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E A Arbas

Publications and source records attributed to E A Arbas.

16 recordsLinked to original sources

Variability in odor-modulated flight by moths.

Based on previous studies of odor-modulated flight where track parameter data was lumped and averaged, the speed and orientation of the moths' movement along their flight tracks have been said to be controlled to maintain certain "preferred" values. The results from our fine-scaled analysis of this behavior show that none of the track parameters typically measured are held constant. The moths' speed along the flight track is modulated substantially and predictably: fastest along the straight legs and slowest around the turns. In addition, about half of the individuals studied progressively reduced the peak speed along the straight legs as they approached the pheromone source. While most of the track legs between the turns were directed upwind, their orientations were widely distributed, indicating no preferred direction. Small fluctuations of orientation along some straight legs suggest corrective maneuvers to stabilize flight direction about an internal set point. The visual inputs hypothesized to control steering and speed, transverse and longitudinal image flow, changed continuously during upwind flight in pheromone, but no regular relationship between them was observed. We found that the orientation of the longitudinal body axis and the direction of thrust (course angle) were only rarely coincident during upwind flight to the odor source, suggesting that moths receive sensory input which differs quantitatively from that calculated by conventional methods. Our results strongly suggest that the long-accepted hypothetical mechanisms of control for this behavior do not operate in the manner in which they have been proposed.

Animals

Morphology of the vasopressin-like immunoreactive (VPLI) neurons in many species of grasshopper.

It has previously been shown that the pair of vasopressin-like immunoreactive (VPLI) neurons of the locust, Locusta migratoria, have cell bodies on the ventral midline of the suboesophageal ganglion and extensive arborisations in all ganglia of the central nervous system. In the present study, we have stained vasopressin-like immunoreactive neurons in 16 additional species of grasshopper, and consistently find this pair of extensive neurons: we assume these to be interspecies homologues. However, the anatomy of these neurons falls into two morphological types: the first, typified by Schistocerca gregaria, has most of its processes distributed in dorsal and lateral neuropil of all ganglia; the second, typified by Locusta migratoria, is equally extensive in its arborisation, but the distribution of branches is shifted peripherally into the optic lobes and the proximal portions of peripheral nerves. It has been suggested that the peripheral fibres in Locusta migratoria are neurohaemal organs for the release of a vasopressin-like diuretic peptide. Our sample of 17 Acridoid species has deliberately selected animals from very different habitats, but our extensive survey of VPLI anatomy shows that peripheral fibres are only present in species from the subfamily Oedipodinae (of which Locusta migratoria is a member) and that no peripheral fibres are present in any of the species from the 4 other subfamilies of the Acridoidea that we have examined. The presence of peripheral fibres is therefore determined by phylogeny and not by habitat. The absence of peripheral VPLI fibres in most grasshopper species examined in this study probably means that the release of putative diuretic hormone from VPLI to control water homeostasis cannot be a conserved function of this ubiquitous neuron. In contrast, the extensive central arborisations and rare antigenicity, which are highly conserved features of the VPLI neuron in all those grasshoppers we have examined, suggests that any conserved role is more likely to be central. A central role for the VPLI neuron has yet to be determined.

Animals

Transneuronal induction of muscle atrophy in grasshoppers.

Autotomy is a process in grasshoppers whereby one or both hindlimbs can be shed to escape a predator or can be abandoned if damaged. It occurs between the trochanter and the femur (second and third leg segments) and once lost, the legs never regenerate. Autotomy severs branches of the leg nerve (N5) but damages no muscles since none span the autotomy plane. We find, however, that undamaged muscles intrinsic to the thorax of grasshoppers, Barytettix psolus, atrophy to less than 15% of their normal mass after autotomy of a hindlimb. These muscles operate the coxa and trochanter (first and second leg segments) and are innervated by branches of nerves 3 and 4; nerve branches that are not damaged by autotomy. Atrophy is localized to the side and body segment where autotomy occurs. Atrophy is evident 7-10 days after loss of a limb, is complete by about 30 days, and follows a similar time course whether induced in young adult, or sexually mature grasshoppers. During autotomy, leg nerve 5 is served distal to the trochanter, the thoracic muscles lose their normal static and dynamic load, and these muscles are subsequently no longer used to support the weight of the insect during posture and locomotion. Experimental loading and unloading of the affected muscles, and cutting of nerves indicated that it is the severing of leg nerve 5 during autotomy that transneuronally induces muscle atrophy.

