The concept of stressAnimals rely on behavioural <strong>and</strong> physiological mechanisms, which enable them to maintainhomeostasis in response to external <strong>and</strong> internal stimuli. These regulatory mechanisms have optimum<strong>and</strong> maximum tolerance ranges, which depend largely on a species’ evolutionary history, but can bemoderated by individual genetic make-up, as well as short <strong>and</strong> long-term history. Environmentalstimuli, which fall outside an animal’s adaptive range with regard to duration, intensity or frequency,or because of the nature of the stimulus itself, are associated with pathology <strong>and</strong> reduced survival.These ‘overtax’ behavioural <strong>and</strong> physiological control systems <strong>and</strong> are referred to as ‘stress’ (Broom<strong>and</strong> Johnston 1993).Stressful conditions, such as confrontation with a predator or rival, disturb homeostasis <strong>and</strong> result inprofound physiological <strong>and</strong> behavioural changes, which involve complex interrelated hormonal,metabolic, neural <strong>and</strong> neuroendocrine responses (e.g., Toates 1995). The main transmitter substances<strong>and</strong> hormones involved include glucocorticoids (cortisol, corticosterone), the mineralocorticoidaldosterone, catecholamines (adrenaline <strong>and</strong> noradrenaline), insulin, thyroid <strong>and</strong> growth hormone.During stress the body mobilises carbohydrates <strong>and</strong> fatty acids to provide energy. At the same timeblood pressure, cardiac <strong>and</strong> respiratory rate increase. This provides the efficient transport of vitalnutrients to the skeletal <strong>and</strong> cardiac muscles. Less immediately important processes such as digestion,immune defence, reproduction <strong>and</strong> growth are inhibited to further maximise available energy. Thesechanges are independent of physical activity.Psychological stimuli, including fear, elicit strong adrenal responses <strong>and</strong> an assessment of how stimuliare perceived is therefore critical. Mason (1971) emphasised the psychological dimensions of allanimal treatments (see also Toates 1995 <strong>and</strong> von Holst 1998) <strong>and</strong> considers it virtually impossible toavoid the psychological element of physical stressors.The potential stress effects of whaling <strong>and</strong> the welfare implications for hunted cetaceans71Physiological indicators of stress <strong>and</strong> their interpretationStressful situations cause behavioural <strong>and</strong> physiological changes that can be gauged through a rangeof biological indicators. Measurements of cardiac <strong>and</strong> respiratory rate, body temperature, as well as anumber of physiological, haematological <strong>and</strong> biochemical profiles can provide important informationabout whether or not an animal is stressed. Interpreting biological parameters used to assess theimpact of potentially stressful conditions is not always straightforward, <strong>and</strong> several indicators shouldbe employed to avoid misleading results. The importance of accurate baselines against whichexperimental measurements can be compared is critical. Even without visible signs of stress,biochemical <strong>and</strong> physiological profiles may be affected, <strong>and</strong> haematological assessment should,therefore, consider sex, nutritional state, circadian rhythms, seasonal variation, <strong>and</strong> physiologicalstate. Sampling itself can have effects <strong>and</strong> lead to persistently overestimated baseline levels. The sameis true for animals that are already stressed when samples are taken. Sampling method, samplepreparation <strong>and</strong> storage may affect samples.Stress associated with pursuitAcute stress on capture may bring about short <strong>and</strong> long-term morbidity <strong>and</strong> mortality in bothdomestic <strong>and</strong> wild species (Mitchel et al. 1988). Hyperthermia, profuse sweating, hyperventilation,hypotension <strong>and</strong> degrees of skeletal <strong>and</strong> cardiac muscle damage are common post-chase <strong>and</strong> postcaptureconditions.
