Breathing through a diving snorkel; theory and experiment of air flow resistance and cost of breathing.

The snorkel allows a surface swimmer to observe the underwater world through the face mask without being disturbed by inhaling. The effect of a snorkel on breathing resistance and cost is widely held to be substantial. This study aims to model these parameters and to measure indirectly the actual increases. Further, resistances of differing designs and dimensions were assessed and recommendations were made concerning use and choice. Maximal voluntary ventilation in 12 seconds (MVV12) was measured in 19 volunteers seated on dry land with and without a classic J-type snorkel (inner diameter 20.5 mm). The extra and total resistances and costs were calculated using the MVV12 data and using estimated airways resistance extrapolated from subject's demography and spirometric literature data. MVV12 measurements with snorkel showed a minute volume of 152 ±38 L∙min-1, 6.0 ±3.7% lower than without snorkel (p = 7.0x10-6). The theoretical MVV12, calculated from snorkel and airways resistances, decreased by 3.2%. Experimental total breathing resistance (457±83 Pa∙s∙L-1) was 6.5 ± 3.2% higher than without snorkel (p = 2.6x10-7), but the total mechanical breathing cost was unaffected by the snorkel (13.58 Watts with; 13.64 Watts without). Divers' estimations of resistance increase were exaggerated (8.8% at rest, 23% swimming). Classical J-type snorkels with an inner diameter ≥19.5 mm add 3-16% resistance . There is no risk of hypercapnia. Scuba divers are recommended to use their snorkel to breathe more comfortably on the surface. It is recommended the snorkel be made a mandatory safety accessory. The best multipurpose snorkel (19-21 mm) has no top appendages and no water release valve.

Mental stress may cause high gas consumption and accelerated heart rate in fast-descending divers.

The descent is a critical part of a dive, both physically and mentally. Pulmonary ventilation, expressed as respiratory minute volume (RMV) and heart rate (HR) were recorded during fast and slow descents to 35 meters (m) in open water while breathing compressed air, and during swimming horizontally at moderate velocity at an 11-m depth. Values of both types of descents were compared with reference values recorded at 11 m, the "plateau" phase, halfway through the 35-m dives. It is hypothesized that the "slow-descent" and "plateau-phase" values will be less than 'fast-descent values. Depth, cylinder pressure, water temperature and HR were recorded with a dive computer yielding time-averaged means (mRMV and mHR) for the descent and for plateau. Of the 18 divers included, 16 performed the fast descents and 11 made the slow descents. The fast descents (23 m·min-1 vertically), performed with 0-8 fin kicks, yielded mRMVDescent=28 ambient L (aL)·min-1, which is 82% higher (P⟨0.001) than mRMVPlateau of 15 aL·min-1. Further, mHRDescent was121 beats·min-1 23% higher (P⟨0.001), than mHRPlateau of 100 bpm. Slow descents (2.4 m·min-1 vertically) yielded 17 aL·min-1 with mHR=101 beats·min-1, values only slightly higher than at Plateau. The 11-m dive (swimming horizontally) yielded 24 m·min-1 with 32 fin kicks·min-1, mRMV=35 aL·min-1 and mHR=115 beats·min-1. Fast descents cause a higher RMV and HR that cannot be explained by physiology alone. Presumably mental stress is a main contributor. For dives deeper than 20 m, a descent velocity of 10 m·min-1 is recommended to reduce cardiac stress, in particular for older divers.

Venous gas embolism after an open-water air dive and identical repetitive dive.