Animals

Odor-modulated upwind flight of the sphinx moth, Manduca sexta L.

1. Male and female Manduca sexta flew upwind in response to the odor of female sex-pheromone gland extract or fresh tobacco leaf respectively, and generated very similar zigzagging tracks along the odor plume. 2. After loss of odor during flight, males and females alike: (1) first flew slower and steered their flight more across the wind, then (2) stopped moving upwind, and finally (3) regressed downwind. 3. Males flying upwind in a pheromone plume in wind of different velocities maintained their ground speed near a relatively constant 'preferred' value by increasing their air speed as the velocity of the wind increased, and also maintained the average angle of their resultant flight tracks with respect to the wind at a preferred value by steering a course more precisely due upwind. 4. The inter-turn duration and turn rate, two measures of the temporal aspects of the flight track, were maintained, on average, with remarkable consistency across all wind velocities and in both sexes. The inter-turn durations also decreased significantly as moths approached the odor source, suggesting modulation of the temporal pattern of turning by some feature of the odor plume. This temporal regularity of turning appears to be one of the most stereotyped features of odor-modulated flight in M. sexta.

Animals

Physiology and morphology of descending neurons in pheromone-processing olfactory pathways in the male moth Manduca sexta.

1. We have characterized the responses and structure of olfactory descending neurons (DNs) that reside in the protocerebrum (PC) of the brain of male sphinx moths Manduca sexta and project toward thoracic ganglia. 2. Excitatory responses of DNs to pheromone blends were of two general types: (a) brief excitation (BE) that recovered to background in less than 1 s after the stimulus, and (b) long-lasting excitation (LLE) that outlasted the stimulus by greater than or equal to 1 s and, in many cases, as long as 30 s. Individual pheromone components were ineffective in eliciting LLE. 3. Some neurons showing LLE also exhibited state-dependent responses to pheromonal stimuli. When such neurons were in a state of low background firing, stimulation with pheromone blend elicited LLE. When they were in a state of LLE, an identical stimulus reduced firing for 5-10 s after which firing gradually increased to the initial higher level. 4. Thirteen stained DNs were reconstructed from serial sections for detailed analysis of their morphology in the brain. DNs exhibiting LLE had neurites concentrated in the lateral accessory lobes (LALs) in the protocerebrum and adjacent neuropil. Most DNs exhibiting only BE to pheromonal stimuli and other DNs showing responses only to visual or mechanosensory stimuli did not have branches in the LALs.

Animals

Physiology and morphology of protocerebral olfactory neurons in the male moth Manduca sexta.

1. We have used intracellular recording and staining with Lucifer Yellow, followed by reconstruction from serial sections, to characterize the responses and structure of olfactory neurons in the protocerebrum (PC) of the brain of the male sphinx moth Manduca sexta. 2. Many olfactory protocerebral neurons (PCNs) innervate a particular neuropil region lateral to the central body, the lateral accessory lobe (LAL), which appears to be important for processing olfactory information. 3. Each LAL is linked by its constituent neurons to the ipsilateral lateral PC, where projection neurons from the antennal lobe terminate, as well as to other regions of the PC. The LALs are also linked to each other by bilateral neurons with arborizations in each LAL. 4. Some PC neurons showed long-lasting excitation (LLE) that outlasted the olfactory stimuli by greater than or equal to 1 s, and as long as 30 s in some preparations. LLE was more frequently elicited by the sex-pheromone blend than by individual pheromone components. All bilateral neurons that showed LLE had arborizations in the LALs. LLE responses were also recorded in a single local neuron innervating the mushroom body. 5. In some other PC neurons, pheromonal stimuli elicited brief excitations that recovered to background firing rates less than 1 s after stimulation.

Animals

Leydig neuron activity modulates heartbeat in the medicinal leech.

