The elasmobranch spiracular organ

Summary The spiracular organ is a lateral line derived receptor associated with the first gill cleft (spiracle). Its functional morphology was studied in the little skate,Raja erinacea, and a shark, the smooth dogfish,Mustelus canis, with light and electron microscopy. The spiracular organ is a tube (skate) or pouch (shark) with a single pore opening into the spiracle. The lumen is lined with patches of sensory hair cells, and filled with a gelatinous cupula. In the little skate, hair cells form synapses with afferents but apparently not with efferent fibers. In both species, the spiracular organs are deformed by flexion of the hyomandibular cartilage at its articulation with the cranium. The hyomandibula is a suspensory element of the jaws; hyomandibular flexion results in jaw protrusion. The little skate spiracular organ is anchored at one end to the cranium and at the other to the hyomandibula so that it is stretched or relaxed during hyomandibular extension and flexion, respectively. InMustelus, the effects of hyomandibular flexion on the spiracular organ are mediated indirectly by the superior post-spiracular ligament which inserts on the distal end of the hyomandibula. Deformation of the dogfish shark cupula during hyomandibular movement was observed. In the little skate, as revealed by transmission electron microscopy, there is a measurable deflection of the hair cell ciliary bundles from spiracular organs fixed with the hyomandibula in the flexed relative to the extended positions. In both species, hyomandibula flexion should result in hair cell depolarization, and sensory afferent excitation, based on the direction of the observed (skate) or expected (shark) deflection of hair cell cilia.     

Interaction of apical and basal membrane ion channels underlies electroreception in ampullary epithelia of skates.

The exquisite sensitivity of elasmobranch fishes to electric fields is thought to reside in electroreceptive organs called ampullae of Lorenzini. We measured the stimulus-response behavior of ampullary organs excised from skates. Under open-circuit conditions, the ampullary organ showed three distinct response states: spontaneous repetitive spikes, evoked spikes, and small, damped oscillatory responses. Under short-circuit conditions, the amplitude range for a linear current response to a sinusoidal (0.5 Hz) voltage clamp of an organ (assessed by spectral analysis of the harmonics generated) was 7-200 microV rms. Changes in the spike firing rate of the afferent nerve innervating the organ were evident for voltage clamps of the ampullary epithelium of 3 microV and the spike rate saturated for clamp steps exceeding 100 microV. Thus, the linear response range of the ampullary epithelium exceeded the range in spike firing rate of the afferent nerve. The steady-state transorgan electrical properties under voltage clamp conditions were obtained by analysis of complex admittance determinations in the frequency range 0.05-20 Hz for perturbations (< 100 microV rms) in the linear range. Admittance functions were distinctly related to the preparation states observed under open-circuit conditions. A negative real part in the organ admittance (i.e., a steady-state negative conductance generated by the preparation) was a common characteristic of the two (open-circuit) excitable states. The negative conductance was also confirmed by the direction of current flow through the ampullary epithelium in response to step voltage clamps.(ABSTRACT TRUNCATED AT 250 WORDS)     

Single unit activity of dopaminergic neurons in freely moving cats

Single unit activity was recorded from the area of the substantia nigra in freely moving cats. A sub-population of these neurons had the following characteristics: long action potential durations (2–4 msec); relatively slow discharge rates (2–6 spikes/sec); firing as single spikes along with periods of bursting activity in which spike amplitude successively decreased; suppression of unit activity by systemic injection of apomorphine and increased activity after systemic injection of haloperidol. These characteristics are similar to those of identified dopamine neurons recorded in chloral hydrate anesthetized or peripherally paralyzed rats. Therefore, based upon these physiological and pharmacological similarities, this study represents the first systematic report providing evidence for recording the activity of dopaminergic neurons in freely moving cats. In addition, when these cells were studied across the sleep-waking cycle they displayed little variation in firing rates between waking, slow wave sleep and REM sleep.     

