Decompression illness: a comprehensive overview.

Decompression illness is a collective term for two maladies (decompression sickness [DCS] and arterial gas embolism [AGE]) that may arise during or after surfacing from compressed gas diving. Bubbles are the presumed primary vector of injury in both disorders, but the respective sources of bubbles are distinct. In DCS bubbles form primarily from inert gas that becomes dissolved in tissues over the course of a compressed gas dive. During and after ascent ('decompression'), if the pressure of this dissolved gas exceeds ambient pressure small bubbles may form in the extravascular space or in tissue blood vessels, thereafter passing into the venous circulation. In AGE, if compressed gas is trapped in the lungs during ascent, pulmonary barotrauma may introduce bubbles directly into the pulmonary veins and thence to the systemic arterial circulation. In both settings, bubbles may provoke ischaemic, inflammatory, and mechanical injury to tissues and their associated microcirculation. While AGE typically presents with stroke-like manifestations referrable to cerebral involvement, DCS can affect many organs including the brain, spinal cord, inner ear, musculoskeletal tissue, cardiopulmonary system and skin, and potential symptoms are protean in both nature and severity. This comprehensive overview addresses the pathophysiology, manifestations, prevention and treatment of both disorders.

Oxy-Inflammation in Humans during Underwater Activities.

Underwater activities are characterized by an imbalance between reactive oxygen/nitrogen species (RONS) and antioxidant mechanisms, which can be associated with an inflammatory response, depending on O2 availability. This review explores the oxidative stress mechanisms and related inflammation status (Oxy-Inflammation) in underwater activities such as breath-hold (BH) diving, Self-Contained Underwater Breathing Apparatus (SCUBA) and Closed-Circuit Rebreather (CCR) diving, and saturation diving. Divers are exposed to hypoxic and hyperoxic conditions, amplified by environmental conditions, hyperbaric pressure, cold water, different types of breathing gases, and air/non-air mixtures. The "diving response", including physiological adaptation, cardiovascular stress, increased arterial blood pressure, peripheral vasoconstriction, altered blood gas values, and risk of bubble formation during decompression, are reported.

A novel method for tracking nitrogen kinetics in vivo under hyperbaric conditions using radioactive nitrogen-13 gas and positron emission tomography.

Decompression sickness (DCS) is caused by gaseous nitrogen dissolved in tissues forming bubbles during decompression. To date, no method exists to identify nitrogen within tissues, but with advances in positron-emission tomography (PET) technology, it may be possible to track gaseous radionuclides into tissues. We aimed to develop a method to track nitrogen movement in vivo and under hyperbaric pressure that could then be used to further our understanding of DCS using nitrogen-13 (13N2). A single anesthetized female Sprague-Dawley rat was exposed to 625 kPa, composed of air, isoflurane, and 13N2 for 10 min. The PET scanner recorded 13N2 during the hyperbaric exposure with energy windows of 250-750 keV. The PET showed an increase in 13N2 concentration in the lung, heart, and abdominal regions, which all reached a plateau after ∼4 min. This showed that it is possible to gain noninvasive in vivo measurements of nitrogen kinetics through the body while at hyperbaric pressures. Tissue samples showed radioactivity above background levels in the blood, brain, liver, femur, and thigh muscle when assessed using a γ counter. The method can be used to evaluate an array of challenges to our understanding of decompression physiology by quantifying nitrogen load through γ counts of 13N2, and signal intensity of the PET. Further development of the method will improve the specificity of the measured outcomes, and enable it to be used with larger mammals, including humans.NEW & NOTEWORTHY This article describes a method for the in vivo quantification and tracking of nitrogen through the mammalian body whilst exposed to hyperbaric pressure. The method has the potential to further our understanding of decompression sickness, and quantitatively evaluate the effectiveness of both the treatment and prevention of decompression sickness.

Stress biomarker changes following a series of repeated static and dynamic apneas in non-divers.

