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S Coombs

Publications and source records attributed to S Coombs.

At least 19 recordsLinked to original sources

Evaluation by respirometry of the degradability of retort water using a shale ash and overburden packed column.

Oil shale processing produces an aqueous wastewater stream known as retort water. The fate of the organic content of retort water from the Stuart oil shale project (Gladstone, Queensland) is examined in a proposed packed bed treatment system consisting of a 1:1 mixture of residual shale from the retorting process and mining overburden. The retort water had a neutral pH and an average unfiltered TOC of 2,900 mg 1(-1). The inorganic composition of the retort water was dominated by NH4+. Only 40% of the total organic carbon (TOC) in the retort water was identifiable, and this was dominated by carboxylic acids. In addition to monitoring influent and effluent TOC concentrations, CO2 evolution was monitored on line by continuous measurements of headspace concentrations and air flow rates. The column was run for 64 days before it blocked and was dismantled for analysis. Over 98% of the TOC was removed from the retort water. Respirometry measurements were confounded by CO2 production from inorganic sources. Based on predictions with the chemical equilibrium package PHREEQE, approximately 15% of the total CO2 production arose from the reaction of NH4+ with carbonates. The balance of the CO2 production accounted for at least 80% of the carbon removed from the retort water. Direct measurements of solid organic carbon showed that approximately 20% of the influent carbon was held-up in the top 20cm of the column. Less than 20% of this held-up carbon was present as either biomass or as adsorbed species. Therefore, the column was ultimately blocked by either extracellular polymeric substances or by a sludge that had precipitated out of the retort water.

Biodegradation, Environmental↗

The orienting response of Lake Michigan mottled sculpin is mediated by canal neuromasts.

Lake Michigan mottled sculpin, Cottus bairdi, exhibit a naturally occurring and unconditioned orienting response that can be triggered by both live prey and chemically inert vibrating spheres, even in blinded animals. CoCl(2)-induced reductions of the orienting response demonstrate that the lateral line is required for this behavior in the absence of non-mechanosensory cues (such as vision), but shed no light on the relative contributions of superficial and canal neuromasts to this behavior. To determine the relative roles of these two subsystems, we measured the frequency with which mottled sculpin oriented towards a small vibrating sphere before and after two treatments: (i) immersion of fish in a solution of gentamicin, an aminoglycoside antibiotic that damages hair cells in canal, but not superficial, neuromasts; and (ii) scraping the skin of the fish, which damages the superficial, but not the canal, neuromasts. To ensure that both superficial and canal neuromasts were adequately stimulated, we tested at different vibration frequencies (10 and 50 Hz) near or at the best frequency for each type of neuromast. At both test frequencies, response rates before treatment were greater than 70 % and were significantly greater than 'spontaneous' response frequencies measured in the absence of sphere vibration. Response rates fell to spontaneous levels after 1 day of gentamicin treatment and did not return to pre-treatment levels for 10-15 days. In contrast, response rates stayed approximately the same after superficial neuromasts had been damaged by skin abrasion. Scanning electron microscopy confirmed hair cell damage (loss of apical cilia) in canal, but not superficial, neuromasts of gentamicin-treated animals after as little as 24 h of treatment. The sensory epithelium of canal neuromasts gradually returned to normal, following a time course similar to behavioral loss and recovery of the orienting response, whereas that of superficial neuromasts appeared normal throughout the entire period. This study shows that the orienting response of the mottled sculpin is mediated by canal neuromasts.

Animals↗

Hydrodynamic image formation by the peripheral lateral line system of the Lake Michigan mottled sculpin, Cottus bairdi.