Morphological variation in the electric organ of the little skate (Leucoraja erinacea) and its possible role in communication during courtship

Skates discharge an electrical current too weak to be used for predation or defense, and too infrequent and irregular to be used for electrolocation. Additionally, skates possess a specialized sensory system that can detect electrical stimuli at the same strength at which they discharge their organs. These two factors are suggestive of a communicative role for the electric organ in skates, a role that has been demonstrated in similarly weakly electric teleosts (e.g., mormyrids and gymnotiforms). There is evidence that the sexual and ontogenetic variations in the electric organ discharge (EOD) in these other weakly electric fishes are linked to morphological variations in electric organs and the electrogenerating cells of the organs, the electrocytes. Little work has been done to examine possible sexual and ontogenetic variations in skate EODs or variations in the electrocytes responsible for those discharges. Electric organs and electrocyte morphology of male and female, and mature and immature little skates, Leucoraja erinacea, are characterized here. Female electric organs were bigger than male electric organs. This is suggestive of a sexually dimorphic EOD waveform or amplitude, which might be used as a sex-specific identification signal during courtship. The shapes of electrocytes that make up the organ were found to be significantly different between mature and immature individuals and, in some cases, posterior membrane surface area of the electrocytes increased at the onset of maturity due to the formation of membrane surface invaginations and papillae. This is evidence that the EOD of skates may differ in its waveform or amplitude or frequency between mature and immature skates, and act as a signal for readiness to mate. This study supports a communicative role during courtship for the weak electric organs of little skates, but studies that characterize skate EOD dimorphisms are needed to corroborate this speculation before conclusions can be drawn about the role the electric organ plays in communication during courtship.     

Motor unit firing rates during isometric voluntary contractions performed at different muscle lengths

Firing rates of motor units and surface EMG were measured from the triceps brachii muscles of able-bodied subjects during brief submaximal and maximal isometric voluntary contractions made at 5 elbow joint angles that covered the entire physiological range of muscle lengths. Muscle activation at the longest, midlength, and shortest muscle lengths, measured by twitch occlusion, averaged 98%, 97%, and 93% respectively, with each subject able to achieve complete activation during some contractions. As expected, the strongest contractions were recorded at 90 degrees of elbow flexion. Mean motor unit firing rates and surface EMG increased with contraction intensity at each muscle length. For any given absolute contraction intensity, motor unit firing rates varied when muscle length was changed. However, mean motor unit firing rates were independent of muscle length when contractions were compared with the intensity of the maximal voluntary contraction (MVC) achieved at each joint angle.     

Firing properties of substantia nigra dopaminergic neurons in freely moving rats

Single unit recordings were obtained from putative dopaminergic neurons in the substantia nigra of awake, freely moving rats. The cells exhibited waveforms, range of firing rates and types of firing patterns identical to those of identified DA neurons of anesthetized or paralyzed rats. Two firing patterns were observed: single spike activity and a bursting mode with spikes of progressively diminished amplitude and increased duration within each burst. The degree of burst firing varied considerably among the cells and individual cells sometimes switched from one pattern of firing (e.g. predominantly single spike) to another (e.g. bursting), although the determinants of these transitions are, at this time, unclear. Putative DA neurons were inhibited by i.v. apomorphine and excited by i.v. haloperidol. Haloperidol also reversed the apomorphine-induced inhibition of firing. Inhibitions and excitations were associated with a reduction and elevation, respectively, in burst firing. The effects of the two drugs were identical to their effects in immobilized rats. In several cases, a putative DA neuron was observed to fire all of its spikes in near coincidence with at least one other cell with identical electrophysiological characteristics. This form of interaction (i.e. presumed electrical coupling) between DA cells is only rarely observed in anesthetized or paralyzed rats and may play a significant role in the normal functioning of the nigrostriatal DA system.     

