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ICES Journal of Marine Science: Journal du Conseil 2005 62(5):984-995; doi:10.1016/j.icesjms.2005.03.010
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© 2005 International Council for the Exploration of the Sea

Acoustic backscattering by Atlantic mackerel as being representative of fish that lack a swimbladder. Backscattering by individual fish

Natalia Gorskaa,*, Egil Onab and Rolf Korneliussenb

a Institute of Oceanology of Polish Academy of Sciences ul. Powstanców Warszawy 55, PL-81-712 Sopot, Poland
b Institute of Marine Research PO Box 1870, 5817 Bergen, Norway

*Correspondence to N. Gorska: tel: +48 58 551 72 81; fax: +48 58 551 21 30. e-mail: gorska{at}iopan.gda.pl.

Developing acoustic methods for the identification of fish remains a long-term objective of fisheries acoustics. The accuracy of abundance estimation may be increased when the acoustic-scattering characteristics of the fish are known, including their expected variability and uncertainty. The modelling approach is valuable during the process of interpreting multi-frequency echograms. This paper attempts to improve the understanding of sound backscattering of fish without a swimbladder, here represented by Atlantic mackerel (Scomber scombrus). Our approach includes results from modelling as well as comparisons with field data. There will be two papers. The first is a study of the non-averaged backscattering characteristics. This initial analysis is important for the understanding of the averaged backscattering cross-section, which will be considered in the second paper. In that paper the relative importance of bones in acoustic backscattering at higher frequencies will be verified.

Keywords: backbone, fish flesh, modelling, sound backscattering by mackerel

Received 30 May 2004; accepted 20 March 2005.


    Introduction
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 
Acoustic surveys are widely used for the stock assessment of many pelagic fish species (MacLennan, 1990). A thorough understanding of the mechanisms of sound scattering by fish, including the understanding of the contribution of the various anatomical features to the overall backscattering, is required to improve present acoustic methods of fish species identification (Horne, 2000; Reeder et al., 2004). Numerical modelling of sound backscattering by fish (see the review presented by Horne and Clay, 1998; Reeder et al., 2004) and controlled accurate laboratory measurements (Sun et al., 1985; Nash et al., 1987; Barr, 2001; Reeder et al., 2004) have been carried out to gain more knowledge of the process by which sound is scattered by selected fish species.

The paper stems from the need for proper assessment from acoustic backscattering data of the abundance of the economically important fish species, Atlantic mackerel (Scomber scombrus). Multi-frequency measurements on mackerel made at 18, 38, 70, 120, 200, and 364 kHz demonstrated that backscatter intensities at different frequencies (i.e. frequency response) have a specific pattern. Korneliussen and Ona (2002) found that the frequency response for mackerel was flat at 18, 38, and 120 kHz and then increased in intensity towards 200 kHz. Unpublished backscatter measurements made in net pens and at sea confirmed frequency-independent backscatter at 18, 38, and 70 kHz, and stronger backscatter at 200 kHz. In the measurement series, backscatter intensities at 120 kHz were more variable. In some series it was similar to that at 38 kHz, and in others the increase in intensity lay anywhere in the range up to double the intensity at 38 kHz. To gain an understanding of this peculiar frequency response and to study its stability, we wanted to see if this particular frequency spectrum was consistent for Atlantic mackerel and could be used for acoustic identification.

In order to begin to answer this question the backscattering needed investigation. Most modelling to date has been done for fish with swimbladders where that organ is the source of most backscatter per se (Reeder et al., 2004). Backscattering by other anatomical features, which can be important for sound incidence, are not normal to the surface of swimbladder, as well as for fish without swimbladders, are still not well known. In this paper the need to understand the impact of the different anatomical components on total backscattering and the factors which could modify mackerel target strength is addressed. Furthermore, the contribution of mackerel body and backbone to the overall backscattering characteristics across a selected frequency range is examined. The effects of orientation and the morphological condition of the fish are also considered. The modelling was done using the Distorted Wave, Born-Approximation (DWBA) (Chu et al., 1993; Stanton et al., 1993, 1998; Stanton and Chu, 2000) and the Modal-Based, Deformed-Cylinder Model (MB-DCM) (Stanton, 1988a, b, 1989). A high-resolution morphology of flesh, based on measurements of the mackerel body, was considered. To study mackerel-backbone backscatter, straight and uniformly bent cylinders were used to describe its shape. The typical observed backscattering-frequency responses for mackerel are then explained in theoretical terms. The use of the frequency response in mackerel identification is justified.

