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Fisheries Acoustics Session 6: Fish behaviour and physiology with respect to target strength Rapp. P.-v. Rcun. Cons. int. Explor. Mer, 189: 233—244. 1990 Swimbladder “behaviour” and target strength J. H. S. Blaxter and R. S. Batty Blaxter, J.H .S., and Batty, R.S. 1990. Swimbladder "behaviour” and target strength. - Rapp. P.-v. Réun. Cons. int. Explor. Mer, 189: 233-244. The target strength (TS) of fish is higher when a swimbladder is present. TS depends on the swimbladder volume which increases with fish size and varies with depth. Changes of volume occur during vertical migration; the volume then remains rather stable in physostomes apart from slow losses of gas by diffusion, but physoclists secrete or absorb gas, so tending to restore the original volume. Such adaptation to a depth change is slow compared with the speed of movement of vertical migrants whose TS will change at dusk and dawn. TS also varies with the orientation of the fish to the echo-sounding beam and may vary with its swimming speed. The highest TS is found with the head tilted down 5-10°. The swimbladder axis is, however, tilted up by the same amount so that it lies normal to the beam. In many species there are day:night differences in activity, swimming speed, tilt angle, density, and vertical stacking, all affecting TS. It is unfortunate that TS depends on a labile organ with a role in buoyancy and posture. J. H. S. Blaxter and R. S. Batty: Dunstaffnage Marine Laboratory, P. O. Box 3, Oban, Argyll, Scotland. (3) physoclists which have a closed swimbladder, a gas Introduction gland for gas secretion, and an occlusible oval for Since the bodies of fish have an acoustic impedance gas resorption; close to that of water as much as 90 % of the backscat- (4) some bathypelagic species which have a swimblad­ tering of sound from echo sounders or sonars depends der filled with oil or invested with fat. on gas-filled structures such as the swimbladder. The extent of backscattering also depends, amongst other In the past it has been supposed that the swimbladder’s things, on the cross-sectional area of the gas phase main role is to confer neutral buoyancy on the fish and perpendicular to the incident acoustic beam, and it so mechanisms have evolved to maintain its volume at would be enhanced if the gas-filled structure were of a some optimum value, with a clear requirement for a size and shape to resonate at the frequency of enson- compensation mechanism to allow the fish to make ification. vertical movements involving changes of hydrostatic Although the swimbladder is the main gas-filled pressure. A neat solution to this problem would seem to structure, gas could be present in the gut as a result of be a non-compressible flotation organ such as the squa- the activity of micro-organisms while in all clupeoids lene-filled liver of sharks or the oil-filled or fat-invested there are gas-filled paired otic bullae within the skull. It swimbladder of some bathypelagic species (Marshall, is likely that the volumes concerned will be small com­ 1960; Butler and Pearcy, 1972) or to reduce ossification pared with the swimbladder. and increase the water content of the flesh (Denton and For our purpose fish can be grouped into four cate­ Marshall, 1958; Blaxter el al., 1971) or to increase the gories: oil content of the bones (Bone, 1972; Lee et al., 1975). The swimbladder may, however, have other roles to (1) those with no swimbladder, especially flatfish, sand- play in hearing and sound production (Blaxter and Tyt- eels, Ammodytes spp., and the Atlantic mackerel ler, 1978) that require it to be gas-filled. (Scomber scombrus) ; Most physostomes obtain gas by swallowing air at the (2) physostomes which have an open swimbladder with surface. During a subsequent descent the swimbladder a duct or ducts to the exterior and which usually contracts, and it is unlikely that a fish will be neutrally lack a gas secretion mechanism, e.g., clupeoids, buoyant at any depth unless it has positive buoyancy at salmonids, and cyprinids; the surface. There is no evidence for such a phenom- 233 Table 1. Swimbladder volumes. Species Stage/size Volumes as % of fish weight Reference ± s.d. (n) Cod 13-50 cm 3.6±0.46 (11) Hawkins (1977) Cod 25-50 cm 4.5-5.4 Harden Jones and Scholes (1985) Pollack 32-45 cm 2.8±0.33 (13) Foote (1985a) Whiting 24 g, 62 g 4.4, 2.8 Alexander (1959b) Pout 69 g 4.9 Alexander (1959b) Herring 9-25 g 4.1a±0.6 (16) Brawn (1962) Herring 45-200 g 4.3-2.31’ Ona (1984a) Herring Larvae 0.2-1.6 (13) Blaxter and Batty (1984) (see Fig. 1) Juveniles 1 Adults J 1.5-5 (46) Spawners 0(6) “4.2 % expressed as % of fish volume. hsb vol. = 0.017 wt + 1.157 (226). enon although some additional buoyancy may be ob­ Swimbladder