Thyroid function in cetaceans can also be affected by stressful conditions, but interpretation shouldtake account of sex <strong>and</strong> age (St. Aubin et al. 1996). For examples see; a) beluga whales(Delphinapterus leucas): St. Aubin <strong>and</strong> Geraci 1988, 1992; b) captive white-sided dolphins(Lagenorhynchus obliquidens): Ridgeway <strong>and</strong> Patton 1971; c) bottlenose dolphins (Tursiops truncatus):Orlov et al. 1988; see also St. Aubin et al. 1996.As in most vertebrates, stress can alter immunocompetence in cetaceans (see Romano et al. 1992,1993, 1994; St. Aubin et al. 1989, 1990; Medway et al. 1970; Townsend 1999; reviewed in Curry1999). Suppression of natural killer cell activity is reported in beluga whales (De Guise et al. 1997),<strong>and</strong> capture stress can suppress leukocytes <strong>and</strong> blood iron levels (Geraci <strong>and</strong> Medway 1973, St. Aubin<strong>and</strong> Geraci 1989). Thomson & Geraci (1986) report high mortality associated with capture <strong>and</strong>confinement stress in various small cetaceans, including Phococoena phococoena (Dudock van Heel,1962), Phocoenoides dalli (Ridgeway, 1966) <strong>and</strong> Delphinus delphis (Walker, 1975).Muscle damage <strong>and</strong> exertional myopathy in cetaceansMuscle damage was found in dolphins after capture experiments <strong>and</strong> is likely, therefore, to arise inother cetaceans. Muscle activity during pursuit <strong>and</strong> capture can affect blood enzymes – creatininekinase (CK), aspartate aminotransferase (AST), lactate dehydrogenase (LDH), <strong>and</strong> blood ureanitrogen (BUN) <strong>and</strong> potassium levels in cetaceans. Colgrove (1978) diagnosed EM following thetransportation of a bottlenose dolphin, with first signs appearing 22 hours post-transport. Young etal. (1997) found 11 blood <strong>and</strong> serum parameters linked to stress in net-caught wild <strong>and</strong> healthycaptive bottlenose dolphins, which matched those for terrestrial animals suffering from EM. Theauthors conclude that dolphins are susceptible to EM. Thurnbull <strong>and</strong> Cowan (1998) speculate thatthe deaths of small cetaceans following capture for marine collections may be linked to EM.The potential stress effects of whaling <strong>and</strong> the welfare implications for hunted cetaceans73<strong>Dolphin</strong>s may be particularly prone to stress-related cardiomyopathy or contraction b<strong>and</strong> necrosis(CBN). CBN is characterized by lesions associated with hypercontracted myocardial cells, which inturn have been linked to elevated catecholamine concentrations (Reichenach <strong>and</strong> Benditt, quoted inThurnbull <strong>and</strong> Cowan 1998). They can occur following traumatic circumstances ranging frompsychological stress <strong>and</strong> drowning to exertional myopathy <strong>and</strong> are considered to contribute to theirfatal outcome. Identical lesions occurred in a sample of str<strong>and</strong>ed cetaceans of nine species. Theauthors attributed these lesions to physiological <strong>and</strong> psychological stress linked to str<strong>and</strong>ing, disease<strong>and</strong> injury. Elevated CK levels in some dolphins killed during purse-seining are considered indicativeof “muscular exertion or muscle damage” which may result in unobserved post-release mortality(Stuntz <strong>and</strong> Shay 1979 quoted in Curry 1999). More recent evidence for potential stress-relatedinjury or unobserved mortality emerged for dolphins involved in purse seine fishing operations(NOAA 2002).ConclusionsThe degree of exercise imposed on whales during whaling may fall outside the species’ adaptive range.“Even species that have evolved for efficient running, either for predator avoidance or for predation, maydevelop exertional myopathy following intense or prolonged muscular activity associated with extreme stressduring air or ground pursuit.” (Williams <strong>and</strong> Thorne 1996). <strong>Whale</strong>rs depend on gaining a minimumdistance between themselves <strong>and</strong> whales for successful harpooning. From what is known about thebehavioural response of cetaceans to ships, this range is likely to fall below what would naturally bemaintained by wild cetaceans in many cases. Pursuit as part of whaling may, therefore, be stressful
