ICES Journal of Marine Science Advance Access published February 12, 2009

Page 1 of 8

A Bayesian approach to estimating target strength

Sascha M. M. Fassler, Andrew S. Brierley, and Paul G. Fernandes

Fassler, S. M. M., Brierley, A. S., and Fernandes, P. G. 2009. A Bayesian approach to estimating target strength. ICES Journal of Marine Science,
66: 000 000.
Currently, conventional models of target strength (TS) vs. sh length, based on empirical measurements, are used to estimate sh
density from integrated acoustic data. These models estimate a mean TS, averaged over variables that modulate sh TS (tilt angle,
physiology, and morphology); they do not include information about the uncertainty of the mean TS, which could be propagated
through to estimates of sh abundance. We use Bayesian methods, together with theoretical TS models and in situ TS data, to deter-
mine the uncertainty in TS estimates of Atlantic herring (Clupea harengus). Priors for model parameters (surface swimbladder volume,
tilt angle, and s.d. of the mean TS) were used to estimate posterior parameter distributions and subsequently build a probabilistic TS
model. The sensitivity of herring abundance estimates to variation in the Bayesian TS model was also evaluated. The abundance of
North Sea herring from the area covered by the Scottish acoustic survey component was estimated using both the conventional
TS length formula (5.34109 sh) and the Bayesian TS model (mean 3.17109 sh): this difference was probably because of
the particular scattering model employed and the data used in the Bayesian model. The study demonstrates the relative importance
of potential bias and precision of TS estimation and how the latter can be so much less important than the former.
Keywords: acoustic target strength, Bayesian statistics, herring, survey uncertainty.
Received 9 August 2008; accepted 7 November 2008.
S. M. M. Fassler and A. S. Brierley: Gatty Marine Laboratory, University of St Andrews, St Andrews, Fife KY16 8LB, Scotland, UK. P. G. Fernandes: FRS
Marine Laboratory, PO Box 101, 375 Victoria Road, Torry, Aberdeen AB11 9DB, Scotland, UK. Correspondence to S. M. M. Fassler: tel: 44 1224
295538; fax: 44 1224 295511; e-mail: s.faessler@marlab.ac.uk.

Introduction
Acoustic-survey techniques are widely used to provide data on
fish abundance and distribution (Simmonds and MacLennan,
2005). Such techniques lend themselves particularly well to
pelagic fish, which often form well-defined schools in midwater
that can be detected efficiently by echosounders. The total error
(i.e. random and systematic components of measurements and
sampling error) in acoustic surveys is generally not quantified,
though work by Tesler (1989), Demer (1994, 2004), Aglen
(1994), and Simmonds and MacLennan (2005) are exceptions
to this statement. Consequently, the resulting estimates of fish
abundance are usually treated as indices to tune population
assessment models (Simmonds, 2003). Often, where errors are
reported, they are based on random-sampling errors (Rose
et al., 2000) and do not account for various sources of systematic
measurement and sampling error specific to acoustic surveys
(Simmonds et al., 1992). Systematic errors result from the equip-
ment (Toresen et al., 1998), acoustic shadowing (Zhao and Ona,
2003), vessel avoidance (Vab et al., 2002), and most notably, the
uncertainty in the fish target strength (TS; Simmonds et al.,
1992). Fish TS quantifies the proportion of incident sound
energy that is scattered back towards the transducer and is
used to convert the collected echo-intensity data to fish density
(Simmonds and MacLennan, 2005). TS is mainly dependent
on acoustic frequency (Foote, 1985; Horne and Jech, 1998),
fish size (Love, 1977) and fish orientation (Nakken and Olsen,
1977). A simple relationship of the form TS = 20 log10(L) 2 b20
is conveniently and commonly used to estimate mean TS from
fish length (L) using a species-specific value for the intercept
b20 (Simmonds and MacLennan, 2005).
The TS L relationship currently applied to estimate abun-
dances of various herring (Clupea harengus) and other clupeid
stocks around the world was determined empirically 2530
years ago. Nakken and Olsen (1977) insonified 41 stunned and
tethered herring at the commonly used frequency of 38 kHz and
determined the maximum dorsal-aspect TS at a given fish
length. Based on observations of tilt-angle distributions of small
herring (mean length = 13 cm), the maximum TS values were
reduced by 6 dB to account for the various incidence angles of
acoustic waves on the fish, giving a final TS L relationship of
TS = 13.6 log10(L) 2 62.8. Edwards and Armstrong (1981)
measured the TS at 38 kHz of a total of 565 live herring (L =
2125 cm) in a cage at 17.5-m depth over a period of 340 h.
They obtained a mean TS of 33.8 dB kg21 of fish. The
Planning Group on ICES-Coordinated Herring and Sprat
Acoustic Surveys (ICES, 1982) subsequently converted Nakken
and Olsens (1977) TS L relationship into TS per kg for a
23 cm herring (34.6 dB kg21) to facilitate comparison with
Edwards and Armstrongs (1981) results. It was decided that the
mean of the two TS estimates (34.2 dB kg21) should be used
together with lengthweight data from northwestern North Sea
herring and the groups final recommendation was a TS L
relationship of TS = 20 log10(L) 2 71.2 dB.
There is evidence that pressure, and therefore fish depth, modu-
lates the TS of Atlantic herring. This dependence could bias survey
results if not taken into account (Ona, 1990, 2003; Ona et al., 2003;

