[Bithynia (Gabbia) kiusiuensis in Takamatsu. 2. The operculum and radula].
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[Parafossarulus manchouricus in Miyagi Prefecture. 2. Radula].
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[Observation on the radula of Oncomelania from four counties in Guangxi].
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Costorage and corelease of modulatory peptide cotransmitters with partially antagonistic actions on the accessory radula closer muscle of Aplysia californica.
Many neurons that contain a classical neurotransmitter also contain modulatory peptides, but it has been difficult to establish unequivocally that these peptides are functional cotransmitters. Here, we provide evidence for functional cotransmission in a neuromuscular system of Aplysia. Using immunocytochemical techniques, we localize members of two peptide families, the small cardioactive peptides (SCPs) and the buccalins (BUCs), to a single subset of dense-core vesicles in the terminals of the cholinergic motorneuron B15. We describe a new preparation and method for the direct detection of released peptides and show that the SCPs and BUCs are released when neuron B15 is intracellularly stimulated. Consistent with their subcellular localization, the SCPs and BUCs are released in a stoichiometric ratio that is constant across conditions that change the absolute amount of peptides released. Peptide release is calcium-dependent but does not require muscle contractions. Thus, the release cannot be attributed to a displacement of peptides that may be present in the extracellular space. In previous studies, we characterized the physiological firing patterns of neuron B15. Here, we simulate these firing patterns and show that peptide release occurs. Additionally, we find that significant quantities of material are released under behaviorally relevant conditions. We find that concentrations of released peptides in the muscle are in the concentration range in which exogenously applied peptides exert characterized modulatory actions on muscle contractions. Together, our findings provide strong support for the hypothesis that peptides contained in neuron B15 are functional cotransmitters.
Animals↗
Phase-locked coordination between two rhythmically active feeding structures in the mollusk Clione limacina. I. Motor neurons.
Coordination between different motor centers is essential for the orderly production of all complex behaviors, in both vertebrates and invertebrates. The current study revealed that rhythmic activities of two feeding structures of the pteropod mollusk Clione limacina, radula and hooks, which are used to extract the prey from its shell, are highly coordinated in a phase-dependent manner. Hook protraction always coincided with radula retraction, while hook retraction coincided with radula protraction. Thus hooks and radula were always moving in the opposite phases, taking turns grabbing and pulling the prey tissue out of the shell. Identified buccal ganglia motor neurons controlling radula and hooks protraction and retraction were rhythmically active in the same phase-dependent manner. Hook protractor motor neurons were active in the same phase with radula retractor motor neurons, while hook retractor motor neurons burst in phase with radula protractor motor neurons. One of the main mechanisms underlying the phase-locked coordination was electrical coupling between hook protractor and radula retractor motor neurons. In addition, reciprocal inhibitory synaptic connections were found between hook protractor and radula protractor motor neurons. These electrical and inhibitory synaptic connections ensure that rhythmically active hooks and radula controlling motor neurons are coordinated in the specific phase-dependent manner described above. The possible existence of a single multifunctional central pattern generator for both radula and hook motor centers is discussed.
Action Potentials↗
In vivo buccal nerve activity that distinguishes ingestion from rejection can be used to predict behavioral transitions in Aplysia.
1. We are studying the neural basis of consummatory feeding behavior in Aplysia using intact, freely moving animals. 2. Video records show that the timing of radula closure during the radula protraction-retraction cycle constitutes a major difference between ingestion (biting or swallowing) and rejection. During ingestion, the radula is closed as it retracts. During rejection, the radula is closed as it protracts. 3. We observed two patterns of activity in nerves which are likely to mediate these radula movements. Patterns I and II are associated with ingestion and rejection, respectively, and are distinguished by the timing of radula nerve activity with respect to the onset of buccal nerve 2 activity. 4. The association of ingestion with pattern I is maintained when the animal feeds on a polyethylene tube, the same food substrate used to elicit rejection responses. Under these conditions, pattern I is associated with either swallowing or no net tube movement. 5. Most transitions from swallowing to rejection were preceded by one or more occurrences of pattern I in which there was no net tube movement, suggesting that these transitions can be predicted. 6. Our data suggest that these two patterns can be used to distinguish ingestion from rejection.
