Integration of visual stimuli by the crayfish central nervous system

For the crayfish, properties of visually reacting interneurone types in the optic nerve are described on the basis of single-unit analysis. Sustaining fibres show: (a) ‘surround’ inhibition over the whole retina, including the excitatory field; (b) ‘on’ and ‘off’ effects at field boundaries; (c) a dark discharge in deteriorated preparations; (d) an increase in adapted firing rate and response to flashes in an ‘excited state’ of the preparation; (e) increased overall impulse frequency to fast-moving shadows at frequencies of 2-10/sec. which elicit short bursts. Dimming fibres, having mainly reversed properties, show: (a) bursts followed by adaptation to a lower ferquency level of spikes on light dimming; (b) total inhibition by illumination for times proportional to light intensity; (c) responsiveness to quickly moving shadows. 'Jittery’ movement fibres lack directional sensitivity. For them: (a) total illumination and contrast have, within a wide range, no effect on discharges to moving targets; (b) large, dark cards moved at constant speed are seen only near their visual field boundary; (c) the less predictable the movement of a small dark object is, the longer its excitatory effect lasts; (d) responsiveness of field parts exposed is decreased for long durations; (e) no firing is caused by active or passive eye motions. Space-constant fibres show changed location and size of their excitatory fields with eye position. Their potential visual fields below the horizontal plane are unresponsive, due to inhibition caused mainly by statocyst input. Two eye muscle motor fibres, also under statocyst control, are influenced by light on the limited sensory fields of two pairs of identifiable sustaining fibres. The two motor fibres, which innervate antagonistic muscles, are reversely excited and inhibited by these fields

The effect of the spacing of background elements upon optomotor memory responses in the crab: the influence of adding or deleting features during darkness

1. Study was made of the effect of separation between stripes in the visual field upon responses which indicate memory of those stripes. 2. The amount of separation between white stripes had very little effect, whereas response strength and the amount of separation between black stripes were directly proportional. 3. The presence of extra, non-displaced black stripes prior to or following displacement reduced the size of the memory responses. 4. The effects of the amount of separation in the two cases were comparable. In both situations the separation affected only the responses to displacement of the stripe borders nearest to the extra stripe. 5. The effect of extra stripes present prior to displacement was in turn affected by the duration of the dark period, whilst the effect of those present during the post-displacement period was not. This accounts for the larger effect of extra stripes present during the psot-displacement period. 6. By expanding stripe width during darkness it was possible to distinguish between the effects of distance between stripes and the amount of white space separating them. Reducing white space while distance remains constant causes reductions in response strength, whereas reducing the distance between a memory zone and the white space between it and the neighboring stripe increased the size of the memory response

Unidirectional Rotation Neurones in the Optomotor System of the Crab, Carcinus

1. Among the optomotor fibres to the eye muscles in Carcinus a class was found which responds to unidirectional fast rotations around various body axes. All had large signals and are therefore of large diameter. 2. In one set of these fibres which fires especially for rotations around the dorsoventral axis, it could be shown that discharges take place especially during accelerations and that, when a rotation in the null direction is suddenly stopped, a short discharge occurs. The fibres for other axes behave in a similar manner. 3. For rotations around the ventro-dorsal axis, but not for other directions, mediumsized fibres are present which, in contrast to the fast fibres, respond to visual stimulation, as well as to body rotations in darkness, thus combining the input properties of the unidirectional fast rotatory and the unidirectional purely optokinetic small fibres. Their sensitivity to visual input is for low rotation velocities, to body rotations is for high rotation velocities

Factors Regulating the Discharge Frequency in Optomotor Fibres Of Carcinus Maenas

The influence of the excited state of the animal on various motor neurone discharges and accompanying muscle action potentials was studied in the eyestalk of the crab, Carcinus maenas. In most cases large increases in firing frequency could be obtained during such states. An exception is the tonic eye-withdrawal system in which an inhibitory effect is caused. A pronounced difference in habituation to constant stimuli between spring and summer was found for the position fibres; in spring it was slow and in summer much quicker

Unidirectional Movement Fibres from A Proprioceptive Organ of the Crab, Carcinus Maenas