The final outcome of the study should be to explain the frequency response of mackerel shoals. The frequency response is defined by the averaged backscattering cross-section of mackerel at different frequencies. However, for a better understanding of the averaged characteristics the modelling has to be done first at an individual level, i.e. for the backscattering cross-section before averaging takes place. The modelling section has, therefore, been divided into two aspects; first, backscattering by individual mackerel (this paper), and second, average backscattering by mackerel (in a second paper).


    Material and methods
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 
Main backscattering equation: backscattering by fish flesh
The DWBA-based, deformed-cylinder model (Chu et al., 1993; Stanton et al., 1993, 1998; Stanton and Chu, 2000) has been used to describe backscatter by mackerel flesh. This model is assumed to be applicable because: (i) mackerel flesh has material properties that are similar, within a few per cent, to those of the surrounding water, so mackerel can be referred to as a weak scattering target; and (ii) the mackerel body has a cross-section that can be described, at first order, as circular.

Using the analytical solution (Equation (6) from Stanton and Chu, 2000), some derivations have been made to apply it to mackerel. A solution has been obtained for mackerel backscattering length, fbscfl, normalized by length, lfl.


Formula 1

(1)
where the F and G functions can be written as


Formula 2

(2)
and


Formula 3

(3)
Here the parameter {gamma} is expressed as {gamma} = 2(kfla0). Ratio values of f(u) = a(u)/a0 describe the variability of the cross-sectional radius of mackerel body a(u), normalized by the maximum radius a0, along the longitudinal axis of the fish (see Figure 1a). The variable u denotes the x-variable along the axis, normalized by fish length lfl (u = x/lfl). The symbol a(x) in the argument of the F function in Equation (1) refers to the sensitivity of the function to the shape of the mackerel body, or more precisely, to the dependence of the normalized radius f(u) on u.


Figure 1
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Figure 1 Backscattering geometry. Backscattering by fish flesh (a), by backbone, modelled as straight (b), and bent cylinders (c).

 
Equations (1), (2), (3) incorporate the fact that the cross-sectional radius varies only along the axis of the body. Although fat content is known to differ from dorsal to anterior (i.e. back to stomach), sufficient data on the properties of mackerel-body components are not available and have been treated as being homogeneous.

Backscatter from the backbone
The Modal-Based, Deformed-Cylinder Model (Stanton, 1988a, b, 1989) was used to study backscattering by the backbone. This model is applicable to elongated deformed cylinders with large aspect ratios and where the direction of incidence and scattering is normal or near-normal to the tangent of the axis of the cylinder. The measurements, made on the mackerel cruise by RV "G. O. Sars", October 2002, demonstrated the near-circularity of backbone cross-section and the large backbone elongation: the aspect ratio of backbone, i.e. the length/radius ratio of the backbone, can reach values of 140–160, so justifying the use of the model.

The backbone was modelled in two different ways: (i) as an elastic, solid, straight cylinder of uniform composition (see Figure 1b); and (ii) as an elastic, solid, uniformly bent cylinder of constant radius of curvature of its axis, constant cross-section radius, and constant composition (see Figure 1c).

These simple geometric shapes were chosen because approximate analytical–numerical MB-DCM solutions have been obtained in similar cases (cf. Stanton, 1988b, 1989).

Equation (7) of Stanton (1988b) was used to model the backscattering length of a straight cylinder. Equation (8) of Stanton (1989) was employed to derive solutions for the backscattering length of a bent cylinder. The modal coefficients, defined by Equation (1) of Stanton (1988b) or by Equations (22)–(25) of Faran (1951), were used in both cases.

