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Plasma catecholamines have been measured in chronically adrenalectomised (ADX) ducks, in chronically adrenal denervated ducks (DNX), in their respective sham-operated controls (SH-adx, SH-dnx) as well as in intact ducks after 3 minutes forced submergence. The results showed that 100% of the plasma Epinephrine (EP) and 40 to 80% of plasma Norepinephrine (NE) released during the dive came from the adrenal glands. 20 to 60% of plasma NE came from endings of the autonomic vascular sympathetic nerves which are strongly stimulated during diving. Adrenal catecholamines were released by nerve activation only; non neural mechanisms did not play any role in their release. Maximum dive times (MDT) in chronically adrenalectomised ducks (ADX: 5 min. 19 ± 20 sec.) and in chronically adrenal denervated ducks (DNX: 7 min. 10 ± 13 sec.) were significantly lower than in sham-operated controls (respectively SH-adx: 9 min. 58 ± 45 sec., SH-dnx: 12 min. 10 ± 28 sec). Venous infusion of catecholamines in ADX and DNX during the dive increased MDT: MDT of DNX ducks perfused with catecholamines (9 min. 46 ± 20 sec.) were significantly higher than in DNX perfused with saline (7 min. 21 ± 17 sec.), but did not reach the MDT observed in the SH-dnx: other adrenal products must be involved. Diving heart rates of ADX and DNX (at 4 min. dive respectively: 62 ± 16 and 31 ± 2 beats/min.) were significantly higher than in their sham-operated controls (23 ± 3 and 17 ± 2 beats/min.) . Blood pressure during the dive was signifi- cantly lower in ADX and DNX (at 4 min. dive respectively: 93 ± 8 and 98 ± 4 mmHg) compared with their sham-operated controls (131 ± 12 and 118 ± 6 mmHg). Infusion of catecholamines in DNX raised blood pressure towards SH-dnx values, but there was no change in heart rate. PaO₂, CaO₂, pHa and lactate levels in DNX (respectively: 42 ± 2 mmHg, 4.5 ± 0.8 ml 02 /100ml blood, 7.233 ± 0.016, 3.1 + 0.3 mM) were significantly lower than in SH-dnx after 5 minutes submergence (53 ± 1 mmHg, 6.8 ± 0.4 ml 02 /100 ml blood, 7.301 ± 0.007, 4.8 + 0.4 mM). There was also a significant increase of plasma N⁺ (+ 5.4 ± 1.7 mEq/L) in SH-dnx after 5 minutes submergence, but this was not the case in DNX where it was K⁺ (+ 1.1 ± 0.4 mEq/L) which increased. This suggested that adrenal catecholamines increase tolerance to underwater submersion by enhancing peripheral vasoconstriction, thus preserving the O₂ stores for the heart and brain. Moreover, they may affect the acid-base equilibrium during diving by increasing the activity of the Na⁺K⁺ pump and may also have a direct effect on the rate of glycogenolysis. Preventing the actions of catecholamines on the heart by injecting beta-blocker during forced submersion did not decrease MDT; however the cardiovascular response was markedly affected. During beta-blockade, diving heart rate rose steadily from 24 ± 6 beats/minute after 2 minutes to 52 ± 8 beats/minute after 6 minutes diving. In contrast, heart rates remained close to the levels reached at 2 minutes (17 ± 3 and 19 ± 4 beats/minute) throughout the control dives. Perfusion pressure and blood flow have been recorded simultaneously in both hind limbs of ducks. One leg was perfused with different blood mixtures devoid of catecholamines (Test leg) and compared with the other, perfused with the ducks'own blood (autoperfused leg). This showed that hyper-capnia has a depressant effect on the neural component of the peripheral vasoconstriction. Perfusion of test legs with hypoxic-hypercapnic blood to which catecholamines were added, showed that circulating catecholamines are needed to increase peripheral vasoconstriction during diving. In summary, during forced submergence circulating catecholamines, released mainly by the adrenal glands, compensate for the depressant action of hypercapnia on the neural component of peripheral vasoconstriction. Maintenance of this peripheral vasoconstriction during forced diving ensures that O₂ stores are not wasted on peripheral tissues, and this explains how MDT is prolonged. / Science, Faculty of / Zoology, Department of / Graduate

The enzymatic organization of muscle tissue usually is examined in only a select few muscles of any one animal species. However, because the functional demands placed on individual muscles can vary so widely from muscle to muscle, it is inappropriate to generalize findings from one or two muscles to muscle tissue in general. The differences or similarities in metabolic machinery between skeletal muscles of a wide functional range provides crucial information with respect to a particular animals' whole body metabolism. Nowhere is this understanding more important than in the diving marine mammal which must operate as a closed system (with respect to oxygen supply) while submerged. The goals of this thesis are: 1) to provide a broad body of information on the metabolic organization