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Efferent controls in crustacean mechanoreceptors.

Since the 1960s it has been known that central neural networks can elaborate motor patterns in the absence of any sensory feedback. However, sensory and neuromodulatory inputs allow the animal to adapt the motor command to the actual mechanical configuration or changing needs. Many studies in invertebrates, particularly in crustacea, have described several mechanisms of sensory-motor integration and have shown that part of this integration was supported by the efferent control of the mechanosensory neurons themselves. In this article, we review the findings that support such an efferent control of mechanosensory neurons in crustacea. Various types of crustacean proprioceptors feeding information about joint movements and strains to central neural networks are considered, together with evidence of efferent controls exerted on their sensory neurons. These efferent controls comprise (1) the neurohormonal modulation of the coding properties of sensory neurons by bioamines and peptides; (2) the presynaptic inhibition of sensory neurons by GABA, glutamate and histamine; and (3) the long-term potentiation of sensory-motor synapses by glutamate. Several of these mechanisms can coexist on the same sensory neuron, and the functional significance of such multiple modulations is discussed.

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Architectonics of crayfish ganglia.

The central nervous system of crayfish consists of a chain of segmental ganglia that are linked by cables of intersegmental axons. Each ganglion contains a highly-ordered core of longitudinal tracts, vertical tracts, commissures, and synaptic neuropils. We review from a technical perspective the history of the description of these ganglia, and recognize four episodes of progress. Each major innovation in anatomical methods has led to new insight into the structure and function of this nervous system, and new awareness of the structural patterns that are common to the CNS of all arthropods. Ganglia in different segments of the body differ in size, and appear to differ in anatomy. From a comparison of the structures of the cores of abdominal, thoracic, and subesophageal ganglia, we argue that this apparent difference is illusory. Rather, each of these ganglia is organized on the same plan, a plan also found in insect segmental ganglia. The apparent differences follow from longitudinal compression during development and from allometric growth of particular neuropils associated with innervation of the walking legs. Different authors have described the internal organization of ganglia in different segments, so we provide a cross-reference to the nomenclatures they have introduced. We compare the locations of cell bodies of motor neurons and accessory neurons that innervate different peripheral structures, and demonstrate double-labeling of certain GABAergic peripheral inhibitory neurons. Finally, we describe the construction of digital movies of serial sections of these ganglia, and discuss their utility.

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Development, growth, and plasticity in the crayfish olfactory system.

Decapod crustaceans have a well-defined olfactory system characterised by a set of chemosensitive sensilla grouped together in an array (the olfactory organ) on their antennules. Olfactory receptor neurons in the olfactory organ project exclusively to, and terminate in, cone-shaped olfactory glomeruli in a discrete neuropil in the brain, the olfactory lobe. The olfactory organ appears to be the only afferent input to the olfactory lobe, making the system convenient for the study of its development and growth. The progression of development of the olfactory system is a continuum and can be traced from the first appearance of peripheral receptor cells and central stem cells through to the construction of the tracts and neuropils that constitute the adult system. Cell proliferation leading to the production of peripheral and central olfactory neurons can be observed with mitotic markers in both embryonic stages and in postembryonic growth. Cell proliferation in the olfactory system in crayfish persists throughout the lives of the animals and can be modulated by manipulating the living conditions imposed on growing animals. Large serotonergic neurons that are associated with the olfactory system may play a role in the regulation of cell proliferation.

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Active dendritic properties constrain input-output relationships in neurons of the central olfactory pathway in the crayfish forebrain.