The trigeminofacial innervation of the cephalic lateral line organs and photophores of the lantern fish Tarletonbeania crenularis (Myctophidae)

The trigeminofacial innervation of the cephalic photophores and lateral line organs of Tarletonbeania crenularis has been studied from gross dissections. The facial and trigeminal roots leave the brainstem separately, but later intermingle forming a trigemino‐facial complex. The seventh nerve gives rise to the hyomandibular trunk and sends a branch rostrad to join the trigeminal forming the supra‐ and infraorbital trunks. The supraorbital trunk innervates the Dn photophore, the snout, the iris, the supraorbital lateral line organs and part of the olfactory sacs. The infraorbital trunk supplies the infraorbital lateral line organs, the Vn photophore and the tissues surrounding the premaxillaries. The hyomandibular trunk passes to the opercular photophores and lateral line organs, and together with a branch from the infraorbital trunk supplies the branchiostegal photophores and lateral line organs of the mandible.     

The tarso-pretarsal chordotonal organ as an element in cockroach walking

Many types of sense organs have been demonstrated to show repetitive discharges during walking that could provide informational cues about leg movements and other parameters of locomotion. We have recorded activities of receptors of the distal (tarsal) segments of the cockroach hindleg in restrained and freely moving animals while they were videotaped. These recordings show peaks of activities at the onset and termination of the stance phase. We have morphologically and physiologically identified a joint angle receptor, the tarso-pretarsal chordotonal organ, that contributes to the discharges seen late in stance, prior to the onset of leg flexion in swing. This sense organ encodes the angle and rate of change of the most distal leg joint and specifically discharges when the claws are disengaged from the substrate. Applied displacements of the claws in restrained preparations elicit reflex activation of the tibial flexor muscle and a crossed extensor reflex in the opposite hindleg. These reflexes could function to insure that leg flexion in swing does not occur until the claws are disengaged and to enhance support by the opposite hindleg. Thus, the regular discharges of the chordotonal organ could assure efficient and coordinated muscle contractions and movements during normal, unperturbed walking. Accepted: 2 January 1997     

Differences between motor unit firing rate, twitch characteristics and fibre type composition in an agonistic muscle group in man

A comparison was carried out between the motor unit (MU) firing rate and the characteristics of the twitch and the fibre type composition of anconeus and triceps brachii. Fibre type composition (type I, type II) was determined in whole cross-sections of cadaver specimens. The proportion of type I fibre was 60%-67% in anconeus and 32-40% in the lateral head of triceps brachii. Reflecting these histochemical differences, the contraction time of anconeus and triceps was 92 +/- 9 ms and 68 +/- 9 ms respectively. It follows that anconeus can be classified as a slow muscle, as opposed to the lateral head of triceps. The relationship between MU firing rate and isometric force, tested at 90 degrees elbow flexion, differed between the two muscles for force values below 30% of maximal voluntary contraction. No significant increase in MU firing rate was found in anconeus at forces above 5% of maximal voluntary contraction. It is concluded that even within a single agonistic muscle group acting at a single joint there is an adaptation of MU firing rate to the contractile characteristics of each muscle.     

Motor unit activation patterns during concentric wrist flexion in humans with different muscle fibre composition

Muscle activity was recorded from the flexor carpi radialis muscle during static and dynamic-concentric wrist flexion in six subjects, who had exhibited large differences in histochemically identified muscle fibre composition. Motor unit recruitment patterns were identified by sampling 310 motor units and counting firing rates in pulses per second (pps). During concentric wrist flexion at 30% of maximal exercise intensity the mean firing rate was 27 (SD 13) pps. This was around twice the value of 12 (SD 5) pps recorded during sustained static contraction at 30% of maximal voluntary contraction, despite a larger absolute force level during the static contraction. A similar pattern of higher firing rates during dynamic exercise was seen when concentric wrist flexion at 60% of maximal exercise intensity [30 (SD 14) pps] was compared with sustained static contraction at 60% of maximal voluntary contraction [19 (SD 8) pps]. The increase in dynamic exercise intensity was accomplished by recruitment of additional motor units rather than by increasing the firing rate as during static contractions. No difference in mean firing rates was found among subjects with different muscle fibre composition, who had previously exhibited marked differences in metabolic response during corresponding dynamic contractions. It was concluded that during submaximal dynamic contractions motor unit firing rate cannot be deduced from observations during static contractions and that muscle fibre composition may play a minor role. Accepted: 5 May 1998     