PURPOSE: This study examined the magnitude of physiological strain imposed by repeated maximal static and dynamic apneas through assessing a panel of stress-related biomarkers. METHODS: Eleven healthy men performed on three separate occasions (≥72-h apart): a series of five repeated maximal (i) static (STA) or (ii) dynamic apneas (DYN) or (iii) a static eupneic protocol (CTL). Venous blood samples were drawn at 30, 90, and 180-min after each protocol to determine ischaemia modified albumin (IMA), neuron-specific enolase (NSE), myoglobin, and high sensitivity cardiac troponin T (hscTnT) concentrations. RESULTS: IMA was elevated after the apnoeic interventions (STA,+86%;DYN,+332%,p ≤ 0.047) but not CTL (p = 0.385). Myoglobin was higher than baseline (23.6 ± 3.9 ng/mL) 30-min post DYN (+70%,38.8 ± 13.3 ng/mL,p = 0.030). A greater myoglobin release was recorded in DYN compared with STA and CTL (p ≤ 0.035). No changes were observed in NSE (p = 0.207) or hscTnT (p = 0.274). CONCLUSIONS: Five repeated maximal DYN led to a greater muscle injury compared with STA but neither elicited myocardial injury or neuronal-parenchymal damage.

Blood oxygen transport and depletion in diving emperor penguins.

Oxygen store management underlies dive performance and is dependent on the slow heart rate and peripheral vasoconstriction of the dive response to control tissue blood flow and oxygen uptake. Prior research has revealed two major patterns of muscle myoglobin saturation profiles during dives of emperor penguins. In Type A profiles, myoglobin desaturated rapidly, consistent with minimal muscle blood flow and low tissue oxygen uptake. Type B profiles, with fluctuating and slower declines in myoglobin saturation, were consistent with variable tissue blood flow patterns and tissue oxygen uptake during dives. We examined arterial and venous blood oxygen profiles to evaluate blood oxygen extraction and found two primary patterns of venous hemoglobin desaturation that complemented corresponding myoglobin saturation profiles. Type A venous profiles had a hemoglobin saturation that (a) increased/plateaued for most of a dive's duration, (b) only declined during the latter stages of ascent, and (c) often became arterialized [arterio-venous (a-v) shunting]. In Type B venous profiles, variable but progressive hemoglobin desaturation profiles were interrupted by inflections in the profile that were consistent with fluctuating tissue blood flow and oxygen uptake. End-of-dive saturation of arterial and Type A venous hemoglobin saturation profiles were not significantly different, but did differ from those of Type B venous profiles. These findings provide further support that the dive response of emperor penguins is a spectrum of cardiac and vascular components (including a-v shunting) that are dependent on the nature and demands of a given dive and even of a given segment of a dive.

First Real-Life Data on the Diving Response in Healthy Children.

Swimming and diving are popular recreational activities, representing an effective option in maintaining and improving cardiovascular fitness in healthy people. To date, only little is known about the cardiovascular adaption to submersion in children. This study was conducted to improve an understanding thereof. We used a stepwise apnea protocol with apnea at rest, apnea with facial immersion, and at last apnea during whole body submersion. Continuous measurement of heart rate, oxygen saturation, and peripheral resistance index was done. Physiologic data and analysis of influencing factors on heart rate, oxygen saturation, and peripheral vascular tone response are reported. The current study presents the first data of physiologic diving response in children. Data showed that facial or whole body submersion leads to a major drop in heart rate, and increase of peripheral resistance, while the oxygen saturation seems to be unaffected by static apnea in most children, with apnea times of up to 75 s without change in oxygen saturation.

Do freedivers and spearfishermen differ in local muscle oxygen saturation and anaerobic power?

BACKGROUND: Freediving is defined as an activity where athletes repetitively dive and are exposed to long efforts with limited oxygen consumption. Therefore, anaerobic features are expected to be an important facet of diving performance. This study aimed to investigate differences in anaerobic capacity and local muscle oxygenation in spearfisherman and freedivers. METHODS: The sample of participants included 17 male athletes (nine freedivers, and eight spearfishermen), with an average age of 37.0±8.8 years, training experience of 10.6±9.5 years, body mass of 82.5±9.5 kg and height of 184.2±5.7 cm. Anthropometric characteristics included: body mass, body height, seated height, and body fat percentage. Wingate anaerobic test was conducted, during which local muscle oxygenation was measured with a NIRS device (Moxy monitor). Wingate power outputs were measured (peak power [W/kg] and average power [W/kg]), together with muscle oxygenation variables (baseline oxygen saturation [%], desaturation slope [%/s], minimum oxygen saturation [%], half time recovery [s], and maximum oxygen saturation [%]). RESULTS: The differences were not obtained between freedivers and spearfisherman in power outputs (peak power (9.24±2.08 spearfisherman; 10.68±1.04 freedivers; P=0.14); average power (6.85±0.95 spearfisherman; 7.44±0.60 freedivers; P=0.15) and muscle oxygenation parameters. However, analysis of effect size showed a moderate effect in training experience (0.71), PP (0.89), AP (0.75), Desat slope mVLR (0.66), half time recovery mVLR (0.90). CONCLUSIONS: The non-existence of differences between freedivers and spearfishermen indicates similar training adaptations to the anaerobic demands. However, the results show relatively low anaerobic capacities of our divers that could serve as an incentive for the further development of these mechanisms.

