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T Reuter

Publications and source records attributed to T Reuter.

At least 37 records · Page 2Linked to original sources

What middle ear parameters tell about impedance matching and high frequency hearing.

Acoustic energy enters the mammalian cochlea aided by an anatomical impedance matching performed by the middle ear. The purpose of this paper is to analyse the functional consequences of changes in scale of the middle ear when going from the smallest mammals to the largest. Our anatomical measurements in mammals of different sizes ranging from bats to elephants indicate that middle ear proportions are largely isometric. Thus the calculated transformer ratio is basically independent of animal size, a typical value lying between 30 and 80. Similarly, the calculated specific acoustic input impedance of the inner ear is independent of animal size, the average value being about 140 kPa s/m. We show that if the high frequency hearing limit of isometric ears is limited by ossicle inertia, it should be inversely proportional to the cubic root of the ossicular mass. This prediction is in reasonable agreement with published audiogram data. We then present a three-parameter model of the middle ear where some obvious deviations from perfect isometry are taken into account. The high frequency hearing limits of different species generally agree well with the predictions of this simple model. However, the hearing limits of small rodents clearly deviate from the model calculation. We interpret this observation as indicating that the hearing limit towards very high frequencies may be set by cochlear transduction mechanisms. Further we discuss the exceptional high frequency hearing of the cat and the amphibious hearing of seals.

Acoustic Impedance Tests↗

Dark adaptation of toad rod photoreceptors following small bleaches.

The recovery of toad rod photoreceptors, following exposure to intense lights that bleached 0.02-3% of the rhodopsin, has been investigated using the suction pipette technique. The post-bleach period was accompanied by reduced flash sensitivity, accelerated kinetics, and spontaneous fluctuations (noise). The power spectrum of the fluctuations had substantially the form expected for the random occurrence of single-photon events, and the noise could therefore be expressed as a "photon-noise equivalent intensity". From the level of desensitization at any time, the after-effect of the bleach could also be expressed in terms of a "desensitization-equivalent intensity", and this was found to be at least a factor of 20 times higher than the noise-equivalent intensity at the corresponding time. Our results indicate that a bleach induces two closely-related phenomena: (a) a process indistinguishable from the effect of real light, and (b) another process which desensitizes and accelerates the response in the same way that light does, but without causing photon-like noise. We propose a mechanism underlying these processes.

Animals↗

Visual performance of the toad (Bufo bufo) at low light levels: retinal ganglion cell responses and prey-catching accuracy.

The accuracy of toad snapping towards moving worm dummies under various levels of dim illumination (from absolute threshold to "moonlight") was video-recorded and related to spike responses of retinal ganglion cells exposed to equivalent stimuli. Some toads (at ca. 16 degrees C) successfully snapped at dummies that produced only one photoisomerization per 50 rods per second in the retina, in good agreement with thresholds of sensitive retinal ganglion cells. One factor underlying such high sensitivity is extensive temporal summation by the ganglion cells. This, however, is inevitably accompanied by very long response latencies (around 3 s near threshold), whereby the information reaching the brain shows the dummy in a position where it was several seconds earlier. Indeed, as the light was dimmed, snaps were displaced successively further to the rear of the dummy, finally missing it. The results in weak but clearly supra-threshold illumination indicate that snaps were aimed at the advancing head as seen by the brain, but landed further backwards in proportion to the retinal latency. Near absolute threshold, however, accuracy was "too good", suggesting that the animal had recourse to a neural representation of the regularly moving dummies to correct for the slowness of vision.

Animals↗

Retinal origins of the temperature effect on absolute visual sensitivity in frogs.

1. The absolute sensitivity of vision was studied as a function of temperature in two species of frog (Rana temporaria, 9-21 degrees C, and Rana pipiens, 13-28 degrees C). 2. Log behavioural threshold (measured as the lowest light intensity by which frogs trying to escape from a dark box were able to direct their jumping) rose near-linearly with warming with a regression coefficient of 1.26 +/- 0.03 log units per 10 degrees C (Q10 = 18). Threshold retinal illumination corresponded to 0.011 photoisomerizations per rod per second (Rh* s-1) at 16.5 degrees C. 3. The effect of dim backgrounds on jumping thresholds suggested 'dark lights' of 0.011 Rh* s-1 at 16.5 degrees C and 0.080 Rh* s-1 at 23.5 degrees C, corresponding to Q10 = 17. 4. Response thresholds of retinal ganglion cells were extracellularly recorded in the isolated eyecup of R. temporaria. The thresholds of the most sensitive cells when stimulated with large-field steps of light were similar to the behavioural threshold and changed with temperature in a similar manner. 5. The decrease in ganglion cell 'step' sensitivity with warming consisted of a decrease in summation time (by a factor of 2-3 between 10 and 20 degrees C) and an increase in the threshold number of photoisomerizations (a decrease in 'flash' sensitivity, by a factor of 2-5 over the same interval). No effect of temperature changes on spatial summation was found. 6. Frequency-of-response functions of ganglion cells indicated an 11-fold increase in noise-equivalent dark light between 10 and 20 degrees C (mean values in four cells 0.009 vs. 0.10 Rh* s-1). 7. The temperature dependence of ganglion cell flash sensitivity could be strongly decreased with dim background illumination. 8. It is concluded that the desensitization of dark-adapted vision with rising temperature is a retinal effect composed of shortened summation time and lowered flash sensitivity (increased numbers of photons required for a threshold response) in ganglion cells. The desensitization bears no simple relation to the apparent retinal noise increase.

