Functional Organization of the Neural Control of Circulation in Aplysia

The abdominal ganglion of Aplysia provides a useful model forstudying the functional organization of motor systems. Herewe review studies of the neural network controlling circulation,emphasizing the organizational features it may share with othermotor systems controlled by the abdominal ganglion. We identifiedseven motor neurons to the heart and vascular system. Motorneurons having similar motor effects (e.g. the two heart inhibitors,or the three vasoconstrictors), together with cells of unknownmotor function located near them, make up distinct homogeneouscell groups. The members of each group appear to be nearly identicalwith respect to biophysical and neurochemical properties, sizeand effectiveness of synaplie inputs, and firing patterns. Thereare no interconnections between the members of the groups, butfive interneurons innervate the homogeneous groups in variouscombinations, exciting some groups and inhibiting others. Twoof the interneurons, Interneuron I (cell I₁₀) and InterneuronII, are command cells which produce centrally generated motorprograms in the absence of sensory feedback. Eacli command apparentlycodes for a specific homeostatic function, such as increasedcardiac output. Coordination of the two commands is achievedby mutual inhibitory connections between them, ensuring thatthe motor neurons of the system receive only one command ata time. Some synaptic connections made by the command interneuronsappear to be functionally ineffective; the possible significanceof them is discussed. Available evidence suggests that manyfeatures of the network controlling circulation may be characteristicof other visceromotor systems of the abdominal ganglion.     

Morphology and physiology of auditory and vibratory ascending interneurones in bushcrickets

Auditory/vibratory interneurones of the bushcricket species Decticus albifrons and Decticus verrucivorus were studied with intracellular dye injection and electrophysiology. The morphologies of five physiologically characterised auditory/vibratory interneurones are shown in the brain, subesophageal and prothoracic ganglia. Based on their physiology, these five interneurones fall into three groups, the purely auditory or sound neurones: S-neurones, the purely vibratory V-neurones, and the bimodal vibrosensitive VS-neurones. The S1-neurones respond phasically to airborne sound whereas the S4-neurones exhibit a tonic spike pattern. Their somata are located in the prothoracic ganglion and they show an ascending axon with dendrites located in the prothoracic, subesophageal ganglia, and the brain. The VS3-neurone, responding to both auditory and vibratory stimuli in a tonic manner, has its axon traversing the brain, the suboesophageal ganglion and the prothoracic ganglion although with dendrites only in the brain. The V1- and V2-neurones respond to vibratory stimulation of the fore- and midlegs with a tonic discharge pattern, and our data show that they receive inhibitory input suppressing their spontaneous activity. Their axon transverses the prothoracic ganglion, subesophageal ganglion and terminate in the brain with dendritic branching. Thus the auditory S-neurones have dendritic arborizations in all three ganglia (prothoracic, subesophageal, and brain) compared to the vibratory (V) and vibrosensitive (VS) neurones, which have dendrites almost only in the brain. The dendrites of the S-neurones are also more extensive than those of the V-, VS-neurones. V- and VS-neurones terminate more laterally in the brain. Due to an interspecific comparison of the identified auditory interneurones the S1-neurone is found to be homologous to the TN1 of crickets and other bushcrickets, and the S4-neurone also can be called AN2. J. Exp. Zool. 286:219-230, 2000.     

Distribution of NADPH-diaphorase-positive ascending interneurones in the crayfish terminal abdominal ganglion