Cardiorespiratory adaptations in small cetaceans and marine mammals.

The dive response, or the 'master switch of life', is probably the most studied physiological trait in marine mammals and is thought to conserve the available O2 for the heart and brain. Although generally thought to be an autonomic reflex, several studies indicate that the cardiovascular changes during diving are anticipatory and can be conditioned. The respiratory adaptations, where the aquatic breathing pattern resembles intermittent breathing in land mammals, with expiratory flow exceeding 160 litres s-1 has been measured in cetaceans, and where exposure to extreme pressures results in alveolar collapse (atelectasis) and recruitment upon ascent. Cardiorespiratory coupling, where breathing results in changes in heart rate, has been proposed to improve gas exchange. Cardiorespiratory coupling has also been reported in marine mammals, and in the bottlenose dolphin, where it alters both heart rate and stroke volume. When accounting for this respiratory dependence on cardiac function, several studies have reported an absence of a diving-related bradycardia except during dives that exceed the duration that is fuelled by aerobic metabolism. This review summarizes what is known about the respiratory physiology in marine mammals, with a special focus on cetaceans. The cardiorespiratory coupling is reviewed, and the selective gas exchange hypothesis is summarized, which provides a testable mechanism for how breath-hold diving vertebrates may actively prevent uptake of N2 during routine dives, and how stress results in failure of this mechanism, which results in diving-related gas emboli.

Physiological responses during static apnoea efforts in elite and novice breath-hold divers before and after two weeks of dry apnoea training.

This study examined the effect of breath-hold (BH) training on apnoeic performance in novice BH divers (NBH:n = 10) and compared them with data from elite BH divers (EBH:n = 11). Both groups performed 5-maximal BHs (PRE). The NBH group repeated this protocol after two weeks of BH training (POST). The NBH group during BH efforts significantly increased red blood cell concentration (4.56 ± 0.16Mio/µl) by 5.06%, hemoglobin oxygen saturation steady state duration (110.32 ± 29.84 s) by 15.48%, and breath-hold time (BHT:144.19 ± 47.35 s) by 33.77%, primarily due to a 59.70% increase in struggle phase (71.85 ± 30.89 s), in POST. EBH group exhibited longer BHT (283.95 ± 36.93 s) and struggle-phase (150.10 ± 34.69 s) than NBH (POST). Elite divers recorded a higher peak MAP (153.18 ± 12.28 mmHg) compared to novices (PRE:123.70 ± 15.65 mmHg, POST:128.30 ± 19.16 mmHg), suggesting that a higher peak MAP is associated with a better BHT. The concurrent abrupt increase of diaphragmatic activity and MAP, seen only in the EBH group, suggests a potential interaction. Additionally, apnoea training increases red blood cells concentration in repeated apnoea efforts and increases BH stamina.

Breath-holding as model for the evaluation of EEG signal during respiratory distress.

PURPOSE: Research describes the existence of a relationship between cortical activity and the regulation of bulbar respiratory centers through the evaluation of the electroencephalographic (EEG) signal during respiratory challenges. For example, we found evidences of a reduction in the frequency of the EEG (alpha band) in both divers and non-divers during apnea tests. For instance, this reduction was more prominent in divers due to the greater physiological disturbance resulting from longer apnea time. However, little is known about EEG adaptations during tests of maximal apnea, a test that voluntarily stops breathing and induces dyspnea. RESULTS: Through this mini-review, we verified that a protocol of successive apneas triggers a significant increase in the maximum apnea time and we hypothesized that successive maximal apnea test could be a powerful model for the study of cortical activity during respiratory distress. CONCLUSION: Dyspnea is a multifactorial symptom and we believe that performing a successive maximal apnea protocol is possible to understand some factors that determine the sensation of dyspnea through the EEG signal, especially in people not trained in apnea.