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Page 2 of 8
Lland et al., 2007) Herring are physostomes and therefore do not
have a gas gland that allows them to alter the swimbladder volume
actively. As a result, the swimbladders, which can reflect between 90
and 95% of incident sound energy (Foote, 1980a), will change
volume with water pressure according to Boyles law (Fassler
et al., 2009), leading to a steady decrease in TS with increasing
depth. Maturity stage or stomach fullness may also have important
effects on swimbladder size, shape, and ultimately, TS (Ona, 1990).
Additionally, it has been demonstrated that the physical environ-
ment can influence the physiology and morphology of herring,
leading to geographic variation in swimbladder sizes and mean
TS. For instance, herring living in the Baltic Sea have larger swim-
bladders than their conspecifics in the Northeast Atlantic (Didrikas
and Hansson, 2004; Peltonen and Balk, 2005; Fassler et al., 2008).
This is linked to their lower fat content and the low salinity of
the Baltic Sea, which taken together place increasing importance
on the swimbladder for buoyancy regulation.
In this paper, we further develop the backscattering model
described in Fassler et al. (2008) that predicts mean TS as a function
of fish orientation, size, physical properties, and acoustic frequency,
applying Bayesian methods. TS data from Norwegian spring-
spawning herring at depth (Ona, 2003) were used to estimate the dis-
tribution of the model parameters; namely the relative swimbladder
volume at the surface, the tilt-angle distribution, and the s.d. of the
estimated mean TS. Bayesian methods were used because these
allow parameter uncertainty to be incorporated into estimates of
mean TS. The TS model comprised two separate parts: (i) the
swimbladder was approximated as a gas-filled, prolate spheroid
using the modal-series-based deformed-cylinder model (MSB
DCM; Stanton, 1988); and (ii) the fish body was assumed to be a
fluid-filled ellipsoid and modelled using the distorted-wave Born
approximation (DWBA; Morse and Ingard, 1968; Stanton et al.,
1993; Chu et al., 1993; McGehee et al., 1998). Posterior distributions
of model parameters were then used with the error-propagation
properties of the Bayesian framework to simulate backscattering by
herring, while quantifying the precision of the TS estimate. The
resulting length- and depth-dependent TS model was applied to esti-
mate random and systematic errors in TS estimation and ultimately
to estimate North Sea herring abundance.
This study is an important step towards incorporating the error
associated with TS in estimates of total error in acoustically
derived abundance estimates of fish. The methods will be advan-
tageous for ecosystem studies and stock assessment where esti-
mates of fish abundance with estimates of total error are required.
Material and methods
Bayesian methods
In Bayesian methods, posterior density functions of model par-
ameters (u) are derived based on the goodness-of-fit of a model
to data and on prior information not contained in those data
(MacAllister and Kirkwood, 1998). The posterior probability dis-
tributions p(uijdata) of a set of continuous model parameter (ui)
given the data are determined using Bayes theorem (Bayes,
1763) according to
pui ; data pui  Ldatajui prior was chosen for SDTS. Because of the complexity of the