Previous neuropharmacological studies have described the presence of a nitric oxide-cGMP signalling pathway in the crayfish abdominal nervous system. In this study we have analysed the distribution of putative nitric oxide synthase (NOS)-containing ascending interneurones in the crayfish terminal abdominal ganglion using NADPH-diaphorase (NADPHd) histochemistry. Ascending intersegmental interneurones were stained intracellularly using the fluorescent dye Lucifer yellow and the ganglia containing the stained interneurones subsequently processed for NADPHd activity. Fluorescence persisted throughout histochemical processing. These double-labelling experiments showed that 12 of 18 identified ascending interneurones were NADPHd positive. Thus many ascending interneurones that process mechanosensory signals in the terminal ganglion may contain NOS, and are themselves likely sources of NO which is known to modulate their synaptic inputs. Three clear relationships emerged from our analysis between the effects of NO on the synaptic inputs of interneurones, their output properties and their staining for NADPH-diaphorase. First were class 1 interneurones with no local outputs in the terminal ganglion, the NE type interneurones, which had sensory inputs that were enhanced by NO and were NADPHd positive. Second were class 1 interneurones with local and intersegmental output effects that had sensory inputs that were also enhanced by NO but were NADPHd negative. Third were class 2 interneurones with local and intersegmental outputs that had synaptic inputs that were depressed by the action of NO but were NADPHd positive. These results suggest that NO could selectively enhance specific synaptic connections and sensory processing pathways in local circuits.     

The Role of the Escape Swim Motor Network in the Organization of Behavioral Hierarchy and Arousal in Pleurobranchaea

The escape swimming pattern generator of the notaspid opisthobranchPleurobranchaea drives a high threshold, override behavior.The pattern generator is integrated with neural networks ofother behaviors so as to coordinate unitary behavioral expressionand to promote general behavioral arousal. These functions areseparately produced by different swim network elements. Oneset of swim premotor neurons, the A1/A10 ensemble, A3 and I_VS,generate the swim pattern and, through corollary activity, suppresspotentially conflicting feeding behavior by exerting broad inhibitionat major feeding network interneurons. A second set of swimneurons, the serotonergic As1–4 neurons, provides intrinsicneuromodulatory excitation to the swim pattern generator thatsustains the escape swim episode through multiple cycles. TheAs1–4 also provide neuromodulatory excitation to importantmodulatory, serotonergic cells in the feeding motor networkand locomotor network, and may have a general regulatory rolein the distributed serotonergic arousal network of the mollusk.The As1–4 appear to be also necessary to both avoidanceand orienting turning, and are therefore likely to be critical,multi-functional components upon which much of the organizationof the animal's behavior rests.     

Interactions between the tonic and cyclic postural motor programs in the crayfish abdomen

1. In the crayfish (Procambarus clarkii) abdomen, the superficial flexor and extensor muscles and the motoneurons that innervate them are employed during two completely different modes of behavior: (1) tonic postural adjustments and (2) cyclic movements associated with backwards terrestrial walking. We have tested the possibility that these two behavioral subsystems share at least some of the same tonic premotor interneurons. 2. Of the 108 tonic flexion- and extension-producing interneurons monitored during cyclic pattern generation, only 25 were recruited while 36 were inhibited. None of the recruited interneurons made a measurable contribution to the cyclic motor output. Similarly, none of the 20 inhibitory interneurons of the tonic subsystem recorded in this study was found to play a role in shaping the cyclic motor pattern. 3. Simultaneous activation of single tonic postural interneurons with the cyclic motor pattern revealed that the two behavioral subsystems interact in complex ways. Some tonic interneurons produced motor outputs that overrode the cyclic motor outputs while the motor outputs of other tonic interneurons were completely overwhelmed by the cyclic motor program. Still other tonic interneurons generated motor outputs that predominated over cyclic patterned outputs in some ganglia but were masked by the cyclic motor pattern in other ganglia. 4. Although weak interactions between the two subsystems occur at the premotor level, they have little effect on the normal generation of the cyclic pattern. Stronger interactions apparently occur at the level of the motoneurons and these interactions presumably may form the basis of switching from one behavior to the other. We conclude, therefore, that each behavioral subsystem relies upon its own unique set of premotor interneurons. Finally, those interneurons contributing to the cyclic motor pattern have not yet been identified.     