Measuring whole body inert gas wash-out.

Introduction: Quantifying inert gas wash-out is crucial to understanding the pathophysiology of decompression sickness. In this study, we developed a portable closed-circuit device for measuring inert gas wash-out and validated its precision and accuracy both with and without human subjects. Methods: We developed an exhalate monitor with sensors for volume, temperature, water vapor and oxygen. Inert gas volume was extrapolated from these inputs using the ideal gas law. The device's ability to detect volume differences while connected to a breathing machine was analysed by injecting a given gas volume eight times. One hundred and seventy-two coupled before-and-after measurements were then compared with a paired t-test. Drift in measured inert gas volume during unlabored breathing was evaluated in three subjects at rest using multilevel linear regression. A quasi-experimental cross-over study with the same subjects was conducted to evaluate the device's ability to detect inert gas changes in relation to diving interventions and simulate power. Results: The difference between the injected volume (1,996 ml) and the device's measured volume (1,986 ml) was -10 ml. The 95% confidence interval (CI) for the measured volume was 1,969 to 2,003 ml. Mean drift during a 43 min period of unlaboured breathing was -19 ml, (95% CI, -37 to -1). Our power simulation, based on a cross-over study design, determined a sample size of two subjects to detect a true mean difference of total inert gas wash-out volume of 100 ml. Conclusions: We present a portable device with acceptable precision and accuracy to measure inert gas wash-out differences that may be physiologically relevant in the pathophysiology of decompression sickness.

Pulmonary oxygen toxicity breath markers after heliox diving to 81 metres.

Pulmonary oxygen toxicity (POT), an adverse reaction to an elevated partial pressure of oxygen in the lungs, can develop as a result of prolonged hyperbaric hyperoxic conditions. Initially starting with tracheal discomfort, it results in pulmonary symptoms and ultimately lung fibrosis. Previous studies identified several volatile organic compounds (VOCs) in exhaled breath indicative of POT after various wet and dry hyperbaric hypoxic exposures, predominantly in laboratory settings. This study examined VOCs after exposures to 81 metres of seawater by three navy divers during operational heliox diving. Univariate testing did not yield significant results. However, targeted multivariate analysis of POT-associated VOCs identified significant (P = 0.004) changes of dodecane, tetradecane, octane, methylcyclohexane, and butyl acetate during the 4 h post-dive sampling period. No airway symptoms or discomfort were reported. This study demonstrates that breath sampling can be performed in the field, and VOCs indicative of oxygen toxicity are exhaled without clinical symptoms of POT, strengthening the belief that POT develops on a subclinical-to-symptomatic spectrum. However, this study was performed during an actual diving operation and therefore various confounders were introduced, which were excluded in previous laboratory studies. Future studies could focus on optimising sampling protocols for field use to ensure uniformity and reproducibility, and on establishing dose-response relationships.

Scuba diving and the stress response: considerations and recommendations for professional and recreational divers.

Scuba diving is an activity that people engage in both for recreational purposes as well as having professional, commercial, and military applications. Scuba diving has often been considered a high-risk activity but, overall, scuba diving has been shown to be a safe activity when divers participate within their experiential, physical, and psychological limits. However, increased physical and psychological stress can quickly arise during diving activities due to unexpected events and situations and may lead to the onset of panic in an unprepared diver. Dive safety is dependent on the ability of a diver to understand the primary signs of stress and panic and attempt to minimise their potential impacts on the immediate situation. The purpose of this review is to examine the stress response in divers, illustrate the role that panic plays in potential diving accidents and fatalities, and provide recommendations to both help understand and manage stress and panic in the diving community in an effort to further increase the overall safety of scuba diving across all applications.

Field physiology in the aquatic realm: ecological energetics and diving behavior provide context for elucidating patterns and deviations.