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Activity patterns of the B31/B32 pattern initiators innervating the I2 muscle of the buccal mass during normal feeding movements in Aplysia californica.
1. B31 and B32 are pattern-initiator neurons in the buccal ganglia of Aplysia. Along with the B61/B62 neurons, B31/B32 are also motor neurons that innervate the 12 buccal muscle via the I2 nerve. This research was aimed at determining the physiological functions of the B31/B32 and B61/B62 neurons, and of the I2 muscle. 2. Stimulating the I2 muscle in the radula rest position produces radula protraction. In addition, in behaving animals lesioning either the muscle or the I2 nerve greatly reduces radula protraction. 3. During buccal motor programs in reduced preparations, B31/B32 and B61/62 fire preceding activity in neuron B4, whose firing indicates the onset of radula retraction. In addition, during both ingestion-like and rejection-like patterns the activity in the I2 nerve is correlated with protraction. 4. B31/B32 fire at frequencies of 15-25 Hz. Neither B31/B32 nor B61/B62 elicit facilitating end-junction potentials (EJPs) and electromyograms (EMGs) in the I2 muscle. EMGs from B31/B32 are smaller than those from B61/B62. B31/B32 and B61/B62 innervate all areas of the muscle approximately uniformly. 5. In behaving animals, EMGs consistent with B31/B32 activity are seen in the I2 muscle during the protraction phase of biting, swallowing, and rejection movements. In addition, the I2 muscle receives inputs that cannot be attributed to either the B31/B32 or B61/B62 neurons, either because the potentials are too large, firing frequencies are too low, or a prominent facilitation is seen. Such potentials are associated with lip movements, and also with radula retraction. 6. EMGs were recorded from the I2 muscle during feeding behavior after a lesion of the I2 nerve. Animals that had severe deficits in protraction showed no activity consistent with B31/B32 or B61/B62, but did show activity during retraction. 7. Our data indicate that the I2 muscle and the B31/B32 motor neurons are essential constituents contributing to protraction movements. Activity in these neurons is associated with radula protraction, which occurs as a component of a number of different feeding movements. The I2 muscle may also contribute to retraction, via activation by other motor neurons.
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Interneuronal basis of the generation of related but distinct motor programs in Aplysia: implications for current neuronal models of vertebrate intralimb coordination.
Coordination of two sets of movements, protraction-retraction versus opening-closing, of the feeding apparatus (the radula) in ingestive and egestive motor programs of Aplysia resembles vertebrate intralimb coordination in that the relative timing of the two sets of movements differs in the two motor programs. In both ingestion and egestion, radula protraction and retraction alternate, whereas radula closure shifts its phase relative to protraction-retraction. In egestion, the radula closes in protraction; in ingestion, the radula closes in retraction. In both ingestive and egestive motor programs elicited by the command-like neuron, cerebral-buccal interneuron-2 (CBI-2), the protraction and retraction movements are mediated by the same sets of controller interneurons. In contrast, radula closure is mediated by two controller interneurons, B20 and B40, that are preferentially active in egestion and ingestion, respectively. In egestion, B20, active in protraction, drives closure motorneuron B8 in protraction, whereas in ingestion, B40, also active in protraction, uses a functionally novel mechanism, fast inhibition and slow excitation, to drive B8 in retraction. Our findings are summarized in a neural model that permits a conceptual comparison of our model with two previous hypothetical models of intralimb coordination in spinal circuits that were proposed by Grillner (1981, 1985) and Berkowitz and Stein (1994). Although our model supports the existence of separate controllers for different movements as in the Grillner (1981, 1985) model; in terms of basic mechanisms, our model is similar to the Berkowitz and Stein (1994) model because the closure movement is mediated by separate controllers in different programs, and thus both models can be classified as recruitment models.