硕螽(Deracantha onos)听觉中间神经元的声反应特征

本文报道了硕螽听通路单个听觉中间神经元的声反应特征。依据动作电位发放模式的不同,听觉中间神经元可分为两类,即紧张型与相位型。紧张型听觉中间神经元属于窄凋谐带神经元,敏感的频率范围8—18千赫,反应最佳频率在12千赫附近,与同种雄硕螽叫声的主能峰相匹配。相位型听觉中间神经元属于宽调谐带神经元,有二个敏感频率范围,分别为5—8千赫和12—18千赫。它们对声强度的编码方式也不一样:分别以动作电位的数目与反应潜伏期对声强编码。本文还讨论了不同类型听觉中间神经元的功能意义。     

Heartbeat control in the medicinal leech: A model system for understanding the origin,coordination, and modulation of rhythmic motor patterns

We have analyzed in detail the neuronal network that generates heartbeat in the leech. Reciprocally inhibitory pairs of heart interneurons form oscillators that pace the heartbeat rhythm. Other heart interneurons coordinate these oscillators. These coordinating interneurons, along with the oscillator interneurons, form an eight-cell timing oscillator network for heartbeat. Still other interneurons, along with the oscillator interneurons, inhibit heart motor neurons, sculpting their activity into rhythmic bursts. Critical switch interneurons interface between the oscillator interneurons and the other premotor interneurons to produce two alternating coordination states of the motor neurons. The periods of the oscillator interneurons are modulated by endogenous RFamide neuropeptides. We have explored the ionic currents and graded and spike-mediated synaptic transmission that promote oscillation in the oscillator interneurons and have incorporated these data into a conductance-based computer model. This model has been of considerable predictive value and has led to new insights into how reciprocally inhibitory neurons produce oscillation. We are now in a strong position to expand this model upward, to encompass the entire heartbeat network, horizontally, to elucidate the mechanisms of FMRFamide modulation, and downward, to incorporate cellular morphology. By studying the mechanisms of motor pattern formation in the leech, using modeling studies in conjunction with parallel physiological experiments, we can contribute to a deeper understanding of how rhythmic motor acts are generated, coordinated, modulated, and reconfigured at the level of networks, cells, ionic currents, and synapses. © 1995 John Wiley & Sons, Inc.     

Upper airway muscle activity in normal women: influence of hormonal status

Obstructive sleep apnea is a disorder with astrong male predominance. One possible explanation could be an effectof female hormones on pharyngeal dilator muscle activity. Therefore, we determined the level of awake genioglossus electromyogram (EMGgg) andupper airway resistance in 12 pre- and 12 postmenopausal women underbasal conditions and during the application of an inspiratory resistiveload (25 cmH₂O · l¹ · s).In addition, a subgroup of eight postmenopausal women were studied asecond time after 2 wk of combined estrogen and progesterone replacement in standard doses. Peak phasic and tonic genioglossus activity, expressed as a percentage of maximum, were highest in theluteal phase of the menstrual cycle (phasic 23.9 ± 3.8%, tonic 10.2 ± 1.0%), followed by the follicular phase (phasic 15.5 ± 2.2%, tonic 7.3 ± 0.8%), and were lowest in the postmenopausal group (phasic 11.3 ± 1.6%, tonic of 5.0 ± 0.6), whereas upper airway resistance did not differ. There was a weak but significant positive correlation between progesterone levels and both peak phasic(P < 0.05) and tonic(P < 0.01) EMGgg. Finally, there was a significant increase in EMGgg in the postmenopausal group restudied after hormone therapy. In conclusion, female hormones (possibly progesterone) have a substantial impact on upper airway dilator muscleactivity.

     

Co-ordinating interneurones of the locust which convey two patterns of motor commands: their connexions with flight motoneurones.

1. Some flight motoneurones receive two superimposed rhythms of depolarizing synaptic potentials when the locust is not flying; a slow rhythm which is invariably linked to the expiratory phase of ventilation, and a fast rhythm with a period of about 50 ms which is similar to the wingbeat period in flight. 2. By recording simultaneously from groups of motoneurones, the synaptic potentials which underly these rhythms have been revealed in 30 flight motoneurones in the three thoracic ganglia. The inputs occur in elevator motoneurones and some depressors but are of lower amplitude in the latter. The inputs have not been found in leg motoneurones. 3. The rhythmic depolarizations are usually subthreshold but sum with sensory inputs to evoke spikes in flight motoneurones at intervals equal to or multiples of the wingbeat period in flight. 4. Both rhythms originate in the metathoracic ganglion and are mediated by the same interneurones. They can be adequately explained by supposing that there are two symmetrical interneurones which each make widespread connexions with left and right flight motoneurones in the three ganglia. 5. The slow rhythm is coded in the overall burst of interneurone spikes during expiration and the fast rhythm in the interval between the spikes of a burst.     