1. The proprioceptive organ of the shore crab Carcinus maenas, which signals all proprioceptive sensation from the joint between propodite and dactylopodite, has been shown to contain sense cells receptive to position as well as to movement. 2. Certain cells of the organ increase their discharge frequency in response to greater degrees of flexion, while other endings respond to greater degrees of extension. Movement has little effect on the frequency of the position fibres. 3. Cells responding to movement in one direction only are present, certain ones signalling fiexion and others extension. The most sensitive undirectional movement fibres are almost completely independent of position and velocity. They provide the crab with an extremely sensitive movement sense not as yet reported in other proprioceptive mechanisms. 4. Both position-sensitive and movement-sensitive cells show a wide range of thresholds. 5. Cells which cannot be strictly classified as either position or movement sensitive have also been found. 6. By tracing the origin of the signals to the cell bodies, it appears that movement fibres have on the whole larger cell bodies than position fibres and are more proximally located. 7. Flexor and extensor movement cells appear to lie on different sides of elastic strand. 8. The question of how unidirectional sensitivity may be achieved is discussed

A Comparative Study Of The Double Motor Innervation In Marine Crustaceans

A double motor innervation has been shown for several muscles of marine crustaceans. The adductors of the claws of Randallia and Blepharipoda and the adductor of the dactylopodite of the walking leg of Cancer were studied physiologically. The two motor axons which innervate these muscles have a different diameter (ratio 1.4:1). Stimulation of the thick fibre causes a response, which, though it is not always faster than the response of the thin fibre, must be considered as a "fast" contraction. In Randallia and in Blepharipoda the slow contraction is higher than the fast with frequencies of less than ± 50 per sec., in Cancer with frequencies less than 100 per sec. The action currents of the two kinds of contraction are different. Both show facilitation, but under the same conditions of stimulation the fast-action currents are higher. The first stimulus of the thick fibre causes an action current top which is clearly distinguishable, the action currents of the slow contraction show up only after a number of stimuli. Even when the mechanical reaction on stimulation of the thick fibre is smaller than on similar stimulation of the thin fibre, the action currents are higher in the first case. A single impulse in the thick fibre does not cause a contraction, but sets up a muscle-action current. The chronaxie of this action current in Blepharipoda and Randallia is 0.8{sigma} and is about the same as that found for the action current of the nerve. Two impulses in the thick fibre may cause a mechanical response, as is shown by summation experiments. The pseudo-chronaxie of this contraction was measured as 3.5 {sigma}. The second action current shows facilitation, when it follows the first within 1 sec.; a mechanical reaction results with summation intervals of two stimuli of less than 10{sigma}. The facilitation of the action current increases with decrease of the time interval between the two impulses; with the shortest intervals that give summation the resulting action current is a smooth high spike

The Co-ordination of Swimmeret Movements in the Crayfish, Procambarus Clarkii (Girard)

1. Electrical activity recorded in the first roots of the abdominal nerve cord show bursts of activity which accompany rhythmic movements of the swimmerets. These bursts persist when a root is cut distal to the electrodes. 2. Proprioceptive impulses have been recorded from the distal portion of these nerves during passive movements of the swimmerets. They have also been recorded in the connectives during both passive and active movements. 3. Cutting all second and third roots of the abdominal ganglia does not interfere with the rhythmic swimmeret movements of the isolated abdomen. Rhythmic efferent bursts persist in first roots even when only a single first root remains intact in such a preparation. 4. Intermittent bursts are also found in the first roots of a completely isolated cord but their pattern, frequency and phase relationships differ from those recorded in the same roots before isolation was completed. Such bursts are very rare in second and third roots. 5. Stimulation of small bundles of fibres in the circumoesophageal commissure produces well-defined inhibitory and excitatory effects on the discharge of single units in the first roots. 6. One specific bundle found in many preparations, when stimulated at 50/sec., caused a tonic retraction of all the swimmerets which then began to beat rhythmically and continued to do so when stimulation was discontinued. 7. Intermittent bursts in the first roots took place with correct phasing during this stimulation even when the abdominal ganglia were isolated except for their connexion with the last thoracic ganglion. Again, such bursts were absent in the second and third roots. 8. It is concluded that both the inflow from peripheral proprioceptors and intrinsic properties of the central ganglia play essential parts in the co-ordination of the metachronal movements of the swimmerets