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Neural mechanisms of motor program switching in Aplysia.
The Aplysia multifunctional feeding central pattern generator (CPG) produces at least two types of motor programs, ingestion and egestion, that involve two sets of radula movements, protraction-retraction and opening-closing movements. In ingestion, the radula closes during retraction to pull food in, whereas in egestion, the radula closes during protraction to push inedible objects out. Thus, radula closure shifts the phase in which it occurs with respect to protraction-retraction in the two programs. To identify the central switching mechanisms, we compared activity of CPG neurons during the two types of motor programs elicited by a higher-order interneuron, cerebral-buccal interneuron-2 (CBI-2). Although CPG elements (B63, B34, and B64) that mediate the protraction-retraction sequence are active in both programs, two other CPG elements, B20 and B4/5, are preferentially active in egestive programs and play a major role in mediating CBI-2-elicited egestive programs. Both B20 and B4/5 control the phasing of radula closure motoneurons (B8 and B16) to ensure that, in egestive programs, these motoneurons fire and produce radula-closing movements only during protraction. Elsewhere, another higher-order interneuron, CBI-3, was shown to convert CBI-2-elicited egestion to ingestion. We show that CBI-3 switches the programs by suppressing the activity of B20 and B4/5. CBI-3, active only during protraction, accomplishes this through fast inhibition of B20 during protraction and slow inhibition of B4/5 during retraction. The slow inhibition is mimicked and occluded by APGWamide, a neuropeptide contained in CBI-3. Thus, fast conventional and slow peptidergic transmissions originating from the same interneuron act in concert to meet specific temporal requirements in pattern switching.
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Control of feeding movements in the pteropod mollusc, Clione limacina.
(1) The buccal apparatus of the pteropod marine mollusc Clione limacina, isolated together with buccal ganglia, could perform rhythmic feeding movements. Movements of the radula and the hooks (which the Clione inserts into the body of its prey) as well as the electroneurogram of the radular nerve were recorded. Usually one could observe rhythmic radula movements alone, while the hooks were motionless. But sometimes the hooks also performed rhythmic movements which were more or less synchronous with those of the radula. The radula movement cycle consisted of the protraction and the retraction phases, which were occasionally followed by a quiescent phase. Corresponding to each radula movement was a burst of activity in the radular nerve, consisting of the protractor and the retractor components. (2) Isolated buccal ganglia were capable of feeding rhythm generation. Most of the buccal neurons exhibited rhythmic activity correlating with the activity in the radular nerve. According to the phase of activity in the feeding cycle, rhythmic neurons were divided into two groups - the protractor and the retractor ones. The neurons within each of the groups were electrically coupled with each other. The protractor and retractor neurons inhibited each other. (3) Protractor and retractor neurons were extracted from buccal ganglia by means of a microelectrode. Many isolated cells generated slow oscillations of membrane potential and bursts of spikes, the pattern of this activity being similar to that before isolation. (4) A model of the feeding rhythm generator is discussed. It consists of two (protractor and retractor) groups of neurons with mutual inhibitory connections, neurons of each group being endogenous bursters.
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Synaptic modulation contributes to firing pattern generation in jaw motor neurons during rejection of seaweed in Aplysia kurodai.