Comparative physiology has developed a rich understanding of the physiological adaptations of organisms, from microbes to megafauna. Despite extreme differences in size and a diversity of habitats, general patterns are observed in their physiological adaptations. Yet, many organisms deviate from the general patterns, providing an opportunity to understand the importance of ecology in determining the evolution of unusual adaptations. Aquatic air-breathing vertebrates provide unique study systems in which the interplay between ecology, physiology and behavior is most evident. They must perform breath-hold dives to obtain food underwater, which imposes a physiological constraint on their foraging time as they must resurface to breathe. This separation of two critical resources has led researchers to investigate these organisms' physiological adaptations and trade-offs. Addressing such questions on large marine animals is best done in the field, given the difficulty of replicating the environment of these animals in the lab. This Review examines the long history of research on diving physiology and behavior. We show how innovative technology and the careful selection of research animals have provided a holistic understanding of diving mammals' physiology, behavior and ecology. We explore the role of the aerobic diving limit, body size, oxygen stores, prey distribution and metabolism. We then identify gaps in our knowledge and suggest areas for future research, pointing out how this research will help conserve these unique animals.

The Effect of Acute Breath-Holding Exercise in Different Conditions on Cognitive Performance in Free Divers / El efecto del ejercicio agudo de contener la respiración en diferentes condiciones sobre el rendimiento cognitivo en buceadores libres

There is a growing trend towards studying human cognition in aquatic environments. At present, there exists a dearth of scholarly investigations pertaining to the immediate effects on executive functions subsequent to a solitary breath-holding training session within the cohort of professional free divers who engage in highly strenuous activities that test their physiological boundaries. The objective of this study was to investigate the immediate impact of breath-holding exercises conducted in varying environments (water and land) on the executive functions of professional diving athletes. The research comprised a sample of 18 male individuals engaged in competitive free diving. The experimental design encompassed four distinct sessions: i) an initial phase dedicated to familiarising participants with the N-back test; ii) a subsequent phase involving the collection of baseline and control measurements for the N-back test; iii) a session focused on obtaining anthropometric measurements; and iv) a final session dedicated to measuring lung capacity. The cognitive assessments were conducted subsequent to the breath-holding exercise protocol, which occurred subsequent to both the land and water sessions. The results indicated a significant difference in reaction times between breath-holding exercises conducted on land and in water (p =.021). The computation of delta values was employed to ascertain alterations in cognitive test outcomes under distinct conditions (water and land) in comparison to the control condition. The findings revealed a statistically significant decline in cognitive performance in the water condition relative to the land condition (z:-2.025, p= 0.043, r= -0.544). This study claims that the implementation of breath-holding exercise training in surface water conditions among divers could potentially result in adverse effects on executive functions. Moreover, it has been observed that the identical breath-holding exercises, when executed in terrestrial environments, exhibit a moderate enhancement of executive functions. The present study posits that the aforementioned findings will make a valuable contribution to the development of training methodologies for athletes and coaches involved in the discipline of freediving. Additionally, these findings are anticipated to offer valuable insights into the physiological well-being of divers.(AU)

Does Decreased Diffusing Capacity of the Lungs for Carbon Monoxide Constitute a Risk of Decompression Sickness in Occupational Divers?

Long-term alterations of pulmonary function (mainly decreased airway conductance and capacity of the lungs to diffuse carbon monoxide (DLCO)) have been described after hyperbaric exposures. However, whether these alterations convey a higher risk for divers' safety has never been investigated before. The purpose of the present pilot study was to assess whether decreased DLCO is associated with modifications of the physiological response to diving. In this case-control observational study, 15 "fit-to-dive" occupational divers were split into two groups according to their DLCO measurements compared to references values, either normal (control) or reduced (DLCO group). After a standardized 20 m/40 min dive in a sea water pool, the peak-flow, vascular gas emboli (VGE) grade, micro-circulatory reactivity, inflammatory biomarkers, thrombotic factors, and plasmatic aldosterone concentration were assessed at different times post-dive. Although VGE were recorded in all divers, no cases of decompression sickness (DCS) occurred. Compared to the control, the latency to VGE peak was increased in the DLCO group (60 vs. 30 min) along with a higher maximal VGE grade (p < 0.0001). P-selectin was higher in the DLCO group, both pre- and post-dive. The plasmatic aldosterone concentration was significantly decreased in the control group (-30.4 ± 24.6%) but not in the DLCO group. Apart from a state of hypocoagulability in all divers, other measured parameters remained unchanged. Our results suggest that divers with decreased DLCO might have a higher risk of DCS. Further studies are required to confirm these preliminary results.

Scuba diving after a heart transplant: excessive daring or calculable risk?