of a large cross-section of marine mammal muscles, both functionally and with regard to location, 2) to assess the implications of the enzyme differences between muscles to the diving habit, and 3) to compare the metabolic organization of skeletal muscle among several species of marine mammal with different diving abilities and habits. A series of 13 enzymes were measured in 21 skeletal muscles of the harbor seal, Phoca vitulina. In addition, 23 enzyme activity ratios were calculated and analyzed for these muscles. A similar analysis of 22 muscles from fin whales, Balaenoptera physalis. was conducted --including 7 key enzymes and 15 activity ratios. Overall, both the maximum activities and the enzyme activity ratios are consistent with the idea that marine mammal muscle is typical mammalian muscle, exhibiting few significant differences from terrestrial species with respect to catabolic enzymes. The only obvious exception to this in the species examined is observed with fin whale locomotory muscle which has extremely high activities of lactate dehydrogenase (over 2000 units/gm wet wt at 25°C) due to an apparent scaling phenomenon. Tight control of this high potential glycolytic flux is indicated by pyruvate kinase activities that scale downward. Comparisons of enzyme relationships between muscles of harbor seals seem to indicate a very aerobically poised metabolic make-up. This is especially true with respiratory and locomotory muscles, which also show a high tendency to utilize fat. This pattern of enzyme activities and activity ratios in the locomotory muscles of harbor seal is evidence that muscle contractile activity while diving is powered primarily through oxidative pathways and largely based on fat as fuel. The majority of non-locomotory muscles appear to be more able to function anaerobically utilizing carbohydrate. This pattern may correlate with circulatory redistributions while diving that preferentially fuel the locomotory muscles with oxygen, leaving the inactive muscles significantly more hypoperfused and, therefore, candidates for energy saving O₂ sparing (metabolic depression). Fin whales exhibit an opposite pattern, with enzyme profiles more typical of "white" muscle. Unlike harbor seals, the locomotory muscles of fin whales are consistently the least oxidatively poised of the muscles examined. This apparently more anaerobic nature of fin whale muscle is possibly complicated by scaling adaptations, but appears to be a real phenomenon. The examination of three to four skeletal muscles from each of three additional phocid seal species from Antarctica, leopard seals (Hydrurga leptonyx). crab-eater seals (Lobodon carcinophagus). and Weddell seals (Leptonychotes weddelli) confirm that the harbor seal pattern of enzyme profiles is fairly consistent among phocid seals. By these criteria skeletal muscles of phocid seals (particularly the locomotory and respiratory muscles) appear to be designed for sustained aerobic metabolism during diving regardless of the habits or diving capabilities of the seal. / Science, Faculty of / Zoology, Department of / Graduate

The extreme elevation in plasma levels of norepinephrine (NE) and epinephrine (EP) which occurs during forced diving of ducks (Anas platyrhynchos) was studied before and after denervation of the adrenal glands. Elevated PaCO₂, decreased arterial pH, decreased blood glucose as well as low PaO₂ have been described as causal factors in this response. These variables, as well as blood pressure, heart rate and breathing frequency were measured in ducks dived after breathing air or pure 0₂ to clarify and quantify the mechanism involved and its physiological function. Both NE and EP concentration increased by up to 2 orders of magnitude in the 4 minute dive period, but by a significantly lesser amount if the duck breathed 0₂, before the dive. While pH and PaCO₂ were well correlated with the changes in plasma NE and EP levels during both air and 0₂ dives, both pH and PaCO₂ changed more in the 0₂ trials, indicating that they are not the primary cause of the response. Plasma glucose levels were variable. PaO₂ values less than normal correlated well with increasing NE and EP concentrations, but at high PaO₂s, there was no correlation, suggesting that hypoxia is the permissive state for the full response. Compared with breathing air, breathing O₂ before the dive attenuated the diving bradycardia, eliminated the decrease in blood pressure normally observed during dives, and caused more extreme changes in pH, PaC0₂, and of course, PaO₂. Denervating the adrenals decreased the amounts of both catecholamines released during dives after breathing air and 0₂, EP significantly more than NE. Adrenal denervation per se did not cause a significant change in heart rate, blood pressure, arterial gas tensions, pH, or plasma glucose changes during dives although the operation caused increased variation in some of the parameters. In ducks, the cause for the catecholamine release is decreasing PaO₂ and full expression of the response is dependent on intact innervation of the adrenal gland, although there is a component that is unaffected by denervation. While possible roles for this response are discussed, the true physiological function of this response remains cast in shadows. / Science, Faculty of / Zoology, Department of / Graduate