Chemical changes in the adductor muscle of the cheliped of the crayfish in relation to the double motor innervation

Work from this laboratory has shown that a number of the striated muscles of the legs of the crayfish show a typical motor innervation (1). Such muscles are innervated by two motor fibers only, each causing a different type of contraction. One of these, resulting from stimulation of the thicker fiber, is termed the "fast" contraction, while the other is called the "slow." Both anatomical evidence and physiological experiments indicate that each muscle fiber is innervated by these two axons and that both contractions occur in the same muscle fibers (2, 3). The most striking difference between the two contractions is found in the adductor muscle of the cheliped of the crayfish. In this case, a single impulse in the thicker motor fiber causes a twitch-like contraction, and the system behaves in almost all respects as does a single motor unit of a vertebrate muscle: single shocks of different strengths give twitches of a considerable strength and a constant height, and during tetanic contraction the action currents are all of the same magnitude. In contrast to this, single shocks given to the thinner fiber produce no visible response in the muscle, but faradic stimulation causes action currents which grow in height and a contraction with a long latent period. The action currents in this case are always much smaller than those of the fast contraction. The chemical changes occurring in the adductor muscle of the cheliped of the crayfish Cambarus clarkii during these two types of contraction were investigated in order to obtain evidence on two possible mechanisms by which the two contractions might occur in the same muscle fiber. In the first place, two contractile substances might be present, a "phasic" and a "tonic" one, analogous to those supposed by Botazzi (4) in the vertebrate striated muscle, or the same substance might contract in both cases, the difference between the contractions being due solely to differences in the transmission mechanism between the nerve impulse and the contractile substance. Methods were devised for the rapid removal of the stimulated cheliped to liquid air and for the analysis of the frozen muscle tissue. The changes in phosphate and in lactic acid content were used as indices of the chemical changes occurring

Command Interneurons in the Crayfish Central Nervous System

The motor effects evoked by stimulation of each of eight command fibres in the circumoesophageal commissures of the crayfish are described. The fibres obtained appeared to have widespread connexions in all or most of the lower ganglia. For certain fibres the response was stronger on the homolateral side of the animal; for others it was symmetrical. The frequency of stimulation of a command fibre generally had a pronounced influence on the speed of the evoked response. In addition, segments of the total response could be elicited selectively by alteration of the frequency and duration of stimulation. Although the responses associated with most of the fibres were not sensitive to the fine temporal pattern of the applied stimulation, for one fibre the motor output depended clearly on the spacing of the stimulating pulses

Neuronal Pathways and Synaptic Connexions in the Abdominal Cord of the Crayfish

1. An investigation has been made into the function and distribution of nerve fibres in the abdominal ganglion chain and its roots in the crayfish, Procambarus clarkii, by leading off action potentials from small prepared bundles following sensory stimulation. 2 .The sensory fields belonging to the first and second roots of each abdominal ganglion were determined, and the antero-posterior pathway of sensory fibres within the cord noted. It was found that the primary sensory fibres of the dorsal muscle receptor organs, entering through the second root, send out an anterior branch to the brain and a posterior one to the last ganglion. For most other sensory fibres much shorter intracentral branches are indicated, though some of them extend for two ganglia in the anterior direction and for one posteriorly. All sensory fibres in the connectives run on the same side as they enter. 3. The segmental divisions of the external skeleton and of the nervous system do not coincide, the neural segment slants in a posterior dorsal direction with respect to the skeletal one. 4. For the majority of the interneurones which innervate more than two abdominal segments it has been proved that they synapse with primary sensory fibres in each of the ganglia that these enter. Depending on the segment stimulated with respect to the leading-off position, both ascending and descending impulses are obtained in such interneurones and collision of the impulses has been observed. Some consequences of this type of integration are discussed. 5. For interneurones responding to bilateral or heterolateral stimulation the course of the impulses proved to be of at least two types. In some, cutting the fibre prevents the arrival of impulses except those set up on the side of the cut from which the recording is made. In others, recording from either side of the cut fibre does not exclude any of the sensory fields to which the fibre normally responded. 6. At least one interneurone is present in which all primary sensory fibres from the different segments to whose activity it responds collect in one ganglion