Lake Michigan mottled sculpin (Cottus bairdi) have a lateral-line-mediated prey-capture behaviour that consists of an initial orientation towards the prey, a sequence of approach movements, and a final strike at the prey. This unconditioned behaviour can be elicited from blinded sculpin in the laboratory by both real and artificial (vibrating sphere) prey. In order to visualize what Lake Michigan mottled sculpin might perceive through their lateral line when approaching prey, we have combined anatomical, neurophysiological, behavioural and computational modelling techniques to produce three-dimensional maps of how excitation patterns along the lateral line sensory surface change as sculpin approach a vibrating sphere. Changes in the excitation patterns and the information they contain about source location are consistent with behavioural performance, including the approach pathways taken by sculpin to the sphere, the maximum distances at which approaches can be elicited, distances from which strikes are launched, and strike success. Information content is generally higher for laterally located sources than for frontally located sources and this may explain exceptional performance (e.g. successful strikes from unusually long distances) in response to lateral sources and poor performance (e.g. unsuccessful strikes) to frontal sources.

Animals↗

The overlapping roles of the inner ear and lateral line: the active space of dipole source detection.

The problems associated with the detection of sounds and other mechanical disturbances in the aquatic environment differ greatly from those associated with airborne sounds. The differences are primarily due to the incompressibility of water and the corresponding increase in importance of the acoustic near field. The near field, or hydrodynamic field, is characterized by steep spatial gradients in pressure, and detection of the accelerations associated with these gradients is performed by both the inner ear and the lateral line systems of fishes. Acceleration-sensitive otolithic organs are present in all fishes and provide these animals with a form of inertial audition. The detection of pressure gradients, by both the lateral line and inner ear, is the taxonomically most widespread mechanism of sound-source detection amongst vertebrates, and is thus the most likely primitive mode of detecting sound sources. Surprisingly, little is known about the capabilities of either the lateral line or the otolithic endorgan in the detection of vibratory dipole sources. Theoretical considerations for the overlapping roles of the inner ear and lateral line systems in midwater predict that the lateral line will operate over a shorter distance range than the inner ear, although with a much greater spatial resolution. Our empirical results of dipole detection by mottled sculpin, a benthic fish, do not agree with theoretical predictions based on midwater fishes, in that the distance ranges of the two systems appear to be approximately equal. This is almost certainly as a result of physical coupling between the fishes and the substrate. Thus, rather than having a greater active range, the inner ear appears to have a reduced distance range in benthic fishes, and the lateral line distance range may be concomitantly extended.

Animals↗

Signal detection theory, lateral-line excitation patterns and prey capture behaviour of mottled sculpin.

The frequency with which blinded mottled sculpin, Cottus bairdi, oriented towards a dipole current source (50-Hz vibrating sphere) was measured as a function of source distance (2-18 cm) and azimuth (either 0 degrees in front or 90 degrees to the side of the fish). The orienting frequency declined from over 70% to under 50% as source distance increased from 4 to 12 cm for both frontal and lateral sources. When response biases (frequency of responding in the absence of the signal) were taken into account with the performance metric d', threshold distances (distances at which d' fell to 1) for frontal (12.5 cm) and lateral (11.6 cm) sources were 1.35-1.45 times the mean standard length of fish used in this study. At distances less than 8 cm, d' values were considerably higher (i.e. performance was better) for the lateral source, despite the fact that peak stimulus levels at the fish were twice as high for frontal as for lateral sources at any given distance. Performance differences may be related to differences in spatial excitation patterns, in particular the distribution of opposing pressure gradient directions along the lateral-line system, present for lateral sources, but absent for frontal sources. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Transformation of peripheral inputs by the first-order lateral line brainstem nucleus.