Cardiovascular responses evoked during forced submersion enable the Pekin duck (Anas platyrhynchos) to survive protracted periods of asphyxia. The responses include an extraordinary bradycardia and intense peripheral vasoconstriction with the result that blood flow is favoured to those organs most susceptible to lack of oxygen. These adjustments appear to be mediated via the caudal brainstem following stimulation of peripheral and central arterial chemoreceptors. The minor role that baroreceptors play in the generation of these responses was demonstrated by the persistence of the cardiovascular changes following peripheral arterial baroreceptor denervation. Isolation of the cephalic circulation from the systemic circulation enabled a series of experiments to assess the relative contributions from peripheral chemoreceptors, located in the carotid bodies, and from unidentified central chemoreceptors within the cranial circulation. A declining arterial P0₂ in the systemic circulation appeared especially potent in evoking bradycardia during submersion. Increased arterial PC0₂, likewise, resulted in a reduced heart rate. Similar changes in the blood gas levels of the cephalic circulation did not elicit significant bradycardia. However, both receptor groups responded to arterial hypoxic hypercapnia by activating substantial reduction in peripheral blood flow, as reflected by the rise in hind limb vascular resistance. Although baroreceptors may continue to mitigate changes in arterial blood pressure and cause some change in heart rate and vascular resistance, chemoreceptors appear to be predominantly responsible for the changes during submersion. The cardiac response to chemoreceptor stimulation during submersion was discovered to habituate following repetitive diving. Habituation was so pronounced in some ducks that after several training sessions the bradycardia during 40-second forced dives was abolished. Habituation of the cardiac response appeared dependent on the intensity of chemoreceptor stimulation. With severe arterial hypoxia, produced by either prolonging dive times or by reducing the pre-dive inspired oxygen content, little or no cardiac habituation was observed. Tests were conducted to demonstrate efficacy of the cardioinhibitory efferent discharge. Maintained sensitivity of chemoreceptors was suggested by the lack of change in oxygen breathing tests before and after training. Furthermore, the persistence of stimulus intensity was established and these observations led to the suggestion that the locus of habituation is within the CNS. The demonstration that the level of bradycardia was dependent on arterial P0₂ in both naive and habituated animals argues against the contention that the diving response is a fear response. Further evidence against this view was provided by the demonstration that the diving response remains essentially intact following transection in the rostral mesencephalon below the level of the hypothalamus. It is concluded that chemoreceptor-driven cardiovascular changes evoked as part of the diving response are mediated by regions of the CNS below the rostral brainstem. Modification of these responses can be produced in the intact animal by simple forms of learning. However, it remains uncertain at what level this influence arises. / Science, Faculty of / Zoology, Department of / Graduate

Force dived animals undergo cardiovascular changes characterized by bradycardia, increased total peripheral resistance, and changes in blood flow distribution. Since these changes occur in decerebrated animals, the dive response must be a brainstem reflex. However, in voluntary dives, animals may show anticipatory bradycardia and may also adjust their cardiovascular responses according to anticipated dive duration, indicating suprabulbar influences upon dive responses. Studies of heart rate using telemetry have shown that there can be substantial differences in the dive response of voluntarily and force dived animals. Furthermore, some animals show a "fear bradycardia" when trapped in a stressful situation, leading some researchers to suggest that bradycardia during forced submersion is an artifact of the stress of the situation. Muskrats (Ondatra zibethica) were observed freely diving for food in an indoor tank using a video camera and VCR unit. EKG was telemetered from the animals and recorded on the audio channel of the VCR tape. Heart rate responses to voluntary dives were analyzed and compared with