
Section: Pavlovian Conditioningmentioning
confidence: 99%
“…Additional evidence implicating 5-HT in the US pathway and in conditioning of Hermissenda comes from studies showing that pharmacological agents that affect 5-HT neurotransmission (imipramine, bufotenine, and 5,7-DHT) attenuate in vitro conditioning correlates in type B-photoreceptors ). In addition, 5-HT modulates generator potentials and membrane conductances in type B photoreceptors, modifications that have been identified as neural correlates of Pavlovian conditioning (Crow and Bridge 1985;Farley and Wu 1989;Acosta-Urquidi and Crow 1993;Rogers and Matzel 1995;Crow 1995, 1996). A computational model of the type B photoreceptor used to investigate the contribution of different ionic conductances modulated by 5-HT to the enhanced excitability produced by 5-HT suggested that changes in I A , I K,Ca , or I h (Yamoah et al 1998) would produce excitability changes comparable to experimental findings (Cai et al 2003).…”
Section: Convergence Of the Cs And Us Pathwaysmentioning
The less-complex central nervous system of many invertebrates make them attractive for not only the molecular analysis of the associative learning and memory, but also in determining how neural circuits are modified by learning to generate changes in behavior. The nudibranch mollusk Hermissenda crassicornis is a preparation that has contributed to an understanding of cellular and molecular mechanisms of Pavlovian conditioning. Identified neurons in the conditioned stimulus (CS) pathway have been studied in detail using biophysical, biochemical, and molecular techniques. These studies have resulted in the identification and characterization of specific membrane conductances contributing to enhanced excitability and synaptic facilitation in the CS pathway of conditioned animals. Second-messenger systems activated by the CS and US have been examined, and proteins that are regulated by one-trial and multi-trial Pavlovian conditioning have been identified in the CS pathway. The recent progress that has been made in the identification of the neural circuitry supporting the unconditioned response (UR) and conditioned response (CR) now provides for the opportunity to understand how Pavlovian conditioning is expressed in behavior.The analysis of learning in several vertebrate and invertebrate nervous systems has generated a number of candidate mechanisms of Pavlovian conditioning involving changes in both cellular excitability and synaptic strength. In general, studies of associative learning in these model systems have been dominated by the search for mechanisms of Pavlovian conditioning that provide an explanation for temporal contiguity between the conditioned stimulus (CS) and unconditioned stimulus (US). However, a comprehensive analysis of associative learning requires an understanding of all aspects of the associative process, including the generation of behavior, in addition to providing insights into mechanisms of temporal contiguity. The issue of how learning is expressed in behavior may be effectively addressed by studying conditioning from a cellular and synaptic perspective in the relatively simple nervous system of invertebrates. The analyses of learning in several model systems have used a combination of cellular and synaptic physiology in conjunction with a neural circuit analysis to examine how Pavlovian conditioning is expressed in behavior, or how learning results in the generation of a conditioned response (CR). Some invertebrate preparations are especially attractive for this type of analysis, as the neural circuitry supporting behaviors involving muscular contraction, respiration, locomotion, and feeding is known in considerable detail (for review, see Sahley and Crow 1998).One animal that has contributed to an understanding of the physiology of learning and memory at a cellular, synaptic, and systems level of analysis is the nudibranch mollusk Hermissenda crassicornis. Associative learning in Hermissenda has been extensively examined using a Pavlovian conditioning procedure. The Hermissenda central...
A breakthrough for studying the neuronal basis of learning emerged when invertebrates with simple nervous systems, such as the sea slug Hermissenda crassicornis, were shown to exhibit classical conditioning. Hermissenda learns to associate light with turbulence: prior to learning, naive animals move toward light (phototaxis) and contract their foot in response to turbulence; after learning, conditioned animals delay phototaxis in response to light. The photoreceptors of the eye, which receive monosynaptic inputs from statocyst hair cells, are both sensory neurons and the first site of sensory convergence. The memory of light associated with turbulence is stored as changes in intrinsic and synaptic currents in these photoreceptors. The subcellular mechanisms producing these changes include activation of protein kinase C and MAP kinase, which act as coincidence detectors because they are activated by convergent signaling pathways. Pathways of interneurons and motorneurons, where additional changes in excitability and synaptic connections are found, contribute to delayed phototaxis. Bursting activity recorded at several points suggest the existence of small networks that produce complex spatiotemporal firing patterns. Thus, the change in behavior may be produced by a nonlinear transformation of spatiotemporal firing patterns caused by plasticity of synaptic and intrinsic channels. The change in currents and the activation of PKC and MAPK produced by associative learning are similar to those observed in hippocampal and cerebellar neurons after rabbit classical conditioning, suggesting that these represent general mechanisms of memory storage. Thus, the knowledge gained from further study of Hermissenda will continue to illuminate mechanisms of mammalian learning. Anat Rec (Part B: New Anat) 289B:25-37, 2006.
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