pui jdata ; 1
pdata pu  Ldatajudu model, integration of the denominator (i.e. the normalization
where p(ui,data) denotes the joint probability for a set of parameters
ui and the obtaining of the data. The data used here were derived
S. M. M. Fassler et al.
from a set of extensive in situ TS measurements of Norwegian spring-
spawning herring at a range of water depths from 5 to 300 m (Ona,
2003). These were mean TS values standardized to a common fish
length (L) of 32 cm (Ona, 2003) at an observed depth (z), each deter-
mined from .2000 single-fish targets. Suppose that the data rep-
resent j samples of a continuous probability density function
(PDF) that depends on the unknown parameters u, in a known
way that can be described by a model (Lee, 2004). p(u) describes
the prior PDF (i.e. the prior probability) for u. L(datajui) is the
probability of obtaining the data values if ui were the true values.
L(dataju) is commonly referred to as the likelihood function describ-
ing the dependence of the data on u. The likelihood function of the
entire TS dataset is given by the product of the normal density func-
tion over all datapoints:
Y
n  
1 Yj  Uj 2
Ldataju p exp  ; 2
j1 s 2p
2s2
where s is the s.d. of the observation error. Likelihood functions can
result in small numbers and for that reason the log10 (likelihood) was
used in the computer program to reduce calculation time and avoid
rounding issues (MacAllister and Kirkwood, 1998). Here, Yj is the jth
observed TS value in the dataset and Uj is the expected TS, estimated
from a theoretical TS model described below. Uj can be described in
simple terms as a function of model parameters (u), fish length Lj
and depth zj:
Uj f u; Lj ; zj : 3
The prior distribution for a parameter is based on previous
knowledge of the parameter, without incorporating the data
used to calculate the likelihood function (Punt and Hilborn,
1997). Priors are usually obtained by consulting experts or histori-
cal records. We determined prior distributions for the three par-
ameters that were deemed most important for the outcome of
the model. These were the standard deviation of the fish tilt-angle
distribution (SDt), relative volume of the swimbladder at the sea
surface (V0), and the standard deviation of the observed mean
TS (SDTS). Prior distributions of each of these parameters were
constructed independently and consequently the joint prior
p(ui) was simply
pui pSDti pV0i pSDTSi ; 4
where p(SDt), p(V0), and p(SDTS) are the priors for the respective
parameter values. These priors are assumed to be independent of
each other and of both fish length and water depth. A prior can be
either informative or uninformative. An uninformative prior pro-
vides little information relative to the data (Box and Tiao, 1973)
and usually has the form of a uniform distribution or one with a
large variance, giving all reasonable parameter values approxi-
mately equal probabilities. Informative priors provide precise
information based on previous evidence and may have a consider-
able affect on the result. Priors for parameters SDt and V0 were
based on published values (Table 1), whereas an uninformative
constant) in Equation (1) was not feasible. Instead, values were
sampled directly from the posterior distributions of the model
parameters using Markov Chain Monte Carlo (MCMC)
Bayesian approach to estimating TS Page 3 of 8