those from escape and forced dives. Heart rate responses were also recorded from decorticate and sham operated muskrats to elucidate the role that the cerebral cortex plays in the dive response. In all types of dives, muskrats exhibited a rapid and large bradycardia upon submergence (heart rate declined by greater than 55% of the predive heart rate). Obviously diving bradycardia in the muskrat was not due to fear or stress, but occurred as a response to submersion per se. There was no evidence of post-dive tachycardia or anticipatory immersion bradycardia. Disturbing the animal in a non-diving situation resulted in only a 13% decrease in heart rate. In intact animals voluntary, escape, and forced submergence resulted in progressively greater decreases in heart rate. Heart rate fell by 56% in voluntary dives, 65% in escape dives, and 73% in forced dives. Intensification of the bradycardia to a lower heart rate than that seen in voluntary dives was mediated by the cerebral cortex, as heart rate in decorticate muskrats in escape and forced dives did not fall below that seen in voluntary dives. This indicates that the final adjustment of dive heart rate is dependent upon an intact cerebral cortex. However, in decorticate muskrats there appeared to be a recovery of cortical function, as intensification of bradycardia in forced dives was dependent upon the time that had elapsed after surgery. This study shows that there is a cortical influence upon the cardiovascular system during diving. It also indicates that in experiments with unanesthetized animals, the degree of stress of the situation must be taken into account, as this may affect physiological responses. / Science, Faculty of / Zoology, Department of / Graduate

Blood flow distribution during forced and voluntary diving in ducks, and the energetic cost of diving was investigated. It has been suggested that in order for the leg muscles to generate enough power for ducks to dive, blood flow to those tissues must be maintained. A technique to determine blood flow distribution which could be used during voluntary diving was first developed and tested during forced laboratory dives of ducks. This technique was then used to determine the blood flow distribution during voluntary diving. Regional blood flow distribution was visualized by utilizing a radioactive tracer technique (macro aggregated albumin labelled with ⁹⁹ⅿ technetium). The tracer when injected into an animal is trapped and held by capillaries. During forced dives in dabbling (Anas platyrhynchos) and diving (Aythya affinis) ducks the blood flow distribution was found to be restricted to the thoracic and head areas. Whereas during a voluntary dive in A. affinis blood flow distribution was shown to be preferentially directed towards three tissue areas, the heart, brain, and active leg muscles. The work required to dive was determined from the measurement of subsurface drag forces and buoyancy in A. affinis. Subsurface drag increased as a nonlinear function of swimming velocity. At a velocity of 1 m•s⁻¹, the drag force was approximately 1.067 N. The average measured buoyant force of 11 ducks was 0.953 N. The calculated mechanical work done by ducks during a 14.4 s unrestrained dive was 9.34 J. The power output during voluntary was estimated to be 0.751 W (0.0374 ml 0₂•s⁻¹). During diving buoyancy is clearly the dominant force (8.8 J) against which ducks have to work while drag (0.54 J) adds little (~6%) to the energetic cost of diving. / Science, Faculty of / Zoology, Department of / Graduate

When dabbling ducks are involuntarily submerged, arterial vasoconstriction produces a large increase in the peripheral resistance to blood flow which is balanced by a decrease in output of the heart, and arterial blood pressure is maintained. Arterial baroreceptors sense systemic blood pressure, and provide the afferent information to the baroreflex for pressure regulation. The effector limbs of the baroreflex are the same as those involved in the diving responses, and the baroreceptors are likely to be important in the integration of the cardiovascular responses to diving. The purpose of this study was to investigate the role of the arterial baroreceptors in maintaining blood pressure during diving, and in the initiation and maintenance of the diving responses. Baroreceptor function was studied by diving ducks at various times after barodenervation, and by electrically stimulating the central end of one baroreceptor nerve in baroreceptor-denervated animals to simulate a controlled baroreceptor input before and during submersion. Intact baroreceptor innervation was not necessary for the development of peripheral vasoconstriction, but loss of the baroreceptors modified the cardiac response to submersion by impairing the vagally mediated bradycardia. There was no effect of baroreceptor nerve stimulation on peripheral resistance during diving, and the baroreflex