Japanese species, Aplysia kurodai, feeds well on Ulva but rejects Gelidium (Geli.) or Pachydictyon (Pach.) with different rhythmic patterned movements of the jaws and radula. During ingestion the jaws open at the radula-protraction phase and remain half open at the initial phase of the radula-retraction, whereas during rejection the jaws open similarly but start to close immediately after the onset of the radula-retraction. These can be induced not only by the natural seaweed but by the extract solutions. We previously showed that the change of the patterned jaw movements from the ingestion to the rejection may result from the decrease in the delay of the firing onset of the jaw-closing (JC) motor neurons during their depolarization. This diminished delay produces a phase advance relative to the radula-retraction phase. In that study, we showed that during ingestion the buccal multiaction (MA) neurons may generate the delay of firing onset of the JC motor neurons by producing monosynaptic inhibitory postsynaptic potentials (IPSPs) during the initial portion of their depolarization. In the present experiments, the firing patterns in the MA neurons induced by application of the Geli. or Pach. extract to the lips were explored in the semi-intact preparations. During the Pach. response the duration and the firing frequency of the MA firing at each depolarizing phase were decreased in comparison with the Ulva response. No decreases in the MA firing were observed during the Geli. response, suggesting that the similar patterned jaw movements during rejection of Geli. and Pach. may be generated by different neural mechanisms. Moreover, the size of the MA-induced IPSP in the JC motor neurons was largely decreased by application of the Geli. or Pach. extract to the lips in the reduced preparations consisting of the tentacle-lips and the cerebral-buccal ganglia. Voltage-clamp experiments on the JC motor neurons showed that the size of synaptic current induced by the MA spikes was decreased by application of these solutions to the lips. The decrease was induced when the buccal ganglia were bathed in a solution to block polysynaptic pathways. These results suggest that the advance of the onset of the JC firing at each depolarizing phase during the Geli. or Pach. response may be mainly or partly caused by the decrease in the size of the MA-induced IPSP in the motor neurons. Modulatory action of cerebral neurons or peripheral afferent neurons in the lip region may contribute to this synaptic plasticity.
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Feeding neural networks in the mollusc Aplysia.
Aplysia feeding is striking in that it is executed with a great deal of plasticity. At least in part, this flexibility is a result of the organization of the feeding neural network. To illustrate this, we primarily discuss motor programs triggered via stimulation of the command-like cerebral-buccal interneuron 2 (CBI-2). CBI-2 is interesting in that it can generate motor programs that serve opposing functions, i.e., programs can be ingestive or egestive. When programs are egestive, radula-closing motor neurons are activated during the protraction phase of the motor program. When programs are ingestive, radula-closing motor neurons are activated during retraction. When motor programs change in nature, activity in the radula-closing circuitry is altered. Thus, CBI-2 stimulation stereotypically activates the protraction and retraction circuitry, with protraction being generated first, and retraction immediately thereafter. In contrast, radula-closing motor neurons can be activated during either protraction or retraction. Which will occur is determined by whether other cerebral and buccal neurons are recruited, e.g. radula-closing motor neurons tend to be activated during retraction if a second CBI, CBI-3, is recruited. Fundamentally different motor programs are, therefore, generated because CBI-2 activates some interneurons in a stereotypic manner and other interneurons in a variable manner.
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[Classification Of Planorbidae Cellected From Geum River Basin]
The distribution, external morphology, radula, chromosome numbers of Planorbidae snails were studied. 1. The specimens were collected at four stations in Nonsangun, Kongjugun, and Daedukgun which are located around Geum river. Three genera and three species of Planorbidae, Hippeutis cantori, Segmentina hemisphaerula and Gyraulus convexiusculus, were collected. H. cantori was the most abundant species among the three species. G. convexiusculus was the least abundant one. Each species could be identified on the basis of its external characteristic, since the periphery of each species has a peculiar shape. H. cantori was the largest one among the three species. The radula formula of each species was very similar to other species. The size of radula was proportional to the size of shell. The radula formulae of H. cantori, S. hemisphaerula, and G. convexiusculus were 29:1:29, 23:l:23, and 16:1:16 respectively. The difference of radula formula could be found in the total numbers of laternal and marginal teeth. The haploid chromosome number of H. cantori was eighteen (n=18). S. hemisphaerula and G. convexiusculus were assumed to be same in their chromosome numbers (n=18).
Journal Article↗
Proprioceptive input to feeding motor programs in Aplysia.