Decompression tables indicate that a repetitive dive to the same depth as a first dive should be shortened to obtain the same probability of occurrence of decompression sickness (pDCS). Repetition protocols are based on small numbers, a reason for re-examination. Since venous gas embolism (VGE) and pDCS are related, one would expect a higher bubble grade (BG) of VGE after the repetitive dive without reducing bottom time. BGs were determined in 28 divers after a first and an identical repetitive air dive of 40 minutes to 20 meters of sea water. Doppler BG scores were transformed to log number of bubbles/cm2 (logB) to allow numerical analysis. With a previously published model (Model2), pDCS was calculated for the first dive and for both dives together. From pDCS, theoretical logBs were estimated with a pDCS-to-logB model constructed from literature data. However, pDCS the second dive was provided using conditional probability. This was achieved in Model2 and indirectly via tissue saturations. The combination of both models shows a significant increase of logB after the second dive, whereas the measurements showed an unexpected lower logB. These differences between measurements and model expectations are significant (p-values < 0.01). A reason for this discrepancy is uncertain. The most likely speculation would be that the divers, who were relatively old, did not perform physical activity for some days before the first dive. Our data suggest that, wisely, the first dive after a period of no exercise should be performed conservatively, particularly for older divers.

Temporal and spatial congruence of components of motion-onset evoked responses investigated by whole-head magneto-electroencephalography.

Motion-onset related components in averaged whole head co-recorded MEG and EEG responses of 24 adults to a low-contrast checkerboard pattern were studied. The aims were to identify these components, to characterize quantitatively their maps and to localize the underlying sources by equivalent-current-dipole (ECD) analyses with a spherical head model.After a weak P1, a large start-elicited negativity arises, comprising the novel N2a (occipital positive and parieto-central negative, peak-latency 141 ms) and the N2 like N2b (bilateral parieto-temporal, 175 ms) component. It is followed by a large positive stop-related component, P2 (156 ms after motion-offset). The corresponding MEG components N2am and N2bm showed bilateral dipole fields with considerable overlap. P1m has a single dipole field around the midline. N2a(m) and N2b(m) can be modelled with two bilateral ECDs with significant different locations. The study shows that accurate mapping and ECD analyses can distinguish two neighbouring areas of the visual cortex, 21+/-4 (SE) mm separated, which activities are reflected in both spatio-temporally closely related N2(m) components. N2a(m) and N2b(m) originate in the extrastriate cortex, possibly close to or in V3/V3A and MT/V5 respectively. Motion-evoked activity in (near) V3/V3A is novel on the basis of EEG data.

Conduction velocity compensation for afferent fiber length in the trunk lateral line of the trout.

The trout lateral line contains about 122 trunk scales and is tens of centimeters long. The difference in time of arrival in the hindbrain of simultaneously elicited afferent responses from the neuromasts is unknown. Propagation times of single-fiber afferent responses to water motion revealed that their mean conduction velocity was lowest (13 m s(-1)) for fibers innervating a neuromast close to the operculum and highest (33 m s(-1)) for those close to the tail. Histological examination showed that the nerve close to the operculum comprises about 500 afferents and that this number diminishes from operculum to tail with 4/scale. The mean diameter of the fibers changed from 12.5 micro m at the operculum to 7.5 micro m at three-quarters of the operculum-to-tail distance. Comparison of the distributions of diameters indicated that the fibers are tapered with the thick end towards the operculum. A model was developed describing the relationship between tapering and local conduction velocity. We conclude that simultaneous stimulation of all trunk neuromasts causes an average time-of-arrival difference in the hindbrain of 2.8 ms, which is 2.1 times less than the difference expected with a distance-independent conduction velocity. This suggests that tapering and velocity compensation are relevant for central processing of lateral line information.

Transient and maintained changes of the spontaneous occipital EEG during acute systemic hypoxia.