Action Potentials↗

Signal transmission through the dark-adapted retina of the toad (Bufo marinus). Gain, convergence, and signal/noise.

Responses to light were recorded from rods, horizontal cells, and ganglion cells in dark-adapted toad eyecups. Sensitivity was defined as response amplitude per isomerization per rod for dim flashes covering the excitatory receptive field centers. Both sensitivity and spatial summation were found to increase by one order of magnitude between rods and horizontal cells, and by two orders of magnitude between rods and ganglion cells. Recordings from two hyperpolarizing bipolar cells showed a 20 times response increase between rods and bipolars. At absolute threshold for ganglion cells (Copenhagen, D.R., K. Donner, and T. Reuter. 1987. J. Physiol. 393:667-680) the dim flashes produce 10-50-microV responses in the rods. The cumulative gain exhibited at each subsequent synaptic transfer from the rods to the ganglion cells serves to boost these small amplitude signals to the level required for initiation of action potentials in the ganglion cells. The convergence of rod signals through increasing spatial summation serves to decrease the variation of responses to dim flashes, thereby increasing the signal-to-noise ratio. Thus, at absolute threshold for ganglion cells, the convergence typically increases the maximal signal-to-noise ratio from 0.6 in rods to 4.6 in ganglion cells.

Action Potentials↗

Weber and noise adaptation in the retina of the toad Bufo marinus.

Responses to flashes and steps of light were recorded intracellularly from rods and horizontal cells, and extracellularly from ganglion cells, in toad eyecups which were either dark adapted or exposed to various levels of background light. The average background intensities needed to depress the dark-adapted flash sensitivity by half in the three cell types, determined under identical conditions, were 0.9 Rh*s-1 (rods), 0.8 Rh*s-1 (horizontal cells), and 0.17 Rh*s-1 (ganglion cells), where Rh* denotes one isomerization per rod. Thus, there is a range (approximately 0.7 log units) of weak backgrounds where the sensitivity (response amplitude/Rh*) of rods is not significantly affected, but where that of ganglion cells (1/threshold) is substantially reduced, which implies that the gain of the transmission from rods to the ganglion cell output is decreased. In this range, the ganglion cell threshold rises approximately as the square root of background intensity (i.e. in proportion to the quantal noise from the background), while the maintained rate of discharge stays constant. The threshold response of the cell will then signal light deviations (from a mean level) of constant statistical significance. We propose that this type of ganglion cell desensitization under dim backgrounds is due to a post-receptoral gain control driven by quantal fluctuations, and term it noise adaptation in contrast to the Weber adaptation (desensitization proportional to the mean background intensity) of rods, horizontal cells, and ganglion cells at higher background intensities.

Action Potentials↗

Differences between induction effects of 1,4-bis[2-(3,5-dichloropyridyloxy)]benzene and phenobarbitone.

The inductive effects of phenobarbitone (PB) and 1,4-bis[2-(3,5-dichloropyridyloxy)]benzene (TCPOBOP) were compared in C57BL/6J mice. Induction parameters included six substrates: ethylmorphine (EM), benzphetamine (Bph), biphenyl, ethoxycoumarin (EtoC), pentoxyresorufin and dichloro-p-nitroanisole (DPNA). In order to validate this descriptive approach the comparison was extended to diazepam, rifampicin, warfarin, and pregnenolone-16 alpha-carbonitrile (PCN). All inducers were clearly distinguishable from each other. Warfarin was similar to PB, rifampicin was similar to PCN. TCPOBOP differed significantly from PB in relative liver weight, cytochrome P-450 content of liver microsomes, EM-, Bph- and DPNA-demethylations, biphenyl-hydroxylations, EtoC de-ethylation and absorption maximum of reduced CO-cytochrome P-450. TCPOBOP, as an inducer, was less "specific" than PB: total metabolic rates were excessively increased due to microsomal protein (1.5 times) and cytochrome P-450 (4 times) augmentation, whereas cytochrome P-450-related metabolic rates were less increased than those after PB. Thus TCPOBOP does not seem to be as similar to PB as was suggested in the first description of its inducing potency.