The backscattering length of the backbone can be expressed by


Formula 4

(4)
or


Formula 5

(5)

The scattering phase angles {eta}m, {delta}m, {Phi}m, {alpha}m, ßm, {xi}m involved in modal sum coefficients are functions of acoustic frequency, cylinder cross-section radius, sound-speed contrasts for compressional (hcom) and shear (hsh) waves (i.e. parameters x, x1, x2), and the density contrast (g) inside the backbone. They can be expressed as:


Formula 6

(6)


Formula 7

(7)


Formula 8

(8)


Formula 9

(9)


Formula 10

(10)
and


Formula 11

(11)

Modelling parameters
Acoustic backscattering by fish is a complex function of the geometrical shape of various body components, the properties of their materials, the orientation of the fish in space, and the acoustic frequency (Horne, 2003). In the modelling process, we considered the following factors.

Size and frequency
To include a full range of frequencies and lengths of mackerel bodies and backbones, ka0 values from 0 to 40 were used for fish body and ka values from 0 to 2.5 for fish backbones. These ranges were based on the results of a large number of body-size measurements (Korneliussen et al., 2003) and mackerel-morphology studies conducted during October 2002 on the RV "G. O. Sars" (2) and October 2003 on the RV "G.O. Sars" (3).

Animal morphology
The digitizing of fish-body morphology needs to include the acoustic properties of the flesh and backbone (the spatial, three-dimensional distribution of the contrasts gfl, g and hfl, hcom, hsh) and the three-dimensional shape. Two classes of mackerel, differing in body shape, were chosen for analysis – thick and lean mackerel. Examples of these classes are presented in Figure 2a, b. In Figure 2c, the digitizing of the outer boundary of mackerel bodies is shown using light and dark grey lines. The radius of the mackerel body and the x coordinate, both normalized by the fish length lfl, are indicated in the vertical and horizontal axes. Aspect-ratio values for fish flesh were efl= 10.54 for lean fish and efl= 8.5 for thick fish. We assumed independence of the aspect ratio of the fish length for the two classes.


Figure 2
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Figure 2 Mackerel (thick fish (a) and lean fish (b)). The digitized outer boundary of the mackerel body for different classes of mackerel (c).

 
Simple geometric shapes (e.g. straight and uniformly bent cylinders) were considered when modelling the backbone. Backbone dimensions (a and l) and the aspect ratio eb, obtained in mackerel morphology studies conducted during October 2002 on the RV "G. O. Sars" (2) were used in the computations. The information on the change in angle between the main axis of the body and the backbone may be important for the study. A 3° angle was measured between the sagittal axis of the body and the backbone.

There are few data on sound-speed and density contrasts for Atlantic mackerel. According to Lockwood (1988), the fat content of mackerel varies from 10% in June to 25–30% in October, which is in agreement with a typical fat content of 5.5% for mackerel landed in Norway in June 2002, increasing to 20% in July–September (personal communication with the Norwegian Directorate of Fisheries). Accounting for the fat content varying from 5% to 30%, and the empirical relationship between mackerel fat content Ff and the density contrast gfl, gfl = 1.03 – 0.094Ff, found in our own measurements, a range of variability in density contrast of 1.002–1.025 is considered appropriate for mackerel flesh. According to Sigfusson et al. (2001), the contrast varies between 1.002 and 1.025. The measured sound-speed contrast in mackerel flesh hfl varied around 1.025 both in our own measurements and in Sigfusson et al. (2001), where it was found to be hfl = 1.034 – 0.125Ff at 25°C. The measured density contrast of backbone 1.10 ± 0.05 was used in the computations. Sound-speed contrast measurements for shear and compressional waves were not performed. Given the lack of available information on these two parameters, we assumed sound-speed contrasts of 0.1–1.0 for shear waves and 1.3–2.0 for compressional waves.