Inter- and intraspecific variation in the distribution and number of pit organs (free neuromasts) of sharks and rays

The distribution of pit organs (free neuromasts) has previously been documented for several species of pelagic sharks, but is relatively poorly known for rays and bottom-dwelling (demersal) sharks. In the present study, the complete distribution of pit organs was mapped in the demersal sharks Heterodontus portusjacksoni, Orectolobus maculatus, Hemiscyllium ocellatum, Chiloscyllium punctatum, and Asymbolus analis, and the rays Rhinobatos typus, Aptychotrema rostrata, Trygonorrhina sp. A, Raja sp. A, and Myliobatis australis. All of these species had pit organs scattered over the dorsolateral surface. The sharks also had "mandibular" pit organs (and "umbilical" pit organs in C. punctatum and A. analis) on the ventral surface, while pit organs were sparse or absent on the ventral surface of rays. All of the species examined here, except for M. australis, also had a "spiracular" group of pit organs adjacent to the eye and/or spiracle. Spiracular pit organs were also recorded for the sawshark Pristiophorus sp. A and the skate Pavoraja nitida, although the remainder of pit organs were not mapped in these species. The distribution and number of pit organs varied both within and among species. Pit organ distribution was asymmetrical in each individual examined, but no particular trend towards left or right "handedness" was observed in any species. Although rays have been thought to have fewer pit organs than sharks in general, this was not the case in the present study. All of the species examined here had few pit organs compared to the pelagic sharks previously documented, but it is not clear whether this is due to ecological or phylogenetic causes.     

Responses of spiking local interneurones in the locust to proprioceptive signals from the femoral chordotonal organ

Summary The responses of spiking local interneurones of a ventral midline population in the metathoracic ganglion of the locust,Schistocerca gregaria, to controlled movements of a proprioceptor, the femoral chordotonal organ (FCO) in a hindleg, were revealed by intracellular recording. Afferents from the FCO which signal specific features of the movement or angle of the femoro-tibial joint, can make direct excitatory synapses with particular interneurones in this population (Burrows 1987a).Some interneurones in this population are excited only by flexion, some only by extension, but others by both flexion and extension movements of the femoro-tibial joint. Interneurones excited by one direction of movement may be either unaffected, or inhibited by the opposite movement. The balance between excitation and inhibition is determined by the range over which the movement occurs, and can increase the accuracy of a representation of a movement.The response of some interneurones has tonic components, so that the angle of the joint over a certain range is represented in the frequency of their spikes. Different interneurones respond within different ranges of femoro-tibial angles so that information about the position of the joint is fractionated amongst several members of the population. These interneurones respond to repetitive movements, similar to those used by the locust during walking, with bursts of spikes whose number and frequency are determined by the repetition rate and amplitude of the movement. A brief movement of the FCO may induce effects which persist for many seconds and outlast the changed pattern of afferent spikes. The sign of such an effect depends upon the preceding history of stimulation.Other interneurones respond only to movement so that their response is more phasic. The velocities to which they respond fall within the range of those generated by twitches of the flexor and extensor tibiae muscles and the movements of the tibia during locomotion. Some interneurones respond only to a specific range of velocities because they are inhibited by all other movements. Some interneurones respond to repetitive movements with reliable bursts of spikes, whilst in others the frequency of spikes may be raised but may contain no cyclical information. All, however, produce the largest number of spikes during the first cycle of a repetitive movement.Inputs from the FCO may sum either with excitation generated by direct inputs from exteroceptors or with inhibition produced by other local interneurones as a result of afferent signals.These spiking local interneurones are essential elements in the integration of local reflexes initiated by signals from the FCO. For example, one ensures that the levator tarsi motor neurone is reflexly inhibited when the FCO signals an extension movement. Exteroceptive inputs from the ventral tarsus suppress the spikes in this interneurone and would prevent expression of the reflex when the tarsus is in contact with ground.Abbreviation FCO femoral chordotonal organ     

Comparison of acid mucin goblet cell distribution and Hox13 expression patterns in the developing vertebrate digestive tract