运动过程的网络逻辑——从离子通道到动物行为

为了揭示神经网络在脊椎动物运动中所行使的内在功能,作者开发了七鳃鳗这种低等脊椎动物模型。在这套系统中,不仅可以了解到运动模式生成网络以及激活此网络的命令系统,同时还可以在运动中研究方向控制系统和变向控制系统。七鳃鳗的神经系统有较少的神经元,而且运动行为中的不同运动模式可以由分离的神经系统所引发。模式生成神经网络包括同侧的谷氨酸能中间神经元和对侧的抑制性甘氨酸能中间神经元。网络中的突触连接、细胞膜特性和神经递质都也已经被鉴定。运动是由脑干区域的网状脊髓神经元所引起,而这些神经元又是被问脑和中脑分离的一些运动命令神经元群所控制。因此,运动行为最初是由这两个“运动核心”所启动。而这两个运动核心被基底神经节调控,基底神经节即时地做出判断是否允许下游的运动程序启动。在静止情况下基底神经节的输出核团维持对下游不同运动核心的抑制作用,反之则去除抑制活化运动核心。纹状体和苍白球被认为是这个运动抉择系统的主要部件。根据“霍奇金一贺胥黎”模型神经元开发了这套网络模型,不同的细胞具有各自相应的不同亚型的钠、钾、钙离子通道和钙依赖的钾通道。每个模型神经元拥有86个不同区域模块以及其对应的生物学功能,例如频率控制、超极化等等。然后根据已有实验证据,利用突触将不同的模型神经元相连。而系统中的10000个神经元大致和生物学网络上的细胞数量相当。突触数量为760000。突触类型有AMPA、NMDA、glycine型。有了这样大规模的模型,不仅可以模拟肌节与肌节之间的神经网络,还可以模拟到由基底神经节开始的行为起始部分。此外,这些网络模拟还被用于一个神经机械学模型来模拟包含有推进和方向控制部分的真实运动。     

Ganglionic distribution of inputs and outputs of C-PR, a neuron involved in the generation of a food-induced arousal state inAplysia

Cerebral neuron C-PR is thought to play an important role in the appetitive phase of feeding behavior ofAplysia. Here, we describe the organization of input and output pathways of C-PR. Intracellular dye fills of C-PR revealed extensive arborization of processes within the cerebral and the pedal ganglia. Numerous varicosities of varying sizes may provide points of synaptic inputs and outputs.Blocking polysynaptic transmission in the cerebral ganglion eliminated the sensory inputs to C-PR from stimuli applied to the rhinophores or tentacles, indicating that this input is probably mediated by cerebral interneurons. Identified cerebral mechanoafferent sensory neurons polysynaptically excite C-PR. Stimulation of the eyes and rhinophores with light depresses C-PR spike activity, and this effect also appears to be mediated by cerebral interneurons.C-PR has bilateral synaptic actions on numerous pedal ganglion neurons, and also has effects on cerebral neurons, including the MCC, Bn cells, CBIs and the contralateral C-PR. Although the somata of these cerebral neurons are physically close to C-PR, experiments using high divalent cation-containing solutions and cutting of various connectives indicated that the effects of C-PR on other cerebral ganglion neurons (specifically Bn cells and the MCC) are mediated by interneurons that project back to the cerebral ganglion via the pedal and pleural connectives. The indirect pathways of C-PR to other cerebral neurons may help to ensure that consummatory motor programs are not activated until the appropriate appetitive motor programs, mediated by the pedal ganglia, have begun to be expressed.     