Length and orientation statistical distributions
Based on samples from trawl and purse-seine catches, mean total body lengths of 30–40 cm with a standard deviation of 10% were used in the analysis. Given the lack of mackerel tilt-angle measurements, we assumed a narrow distribution of orientations, based on visual observations of behaviour in net pens.


    Results and discussion
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 
Sensitivity analysis for mackerel flesh
The possible effect of changes in mackerel morphology on backscattering was analysed using expressions for the normalized backscattering length, fbscfl/lfl (Equations (1), (2), (3)).

The ka0-dependencies of reduced target strength, RTS = 10 log({sigma}bscfl/lfl2), are presented in Figure 3 by grey and black lines for lean and thick mackerel, respectively. The density contrast gfl = 1.03 was used in both calculations at normal incidence (i.e. ß = 0). Comparison of the two curves in the figure demonstrates that the reduced target strength is slightly dependent on the geometrical shape of the mackerel body. The reduced target strength is sensitive to the sound-speed contrast, which influences both the amplitude of the oscillations and their period. The period of oscillations over ka0 is defined mainly by the difference in speed of sound (hfl), and it is proportional to hfl. The density contrast influences only the value of the RTS, not the periodicity of its oscillations. For the density contrast variation from 1 to 1.03, the reduced target strength increases 6–12 dB, depending on the value of the sound-speed contrast.


Figure 3
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Figure 3 A comparison of the ka0-dependencies for the flesh of mackerel of different geometrical shapes. Maximum dorsal incidence. Calculation parameters: sound-speed of 1.025, density contrast of 1.03 and aspect ratios of 8.5 and 10.54 for thick and lean mackerel, respectively. The arrows indicate the values of ka0-parameter, for which the calculations of the directivity pattern, shown in Figure 4, were made.

 
Sensitivity analysis of the orientation dependence (Figure 4) shows that the shape of the directivity pattern depends strongly on the value of ka0. There is a minimum in the pattern for both curves at ß = 0 (Figure 4a). While the minimum in the mackerel-directivity pattern at ß = 0 is surprising, it is consistent with the shape (minimum) of the ka0-dependence, as indicated by solid arrows in Figure 3, at ka0 = 2.5 and ka0 = 12.15, at which the calculations were made. The possibility of the minimum in the directivity pattern at normal dorsal incidence is also supported by the results of earlier measurements (Foote and Nakken, 1978). A maximum and local minimum at ß = 0 for curves in Figure 4b are explained by the shape of the ka0-dependence at ka0 = 1.5 (ka0 of the first maximum of RTS) and ka0 = 11.45 (ka0 is close to the ka0 of seventh maximum of RTS), indicated by dotted arrows in Figure 3. Comparison between dotted and solid curves in Figure 4 demonstrates that the width of the lobes of the directivity pattern is controlled by the ka0 parameter. For the larger ka0, the individual lobe width is smaller and number of lobes is larger.


Figure 4
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Figure 4 Directivity pattern for flesh of thick mackerel. The corresponding values of ka0 are shown in the legend. The calculations were performed for thick mackerel with an aspect ratio of 8.5, sound-speed contrast of 1.025 and density contrast of 1.03.

 
The calculations of the orientation dependence of the target strength of mackerel flesh TS = 10 log({sigma}bscfl) (Figure 5) show that the dependence is more complex for higher frequencies. The number of the directivity-pattern lobes increases and the width of an individual lobe decreases with frequency.


Figure 5
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Figure 5 The directivity pattern of thick mackerel at frequencies of 18 (a), 38 (b), 120 (c), and 200 kHz (d). The calculations were made for thick mackerel of total length 40 cm and aspect ratio 8.5, sound-speed contrast of 1.025, and density contrast of 1.03.

 
Sensitivity analysis for mackerel backbone
The analysis demonstrates (Figure 6) that the resonance and anti-resonance structure (the ka of maxima (resonances) and minima (anti-resonances) of the target strength, the frequency of the occurrence of resonances and anti-resonances, the width of the resonance and anti-resonance peaks and their amplitudes) are all extremely sensitive to the sound-speed contrast of shear waves in backbone. For the larger contrast the resonances and anti-resonances are more frequent and narrow, and their amplitude is higher. The similar qualitative dependences are observed for sound backscattering by elastic solid spheres (Hickling, 1962).