Extracellular recording techniques were used to record the responses of medial nucleus cells and posterior lateral line nerve fibers in mottled sculpin, Cottus bairdi, and goldfish, Carassius auratus, to a 50-Hz dipole source (vibrating sphere). Responses were characterized in terms of (1) receptive fields that relate responsiveness (spike rate and phase-locking) to the location of the source along the length of the fish, (2) input-output functions that relate responsiveness to vibration amplitude for a fixed source location, and (3) peri-stimulus time histograms that relate responsiveness to time during a sustained period of vibration. Relative to posterior lateral line nerve fibers, medial nucleus cells in both species were similar in showing (1) lower spontaneous and evoked rates of spike activity, (2) greater degrees of adaptation, (3) greater heterogeneity in all response characteristics, and (4) evidence for inhibitory/excitatory interactions. Whereas receptive fields of nerve fibers in both species faithfully reflect both pressure gradient amplitudes (with rate changes) and directions (with phase-angle changes) in the stimulus field, receptive fields of medial nucleus were more difficult to relate to the stimulus field, Some, but not all, receptive fields could be modeled with excitatory center/inhibitory surround and inhibitory center/excitatory surround organizations.

Animals↗

Dipole source localization by mottled sculpin. III. Orientation after site-specific, unilateral denervation of the lateral line system.

To test the hypothesis that spatial excitation patterns along the lateral-line system underlie source localization, we videotaped the orientation behavior of blinded mottled sculpin in response to a small dipole source (50-Hz vibrating sphere) before and after unilateral denervation of the lateral line system on different body regions (head, trunk and head + trunk). Approach pathways were qualitatively similar to those followed by normal intact animals. Abnormal behavior (turning in circles) was not observed. However, the frequency with which fish placed their intact side facing the source increased by 12-89%, depending on the denervation site. The angular accuracy of orientation decreased by 20 degrees to 60 degrees (100% to 370% change) depending on source location and region of lateral line denervated. Deficits tended to be site-specific. For example, unilaterally denervating lateral-line organs on the head resulted in less accurate orienting responses when the source was located on the denervated side of the head, but not on the opposite side of the head or on either side of the trunk. Site-specific deficits and the absence of abnormal approach pathways argue that animals are relying on a point-by-point spatial representation of source location along the sensory surface rather than computations based on bilateral comparisons.

Animals↗

Dipole source localization by mottled sculpin. I. Approach strategies.

Lake Michigan mottled sculpin respond to a chemically-inert vibrating sphere (a dipole source) with an initial orientation towards the source followed by a step-wise progression towards and final strike at the source. An analysis of videotape recordings of this behavior indicate that although pathways to the source varied, they tended to be influenced by the fish's position at signal onset. Fish heading toward the source at signal onset approached the source in an indirect fashion by either (a) keeping the source to one side in a smoothly arching path to the source or (b) alternating between keeping the source to the left and to the right. When the source was to the side of the fish at the time of stimulus onset, fish tended to approach the source in a more direct path. Most (79%) initial orienting responses placed the fish within 45 degrees of the source, but response angles were not strongly correlated with initial source angle. Most (83%) unsuccessful strikes (misses) occurred when the source was directly in front of the fish (+/- 20 degrees) and source angles associated with misses were significantly smaller than source angles associated with successful strikes. Approach strategies used by mottled sculpin in finding dipole sources appear to include (1) moving in a direction that increases the pressure difference along the head while keeping it consistently low (between 1 and 10 Pa) across the head, (2) narrowing the fish-to-source gap with each successive step in the pathway, (3) keeping the source lateralized (on average, 30 degrees to one or the other side of the head) and (4) avoiding approach positions that are perpendicular to the flow line or that place the fish in the pressure null area of the dipole field. These results are consistent with the hypothesis that spatial excitation patterns along the lateral line system play a major role in encoding both source direction and distance.

Animals↗

Dipole source localization by the mottled sculpin. II. The role of lateral line excitation patterns.