Table 1. Prior distributions and values of model parameters.
Description
Physical properties
Sound speed contrast
water-sh body
Density contrast water-sh
body
Sound speed contrast
swimbladder-sh body
Density contrast
swimbladder-sh body
Fat density
Scales density
Bones density
Fish esh density
Sound speed in water
Echosounder frequency
Fish behaviour and dimensions
s.d. of tilt-angle distribution
Angle of swimbladder relative
to snout-tail axis
Ratio sh width/height
Ratio sh width/height
Relative swimbladder volume
at sea surface
Swimbladder minor-axis
compression factor
Swimbladder minor-axis
compression factor
Precision
s.d. of mean TS estimate
Value or prior distribution
(source)
hb = 1.04 (Fassler et al., 2008)
gb = 1.04 (Fassler et al., 2008)
hsb = 0.23 (Fassler et al., 2008)
gsb = 0.00128 (Fassler et al., 2008)
rf = 0.926 g cm23 (Brawn, 1969)
rsc = 1.966 g cm23 (Brawn, 1969)
rb = 1.993 g cm23 (Brawn, 1969)
P = 1.057 g cm23 (Brawn, 1969)
cw = 1500 m s21
f = 38 kHz
SDt Normal (8.6, 3.72) (Nakken
and Olsen, 1977; Ona, 2001; Gorska
and Ona, 2003)
Du = 5.568 (measurements)
b/a = 0.4873 (measurements)
c/a = 4.7100 (measurements)
V0 Normal (5, 0.72) (Harden
Jones and Marshall, 1953; Ona 1990)
a = 0.42 (Fassler et al., 2009)
b = 0 (Fassler et al., 2009)
SDTS Gamma (0.001,1)
(uninformative prior)
Details of survey procedures can be found in individual survey
reports (e.g. ICES, 2008). Acoustic data were collected using
Simrad EK60 echosounders operating at 18, 38, 120, and 200 kHz
with split-beam transducers mounted on the drop keel typically at
depths of 5 m. The transducers had nominal beam widths of 11, 7,
7, and 78, respectively, and were located in proximity to each
other. The maximum offset of 0.6 m was between the centres of
the 18 and 38 kHz transducers. Data were collected between 02:00
and 22:00 GMT from 29 June to 18 July 2007, with integration
starting at a depth of 12 m. Post-processing and analysis were done
using Myriax EchoView software (v 4.40). Biological samples were
collected from some of the more dense aggregations observed
throughout the survey using a PT160 pelagic trawl for identification
purposes and to generate length frequency distributions for the
herring in the various survey subareas. Fish lengths were recorded
in 0.5 cm intervals to the nearest 0.5 cm below total length.
Integrated volume-backscattering coefficients sA were averaged
over distances surveyed each 15 min (2.5 nautical miles).
Numbers of fish in length group i in a given area were estimated by
sA
Ni  pi  A; 7
s bsi
where Ni is the number of herring in length group i and sA the
nautical-area-scattering coefficient (NASC) with units m2 nautical