Co-ordinating interneurones of the locust which convey two patterns of motor commands: their connexions with ventilatory motoneurones.

1. The interneurones which make widespread connexions with flight motoneurones also synapse upon ventilatory motoneurones so that in all 50 motoneurones receive synapses. They influence three aspects of ventilation; (a) the closing and opening movements of the thoracic spiracles, (b) some aspects of abdominal pumping movements and (c) the recruitment of some motoneurones controlling head pumping. 2. The two closer motoneurones of a particular thoracic spiracle receive the same excitatory synaptic inputs (EPSPs) during expiration. The EPSPs match those in appropriate flight motoneurones. 3. The closer motoneurones of each thoracic spiracle whose somata are in the pro-, meso- or metathoracic ganglia all receive the same excitatory synaptic inputs. These inputs are an adequate explanation of the pattern of spikes in the closer motoneurones. Both the slow ventilatory and fast rhythms of synaptic potentials are expressed as spikes; the slow as the overall expiratory burst of spikes and the fast as the groups of spikes within that burst. This establishes a ventilatory function for the interneurones. All thoracic closer motoneurones therefore receive the same excitatory commands which will tend to synchronize the movements of each spiracle. 4. Spiracular opener motoneurones are inhibited during expiration, their IPSPs matching the EPSPs in flight or closer motoneurones. Therefore the interneurones have reciprocal effects on the antagonistic motoneurones of the spiracles. 5. The interneurones synapse upon some motoneurones which control the pumping movements of the abdomen and which have their somata in the metathoracic or first unfused abdominal ganglion. Motoneurones in four separate ganglia therefore receive inputs from these interneurones. 6. The interneurones also synapse upon motoneurones which control an auxiliary form of ventilation, head pumping.     

Integrative Neurophysiology of the Lobster Cardiac Ganglion

The synchronized bursts of impulses produced by the nine neuronsof the isolated Homarus cardiac ganglion are usually initiatedby Cell 7. Activity in all other cells commences with very shortlatency thereafter. Impulses in most cells originate in triggerzones located 1–2 mm from the cell body, but the firstseveral impulses in Cells 8 and 9 frequently originate in distaltrigger zones some distance from the somata. Large cells fireat a high initial frequency, dropping rapidly to a low frequencyplateau. Small cells exhibit a more tonic behavior and fireat intermediate rates. More anterior small cells tend to firefaster than more posterior ones. The major synaptic interactionsare the impulse-mediated excitatory ones from small cells tolarge cells, and possibly to more anterior small cells. Thereare weak interactions from large cells back onto small cells,and very specific interactions from Cells 1 and 2 onto 3_A, 4_A,5_A, and 3_B 4_B 5_B respectively. The large discrete EPSPs generatedin large cells by small cell impulses appear to be the explanationfor "discrete positioning" in large-cell firing patterns. Inthis situation, large-cell impulses only fire at discrete timesduring the burst, regardless of the actual large-cell pattern. The overall view is of a two-layered neural system in whichthe small cells possess an endogenous oscillatory driver potential,synchronized by synaptic and electrotonic interactions, anddriving a train of impulses in each cell. This activates excitatorysynapses on the large cells, which combined with a triggereddriver potential in each large cell, produces synchronized trainsof motor impulses which activate the heart muscle, causing theheartbeat.     

Coexistence of lateral and co-tuned inhibitory configurations in cortical networks

The responses of neurons in sensory cortex depend on the summation of excitatory and inhibitory synaptic inputs. How the excitatory and inhibitory inputs scale with stimulus depends on the network architecture, which ranges from the lateral inhibitory configuration where excitatory inputs are more narrowly tuned than inhibitory inputs, to the co-tuned configuration where both are tuned equally. The underlying circuitry that gives rise to lateral inhibition and co-tuning is yet unclear. Using large-scale network simulations with experimentally determined connectivity patterns and simulations with rate models, we show that the spatial extent of the input determined the configuration: there was a smooth transition from lateral inhibition with narrow input to co-tuning with broad input. The transition from lateral inhibition to co-tuning was accompanied by shifts in overall gain (reduced), output firing pattern (from tonic to phasic) and rate-level functions (from non-monotonic to monotonically increasing). The results suggest that a single cortical network architecture could account for the extended range of experimentally observed response types between the extremes of lateral inhibitory versus co-tuned configurations.     