Extracellular, single unit recording techniques were used to measure the responses of posterior lateral line nerve fibers to a 50-Hz dipole source that slowly changed its location along the length of the fish. The flow-field equations for a dipole source were used to model the pressure gradient pattern and thus, the expected excitation pattern along a linear array of lateral line receptor organs for different source locations. Finally, excitation patterns were similarly modeled along the left and right side of the fish's head for actual steps taken by sculpin in approach pathways to the 50-Hz dipole source. Spatial histograms of posterior lateral line nerve fiber responses to different locations of the dipole source could be predicted from pressure gradient patterns modeled from the flow-field equations, confirming that the modeling approach applied to behavioral results was a good predictor of excitation patterns likely to be encoded by the lateral line periphery. An examination of how modeled excitation patterns changed from one position to the next in typical approach pathways and how patterns differed between positions from which successful and unsuccessful strikes were launched suggests that approach and strike strategies can indeed be explained by the information available in excitation patterns. In particular, changes in the spatial distribution of pressure gradient directions (polarities), available only when the source is lateral (as opposed to directly in front of the fish), appear to enhance the ability of sculpin to determine source distance. Without such information, misses are more likely to occur and successful strikes are more likely to be launched from short distances only.

Acoustic Stimulation↗

Cytoarchitecture of the medial octavolateralis nucleus in the goldfish, Carassius auratus.

The medial octavolateralis nucleus (MON) is the principal first-order medullary lateral line sensory nucleus found in the majority of anamniotic vertebrates. Although its presence has been confirmed in numerous taxa, the cytoarchitecture of this region has not been extensively studied in any species. The purpose of this study was to examine in detail the cytoarchitecture of the MON in the goldfish using Golgi staining and HRP histochemical techniques. The results of this study demonstrated the presence of a number of cell types with distinct cellular morphologies, several of which strongly resemble those described in octavolateralis nuclei dedicated to audition and electroreception. The most prominent of these MON neurons included crest cells of two varieties, either possessing or lacking basilar dendrites. Additionally, we described stellate and cristal interneurons and granule-like cells in the molecular layer, and lateral interneurons and granule-like neurons in deeper MON layers. These morphological similarities together with similarities in functional organization, and the probable close phyletic relationships of this "family" of hair cell sensory systems, argue for parallels in mechanisms of sensory processing and analysis in strongly divergent sensory modalities.

Afferent Pathways↗

Modeling and measuring lateral line excitation patterns to changing dipole source locations.

In order to determine excitation patterns to the lateral line system from a nearby 50 Hz oscillating sphere, dipole flow field equations were used to model the spatial distribution of pressures along a linear array of lateral line canal pores. Modeled predictions were then compared to pressure distributions measured for the same dipole source with a miniature hydrophone placed in a small test tank used for neurophysiological experiments. Finally, neural responses from posterior lateral line nerve fibers in the goldfish were measured in the test tank to demonstrate that modeled and measured pressure gradient patterns were encoded by the lateral line periphery. Response patterns to a 50 Hz dipole source that slowly changed location along the length of the fish included (1) peaks and valleys in spike-rate responses corresponding to changes in pressure gradient amplitudes, (2) 180 degrees phase-shifts corresponding to reversals in the direction of the pressure gradient and (3) distance-dependent changes in the locations of peaks, valleys and 180 degrees phase-shifts. Modeled pressure gradient patterns also predict that the number of neural amplitude peaks and phase transitions will vary as a function of neuromast orientation and axis of source oscillation. The faithful way in which the lateral line periphery encodes pressure gradient patterns has implications for how source location and distance might be encoded by excitation patterns in the CNS. Phase-shift information may be important for (1) inhibitory/excitatory sculpting of receptive fields and (2) unambiguously encoding source distance so that increases in source distance are not confused with decreases in source amplitude.

Acoustic Stimulation↗

Function and evolution of superficial neuromasts in an Antarctic notothenioid fish.