Figure 6
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Figure 6 Backscattering by the backbone of individual mackerel. Sensitivity to the sound-speed contrast of shear waves. Maximum dorsal incidence ({Theta} = 0). The sound-speed contrast of compressional wave of 1.5, density contrast of 1.1, and aspect ratio 80 were taken for the calculations, which were made for a backbone modelled as a straight cylinder.

 
The dependence of the reduced target strength on ka is illustrated for various sound-speed contrasts of compressional waves in Figure 7. Individual plots refer to different sound-speed contrasts of shear waves: 0.3, 0.5, and 0.9 – from top to bottom. The sound-speed contrast of compressional waves does not impact the width of the peaks of resonances and anti-resonances and their positions over ka-axis, but influences the value of the reduced target strength of backbone. The value increases with the contrast. The impact depends on the ka parameter.


Figure 7
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Figure 7 Backscattering by the backbone of individual mackerel. Sensitivity to the sound-speed contrast of compressional waves, which are indicated in the legend. Maximum dorsal incidence ({Theta} = 0). Different plots are obtained for various sound-speed contrast of shear waves, presented in plots 0.3 (a), 0.5 (b), 0.9 (c). The density contrast 1.1 and aspect ratio 80 were used for the calculations that were made for a backbone modelled as a straight cylinder.

 
Figure 8 illustrates the sensitivity of the reduced target strength of backbone to the backbone-density contrast. The density contrast does not influence the resonance and anti-resonance structure of the curves and defines only the magnitude of the reduced target strength. The magnitude increases with increasing density contrast.


Figure 8
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Figure 8 Backscattering by the backbone of individual mackerel. Influence of the density contrast. Maximum dorsal incidence ({Theta} = 0). The values are shown in the legend. The calculations were made for a backbone modelled as a straight cylinder with a sound-speed contrast for compressional waves of 1.5 and shear waves of 0.3, and an aspect ratio of 80.

 
The sensitivity of backscatter to the backbone shape is demonstrated in Figure 9. The three curves shown refer to various backbone geometries viz. from top to bottom, straight cylinder and bent cylinders with ratios of l/(2{rho}c) 0.1 and 0.2, respectively. Only the magnitude of the reduced target strength is sensitive to the shape of the backbone. The more curved the backbone, the smaller the level.


Figure 9
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Figure 9 Backscattering by the backbone of individual mackerel. The influence of the geometrical shape of backbone. Maximum dorsal incidence ({Theta} = 0). The calculations were made for a sound-speed contrast for shear waves of 0.3, compressional waves of 1.3, density contrast of 1.1, and aspect ratio 100. For the bent-cylinder backbone the values of l/(2{rho}c) parameter are shown in the legend.

 

    Conclusions
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 
A model has been developed to describe sound backscattering by Atlantic mackerel flesh and backbone. It has been shown that:
  1. in the case of normal sound incidence, the value of RTS is defined mainly by the sound-speed and density contrasts, and to a lesser extent by the geometrical shape of the body, while the periodicity of its oscillations over ka0 depends only on the sound-speed contrast: period is proportional to hfl.
  2. the features of the directivity pattern of a fish body are highly dependent on the ka0 parameter: the ka0-value in regard to the ka0 of maxima and minima of the ka0-dependence of RTS is important. Minimum (maximum) of the directivity pattern is observed for the normal incidence at the ka0 of minima (maxima).
  3. the ka of maxima (resonances) and minima (anti-resonances) of the backbone target strength, the frequency of the occurrence of resonances and anti-resonances, the width of the resonance and anti-resonance peaks, and their amplitudes are mainly defined by the sound-speed contrast of shear waves. Sound-speed contrasts of compressional waves and density contrast influence the value of the target strength of fish backbone. The value is also sensitive to the degree of curvature of the cylinder backbone.
The results obtained for individual mackerel will be useful in the analysis of the averaged backscattering cross-section of mackerel aggregations and thus in the explanation of the observed frequency response. Since the lack of a swimbladder is the main acoustic feature of mackerel, modelling and measurements of sound backscattering by mackerel can increase our knowledge of scattering from other pelagic species that do not have swimbladders. Moreover, since the swimbladder of physostomous fish is compressed during descent, there is a possibility that backscatter from physostomous fish at depth is closer to that of mackerel.