Parasol cells are multimodal sensory interneurons of the hemi-ellipsoid body in the decapod forebrain. In reptant crustaceans, the hemi-ellipsoid body resides in the base of the eyecup, as an appendage to the terminal medulla. Parasol cells exhibit periodic depolarizations at a frequency of 0.5-1.0 Hz. I have investigated the role of these periodic depolarizations and their superimposed impulse bursts in affecting the input/output properties of these neurons. Parasol cells receive input from photic, olfactory, and mechanosensory pathways. Strong stimulation over any one of these pathways can lead to the generation of one or more impulse bursts in a subset of parasol cells, timed to occur at the peak of successive periodic depolarizations. A role for the periodic depolarizations in the function of the parasol cells has yet to be established. I suggest the possibility that they may act as a nonlinear amplifier that boosts spatially-summated excitatory synaptic potentials from strong or appropriate stimuli above threshold for burst generation. Another possibility includes modification of voltage-sensitive ion channels in the dendritic membrane, permitting a more effective spread of excitatory synaptic currents to impulse or burst initiating zones. Impulse bursts may be a highly effective mode of output for these neurons, especially so as they occur synchronously in a subset of cells in response to strong sensory input. Furthermore, backfiring of bursts into the dendritic tree has a brief (2-3 second) but effective suppressive action upon weak sensory input, which can thereby be masked by stronger, burst-generating input. This masking phenomenon is seen in other arthropod sensory interneurons, where its physiological basis appears to be a transient accumulation of intracellular Ca(++) ions that open calcium-sensitive potassium channels.

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The circadian system of crayfish: a developmental approach.

Adult crayfish exhibit a variety of overt circadian rhythms. However, the physiological mechanisms underlying the overt rhythms are controversial. Research has centered on two overt rhythms: the motor activity and the retinal sensitivity rhythms of the genus Procambarus. The present work reviews various studies undertaken to localize pacemakers and mechanisms of entrainment responsible for these two rhythms in adult organisms of this crustacean decapod. It also describes an ontogenetic approach to the problem by means of behavioral, electrophysiological, and neurochemical experiments. The results of this approach confirm previous models proposed for adult crayfish, based on a number of circadian pacemakers distributed in the central nervous system. However, the coupling of rhythmicity between these independent oscillators might be complex and dependent on the interaction between serotonin (5-HT), light, and the crustacean hyperglycemic hormone (CHH). The latter compound has, up until now, not been considered as an agent in the genesis and synchronization of the retinal sensitivity rhythm.

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Neuropeptides in the crayfish stomatogastric nervous system.

Neuropeptides are peptides with profound effects on the nervous system. The function of neuropeptides can be studied in detail in the stomatogastric nervous system (STNS). Neuropeptides are ubiquitously distributed in the STNS and it contains well-studied neural circuits that are strongly modulated by neuropeptides. The STNS controls the movements of the foregut in crustaceans and has been studied intensively in a variety of decapod crustaceans including crayfish. This article reviews our knowledge of neuropeptides in the crayfish STNS. Within crayfish, peptides reach the circuits of the STNS as neurohormones released by neurohaemal organs or by putative neurohemal zones located within the STNS. As transmitters, neuropeptides are present in identified motoneurons, interneurons, and sensory neurons (mainly shown by immunocytochemistry), indicating a multiple role of peptides in the plasticity of neural networks. Neuropeptides are not only present in varicosities within the neuropil of ganglia, but also in varicosities on muscles and within small neuropil patches along nerves. This suggests that the muscles of the stomach are under a more direct modulatory control than previously thought, and that information processing can also occur within nerves. In addition to anatomical studies, biochemical and electrophysiological methods were used. For example, MALDI-TOF MS (matrix-assisted laser desorption ionization time of flight mass spectrometry) revealed the presence of four different peptides of the orcokinin family within a single neuron, and electrophysiological experiments demonstrated that the networks of the STNS are not only under excitatory but also inhibitory peptidergic influence. Comparing the similarities and differences between the STNS of crayfish and that of other decapod crustaceans has already contributed to our knowledge about peptides and will further help to unravel peptide function in the plasticity of neural circuits. For example, the identified neurons in the STNS can be used to study co-transmission because neuropeptides are co-localized with classical transmitters, biogenic amines, or other peptides in these neurons.

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