Extracellular recording techniques were used to measure frequency response functions of anterior and posterior lateral line nerve fibers innervating superficial neuromasts at five different locations on the head and trunk of an antarctic notothenioid fish, Trematomus bernacchii. Scanning electron microscopy was used to measure neuromast size according to location. Fibers innervating neuromasts from all locations were similar in showing equal responsiveness in the 10-30 Hz range to equal pk-pk velocity levels of a sinusoidally vibrating sphere. The mean cut-off frequency (CF) at which responsiveness declined to 50% of maximum was 46 Hz for all fibers combined. Superficial neuromasts located on the ventral trunk line were three to six times larger in surface area than most other neuromasts. The mean CF for fibers innervating these large neuromasts was 7-18 Hz lower than mean CF's corresponding to other superficial neuromast locations, but small differences in mean CF's were not consistently related to neuromast size. It is argued that fiber responses from different superficial neuromasts are more similar than dissimilar and that the evolution of large superficial neuromasts on the ventral trunk line is linked to a general paedomorphic trend among notothenioid fishes that may be essentially non-adaptive for the lateral line.

Afferent Pathways↗

Form and function relationships in lateral line systems: comparative data from six species of Antarctic notothenioid fish.

The structure and physiology of the anterior lateral line canal systems were studied in six species of fish belonging to two different families within the suborder of antarctic fish Notothenioidei. Many of the canals within the species belonging to the genus Trematomus are relatively straight sided tubes with diameters around 0.4 mm. Some of the canals in Trematomus, and most of the canals in the icefishes (family Channichthyidae) are more complex. Relatively small pores lead into large tubules, the walls of which appear partially membranous, and the canals not much more than constrictions between adjacent tubules. Dissostichus mawsoni, a large species, has canals with distinctive wide and narrow sections, 1.8 mm and 0.48 mm, respectively. Despite these morphological differences the frequency response characteristics of anterior lateral line units are remarkably similar in all six species. In the case of D. mawsoni, this functional similarity results from narrow sections of the canals, which provide the viscous resistance to flow that preserves the mechanical filtering properties of the canal despite the huge size difference between D. mawsoni and the other species. It is argued that the most appropriate way to view canals is as high pass filters which attenuate lower frequencies, and that this effect is best illustrated by comparing the frequency response characteristics of superficial and canal neuromasts using a sinusoidal stimulus that has a constant peak-to-peak velocity. The functional contribution of canals is to attenuate low frequencies and improve the signal-to-noise ratio for biologically important signals in the presence of low frequency noise produced, for example, by the animal's own movements.

Animals↗

Nearfield detection of dipole sources by the goldfish (Carassius auratus) and the mottled sculpin (Cottus bairdi).

Surprisingly few behavioral data exist on dipole source detection by fish, despite the fact that dipole sources more closely approximate biologically relevant signals than do more nearly monopole sources such as loudspeakers, the stimulus used in nearly all fish auditory studies. In this study, dipole source detection is investigated for two fish species that differ in both their auditory and lateral line systems, the two systems capable of detecting dipole sources. Conditioned suppression of respiration in the goldfish and an unconditioned orienting response in the mottled sculpin were used to measure detection of a 6 mm diameter, sinusoidally vibrating sphere as a function of vibration frequency and source distance. Sound pressure thresholds for the goldfish were nearly independent of distance (15-60 mm) at 800 Hz, but increased with distance at 50 Hz, as they did for the mottled sculpin. The slopes of 50 Hz source level-distance functions, however, differed between the two species. Slopes for the goldfish were independent of distance, remaining at around 8 dB per distance doubling, which is near the 6 dB per distance doubling measured for sound pressure attenuation away from the source, but less than the 18 dB per distance doubling for incompressible flow, measured with an anemometer. Those for the mottled sculpin increased with increasing distance, approaching 18 dB per distance doubling. The nonlinear increase in source level necessary to reach threshold detection was quite similar to the nonlinear decrease in incompressible flow levels measured with the anemometer. Nonlinear increases with distance for 50 Hz sources near the trunk of the mottled sculpin were also similar to those near the head of the fish, where changes in source frequency had little effect on source level-distance functions. These results indicate that sound pressure detection by the ear is important for dipole detection by the goldfish, but that incompressible flow detection by the lateral line is more important for the mottled sculpin. They also indicate that fish such as the goldfish, with a pressure-sensitive swimbladder, are capable of detecting dipole sources at greater distances than are fish without such structures.