    List of symbols
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 

Formula Backscattering cross-section of flesh
Formula Backscattering cross-section of backbone
Formula Backscattering length of flesh
Formula Backscattering length of backbone
lfl Length of body (from the top of the head to the start of tail) Figure 1a
l Length of backbone
a(x) Cross-sectional radius of body, variable along the longitudinal axis Figure 1a
a0 Maximum of a(x)
a Radius of backbone cylinder
efl = lfl/a0 Aspect ratio of body of fish
eb = l/a Aspect ratio of backbone cylinder
{rho}c Radius of curvature of the axis of backbone bent cylinder
{gamma}max l/(2{rho}c)
hfl = cfl/c Sound-speed contrast in fish flesh
hcom = ccom/c Sound-speed contrast of compressional wave in backbone
hsh = csh/c Sound-speed contrast of shear wave in backbone
c Speed of sound in surrounding seawater
cfl Speed of sound in flesh
ccom Speed of compressional sound wave in backbone
csh Speed of shear sound wave in backbone
gfl = {rho}fl/{rho} Density contrast of fish flesh
{rho} Density of surrounding water
{rho}fl Density of fish flesh
g = {rho}1/{rho} Density contrast of backbone
{rho}1 Density of backbone
f Carrier frequency of sound
k = 2{pi}f/c Acoustic wavenumber in surrounding seawater
kfl = 2{pi}f/cfl Acoustic wavenumber in flesh
kcom = 2{pi}f/ccom Acoustic wavenumber of compressional wave in backbone
ksh =2{pi}f/csh Acoustic wavenumber of shear wave in backbone
x ka cos {Theta}
x1 kcoma
x2 ksha
{varepsilon}m Neumann's number: {varepsilon}0 = 1, {varepsilon}m>0 = 2
{eta}m, {delta}m, {Phi}m, {alpha}m, ßm, {xi}m Scattering phase angles
Jm( ) Bessel function of the first kind of order m
Nm( ) Bessel function of the second kind of order m
Jm'( ), Nm'( ) Derivative with respect to argument of Jm( ) or Nm( )
ß Angle between the direction of incidence and the normal to the longitudinal axis of the fish Figure 1a
{Theta} Angle between the direction of incidence and the normal to the backbone longitudinal axis of the fish Figure 1b
{Delta} kl sin {Theta}
Ff Fat fraction of total weight


    Acknowledgements
 
This work has been partially supported by the Institute of Oceanology, Polish Academy of Sciences (sponsor programme 2.7), the Research Council of Norway (Grant No. 133657/120), and the European project SIMFAMI (Grant No. Q5RS-2001-02054).


    References
 Top
 Introduction
 Material and methods
 Results and discussion
 Conclusions
 List of symbols
 References
 

    Barr R. (2001) A design study of an acoustic system suitable for differentiating between orange roughy and other New Zealand deep-water species. Journal of the Acoustical Society of America 109:164–178.[CrossRef][Web of Science]

    Chu D., Foote K.G., Stanton T.K. (1993) Further analysis of target-strength measurements of Antarctic krill at 38 and 120 kHz. Comparison with deformed-cylinder model and inference of orientation distribution. Journal of the Acoustical Society of America 93:2985–2988.[CrossRef][Web of Science]

    Faran J.J. (1951) Sound scattering by solid cylinders and spheres. Journal of the Acoustical Society of America 23:405–418.[CrossRef][Web of Science]

    Foote K.G. and Nakken O. (1978) Dorsal-aspect, target-strength functions of six fishes at two ultrasonic frequencies. Fisken og Havet Series B 1978:31–96.