The digestive tract of vertebrates is a complex organ system required for the digestion of food and the absorption of nutrients. The colon evolved as a water absorption organ essential for vertebrates to survive on land. In contrast to land vertebrates, the Chondrichthyes (sharks, skates and rays) are nearly iso-osmotic with their ocean environment and do not reabsorb water from food waste. To understand the origin of the vertebrate colon, we examined the distribution of sulfated and sialyated mucus-producing cells in the little skate, Raja erinacea, as an indication of water absorption function in the chondrichthian digestive tract. The percentage of acid mucin producing goblet cells was analyzed in the spiral valve and hindgut of little skate and the small intestine and colon of mouse embryos. Levels of acid mucins in the hindgut of the little skate was comparable to that of the small intestines of terrestrial vertebrates, whereas the distal region of the spiral valve contained high levels of acid mucin producing cells similar to the colon of mouse and chick. The low numbers of acid mucins in the little skate hindgut confirms that a functional colon for water absorption is absent in the Chondrichthyes. Interestingly, the presence of high levels of acid mucins in the posterior spiral valve provides evidence for a possible primordial water-absorbing organ in the elasmobranchs. Hoxd13 patterns acid mucins in the colons of terrestrial vertebrates. Expression of Hoxd13 and Hoxa13 in R. erinacea suggests conserved roles for Hox genes in patterning the early hindgut.     

Morphological aspects of the third instar larva of Haematobia irritans

The main morphological features of the cephalic region of the larva of Haematobia irritans (L.) are the oral grooves, tripartite labium and the antennomaxillary protuberances that have the dorsal, terminal and ventral sensory organs. The total number of sensilla that are found on the terminal organ differs from other cyclorrhaphous-fly larvae. The fan-shaped anterior spiracles usually consist of seven bulbous digits that are unequal in length. The creeping welts consist of notched, convex plates that split into two separate plates as they approach the midline of the venter. This characteristic has not been described previously for this species or other, higher, dipterous larvae. There are two posterior spiracles with an ecdysial scar, four fan-shaped and branching spiracular hairs and irregularly-shaped spiracular openings. The longitudinal anal opening is situated in the cuticular band that is known as the anal organ.     

Ia-afferent input to motoneurons during shortening and lengthening muscle contractions in humans.

The central nervous system employs different strategies to execute specific motor tasks. Because afferent feedback during shortening and lengthening muscle contractions differs, the neural strategy underlying these tasks may be quite distinct. Cortical drive may be adjusted or afferent input regulated. The exact mechanisms are not clear. Here, we examine the control of synaptic transmission across the Ia synapse during shortening and lengthening muscle contractions. Subjects were instructed to maintain isolated activity in a single tibialis anterior (TA) motor unit while muscle length was varied from flexion to extension and back. At a fixed interval after a firing of the active motor unit, a single electrical stimulus was applied to the common peroneal nerve to activate Ia afferents from the TA muscle. We investigated the stimulus-induced change in firing probability of 19 individual low-threshold TA motor units during shortening and lengthening contractions. Any change in firing probability depends on both pre- and postsynaptic mechanisms. In this experiment, motoneuron firing rate was similar during both contraction types. There was no difference in the firing probability between shortening and lengthening contractions (0.23 +/- 0.03 and 0.20 +/- 0.02, respectively). We suggest that there is no contraction type-specific control of Ia input to the motoneurons during shortening and lengthening muscle contractions. Cortical adjustments may have occurred.     

Electrosensory ampullary organs are derived from lateral line placodes in cartilaginous fishes

Ampullary organ electroreceptors excited by weak cathodal electric fields are used for hunting by both cartilaginous and non-teleost bony fishes. Despite similarities of neurophysiology and innervation, their embryonic origins remain controversial: bony fish ampullary organs are derived from lateral line placodes, whereas a neural crest origin has been proposed for cartilaginous fish electroreceptors. This calls into question the homology of electroreceptors and ampullary organs in the two lineages of jawed vertebrates. Here, we test the hypothesis that lateral line placodes form electroreceptors in cartilaginous fishes by undertaking the first long-term in vivo fate-mapping study in any cartilaginous fish. Using DiI tracing for up to 70 days in the little skate, Leucoraja erinacea, we show that lateral line placodes form both ampullary electroreceptors and mechanosensory neuromasts. These data confirm the homology of electroreceptors and ampullary organs in cartilaginous and non-teleost bony fishes, and indicate that jawed vertebrates primitively possessed a lateral line placode-derived system of electrosensory ampullary organs and mechanosensory neuromasts.     