1. Leydig neurons fire spontaneously at low rates (less than 4 Hz), but their activity increases with mechanical stimulation or electrical stimulation of mechanosensory neurons. These conditions also cause acceleration of bursting in heart motor neurons. 2. The firing rate of Leydig cells was found to regulate heart rate in chains of isolated ganglia. When Leydig neurons were made to fire action potentials at relatively high frequencies (ca. 5-10 Hz), however, heart motor neurons ceased bursting and were either silenced or fired erratically. 3. Firing of Leydig neurons at high rates caused bilateral heart interneurons of ganglia 3 or 4 to fire tonically rather than in their normal alternating bursts Tonic firing of these heart interneurons accounts for the prolonged barrages of ipsps recorded in heart motor neurons and the disruption of their normal cyclic activity. 4. Preventing spontaneous activity of Leydig neurons with injected currents in isolated ganglia caused deceleration of the heartbeat rhythm but did not halt oscillation. 5. Electrical stimulation of peripheral nerve roots with Leydig neuron activity suppressed in isolated ganglia caused acceleration of heart rate.

Animals

Physiology and morphology of projection neurons in the antennal lobe of the male moth Manduca sexta.

1. We have used intracellular recording and staining, followed by reconstruction from serial sections, to characterize the responses and structure of projection neurons (PNs) that link the antennal lobe (AL) to other regions of the brain of the male sphinx moth Manduca sexta. 2. Dendritic arborizations of the AL PNs were usually restricted either to ordinary glomeruli or to the male-specific macroglomerular complex (MGC) within the AL neuropil. Dendritic fields in the MGC appeared to belong to distinct partitions within the MGC. PNs innervating the ordinary glomeruli had arborizations in a single glomerulus (uniglomerular) or in more than one ordinary glomerulus of one AL (multiglomerular) or in one case, in single glomeruli in both ALs (bilateral-uniglomerular). One PN innervated the MGC and many or all ordinary glomeruli of the AL. 3. PNs with dendritic arborizations in the ordinary glomeruli and PNs associated with the MGC typically projected both to the calyces of the ipsilateral mushroom body and to the lateral protocerebrum, but some differences in the patterns of termination in those regions have been noted for the two classes of PNs. One PN conspicuously lacked branches in the calyces but did project to the lateral protocerebrum. The PN innervating the MGC and many ordinary glomeruli projected to the calyces of the ipsilateral mushroom body and the superior protocerebrum. 4. Crude sex-pheromone extracts excited all neurons with arborizations in the MGC, although some were inhibited by other odors. One P(MGC) was excited by crude sex-pheromone extract and by a mimic of one component of the pheromone blend but was inhibited by another component of the blend. 5. PNs with dendritic arborizations in ordinary glomeruli were excited or inhibited by certain non-pheromonal odors. Some of these PNs also responded to mechanosensory stimulation of the antennae. 6. The PN with dendritic arborizations in the MGC and many ordinary glomeruli was excited by crude sex-pheromone extracts and non-pheromonal odors and also responded to mechanosensory stimulation of the antenna.

Action Potentials

Morphology and physiological properties of interneurons in the olfactory midbrain of the crayfish.

1. Intracellular recording and staining was used to characterize neurons in the crayfish (Procambarus clarkii) brain that respond to chemical stimuli applied to the major olfactory organs, the antennules. 2. Two distinct morphological types of neurons that have major projections in the olfactory lobes (OLs) of the brain were characterized anatomically (Figs. 1, 2, 3; Table 2) and physiologically (Figs. 4, 5, 6; Table 3). 3. Different individual neurons of one type, with similar 'tree-like' projections in the OLs, have somata distributed in at least 5 different cell body clusters of the brain (Fig. 3) and link different subsets of neuropilar lobes through their distributed arbors (Fig. 1, Table 2). 4. Excitatory, inhibitory and mixed responses were recorded in different neurons when odorant mixtures or individual components of these mixtures were applied to the antennules. Response spectra to individual components were broad and overlapping, but not identical in the neurons tested (Fig. 4; Table 3). Mixture interactions appear to be additive in most of the neurons that we tested, but evidence was obtained for mixture suppression in several cases (Fig. 6). 5. Most of the neurons recorded in this study responded only to stimulation of the ipsilateral antennule (Fig. 5), although subthreshold activity to stimuli applied contralaterally was recorded in several neurons that were strongly excited by ipsilateral stimuli. 6. Chemoresponsive neurons without projections in OL's that have all of their branches confined to the brain, or that project an axon in the circumesophageal connective, are described (Fig. 7).