volume tained from a high tissue fat content. During an ascent there should be no excess gas and no need therefore to There is abundant evidence, which will not be reviewed void gas from the swimbladder. Why some species such here, that the target strength of fish is related to their as the herring do release gas (Sundnes and Bratland. size. Furthermore, size-for-size fish without swimblad- 1972) is something of a mystery. Nevertheless all clupe­ ders such as mackerel and sandeels have lower target oids have one or more ducts from the swimbladder to strengths (Foote, 1980a; Armstrong and Edwards, the exterior (Whitehead and Blaxter, 1989). While the 1985). Implicit in these findings is a relationship be­ pneumatic duct from the gut can be seen as the route for tween fish size, swimbladder volume, and target swallowed gas to enter the swimbladder, the anal duct strength. In order to provide neutral buoyancy the seems to involve gas release to the exterior. A few swimbladder should occupy 5 % of the body volume in a physostomes such as some eels (Fänge, 1983; Kleckner, fully marine fish and 7% in a freshwater fish (Harden 1980) and goldfish Carassius auratus (Overfield and Jones and Marshall, 1953). Although many of their data Kylstra, 1971) can secrete gas. and some of Alexander's (1959a. b) support this as­ The physoclists are much more constrained in their sertion. other data (Horn, 1975; Brooks. 1977) do not. vertical movements. A descent presents no problems There is also considerable evidence from commercial that are different from a physostome’s, i.e., an increas­ marine species recently studied that: ing negative buoyancy. If a physoclist is adapted to a ( 1 ) swimbladder volumes are often much less than 5 % ; particular depth, as it ascends from that depth the swim­ (2) there is a wide variation in swimbladder volume in a bladder expands. Gas is then resorbed, but if the ascent given fish population; is not controlled the swimbladder may rupture. Adapta­ (3) there are effects of age and spawning condition. tions are to be found that increase the depth range by improving the gas secretion/resorption mechanisms. In Some data are given in Table 1 and Figure 1. particular the rete of the gas gland may be elongated in There is other less direct evidence that swimbladder deep-sea species (Marshall, I960, 1972; Horn, 1975; volume might be very variable in terms of its impor­ Kleckner, 1980). tance in target-strength measurement. Sand and Haw­ It is unfortunate that the target strength of fish is kins (1973) found that the resonance frequencies of cod largely determined by such a labile organ. The volume (Gadus morhua) swimbladders were highly variable at of the swimbladder depends on the size of the fish and the depth of adaptation, being 100% to 700% of the the extent of inflation, which in turn depends on the resonance frequency of a free gas bubble at the adapta­ recent depth “history” of the fish. The cross-sectional tion depth. area depends on the shape of the swimbladder and the Other factors may be related to swimbladder volume, angle of tilt of the fish to the horizontal. The resonance in particular the fat content of the fish. Fat content frequency of the swimbladder depends on its shape, varies with gonad maturation state, season, or feeding wall thickness, any differential internal gas pressure, conditions (Love. 1970); in herring (Clupea harengus) and possibly its mode of attachment to the body wall. there is an inverse relationship between fat content and These problems will be considered in the following swimbladder volume (Brawn, 1969; Ona, 1984a, Fig. 2) pages. so that swimbladder volume (and target strength) can 234 Figure 1. Swimbladder volume related to size of herring (redrawn from Blaxter and Batty, 1984). 4- *— 001 01 10 100 9 i---- 1— i— i— i— — 1---- 1------ 1-- 1— I------- 7 10 15 20 25 30 cm Fish size Figure 2. Swimbladder volume as % body weight related to oil content as % body weight of herring, x Atlantic herring and 5 O Pacific herring, from Brawn (1969); oo 0 Norwegian herring, from Ona (1984a). I? 3 »•V 2 0 510 15 20 25 Oil content (%) Table 2. Swimbladder pressure differentials. Species Pressure differential in cm H:0: Reference + above ambient — below ambient Cod + <1.0-2.4a Sand and Hawkins (1974) Cod - 600-900b Sundnes and Gytre (1972) Herring + 15 Brawn (1962) Herring + 17.4± 10.9 Blaxter and Batty (1984) Sockeye salmon + 0.3± 7.5 Harvey el al. (1968) Pinfish + 3.6 McCutcheon (1958) Most cyprinids + 27-41 Alexander (1959a) Bream + 147 Alexander (1959a) Roach + 95 Alexander (1959a) Rudd + 83 Alexander (1959a) “Even if internal pressure is doubled by gas injection, internal excess pressure rapidly falls to +6.2 cm H;0.
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