    Hickling R. (1962) Analysis of echoes from solid elastic sphere in water. Journal of the Acoustical Society of America 34:1582–1592.[CrossRef][Web of Science]

    Horne J.K. (2000) Acoustic approaches to remote species identification: a review. Fisheries Oceanography 9:356–371.[CrossRef][Web of Science]

    Horne J.K. (2003) The influence of ontogeny, physiology, and behaviour on the target strength of walleye pollock (Theragra chalcogramma). ICES Journal of Marine Science 60:1063–1074.[Abstract/Free Full Text]

    Horne J.K. and Clay C.S. (1998) Sonar systems and aquatic organisms: matching equipment and model parameters. Canadian Journal of Fisheries and Aquatic Sciences 55:1296–1306.

    Korneliussen R.J. and Ona E. (2002) An operational system for processing and visualizing multi-frequency acoustic data. ICES Journal of Marine Science 59:293–313.[Abstract/Free Full Text]

    Korneliussen R., Skagen D.W., Slotte A., Knutsen T. (2003) Cruise summary report of survey, 112. 14 pp. (In Norwegian).

    Lockwood S.J. (1998) The Mackerel. Its Biology, Assessment and the Management of a Fishery(Fishing News Books Ltd., Oxford).

    MacLennan D.N. (1990) Acoustical measurement of fish abundance. Journal of the Acoustical Society of America 87:1–15.[CrossRef][Web of Science]

    Nash R.D.M., Sun Y., Clay C.S. (1987) High-resolution acoustic structure of fish. Journal du Conseil International pour l'Exploration de la Mer 43:23–37.

    Reeder D.B., Jech J.M., Stanton T.K. (2004) Broadband acoustic backscatter and high-resolution morphology of fish: measurement and modelling. Journal of the Acoustical Society of America 116:747–761.[CrossRef][Web of Science][Medline]

    Sigfusson H., Decker E.A., McClements D.J. (2001) Ultrasonic characterization of Atlantic mackerel (Scomber scombrus). Food Research International 34:15–23.[CrossRef][Web of Science]

    Stanton T.K. (1988) Sound scattering by cylinders of finite length. I. Fluid cylinders. Journal of the Acoustical Society of America 83:55–63.[CrossRef][Web of Science]

    Stanton T.K. (1988) Sound scattering by cylinders of finite length. II. Elastic cylinders. Journal of the Acoustical Society of America 83:64–67.[CrossRef][Web of Science]

    Stanton T.K. (1989) Sound scattering by cylinders of finite length. III. Deformed cylinders. Journal of the Acoustical Society of America 86:691–705.[CrossRef][Web of Science]

    Stanton T.K. and Chu D. (2000) Review and recommendations for the modelling of acoustic scattering by fluid-like, elongated zooplankton: euphausiids and copepods. ICES Journal of Marine Science 57:793–807.[Abstract/Free Full Text]

    Stanton T.K., Chu D., Wiebe P.H. (1998) Sound scattering by several zooplankton groups. II. Scattering models. Journal of the Acoustical Society of America 103:236–253.[CrossRef][Web of Science][Medline]

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    Sun Y., Nash R., Clay C.S. (1985) Acoustic measurements of the anatomy of fish at 220 kHz. Journal of the Acoustical Society of America 78:1772–1776.[CrossRef][Web of Science]


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Classifying fish schools and estimating their species proportions in fishery-acoustic surveys
ICES J. Mar. Sci., July 1, 2009; 66(6): 1136 - 1142.
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ICES J. Mar. Sci.Home page
H. Pena and K. G. Foote
Modelling the target strength of Trachurus symmetricus murphyi based on high-resolution swimbladder morphometry using an MRI scanner
ICES J. Mar. Sci., December 1, 2008; 65(9): 1751 - 1761.
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ICES J. Mar. Sci.Home page
N. Gorska, R. J. Korneliussen, and E. Ona
Acoustic backscatter by schools of adult Atlantic mackerel
ICES J. Mar. Sci., September 1, 2007; 64(6): 1145 - 1151.
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