BACKGROUND: It has been reported that during systemic hypoxia the spectral power of spontaneous EEG (closed eyes) increases, except for the power in the alpha band, which decreases. HYPOTHESIS: It is surmised that after a sudden decrease in inspired oxygen, the EEG parameters and SaO2 show similar temporal characteristics. METHODS: Normobaric acute hypoxia in 14 healthy subjects was evoked by breathing a hypoxic gas. Frequency spectra were calculated from occipital recordings and quantified together with the parameters of the spectral alpha peak. RESULTS: During the first 20 min of exposure, SaO2 decreases exponentially. With eyes open, the frequency bands show a transient increase in power (110-160%). The peak-time of the transient is shorter for the alpha and beta1 bands (4.5 min) than for the delta2 and theta bands (7 min). After the transient, the bands reach sustained power levels ca. 65% higher than the control value. The frequency of the alpha peak decreases with 0.61 +/- 0.16 Hz (mean +/- SE). In contrast to the eyes-closed condition, the peak amplitude increases (41 +/- 12%). CONCLUSIONS: During systemic hypoxia many characteristics of the spontaneous EEG with open eyes differ from those under normoxia, and alpha activity deviates strongly from that during hypoxia with closed eyes. During the first 20 min of exposure changes in power are most prominent and do not follow the change of SaO2. Monitoring total EEG power with open eyes, SaO2 and respiration gives a good impression of the hypoxia-related neurophysiological state of the subject.

Voluntary breath holding affects spontaneous brain activity measured by magnetoencephalography.

Spontaneous brain activity was measured by multichannel magnetoencephalography (MEG) during voluntary breath holds. Significant changes in the activity are limited to the alpha rhythm: 0.25 Hz frequency increase and narrowing of the peak. The area of alpha activity shifts slightly toward (fronto-) temporal. The topography of other rhythms is unaffected by breath holding. Electroencephalographic (EEG; 36 channels maximal) recordings generally made simultaneously with the MEG recordings show similar effects. However, EEG was inadequate to reveal the small topographic differences. Systemic hypercapnia caused by a long breath hold is unlikely to play an important role in producing the observed effects.

Neuronal encoding of sound direction in the auditory midbrain of the rainbow trout.

Acoustical stimulation causes displacement of the sensory hair cells relative to the otoliths of the fish inner ear. The swimbladder, transforming the acoustical pressure component into displacement, also contributes to the displacement of the hair cells. Together, this (generally) yields elliptical displacement orbits. Alternative mechanisms of fish directional hearing are proposed by the phase model, which requires a temporal neuronal code, and by the orbit model, which requires a spike density code. We investigated whether the directional selective response of auditory neurons in the midbrain torus semicircularis (TS; homologous to the inferior colliculus) is based on spike density and/or temporal encoding. Rainbow trout were mounted on top of a vibrating table that was driven in the horizontal plane to simulate sound source direction. Rectilinear and elliptical (or circular) motion was applied at 172 Hz. Generally, responses to rectilinear and elliptical/circular stimuli (irrespective of direction of revolution) were the same. The response of auditory neurons was either directionally selective (DS units, n = 85) or not (non-DS units, n = 106). The average spontaneous discharge rate of DS units was less than that of non-DS units. Most DS units (70%) had spontaneous activities < 1 spike per second. Response latencies (mode at 18 ms) were similar for both types of units. The response of DS units is transient (19%), sustained (34%), or mixed (47%). The response of 75% of the DS units synchronized to stimulus frequency, whereas just 23% of the non-DS responses did. Synchronized responses were measured at stimulus amplitudes as low as 0.5 nm (at 172 Hz), which is much lower than for auditory neurons in the medulla of the trout, suggesting strong convergence of VIIIth nerve input. The instant of firing of 42% of the units was independent of stimulus direction (shift <15 degrees), but for the other units, a direction dependent phase shift was observed. In the medial TS spatial tuning of DS units is in the rostrocaudal direction, whereas in the lateral TS all preferred directions are present. On average, medial DS units have a broader directional selectivity range, are less often synchronized, and show a smaller shift of the instant of firing as a function of stimulus direction than lateral DS units. DS response characteristics are discussed in relation to different hypotheses. We conclude that the results are more in favor of the phase model.

Effect of dysoxia and moderate air-hyperbarism on red-green color sensitivity.