Animals↗

Ganglion cells in the frog retina: discriminant analysis of histological classes.

Neurons in the ganglion cell layer were studied in Golgi-stained flat-mounted frog (Rana temporaria) retinas. Complementary data were obtained from methylene blue- and HRP-stained retinas. On the basis of qualitative criteria, 55 neurons were ordered into six groups, one class of amacrine cell (A1) and five classes of ganglion cells (G1-G5). A discriminant function analysis based on seven morphological variables resulted in a separation of the cell classes in the space of three axes. The A1 cells are small axonless neurons with knotty and dense dendritic trees. The G1 cells are also small, and apparently very numerous, while the G2 cells are medium-sized neurons with two loose dendritic layers, one vitreal and another (less conspicuous) scleral. The rest of the cells are medium-sized to large neurons with sturdy primary dendrites and more distinct dendritic layers, which in some cells (G3) spread both sclerally and vitreally, in other cells in a single either scleral (G4) or vitreal (G5) layer. The relation between our data and the classification of frog ganglion cells recently presented by Frank and Hollyfield is discussed at length, and in that context problems related to statistical classifications are dealt with. A hypothetical identification of the morphological types with the functional cell classes studied in the Helsinki laboratory is discussed.

Animals↗

[Relation of the family physician to his patient and evaluation of the patient: aspects of willingness to comply?].

This study was conducted to examine the influence of general practitioner's behavior (i.e. the patient-centered behavior) on patients' inferences as well as on patient's compliance. We demonstrated in a simulation study as well as in an inquiry of patients in general practice that doctor's competence, interest and sympathy are estimated higher, if the doctor's behavior is patient-centered. Significant single results are reported in respect to patients' knowledge, their inference on the state of the disease and on the convalescence. Patient-centered behavior did not have a direct influence on patients' compliance.

Adult↗

Low retinal noise in animals with low body temperature allows high visual sensitivity.

The weakest pulse of light a human can detect sends about 100 photons through the pupil and produces 10-20 rhodopsin isomerizations in a small retinal area. It has been postulated that we cannot see single photons because of a retinal noise arising from randomly occurring thermal isomerizations. Direct recordings have since demonstrated the existence of electrical 'dark' rod events indistinguishable from photoisomerization signals. Their mean rate of occurrence is roughly consistent with the 'dark light' in psychophysical threshold experiments, and their thermal parameters justify an identification with thermal isomerizations. In the retina of amphibians, a small proportion of sensitive ganglion cells have a performance-limiting noise that is low enough to be well accounted for by these events. Here we study the performance of dark-adapted toads and frogs and show that the performance limit of visually guided behaviour is also set by thermal isomerizations. As visual sensitivity limited by thermal events should rise when the temperature falls, poikilothermous vertebrates living at low temperatures should then reach light sensitivities unattainable by mammals and birds with optical factors equal. Comparison of different species at different temperatures shows a correlation between absolute threshold intensities and estimated thermal isomerization rates in the retina.

Animals↗

Ganglion cell performance at absolute threshold in toad retina: effects of dark events in rods.

1. The performance of ganglion cells in detecting flashes of light near the absolute threshold was studied in an isolated eye-cup preparation of toad retina. Retinal ganglion cells, through which all visual information from the rods must flow to the brain, are in a key position for evaluating the still unproven hypothesis that the absolute light sensitivity is limited by rod noise (Barlow, 1956). 2. The dark-adapted threshold intensity for these cells, which were selected on the basis of their high sensitivity, averaged 0.029 Rh* flash-1 (range 0.008-0.062), where Rh* signifies one photoisomerization per rod. On average, 46 photoisomerizations were needed per receptive field per flash to evoke a threshold response (range 16-84). 3. In the threshold region, frequency of responses versus mean flash intensity was determined. Threshold performance could be described by theoretical frequency of response curves, allowing intrinsic noise to be estimated in terms of an equivalent rate of photoisomerization-like (dark) events. In two completely characterized cells the rate of dark events corresponded to 0.03 and 0.06 Rh*DS-1, where Rh*D signifies one dark event per rod. 4. Threshold elevations produced by dim backgrounds were studied. The results of these experiments are consistent with a dark event rate equivalent to 0.046 Rh*DS-1, or 0.037 Rh*DS-1 after correcting for a probable decrease in summation time. 5. The rate of actual dark events (0.028 Rh*DS-1, 20 degrees C) measured in Bufo rods (Baylor, Lamb & Yau, 1980) is close to the equivalent rates determined here. Thus, for the ganglion cells signalling the dimmest lights, the dark events in rods appear to be the most significant intrinsic retinal noise source limiting detection.