<strong>and</strong> manifest in a series of lethal <strong>and</strong> sub-lethal pathologies. Some whales, which successfully evadebeing harpooned, or are missed, may still suffer fatal syndromes, such as exertional myopathy (EM).Thus, pursuit may increase anthropogenic mortality levels, <strong>and</strong> this has implications for the welfareof all hunted whales, including those that are pursued but escape the harpoon. Such effects may beexacerbated where gravid females or females accompanied by dependent offspring are targeted.Physiological assessments of stress, as part of currently practised lethal whale research, seem plaguedby technical (manner <strong>and</strong> schedule of sampling, h<strong>and</strong>ling <strong>and</strong> storage etc, see Ishikawa, 1996; Øen<strong>and</strong> Walløe 1999) <strong>and</strong> biological problems, which can significantly distort results. A physiologicalstress assessment carried out during current lethal research is therefore fraught with difficulties. Thisincludes problems with partitioning the physiological <strong>and</strong> biochemical effects of pursuit <strong>and</strong> killing,<strong>and</strong> the fact that the trauma of harpooning will mask any effects of pursuit. With regard to EM thefact that clinical signs may not manifest for hours, days, weeks or even months poses additionalproblems.Rather than focus on harpooning alone, the IWC might consider whale hunting to start from when awhale is first sighted to when it is killed. Factors such as pursuit duration <strong>and</strong> speed, distance coveredas well as direction of approach <strong>and</strong> weather condition should be taken into account <strong>and</strong> recorded.Such information, particularly if combined with behavioural data from target <strong>and</strong> non-targetanimals, <strong>and</strong> easily monitored biological parameters, such as respiratory rate, would provide apractical basis for assessing the potential stress-related risks of whaling.74A REVIEW OF THE WELFARE IMPLICATIONS OF MODERN WHALING ACTIVITIESReferencesBartsch, R.C., E.E. Meconnell, G.D. Imes, <strong>and</strong> JM. Schmidt. 1977. A review of exertional rhabdoyolysis in wild<strong>and</strong> domestic animals <strong>and</strong> man. Vet. Pathol. l4:314-324.Basson, P. A., <strong>and</strong> Hofmeyer, J. M. (1973). Mortalities associated with wildlife capture operations. In: TheCapture <strong>and</strong> Care of Wild Animals (E. Young, ed.), pp. 151-153. Human <strong>and</strong> Rousseau Publishers, Cape Town,South Africa.Brambell, F.W.R. (1965) Report on the Technical Committee to enquire into the welfare of livestock kept underintensive husb<strong>and</strong>ry conditions, HMSO, London.Broom, D. M. <strong>and</strong> Johnson, K. G. (1993) Stress <strong>and</strong> Animal Welfare, Chapman <strong>and</strong> Hall.Broom, D.W. (1985) stress, welfare <strong>and</strong> the state of equilibrium. In: Proc. 2nd Eur. Symp. Poult. Welfare. R.M.Wegner (ed.) World Poultry Science Association, Celle, pp72-81.Chalmers, G. A., <strong>and</strong> Barrett, M. W. (1982). Capture myopathy. In: Noninfectious diseases of wildlife (G. L.Hoff, <strong>and</strong> J. W. Davis, eds.). Iowa State University Press, Iowa, US.Colgrove, G. S. (1978). Suspected transportation-associated myopathy in a dolphin. J. Am. Vet. Med.Assoc.173(9): 1121-1123.Cowan, D.F. <strong>and</strong> Walker, W.A. (1979) Disease factors in Stenella attenuata <strong>and</strong> Stenella longirostris taken in theeastern tropical Pacific yellowfin tuna purse-seine fishery. Southwest Fisheries Science Center, National MarineFisheries Service, NOAA, Administrative Report No. LJ-79-32C.Curry, B.E. (1999) Stress in Marine Mammals: The Potential Influence of Fishery-Induced Stress on <strong>Dolphin</strong>sin the Eastern Tropical Pacific Ocean. NOAA Technical Memor<strong>and</strong>um NMFS, U.S. Dept. of Commerce,NOAA-TM-NMFS-SWFSC-260.
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ForewordWhales are highly evolved a
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1 Executive SummaryThis review exam
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Japan currently whales in the Antar
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Otto, K. 1997. Animal Pain Behaviou
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Social behaviourMother-calf pairsOn
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to store and pass on information to
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15 Whaling and welfarePhilippa Brak
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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