Cellular Mechanisms Underlying Swim Acceleration in the Pteropod Mollusk Clione limacina

The pteropod mollusk Clione limacina swims by dorsal-ventralflapping movements of its wing-like parapodia. Two basic swimspeeds are observed—slow and fast. Serotonin enhancesswimming speed by increasing the frequency of wing movements.It does this by modulating intrinsic properties of swim interneuronscomprising the swim central pattern generator (CPG). Here weexamine some of the ionic currents that mediate changes in theintrinsic properties of swim interneurons to increase swimmingspeed in Clione. Serotonin influences three intrinsic propertiesof swim interneurons during the transition from slow to fastswimming: baseline depolarization, postinhibitory rebound (PIR),and spike narrowing. Current clamp experiments suggest thatneither I_h nor I_A exclusively accounts for the serotonin-inducedbaseline depolarization. However, I_h and I_A both have a stronginfluence on the timing of PIR—blocking I_h increases thelatency to PIR while blocking I_A decreases the latency to PIR.Finally, apamin a blocker of I_K(Ca) reverses serotonin-inducedspike narrowing. These results suggest that serotonin may simultaneouslyenhance I_h and I_K(Ca) and suppress I_A to contribute to increasesin locomotor speed.     

The contribution of the pleural type 12 interneuron to swim acceleration in Clione limacina

The pteropod mollusc, Clione limacina, swims by alternate dorsal–ventral flapping movements of its wing-like parapodia. The basic swim rhythm is produced by a network of pedal swim interneurons that comprise a swim central pattern generator (CPG). Serotonergic modulation of both intrinsic cellular properties of the swim interneurons and network properties contribute to swim acceleration, the latter including recruitment of type 12 interneurons into the CPG. Here we address the role of the type 12 interneurons in swim acceleration. A single type 12 interneuron is found in each of the pleural ganglia, which contributes to fast swimming by exciting the dorsal swim interneurons while simultaneously inhibiting the ventral swim interneurons. Each type 12 interneuron sends a single process through the pleural–pedal connective that branches in both ipsilateral and contralateral pedal ganglia. This anatomical arrangement allowed us to manipulate the influence of the type 12 interneurons on the swim circuitry by cutting the pleural–pedal connective followed by a “culture” period of 48 h. The mean swim frequency of cut preparations was reduced by 19% when compared to the swim frequency of uncut preparations when stimulated with 10⁻⁶ M serotonin; however, this decrease was not statistically significant. Additional evidence suggests that the type 12 interneurons may produce a short-term, immediate effect on swim acceleration while slower, modulatory inputs are taking shape.     

Various transmitters as filters in transferring information in an identified neural network of gastropods

1. The motor and interneurones in the identified neural network regulating the cardio-renal system of Helix pomatia L. produced differing effects with ACh, 5HT, DA and OP. They were less sensitive to glutamate. 2. On the double action interneuron the sensitivity to the neurotransmitters was modified by the source of input and the sequence of their application. 3. The interneuron V21 uses ACh and 5HT as neurotransmitters. Seven afferent excitatory pathways terminate on interneuron V21. 4. The efferent pathways of interneuron V21 can be selectively blocked by haloperidol and ergotamine. 5. The peptide-like factor released spontaneously following haloperidol and ergotamine treatment block the cholinergic and aminergic pathways onto interneuron V21, while the amino acid mechanism remains unaltered.     