Pyramidal unit activity in unanesthetized cats

Pyramidal unit activity in unanesthetized cats at rest and during voluntary movement was recorded by a microelectrode technique from the motor cortex for the forelimb. Some pyramidal neurons were not spontaneously active. The conduction velocity along the axon of these neurons was sometimes high (up to 71.5 m/sec), sometimes low (up to 11.2 m/sec). The remaining pyramidal neurons had spontaneous activity with a mean frequency of 1.29 to 43 spikes/sec. Analysis of interspike interval histograms of spontaneous activity and of autocorrelation histograms showed grouping of the spikes into volleys in most pyramidal neurons (irrespective of the conduction velocity). During voluntary movements the change in the activity of many pyramidal units correlated with changes in the EMG. The firing rate of the pyramidal neurons under these circumstances began to rise at least 50 msec before the increase in amplitude of the EMG and it remained high throughout the movement. The firing rate of most neurons during movement was 40–60/sec. The results are compared with those obtained by other workers who studied pyramidal unit activity of monkeys during voluntary movement.     

Sense organs in the femur of the stick insect and their relevance to the control of position of the femur-tibia-joint

Summary The sensory innervation pattern is described for the femur of the middle and the hind legs ofCarausius morosus. — In one of the nerves (F121) extracellular recordings show a unit which mirrors the tension of the flexor tibiae muscle (tension receptor). The tension receptor increases the firing rate of the slow extensor tibiae motoneuron. It measures the tension of one or more muscle fibres of the anterior side near the distal end of the muscle. The anatomical basis of this receptor is uncertain. — Another receptor was found on the ventral side of the distal end of the apodeme of the extensor tibiae muscle (apodeme receptor). Recordings from this receptor could not be obtained inCarausius. But inExtatosoma tiaratum it responded to stretching of the nerve. In the natural position it shows a minimum of excitation in the 90°-position of the femur-tibia-joint and an increase in firing rate for both flexion and extension. — Tactile hairs react phasically and have no special sensitivity for one direction. Two receptors at the dorsal side of the femur-tibia-joint (RDAL and RDPL), which are situated in the same position as inSchistocerca hind legs, react phasically to extension movements and fire tonically in the most extended position of the joint. — The influence of these receptors on the position of the femur-tibia-joint is only weak.Supported by Deutsche Forschungsgemeinschaft     

Homologies in the fossil record: The middle ear as a test case

This paper examines the middle ear of fossil living animals in terms of the homologies which have been drawn between its parts in different vertebrate groups. Seven homologies are considered: 1, the middle ear cavity/spiracular pouch; 2, the stapes/hyomandibula; 3, the stapedial/hyomandibular processes; 4 the tympanic membrane; 5, the otic notch; 6, the fenestra ovalis; 7, and the stapedial/hyomandibular foramen. The reasons leading to assessments of homology are reviewed. Homologies 1 and 2, based largely on embryological evidence, are fairly robust, though there are arguments about the details. Homologies 3, 4 and 5 stem from ideas about early tetrapod evolution, and were influenced by contingent factors including the order and time of discovery of early fossil taxa, and perceptions of their phylogeny which resulted from this. They were also influenced by ideas of the evolution of terrestriality among tetrapods. Most of the conceptions have been overturned in recent years by new fossil discoveries and new ways of looking at old data. Homology 6 has been little considered. One possible hypothesis, placed in a strictly archetypal theoretical framework has been ignored but deserves consideration on other grounds. Homology 7 depends on how tetrapods are characterised, not a question which has posed difficulties until recently, but which is likely to with the discovery of intermediate fossil forms.