Over the past 50 years, outcomes after heart transplantation (HTX) have continuously and significantly improved. In the meantime, many heart transplant recipients live almost normal lives with only a few limitations. In some cases, even activities that actually seemed unreasonable for these patients turn out to be feasible. This article describes the encouraging example of a patient returning to recreational scuba diving after HTX. So far, there were no scientific experiences documented in this area. We worked out the special hemodynamic features and the corresponding risks of this sport for heart transplant recipients in an interdisciplinary manner and evaluated them using the patient as an example. The results show that today, with the appropriate physical condition and compliance with safety measures, a wide range of activities, including scuba diving, are possible again after HTX. They illustrate again the significant development and the enormous potential of this therapy option, which is unfortunately only available to a limited extent.NEW & NOTEWORTHY Example for shared decision-making process for tricky questions: First scientific publication about heart transplantation (HTX)-recipient restarting scuba diving. As exercise physiology after HTX combined with specific diving medicine aspects is challenging, we formed a multidisciplinary team to identify, evaluate, and mitigate the risks involved. The results show that today, with the appropriate physical condition and compliance with safety measures, a wide range of activities are possible again after HTX.

The pulmonary oxygen toxicity index.

Pulmonary oxygen toxicity (POT) is a major risk in diving while breathing hyperoxic gas and is also considered in clinical hyperbaric oxygen treatment. The POTindex calculated by the power equation K = t2 × PO24.57 with the recovery form Ktr = Ke × e - [- 0.42 + 0.384 × (PO2)ex] × tr which are based on chemical and physiological principles, have a better prediction power than other suggested approaches. Reduction of vital capacity as well as incidence of POT are well predicted by the POTindex. Both the cumulative pulmonary toxic effect and concomitant recovery were suggested to operate at the lower toxic range of PO2 used in saturation diving K = t2 × PO24.57 × e-0.0135 × t, and further experimental support is supplied. The recovery time constant for the full range of PO2 is presented. POTindex is suggested to replace the old method of UPTD for safe diving. Many diving clubs and diving institutes already adopted the POTindex.

Pulmonary Physiology and Medicine of Diving.

Pulmonary physiology is significantly altered during underwater exposure, as immersion of the body and increased ambient pressure elicit profound effects on both the cardiovascular and respiratory systems. Thoracic blood pooling, increased breathing gas pressures, and variations in gas volumes alongside ambient pressure changes put the heart and lungs under stress. Normal physiologic function and fitness of the cardiovascular and respiratory systems are prerequisites to safely cope with the challenges of the underwater environment when freediving, or diving with underwater breathing apparatus. Few physicians are trained to understand the physiology and medicine of diving and how to recognize or manage diving injuries. This article provides an overview of the physiologic challenges to the respiratory system during diving, with or without breathing apparatus, and outlines possible health risks and hazards unique to the underwater environment. The underlying pathologic mechanisms of dive-related injuries are reviewed, with an emphasis on pulmonary physiology and pathophysiology.

Diving behaviour of southern elephant seals: new models of behavioural and ecophysiological adjustments of oxygen store management.

Among pinnipeds, southern elephant seals (SESs, Mirounga leonina) are extreme divers that dive deeply and continuously along foraging trips to restore their body stores after fasting on land during breeding or moulting. Their replenishment of body stores influences their energy expenditure during dives and their oxygen (O2) reserves (via muscular mass), yet how they manage their O2 stores during their dives is not fully understood. In this study, 63 female SESs from Kerguelen Island were equipped with accelerometers and time-depth recorders to investigate changes in diving parameters through their foraging trips. Two categories of dive behaviour were identified and related to the body size of individuals, with smaller SESs performing shallower and shorter dives requiring greater mean stroke amplitude compared with larger individuals. In relation to body size, the larger seals had lower estimated oxygen consumption levels for a given buoyancy (i.e. body density) compared with smaller individuals. However, both groups were estimated to have the same oxygen consumption of 0.079±0.001 ml O2 stroke-1 kg-1 for a given dive duration and at neutral buoyancy when the cost of transport was minimal. Based on these relationships, we built two models that estimate changes in oxygen consumption according to dive duration and body density. The study highlights that replenishing body stores improves SES foraging efficiency, as indicated by increased time spent at the bottom of the ocean. Thus, prey-capture attempts increase as SES buoyancy approaches the neutral buoyancy point.