Although central pattern generators (CPGs) can produce rhythmic activity in isolation, it is now generally accepted that under physiological conditions information from the external and internal environment is incorporated into CPG-induced motor programs. Experimentally advantageous invertebrate preparations may be particularly useful for studies that seek to characterize the cellular mechanisms that make this possible. In these experiments, we study sensorimotor integration in the feeding circuitry of the mollusc Aplysia. We show that a premotor neuron with plateau properties, B51, is important for generating the radula closing/retraction phase of ingestive motor programs. When B51 is depolarized in semi-intact preparations, radula closing/retractions are enhanced. When B51 is hyperpolarized, radula closing/retractions are reduced in size. In addition to being important as a premotor interneuron, B51 is also a sensory neuron that is activated when the feeding apparatus, the radula, rotates backward. The number of centripetal spikes in B51 is increased if the resistance to backward rotation is increased. Thus, B51 is a proprioceptor that is likely to be part of a feedback loop that insures that food will be moved into the buccal cavity when difficulty is encountered. Our data suggest, therefore, that Aplysia are able to adjust feeding motor programs to accommodate the specific qualities of the food ingested because at least one of the neurons that generates the basic ingestive motor program also serves as an on-line monitor of the success of radula movements.
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Distribution of cell wall components in Sphagnum hyaline cells and in liverwort and hornwort elaters.
Spiral secondary walls are found in hyaline cells of Sphagnum, in the elaters of most liverworts, and in elaters of the hornwort Megaceros. Recent studies on these cells suggest that cytoskeletal and ultrastructural processes involved in cell differentiation and secondary wall formation are similar in bryophytes and vascular plant tracheary elements. To examine differences in wall structure, primary and secondary wall constituents of the hyaline cells of Sphagnum novo-zelandicum and elaters of the liverwort Radula buccinifera and the hornwort Megaceros gracilis were analyzed by immunohistochemical and chemical methods. Anti-arabinogalactan-protein antibodies, JIM8 and JIM13, labeled the central fibrillar secondary wall layer of Megaceros elaters and the walls of Sphagnum leaf cells, but did not label the walls of Radula elaters. The CCRC-M7 antibody, which detects an arabinosylated (1-->6)-linked beta-galactan epitope, exclusively labeled hyaline cells in Sphagnum leaves and the secondary walls of Radula elaters. Anti-pectin antibodies, LM5 and JIM5, labeled the primary wall in Megaceros elaters. LM5 also labeled the central layer of the secondary wall but only during formation. In Radula elaters, JIM5 and another anti-pectin antibody, JIM7, labeled the primary wall. The distribution of arabinogalactan-proteins and pectic polysaccharides restricted to specific wall types and stages of development provides evidence for the developmental and functional regulation of cell wall composition in bryophytes. Monosaccharide-linkage analysis of Sphagnum leaf cell walls suggests they contain polysaccharides similar to those of higher plants. The most abundant linkage was 4-Glc, typical of cellulose, but there was also evidence for xyloglucans, 4-linked mannans, 4-linked xylans and rhamnogalacturonan-type polysaccharides.
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Afferent-induced changes in rhythmic motor programs in the feeding circuitry of aplysia.
A manipulation often used to determine whether a neuron plays a role in the generation of a motor program involves injecting current into the cell during rhythmic activity to determine whether activity is modified. We perform this type of manipulation to study the impact of afferent activity on feeding-like motor programs in Aplysia. We trigger biting-like programs and manipulate sensory neurons that have been implicated in producing the changes in activity that occur when food is ingested, i.e., when bites are converted to bite-swallows. Sensory neurons that are manipulated are the radula mechanoafferent B21 and the retraction proprioceptor B51. Data suggest that both cells are peripherally activated during radula closing/retraction when food is ingested. We found that phasic subthreshold depolarization of a single sensory neuron can significantly prolong radula closing/retraction, as determined by recording both from interneurons (e.g., B64), and motor neurons (e.g., B15 and B8). Additionally, afferent activity produces a delay in the onset of the subsequent radula opening/protraction, and increases the firing frequency of motor neurons. These are the changes in activity that are seen when food is ingested. These results add to the growing data that implicate B21 and B51 in bite to bite-swallow conversions and indicate that afferent activity is important during feeding in Aplysia.