operated via the heart rate in modulating blood pressure early in the dive. Later in the dive, stimulation was ineffective in altering either heart rate or blood pressue. Strong chemoreceptor drive results from decreased blood oxygen and increased carbon dioxide levels in the dive, and the results of experiments to determine the mechanism of baroreflex attenuation showed that an interaction between chemoreceptor and baroreceptor inputs may be at least partly responsible for reducing baroreflex effectiveness. The main conclusion from this work is that the arterial baroreceptors contribute to the diving responses through modulation of heart rate, to help balance the fall in cardiac output against the baroreceptor-independent peripheral vasoconstriction in the first minute of forced dives. / Science, Faculty of / Zoology, Department of / Graduate

This purpose of this study was to evaluate the contribution of post-dive peripheral tissue warming to the production of venous gas emboli (VGE) in divers. Inert gas elimination from the tissues is limited by both perfusion and diffusion. If changes in diffusion are matched by corresponding perfusion (vasoactive) changes, decompression should be asymptomatic (within allowable exposure limits). Under conditions when the diffusion of inert gas from the tissues is not matched by blood perfusion, VGE will ensue. Increasing tissue temperature will decrease inert gas solubility and thus diffusion into the blood. It has been demonstrated that problems may arise during rapid changes in peripheral temperature, as often occurs post-dive, when divers previously exposed to cold water actively rewarm themselves in showers or baths. The effect of moderate rewarming, however, may be to increase the rate of inert gas elimination without the formation of VGE since increased perfusion is encouraged. The effect of mild post-dive warming was investigated. Ten male subjects, between the ages of 21 and 29 years completed two dry chamber dives to 70 feet for 35 minutes (no decompression limit of the Canadian Forces Air Diving tables). Each dive was followed by a 30 minute head-out immersion in either a thermoneutral (28°C) or warm (38°C) bath. Non-invasive Doppler ultrasonic monitoring was then carried out at 30 minute intervals for the next 150 minutes to assess measurable VGE. Subjects did not display VGE formation in either the control or experimental conditions. Our findings suggest that: 1) the Canadian Forces table limits (for the profile employed) provide safe no-decompression limits not compromised by mild post-dive warming, and 2) mild peripheral warming, since not bubble generating, may be a useful adjunctive therapy in the management of decompression sickness by increasing the rate of inert gas elimination. / Education, Faculty of / Curriculum and Pedagogy (EDCP), Department of / Graduate

Thesis (MScMedSc)-- Stellenbosch University, 2014. / ENGLISH ABSTRACT: The commercial and military deep diving environment is typically a low visibility environment, where dependence on the visual senses often needs to be replaced by a reliance on tactile senses. This thesis reviewed the current knowledge regarding neuropsychological manifestations of nitrogen narcosis and exposed a number of shortcomings in the current body of knowledge. In particular, the human performance effects of hyperbaric exposure on tactile perception and memory have not been systematically studied. It is further not clear, how exactly psychological factors (e.g. anxiety, mood states) and biographical factors (e.g. age, education, technical exposure, experience) might influence tactile perception and memory performance under conditions of hyperbaric exposure. The correlation between subjective experiences of narcosis, tactile performance, and psychological and biographical variables is also unknown. This study thus set out to investigate certain neuropsychological aspects of nitrogen narcosis, with special reference to tactile perception and memory, and to examine the relationships of tactile performance with other psychological and biographical factors. The effects of experimental hyperbaric exposure (EHE) on tactile (form) perception and tactile shape memory were examined by testing these functions at 6 ATA and 1 ATA, using a cross-over design where two groups completed the same tasks, in opposite sequence. The psychological variables included trait anxiety, transient mood states, and subjective ratings of narcosis, while the biographical variables included age, education, and previous technical exposure. The results demonstrated the detrimental effect of nitrogen narcosis on tactile form perception and manipulation, irrespective of the sequence of testing. It also demonstrated this effect on tactile form memory, although the sequence of testing also played a role here. Higher trait anxiety was associated with poorer recall, and tension was associated