Acoustic Stimulation↗

Source level discrimination by the lateral line system of the mottled sculpin, Cottus bairdi.

A conditioned feeding response of the mottled sculpin was used to measure level discrimination by the lateral line system as a function of source level, frequency, and distance. Level discrimination limens (LDLs) were determined with a 6-mm-diam, sinusoidally vibrating sphere placed in front of the trunk lateral line system at varying distances from the trunk of the fish. LDLs were relatively independent of frequency over most of the detection bandwidth, with mean values ranging from 5-6 dB from 10 to 50 Hz at a fish-to-source distance of 15 mm. The mean LDL at 100 Hz, near the upper end of the frequency range of the lateral line system was 8-9 dB at the same source distance. Mean LDLs at 10 and 20 dB above detection threshold were not significantly different, nor were mean LDLs at distances ranging from 7.5 to 30 mm for a 50-Hz, 10-dB SL source. The level-encoding features of posterior lateral line fibers innervating the trunk were also determined from data previously collected under stimulus conditions nearly identical to those used for behavioral studies. Afferent fibers responded to level increments with increases in both spike rate and phase-locking over most of the conditions used in behavioral experiments. Although it was impossible to eliminate spike rate as a level-encoding mechanism at any stimulus condition, phase-locking appeared to be a strong candidate at low source levels and vibration frequencies.(ABSTRACT TRUNCATED AT 250 WORDS)

Acoustic Stimulation↗

Physiological characterization of lateral line function in the Antarctic fish Trematomus bernacchii.

The Antarctic notothenioids are a monophyletic radiation of fishes that have evolved under conditions of low light and cold, where non-visual sensory systems, such as the mechanosensory lateral line system, would be of importance. As part of a study of the structure and function of the mechanosensory lateral line system in these fishes we systematically characterized the function of the anterior lateral line system in one of the common benthic species, Trematomus bernacchii. Frequency-response and threshold-tuning curve methods yield similar functional characterizations of single afferent fibre responses to vibrational stimulation. Curve fitting of generalized transfer function to frequency-response curves allows an objective splitting of responses into velocity and acceleration sensitive populations thought to correspond to superficial and canal neuromasts, respectively. Both response types are characterized by a low-pass frequency-response curve, with a relatively low upper frequency cut-off when compared with data from temperate species. The sensitivity of the lateral line system of T. bernacchii is comparable to that of the mottled sculpin Cottus bairdi.

Acceleration↗

Fibers innervating different parts of the lateral line system of an Antarctic notothenioid, Trematomus bernacchii, have similar frequency responses, despite large variation in the peripheral morphology.

Regional differences in the architecture and size of lateral line canals and neuromasts were measured in an Antarctic fish, Trematomus bernacchii, and the data were used in models of canal and cupular mechanics to predict the frequency response of these two peripheral structures. These modeled predictions were then compared to frequency response functions measured with single unit recording techniques from anterior and posterior lateral line fibers innervating different canals on the head and trunk of fish of various sizes. Despite large variations in the peripheral morphology of head and trunk canals in fish of different sizes, lateral line fibers were relatively homogeneous in their frequency response properties. In response to stimuli of equal pk-pk acceleration levels, all canal neuromast fibers responded with equal and maximum responsiveness in the 10-45 Hz range, after which responsiveness fell off at about 18 dB/octave. Whereas the biomechanical models of cupular and canal responsiveness predicted the region of equal and maximum responsiveness in the 10-45 Hz range, they did not predict the high frequency cutoff nor the slope. Rather, these models predicted responsiveness out to at least 540 Hz, and a high frequency slope of 12 dB/octave. In terms of the frequency response of peripheral fibers, we conclude that (1) there can be considerable morphological variability, with little consequence for function, as long as some minimum standards for maintaining constant acceleration responsiveness in the 10-45 Hz range are met, and (2) there must be additional filters between the cupula and primary afferent fibers.

Acceleration↗