Action Potentials↗

Retinal noise, the performance of retinal ganglion cells, and visual sensitivity in the dark-adapted frog.

By extracellular recording of action potentials from single ganglion cells in the dark-adapted eyecup of the frog, we show that four different characteristics of a cell, the frequency-of-response function, the variability in responses to low suprathreshold stimulation, the maintained discharge, and the susceptibility to desensitization by weak background lights, all seem to reflect the same intrinsic noise. In a small fraction of reliable cells this noise is close to that expected from dark isomerizationlike events in rods, but it differs by orders of magnitude between cells. We have also determined a threshold intensity for the phototactic jumping behavior of frogs in darkness. This intensity, while below the human threshold, is still above the threshold for an appreciable fraction of frog retinal ganglion cells, and the sensitivity of this visually guided behavior at 16 degrees C is not strictly limited by dark events in rods.

Animals↗

Chromatic properties of the retinal afferents in the thalamus and the tectum of the frog (Rana temporaria).

In order to clarify physiological mechanisms underlying colour-specific visually guided behaviour, we measured spectral sensitivities of On-fibres projecting to the thalamus and class 2 and 3 fibres passing to tectum opticum. In addition we recorded responses of these fibres to moving coloured papers with known spectral reflectancies. The latter method, here called paper colourimetry, allowed us to change the relative stimulations of the blue-, green- and red-sensitive photoreceptors in any direction desired. Under the photopic conditions used the tectal fibres were driven exclusively by red-sensitive receptors, while the thalamic fibres received strong On-inputs from both red- and blue-sensitive receptors. Due to a partly antagonistic interaction between these inputs the On-fibres acted in a dichromatic way, responding with specific extended low-frequency discharges to all relative increases in blue receptor stimulation, e.g. to a great reduction in red stimulation combined with unchanged blue stimulation. Thus they have functional characteristics which could serve a visual system showing colour constancy.

Action Potentials↗

Longitudinal spread of adaptation in the rods of the frog's retina.

1. The stimulus-response function of the red rods in the retina of the common frog (Rana temporaria) was determined in different adaptational states by measuring aspartate-isolated receptor responses. 2. Flash stimuli, background adaptations and bleaches were delivered through the same optical channel forming an oblique light-beam striking the receptor side of the isolated and flat-mounted retina at an angle of 10 degrees. 3. When the light was blue-green and optimally polarized the absorbance of the receptor layer was about 2, from which follows that 70-80% of the light was absorbed in the distal third of the rod outer segments, i.e. the exposure was local. Homogeneous exposures of the whole rod outer segments were obtained with orange and red lights. 4. Combinations of homogeneous and local stimuli with homogeneous and local adaptations were used to investigate the longitudinal spread of background, intermediate and opsin adaptation, i.e. the sensitivity-reducing effect of a background light, and the transient and permanent sensitivity losses following a bleach isomerizing 3.5-26% (usually 10%) of the rhodopsin in the retina. 5. The results obtained were related to predictions based both on the assumption that the adaptation effects spread longitudinally within the rod outer segments and the assumption that they are strictly confined to the disks absorbing the adapting lights. 6. These comparisons reveal that all three types of adaptation spread longitudinally. It is for instance clear that the sensitivity loss observed with homogeneous stimuli and local adaptation (as compared to homogeneous adaptation) is larger than that predicted by the non-spreading hypothesis. 7. The longitudinal spread of background adaptation is largely finished within 10 sec after turning on the background light, while an efficient spread of the intermediate adaptation effect may require minutes. 8. A background light decreasing the sensitivity by about one log unit decreases the time from flash to response maximum from 5 to 1 sec (small responses). Corresponding opsin adaptation effects are accompanied by less dramatic changes in response kinetics. 9. Independent of adaptation type - homogeneous or local, background, intermediate or opsin - it was found that local stimuli are less efficient that homogeneous stimuli in light-adapted retinae. This effect can be explained assuming that the sensitivity-reducing effects are pronounced in the distal than in the proximal parts of the rod outer segments. 10. The opsin adaptation effect following 10% local bleaches decreases the sensitivity to both homogeneous and local stimuli 2-3 times more than corresponding homogeneous bleaches. This means that the strength of the opsin effect is not related to the average percentage bleached but to the fraction bleached in the distal part of the rod, or generally to the fraction bleached in the most affected region. 11...

Adaptation, Ocular↗