Why the ovotestis of Helix aspersa is innervated

Although Schmalz described the innervation of the ovotestis in pulmonate snails as early as 1914, no functions have been attributed to it. In H. aspersa, the intestinal nerve branches profusely within the ovotestis and terminates in the walls of the acini and in the sheath surrounding the early portion of the hermaphroditic duct. We found both sensory and motor functions for this innervation. Significantly, there is a tonic sensory discharge generated by the mechanical pressure of growing oocytes, and the level of tonic afferent activity is strongly correlated with the number of ripe oocytes; this is probably a permissive signal that gates ovulation. Tactile stimulation of the ovotestis causes a phasic sensory discharge and a pronounced cardio activation. Also, an efferent discharge is elicited in the ovotestis branch of the intestinal nerve. To study the motor consequences of efferent activity, the ovotestis branch was electrically stimulated. We found that such stimulation evokes peristaltic contractions of the initial portion of the hermaphroditic duct and increases beat frequencies of the cilia that line the interior of the duct. These effects could facilitate the transport of oocytes down the duct. Still other functions of afferent activity are implied by changes in the spontaneous activity of mesocerebral cells following nerve stimulation. Putative sensory neurons and putative motoneurons have been identified in the visceral and right parietal ganglia.     

Functional and Structural Parallels in Crustacean and Drosophila Neuromuscular Systems

Comparison of morphological and physiological phenotypes ofrepresentative crustacean motor neurons, and selected motorneurons of Drosophila larval abdominal muscles, shows severalfeatures in common. Crustacean motor nerve terminals, and thoseof Drosophila, possess numerous small synapses with well-definedactive zones. In crustaceans, neurons that are more tonicallyactive have markedly varicose terminals; synapses and mitochondriaare selectively localized in the varicosities. Phasic motoraxons have filiform terminals, sometimes with small varicosities;mitochondrial content is less than for tonic axons, and synapsesare distributed along the terminals. Tonic axons generate smallexcitatory potentials which facilitate strongly at higher frequencies,and which are resistant to depression. Thephasic neurons generatelarge excitatory potentials which exhibit relatively littlefrequency facilitation, and depress rapidly. In Drosophila,counterparts of crustacean phasic and tonic motor neurons havebeen found, but the differentiation is less pronounced. It isinferredthat cellular factors regulating the number of participatingsynapses and the probability of quantal release are similarin crustaceans and Drosophila, and that advantage can be takenof this in future to develop experiments addressing the regulationof synaptic plasticity.     

Dual pathways for tactile sensory information to thoracic interneurons in the cockroach

The escape system of the American cockroach is both fast and directional. In response to wind stimulation both of these characteristics are largely due to the properties of the ventral giant interneurons (vGIs), which conduct sensory information from the cerci on the rear of the animal to type A thoracic interneurons (TI_As) in the thoracic ganglia. The cockroach also escapes from tactile stimuli, and although vGIs are not involved in tactile-mediated escapes, the same thoracic interneurons process tactile sensory information. The response of TI_As to tactile information is typically biphasic. A rapid initial depolarization is followed by a longer latency depolarization that encodes most if not all of the directional information in the tactile stimulus. We report here that the biphasic response of TI_As to tactile stimulation is caused by two separate conducting pathways from the point of stimulation to the thoracic ganglia. Phase 1 is generated by mechanical conduction along the animal's body cuticle or other physical structures. It cannot be eliminated by complete lesion of the nerve cord, and it is not evoked in response to electrical stimulation of abdominal nerves that contain the axons of sensory receptors in abdominal segments. However, it can be eliminated by lesioning the abdominal nerve cord and nerve 7 of the metathoracic ganglion together, suggesting that the relevant sensory structures send axons in nerve 7 and abdominal nerves of anterior abdominal ganglia. Phase 2 of the TI_As tactile response is generated by a typical neural pathway that includes mechanoreceptors in each abdominal segment, which project to interneurons with axons in either abdominal connective. Those interneurons with inputs from receptors that are ipsilateral to their axon have a greater influence on TI_As than those that receive inputs from the contralateral side. The phase 1 response has an important role in reducing initiation time for the escape response. Animals in which the phase 2 pathway has been eliminated by lesion of the abdominal nerve cord are still capable of generating a partial startle response with a typically short latency even when stimulated posterior to the lesion. © 1995 John Wiley & Sons, Inc.