with a larger decrement in recall performance, while fatigue was associated with poorer task completion. Subjective experiences also played a role, where feelings of physical anxiety (i.e. increased arousal) were associated with better recall, and feelings of cognitive suppression (decreased arousal) were associated with a larger decrement in recall performance. Lower academic attainment was associated with poorer recall, while higher diving qualification was associated with better recall. Performance on the surface was a good predictor of performance at depth. Qualitative analysis rendered three themes, namely focus vs. distraction, following instructions, and shape memory. Psychometric properties of the subjective narcosis measure were also reported. Theoretical implications include support for the slowed information processing model when completing complex neuropsychological tasks, as well as support for the memory model, thus suggesting that this particular pattern of memory impairment occurs because encoding under narcosis produces a weaker memory trace than normal. Lastly, the study has a number of implications for industry. For example, divers need to compensate for slowed task completion by, firstly, planning more time to complete complex tasks, and secondly, by practicing those tasks prior to the actual deep dive (either on the surface or in shallow water). The need for using additional forms of recording of events or objects at depth, to aid memory encoding and subsequent recall at surface was also emphasised. / AFRIKAANSE OPSOMMING: Kommersieële en militêre duik vind dikwels plaas in ‘n omgewing met swak sig, waar duikers moet staatmaak op taktiele sintuie, eerder as op visuele sintuie. Die tesis begin met ‘n oorsig oor die huidige kennis rakende neurosielkundige verskynsels van stikstof narkose, en het ‘n aantal tekortkominge gevind. Meer spesifiek, die menslike faktor in die effek van hiperbariese druk op taktiese persepsie en geheue is nog nie sistematies bestudeer nie. Dit is verder nie duidelik presies hoe sielkundige faktore (angs, gemoedstoestande) en demografiese faktore (ouderdom, opvoeding, tegniese blootstelling, ondervinding) taktiele persepsie en geheue onder toestande van hiperbariese druk sou beïnvloed nie. The korrelasie tussen die subjektiewe ervaring van narkose, taktiele taakverigting, en sielkundige en biografiese veranderlikes is ook nie bekend nie. Die studie het verskeie neurosielkundige aspekte van stikstof narkose, met spesifieke verwysing na taktiele persepsie en geheue, sowel as die verhouding tussen taktiele prestasie en sielkundige en biografiese faktore ondersoek. Die effek van hiperbariese druk op taktiele persepsie en geheue is ondersoek deur hierdie funksies te toets by 6 en 1 ATA, deur middel van ‘n oorkruis studie ontwerp, waar twee groepe die take voltooi het, in teenoorgestelde volgorde. Die sielkundige veranderlikes het bestaan uit trek-angs, tydelike gemoedstoestande, en die subjektiewe evaluering van narkose, terwyl die biografiese veranderlikes ouderdom, opvoeding, en vorige tegniese blootstelling ingesluit het. Die resultate het die nadelige effek van stikstof narkose op taktiele vorm persepsie en manipulasie gedemonstreer, ongeag die rigting van toetsing. Dit het ook hierdie effek op taktiele vorm geheue gedemonstreer, hoewel die rigting van toetsing wel hier ‘n rol gespeel het. Hoër trek-angs was geassosieër met swakker herroeping, en spanning met ‘n groter agteruitgang in herroeping, terwyl matheid geassosieer was met swakker taakvoltooiing. Subjektiewe ervarings het ook ‘n rol gespeel, met ervarings van fisiese spanning (verhoogde opwekking) geassosieer met beter herroeping, en ervarings van kognitiewe onderdrukking (verlaagde opwekking) met groter agteruitgang in herroeping. Laer akademiese kwalifikasie was geassosieer met swakker herroeping, terwyl hoër duik kwalifikasie geassosieer was met beter herroeping. Taakverrigting op die oppervlak was ‘n goeie voorspeller van prestasie op diepte. Kwalitatiewe analiese het drie temas geidentifiseer, naamlik fokus vs. afleibaarheid, die volg van instruksies, en vorm geheue. Die psigometriese eienskappe van die subjektiewe narkose meetinstrument is ook gerapporteer. Teoretiese implikasies van die studie sluit in ondersteuning vir die vertraagde prosesseringsmodel, wanneer komplekse neurosielkundige take voltooi word, sowel as ondersteuning vir die model vir hierdie spesifieke herroepingspatroon wat ‘n swakker geheuespoor laat wanneer enkodering plaasvind onder toestande van narkose. Die studie het ook praktiese implikasies vir industrie. Dit is byvoorbeeld nodig om te kompenseer vir vertraagde taakvoltooïng deur, eerstens, die beplanning vir meer tyd om komplekse take te voltooi, en tweedens, deur daardie take te oefen voor die diep duik plaasvind. Die noodsaaklikheid vir additionele maniere om gebeure of voorwerpe op diepte vas te lê is ook beklemtoon.