The effect of acute air-hyperbarism (maximal 520 kPa), normobaric low and high FIO2 levels (minimal 0.1, maximal 1.0) and hyperbaric oxygen (HBO2) on the red-green sensitivity ratio (rgSR) and on color discrimination for foveal vision were studied. Effects were quantified by measuring the red-green flicker (16 Hz) fusion point for normals with the Oscar tester (Am J Optom Physiol Opt 1983; 60:892-901). Color discrimination was examined with the Lanthony's Desaturated 15 Hue test After 15-20 min of exposure rgSR is enhanced 4% (relatively increased red sensitivity) by normobaric acute hypoxia, and reduced 4% by normobaric hyperoxia (FIO2 = 1.0), but HBO2 gives a smaller reduction, and air-hyperbarism (FIO2 = 0.21) has no effect. Hypercapnia (increased FICO2), normobaric hypoxia (reduced FIO2), and HBO2 increase the duration of the Lanthony's test about 20-40%, but the number of errors were practically unchanged. The reduced effect during HBO2 upon rgSR is attributed to an opposing effect of hypercapnia. The absence of an effect during air-hyperbarism is probably due to a suppression by nitrogen of the effect of high PO2. In general, during moderate air-hyperbarism and HBO2 color vision seems to be normal, but evaluation of the colored scene is probably less stable and slightly slower.

Mapping of sound direction in the trout lower midbrain.

In the trout lower midbrain 35% of the auditory neurons are directionally selective (DS). Most of these neurons have a higher directional selectivity than the sensory hair cells. DS units and non-DS units occur in vertical clusters, with the former more dorsally. The direction of preference is topographically mapped. Apparently, auditory space mapping is a common feature in the midbrain of vertebrates.

Velocity- and acceleration-sensitive units in the trunk lateral line of the trout.

1. The two main types of lateral line organs of lower vertebrates are the superficial neuromasts (SN), with a cupula that protrudes in the surrounding water, and the canal neuromasts (CN), located in the lateral line canal. The scales of the trunk lateral line canal of fish contain SNs as well as CNs. In this study, we examine whether there exist two functional classes of afferent fibers in the trunk lateral line nerve of the rainbow trout that can be attributed to the SNs and CNs. 2. The response properties of the afferent fibers in the trunk lateral line nerve have been determined during stimulation with sinusoidally varying water motion generated by a small vibrating sphere. Linear frequency response analysis revealed the presence of two distinct populations of afferent fibers in the lateral line nerve. The fibers belonging to the two populations showed significant differences in the frequency at which the sensitivity was maximal, the low-frequency response slope and the low-frequency asymptotic phase angle. 3. One population of fibers has a maximum sensitivity at 36 +/- 13 (SD) Hz (n = 22) and responds up to this frequency to water velocity. The low-frequency slope of the frequency response of these fibers was 20 +/- 3 (SD) dB/decade and the low-frequency phase lead was 121 +/- 11 degrees (mean +/- SD), both with respect to sphere displacement. The fibers of the other population have a maximum sensitivity at 93 +/- 14 (SD) Hz (n = 12) and respond up to this frequency to water acceleration. The low-frequency slope of these fibers was 35 +/- 5 (SD) dB/decade, and the low-frequency phase lead was 188 +/- 13 degrees (mean +/- SD). 4. Analysis of the stochastic properties of the spontaneous activity of both types of fibers revealed that the mean firing rate of the fibers responding to water velocity (26 +/- 12 spikes/s, mean +/- SD; n = 22) was significantly higher than that of the fibers responding to acceleration (36 +/- 11 spikes/s, mean +/- SD; n = 12). The other statistical properties of the spontaneous activity were found to be indistinguishable. 5. From comparison of the results with the available quantitative data on frequency responses of lateral line organs in other species, it has been concluded that the fibers responding (< or = 40 Hz) to water velocity innervate SNs and that the fibers responding (< or = 90 Hz) to water acceleration innervate CNs.(ABSTRACT TRUNCATED AT 400 WORDS)

Contrast sensitivity of air-breathing nonprofessional scuba divers at a depth of 40 meters.