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Coordinated excitatory effect of GABAergic interneurons on three feeding motor programs in the mollusk Clione limacina.
Coordination between different motor centers is essential for the orderly production of all complex behaviors. Understanding the mechanisms of such coordination during feeding behavior in the carnivorous mollusk Clione limacina is the main goal of the current study. A bilaterally symmetrical interneuron identified in the cerebral ganglia and designated Cr-BM neuron produced coordinated activation of neural networks controlling three main feeding structures: prey capture appendages called buccal cones, chitinous hooks used for prey extraction from the shell, and the toothed radula. The Cr-BM neuron produced strong excitatory inputs to motoneurons controlling buccal cone protraction. It also induced a prominent activation of the neural networks controlling radula and hook rhythmic movements. In addition to the overall activation, Cr-BM neuron synaptic inputs to individual motoneurons coordinated their activity in a phase-dependent manner. The Cr-BM neuron produced depolarizing inputs to the radula protractor and hook retractor motoneurons, which are active in one phase, and hyperpolarizing inputs to the radula retractor and hook protractor motoneurons, which are active in the opposite phase. The Cr-BM neuron used GABA as its neurotransmitter. It was found to be GABA-immunoreactive in the double-labeling experiments. Exogenous GABA mimicked the effects produced by Cr-BM neuron on the postsynaptic neurons. The GABA antagonists bicuculline and picrotoxin blocked Cr-BM neuron-induced PSPs. The prominent coordinating effect produced by the Cr-BM neuron on the neural networks controlling three major elements of the feeding behavior in Clione suggests that this interneuron is an important part of the higher-order system for the feeding behavior.
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Different roles of neurons B63 and B34 that are active during the protraction phase of buccal motor programs in Aplysia californica.
The buccal ganglion of Aplysia contains a central pattern generator (CPG) that organizes sequences of radula protraction and retraction during food ingestion and egestion. Neurons B63 and B34 have access to, or are elements of, the CPG. Both neurons are depolarized along with B31/B32 during the protraction phase of buccal motor programs. Both cells excite the contralateral B31/B32 neurons and inhibit B64 and other neurons active during the retraction phase. B63 and B34 also both have an axon exiting the buccal ganglia via the contralateral cerebrobuccal connective. Despite their similarities, B63 and B34 differ in a number of properties, which reflects their different functions. B63 fires during both ingestion and egestion-like buccal motor programs, whereas B34 fires only during egestion-like programs. The bilateral B63 neurons, along with the bilateral B31 and B32 neurons, act as a single functional unit. Sufficient depolarization of any of these neurons activates them all and initiates a buccal motor program. B63 is electrically coupled to both the ipsilateral and the contralateral B31/B32 neurons but monosynaptically excites the contralateral neurons with a mixed electrical and chemical excitatory postsynaptic potential (EPSP). Positive feedback caused by electrical and chemical EPSPs between B63 and B31/B32 contributes to the sustained depolarization in B31/B32 and the firing of B63 during the protraction phase of a buccal motor program. B34 is excited during the protraction phase of all buccal motor programs, but, unlike B63, it does not always reach firing threshold. The neuron fires in response to current injection only after it is depolarized for 1-2 s or after preceding buccal motor programs in which it is depolarized. Firing of B34 produces facilitating EPSPs in the contralateral B31/B32 and B63 neurons and can initiate a buccal motor program. Firing in B34 is strongly correlated with firing in the B61/B62 motor neurons, which innervate the muscle (I2) responsible for much of protraction. B34 monosynaptically excites these motor neurons. B34 firing is also correlated with firing in motor neuron B8 during the protraction phase of a buccal motor program. B8 innervates the I4 radula closer muscle, which in egestion movements is active during protraction and in ingestion movements is active during retraction. B34 has a mixed, but predominantly excitatory, effect on B8 via a slow conductance-decrease EPSP. Thus firing in B34 leads to amplification of radula protraction that is coupled with radula closing, a pattern characteristic of egestion.
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