Both chase <strong>and</strong> pursuit cause stress in terrestrial mammals; this includes stress-related mortality, afactor which may apply to cetaceans. Pursuit-related stress may manifest as a syndrome called:‘exertional myopathy’ (EM), ‘capture myopathy’, ‘stress myopathy’, or ‘exertional rhabdomyolysis’.Myopathies are diseases of the muscle fibres. EM, however, is distinguished from other types ofmyopathy, such as nutritional <strong>and</strong> toxic myopathies by its cause, as it affects both skeletal <strong>and</strong> cardiacmuscles in response to exertion, fear <strong>and</strong> stress.Commonly associated with strenuous or prolonged pursuit, capture, restraint or overexertion, EMdevelops irrespective of capture. Mental stressors, such as fear <strong>and</strong> anxiety, too, have been recognisedas predisposing factors, as have high ambient temperature <strong>and</strong> impeded thermoregulation. Theseelements may act singly or together. The acute stress on capture may bring about short <strong>and</strong> longtermmorbidity <strong>and</strong> mortality. Williams <strong>and</strong> Thorne (1996) stated “even species that have evolved forefficient running, either for predator avoidance or for predation, may develop EM following intense orprolonged muscular activity associated with extreme stress during air or ground pursuit”. These authorsconsider pursuit time a major factor in the development of EM.72A REVIEW OF THE WELFARE IMPLICATIONS OF MODERN WHALING ACTIVITIESAs energy <strong>and</strong> oxygen reserves are depleted during strenuous exercise, muscles switch to anaerobicglycolysis. This leads to either local or systemic build-up of lactic acid, local heat production, muscledegeneration <strong>and</strong> death of areas of muscle tissue (necrosis) (Fowler <strong>and</strong> Boever 1993). Increasedcardiac <strong>and</strong> respiratory rates, elevated body temperature, ataxia, paresis or paralysis <strong>and</strong> acute muscledisruption are some of the symptoms associated with EM (Harthoon <strong>and</strong> Young 1974, Bartsch et al.1977, Chalmers <strong>and</strong> Barrett 1977, Basson <strong>and</strong> Hofmeyr 1978; Fowler <strong>and</strong> Boever 1993). Identifyingor interpreting these factors requires knowledge about the animal’s normal undisturbed behaviour.Harthoorn (1973) describes four syndromes associated with the disease, namely hyperacute, acute,subacute <strong>and</strong> chronic EM, although according to Williams <strong>and</strong> Thorne (1996) these “represent acontinuum of physiologic <strong>and</strong> pathologic changes that occur over time after the initial exertion insult”.Clinical signs, including death, may occur within minutes or hours, or in the case of muscle necrosis<strong>and</strong> nephrosis (destruction of functional kidney tissue), more gradually over days, weeks or evenmonths. Affected animals may initially appear normal (Spraker 1993), <strong>and</strong> even those which recoverfrom acute problems, may die after weeks or months as a result of scar formation in the heart muscle(myocardium) (Jubb et al. 1993).Stress in cetaceansDespite a wealth of evidence from terrestrial species <strong>and</strong> birds, information on the physical <strong>and</strong>behavioural effects of stress in cetaceans, <strong>and</strong> particularly Mysticeti, is limited. However, stress-relatedchanges in adrenal <strong>and</strong> thyroid hormone levels have been documented in cetaceans (reviews inDierauf 1990; St. Aubin <strong>and</strong> Dierauf, 2001<strong>and</strong> Curry 1999).Chase-capture <strong>and</strong> restraint of six captive bottlenose dolphins (Tursiops truncatus) led to 100 per centhigher plasma cortisol levels than under calm-capture conditions. However, plasma cortisolmeasurements increased “even under the calmest conditions of capture”. Unlike most mammals, stressedcetaceans may manifest moderate cortisol elevations, although the physiological consequences ofcortisol secretion in the body are maintained. Aldosterone levels on the other h<strong>and</strong> can increasesubstantially in cetaceans <strong>and</strong> may be a better indicator. Aldosterone moderates effective water <strong>and</strong>sodium resorption <strong>and</strong> elevated levels result in excessive sodium retention (e.g., Townsend 1999).
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ForewordWhales are highly evolved a
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1 Executive SummaryThis review exam
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2 A background to whalingPhilippa B
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y the weapon’s enormous recoil, w
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Japan currently whales in the Antar
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Otto, K. 1997. Animal Pain Behaviou
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Protecting the welfare of animals i
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Toothed whales (Odontoceti)Toothed
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Social behaviourMother-calf pairsOn
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law says, but also the extent to wh
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15 Whaling and welfarePhilippa Brak
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commercial whaling. Times to death
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eath). Using the current criteria t
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16 Summary of conclusionsModern day
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Glossary136A REVIEW OF THE WELFARE
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Appendix IIColour plates©Mark Voti
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142A REVIEW OF THE WELFARE IMPLICAT
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Figure 13. Processing minke whales