Action Potentials

Ionic conductances underlying the activity of interneurons that control heartbeat in the medicinal leech.

Electrical properties of interneurons that control heartbeat in the leech (HN cells) were studied using intracellular recording and stimulation in isolated ganglia bathed by salines of various ionic compositions. Substitution of Na+ ions in the bath by Tris stopped the spontaneous firing of HN cells and led to their gradual hyperpolarization by 15-20 mV. In the absence of Na+, HN neurons produced long-lasting regenerative plateau potentials with thresholds near -55 mV and peaks near -30 mV that were accompanied by an increase in membrane conductance. Elevation of Ca2+ concentration enhanced plateaus, as did replacement of Ca2+ by Ba2+. Plateaus were formed when Sr2+ replaced Ca2+, but were blocked by addition of Mg2+ or Co2+ to the bath, Co2+ being effective at lower concentrations than Mg2+. Hyperpolarization of HN neurons with injected currents revealed a time-dependent change in membrane potential, whereby initial maximum hyperpolarization was followed by a "sag" in potential towards more depolarized values. The sag showed dual voltage dependence, being diminished when HN neurons were hyperpolarized or depolarized outside the normal range of oscillation. The sag was found to depend on the presence of Na+ ions and to be blocked by Cs+ but not by Ba2+. This time-dependent change in membrane potential counters hyperpolarizations of HN neuron membrane potential and may contribute to the escape of these neurons from synaptic inhibition.

Action Potentials

Slow oscillations of membrane potential in interneurons that control heartbeat in the medicinal leech.

In the preceding paper (Arbas and Calabrese, 1987), we identified several properties that contribute to the activity of neurons (HN cells) that control heartbeat in the medicinal leech. Premotor HN (7) interneurons, which do not generate the heartbeat rhythm, exhibit Na+-dependent fast action potentials, Ca2+-mediated plateau potentials in the absence of Na+, and hyperpolarization-activated "restorative" changes in membrane potential that depolarize the membrane potential on hyperpolarization due to injected currents or synaptic inhibition. HN interneurons of ganglia 3 and 4 (i.e., timing oscillator interneurons) exhibit all of the properties described for HN (7) interneurons and have the additional characteristic that they are connected in oscillatory circuits. Reciprocal oscillations in membrane potential occurred in the bilateral HN interneurons (3) and (4) in the presence of elevated Ca2+ that were independent of Na+ -mediated action potentials. Their ability to oscillate in this way is based on 3 parameters: (1) production of a regenerative plateau potential by one of the pair of HN neurons in either ganglion, (2) inhibition of the contralateral HN neuron by the HN neuron in plateau, and (3) a phase transition mediated by escape from inhibition by the hyperpolarized HN neuron. The conductances responsible for restorative membrane potential shifts activated by hyperpolarization during synaptic inhibition may mediate the escape from inhibition that times the phase transition of the 2 HN neurons.

Action Potentials

Presynaptic terminals persist following degeneration of "flight" muscle during development of a flightless grasshopper.

We have studied the development of a neuromuscular system for which mature function has been lost through evolution in the grasshopper, Barytettix psolus (Cohn and Cantrall, 1974). Barytettix is flightless throughout life and has only vestigial wings that are incapable of active movement. Adult Barytettix lack muscles homologous to the indirect flight muscles of locusts and grasshoppers that fly, while other thoracic muscles are similar. We have found, using light and electron microscopic examination of tissues from various developmental stages, that the metathoracic dorsal longitudinal muscle is present and is innervated during nymphal life but is absent in adults. Yet its nerve persists and, in the adult, contains axonal presynaptic specializations opposite inappropriate targets such as glial processes and basal lamina. Our findings indicate that selective muscle death during development is one mechanism underlying the reduction of the flight system of Barytettix through evolution. The finding that presynaptic terminals persist in the absence of the muscle indicates that the muscle and its innervation follow programs of development that are at least partially independent and reinforces the concept that in insects motorneurons, and perhaps neurons in general, are not dependent upon trophic influences from their targets for survival and maintenance of their differentiated phenotype.

Animals

Control of hindlimb posture by wind-sensitive hairs and antennae during locust flight.