Photopic contrast sensitivity of air-breathing scuba divers was measured with a translucent test pattern at depths up to 40 m. The pattern was composed of sine wave gratings with spatial frequency and contrast changing logarithmically. The spatial transfer characteristics were measured at various depths under controlled optical conditions in seawater and in fresh water. Analysis indicates that the visual contrast sensitivity, and therefore probably also acuity, of sport divers is not affected up to depths of 40 m. This holds under ideal as well as poor diving conditions.

The pattern of trunk lateral line afferents and efferents in the rainbow trout (Salmo gairdneri).

The primary projections of the mechanosensory posterior lateral line nerve of the rainbow trout Salmo gairdneri, a teleost without lateral line specializations, were studied by applying horseradish peroxidase (HRP). The afferents project nearly exclusively to the nucleus medialis and a small nucleus caudalis in the hindbrain, and to the eminentia granularis. In addition, there is a sparse projection to the cerebellum, but a projection to the magnocellular nucleus is lacking. The afferent projection to the lateral part of the eminentia granularis is more dense than the projection to the medial nucleus, as shown by image processing of the HRP labeling. The efferent fibers originate from a bilateral pair of large octavolateral nuclei in the hindbrain. Characteristic for these nuclei are their large, fusiform and bipolar neurons. Another specific feature is the oblique orientation of the somata, with the main dendritic shaft coursing ventrolaterally and the axonal shaft coursing dorsomedially. The axons of the efferent neurons bifurcate. The shape, orientation and size of the efferent somata are the same in the entire efferent nucleus. In contrast to other teleosts, the occurrence of efferent somata found contralaterally (21%) is substantial. The part of the octavolateral efferent nucleus innervating the neuromasts of the trunk is estimated to contain at most 150 cell bodies unilaterally.

A model for directional and distance hearing in swimbladder-bearing fish based on the displacement orbits of the hair cells.

It is known that teleosts, without Weberian ossicles but with a swimbladder, can detect the direction of and, under appropriate conditions, the distance to a sound source [e.g., Schuijf and Hawkins, Nature 302, 143-144 (1983)]. It is hypothesized here that the underlying mechanism is the analysis of the parameters of the elliptical movement of the hair cells with respect to the otoliths. This movement results from the displacement wave impinging directly upon the labyrinth and the response displacement wave reradiated by the swimbladder. For a given swimbladder geometry, given the positions of the maculae of both labyrinths with respect to the swimbladder and the damping of the swimbladder, the displacement orbits of the maculae can be calculated [de Munck and Schellart, J. Acoust. Soc. Am. 81, 556-560 (1987)]. These calculations were made for the cod and the trout with the frequency, direction, and distance between the fish and the sound source as parameters with the source within the same horizontal plane as the fish. The orbit model predicts that the utriculus has the most strategic position to detect direction and distance of such a sound source. Moreover, the model predicts that this could basically be done monaurally. A hypothesis is proposed to describe how the utricular system analyzes the orbit parameters. The model is evaluated in relation to the results of behavioral experiments described in the literature.

A model for the nearfield acoustics of the fish swimbladder and its relevance for directional hearing.

It is a well-known fact that fish can determine the direction of an underwater sound source. For bony fish without Weberian ossicles the mechanism proposed here is the analysis of the elliptical movement of the otolith macula relative to the otolith in case a pure tone is emitted. This movement results from the superposition of the direct displacement wave and the displacement wave scattered by the swimbladder acting as a damped harmonic oscillator. A derivation of a closed analytical expression of the scattered wave is given for a prolate spheroidal air bubble (representing the swimbladder) in water. The elliptical displacement orbits of the surrounding medium were calculated. A general numerical approach is given for irregular gas bodies.

Computerized pattern recognition used for grain counting in high resolution autoradiographs with low grain densities.

Using a video-image system coupled to a minicomputer with commercial image handling software, autoradiographic grains displayed in dark-field are counted with a fast (ca. 3.5 min for 120,000 microns 2) and reliable (false scores less than 5%) grain-recognizing FORTRAN program executed in the users memory. The grain counts are printed in a raster of adjustable size overlying a bright-field image, so that the counts can be related directly to the underlying histological structures.