Steering movements of tethered, flying locusts, Schistocerca gregaria, subjected to simulated yaw were examined under open-loop conditions. Lateral movements of hindlimbs or curling of the abdomen were monitored with a capacitive movement transducer and were interpreted as indicating the tendency of the animal to turn. Three responses to simulated yaw were noted: Yaw-correcting upwind turning tendencies (Figs. 1, 2, 3). Downwind turning tendencies (Figs. 2, 3, 4, 5), and transient adjustments of hindlimb position consistent with an upwind turning tendency occurred in animals that made either no sustained postural adjustments of hindlimbs, or that exhibited sustained downwind turning tendencies (Figs. 4, 5). Ablations of certain mechanoreceptors tested their roles in wind detection and wind angle determination. The expression of upwind turning tendencies, whether sustained or transient, depends on inputs from cephalic mechanosensory hairplates (Figs. 2, 3, 4, 5). With hairplates occluded, all locusts exhibited downwind turning tendencies. All downwind turning tendencies depend on inputs from the antennae (Figs. 2, 3). Antennae and hairplates operate in an apparent antagonism in the steering responses they produce, which may provide the control flexibility required for complex flight maneuvering.

Animals

Neural correlates of flight loss in a Mexican grasshopper, Barytettix psolus. I. Motor and sensory cells.

The nervous systems of locusts (Schistocerca gregaria) and flightless grasshoppers (Barytet tix psolus) are compared to evaluate modifications to neurons which are associated with flight loss. Locusts are well known for their powerful flight capability. Barytettix never fly. They lack hindwings, have immobile vestiges of forewings, and are devoid of skeletal specializations for wing movement. Their pterothoracic musculature is similar to that of locusts, except for the absence of those muscles that, in locusts, have the primary function of moving the wings. Individually identified leg motorneurons, the extensors of the tibia, were compared between locusts and Barytettix and were found to have very similar morphologies. Nerve roots which correspond to those supplying wing muscles of locusts were stained by cobalt backfilling in Barytettix to test for presence of counterparts to wing muscle motorneurons. Cobalt backfills of metathoracic nerve 1 reveal the presence in Barytet tix of neurons corresponding to locust dorsal longitudinal motorneurons--neurons which persist in adult Barytettix in the complete absence of peripheral targets. These cells occupy characteristic positions within the CNS but their soma sizes are greatly reduced by comparison to their locust counterparts. Locust metathoracic ganglia bear large flight motorneurons on their ventral anterolateral margin. Viewed in toluidine blue-stained wholemounts, Barytettix ganglia show considerably smaller neuron somata in the corresponding region. In locusts, comparisons of the fast extensor tibiae (FETi) motorneuron soma profile areas with those of the largest anterior cell showed no significant difference between the two, while in Barytettix, the largest anterior cell is 51% smaller than the FETi. A counterpart to the locust wing hinge stretch receptor (SR) was revealed by backfilling metathoracic nerve 1 in Barytettix. Despite its lack of function as a wing movement detector, the central projection of Barytettix SR differs from its locust counterpart only in reduced spread of specific central branches.

Animals

Neural correlates of flight loss in a Mexican grasshopper, Barytettix psolus. II. DCMD and TCG interneurons.

Comparison of the nervous systems of a flightless grasshopper (Barytettix psolus), and locusts (Schistocerca) is extended to include two large interneurons, the descending contralateral movement detector (DCMD) and the tritocerebral commissure giant (TCG). These interneurons are thought to play a role in locust flight. Both were visualized in Barytettix with cobalt staining techniques. Most features of the two neurons are similar in both locusts and Barytettix. Yet, several differences were noted. In locusts, the dorsal DCMD branch in its metathoracic projection terminates in dorsolateral neuropile and synapses with flight motorneurons (Pearson and Goodman, '79). In Barytettix, no branch terminates in the corresponding region. Of the animals examined, 52% lacked branches corresponding to locust dorsal branches. When present, they all had abnormal projections by comparison to their counterparts in locusts. The maintenance of the TCG in Barytettix with so similar a form to that of locusts suggests that the role of the cell in behavior other than flight should be examined. The differences in DCMD projection suggest that a discrete set of output connections may have been modified in Barytettix by the alteration of a single first-order axonal branch.

Animals