simulation with the Random Walk Metropolis algorithm

(Metropolis et al., 1953).
TS model
The theoretical model used to estimate the TS of herring has two
components combined coherently to give the total backscattering
cross section of the whole fish s fbs (cm2) according to

sfbs ssb b
bs z sbs ; 5

where sbbs and ssb

bs (z) denote the backscattering cross section of the
fish body and the swimbladder at depth (z), respectively (Gorska
and Ona, 2003). The output of the model was the expected total
backscattering cross section ksbs l based on model parameters,
which was converted to TS according to MacLennan et al. (2002):

TS 10 log10 ksbs l: 6

The fish-body component was modelled using the DWBA, with

the model routines described in Fassler et al. (2007). The swim-
bladder was modelled with a MSB DCM (see Gorska and Ona,
2003; Fassler et al., 2008, for details). All input parameters for
the model, including their sources, are listed in Table 1.
Survey analysis
Echo-integration data were taken from the Scottish component of
the North Sea herring acoustic survey in summer 2007 (Figure 1).
Figure 1. Map of the northwestern North Sea showing the survey
area (grey) covered by the Scottish component of the North Sea
Herring Acoustic Survey. Numbers represent the ICES acoustic-survey
estimates by statistical rectangle from 2007: absolute numbers
(millions of sh; white, upper part of the gure) and biomass
(thousand tonnes; black, lower part of the gure). The vertical axis
shows degrees of latitude (N), and the horizontal axis shows degrees
of longitude [East and West (2)] of the Greenwich Meridian.
Page 4 of 8
mile 2 (MacLennan et al., 2002). The analysed area (A) covered a
total of 38 769.5 nautical miles2. The survey area was divided into
seven subareas characterized by fish with similar length frequencies.
For each area, pi is the proportion of herring in length group i, and
sbsi is their backscattering cross section, so that
P
sbsi  pi
i
s bsi P : 8
pi
i
Distributions of expected sbs were calculated concurrently in
the Bayesian framework by directly using all the model parameter
values sampled from their posterior distributions in the Markov
Chain. In that way, the uncertainty in the model parameters was
propagated through to the predicted mean backscattering cross
section, and the distribution of the parameters and any corre-
lation between them was taken into account. Samples of sbs
were then drawn from a lognormal distribution with estimated
precision based on SDTS. Hence, the methods applied in this
paper differ from those of previous investigations (Demer, 1994,
2004; Rose et al., 2000) that estimated abundances from acoustic-
survey data by simple Monte Carlo simulation using respective
distributions of model parameters. Computations were done for
all length groups of herring in depth bins of 5 m ranging from 0
to 200 m. Numbers of herring were estimated as follows:
(i) TS = 20 log10(L) 2 71.2 to convert fish length (L) into TS;
(ii) estimated distributions of sbs for every depth bin and length
group; and
(iii) the TS = 20 log10(L) 2 b20 relationship with b20 based on
calculations in (ii) for depths of 10 20 m [following the
methods used by Nakken and Olsen (1977) and Edwards
and Armstrong (1981)].
Results
Computations were done on 45 000 samples from the posterior
distributions. Posterior distributions converged after 200 iter-
ations, but a more conservative burn-in interval (i.e. discarded
values at the beginning of the MCMC chain) of 2000 iterations
was chosen. The plot matrix of samples from posterior distri-
butions of the model parameters revealed no severe autocorrela-
tion between parameters (Figure 2). Possible autocorrelation
could have influenced mixing and convergence of the sampled
posteriors. Figure 3 shows the results from the Bayesian estimation
of the three model parameters. The mean and s.d. of the posterior
distribution for the relative swimbladder volume at the surface
(V0; mean = 4.95%, s.d. = 0.72%) were approximately equal to
those of the prior distribution; this suggests that the prior had a
considerable effect on the parameter estimation. However, the
posterior for SDt had a smaller mean and a considerably narrower
distribution (mean = 7.148, s.d. = 0.508) than the prior. Because of
the uninformative prior assigned to SDTS, all the information used
to construct the posterior distribution came from the data (SDTS;
mean = 1.64 dB, s.d. = 0.12 dB).
Results from the Scottish component of the North Sea Herring
Acoustic Survey can be found in ICES (2008). The herring abun-
dance estimated by ICES (2008) for the survey area (Figure 1),
applying TS = 20 log10(L) 2 71.2, was 5.34  109 fish. Figure 4
shows a comparison of estimated herring abundances using the
S. M. M. Fassler et al.
Figure 2. Plot matrix of sampled posterior values of the
backscattering model parameters. Histograms represent frequency
distributions of parameter values given on the x-axis.
three different methods applied in this study to determine the
herring TS. Using model values based on the Ona (2003) dataset
to determine the TS for herring in surface waters between 10
and 20 m, following the methods applied by Nakken and Olsen
(1977) and Edwards and Armstrong (1981), the mean TS is
4.0 dB higher than the one used to generate the ICES (2008)
biomass estimate. As a result, the estimated herring abundance cal-
culated here is reduced by .50% to 2.45  109 fish. In compari-
son, the estimated normal distribution of abundance based on TS
distributions by length group and depth bin had a mean and s.d. of
3.17  109 and 3.02  107 fish, respectively. This is also less than
the current estimate, but suggests higher estimated abundances
if the depth-dependent TS is applied.
Discussion
When interpreting results from an acoustic survey of fish, it is
important to recognize the stochastic nature of the TS and its poss-
ible dependence on many factors. It has long been known that TS
is influenced significantly by fish length (e.g. Love, 1977; Foote,
1979), fish orientation (e.g. Nakken and Olsen, 1977; Foote,
1980b), and acoustic frequency (e.g. Haslett, 1965; Love, 1977).
Because of a lack of suitable alternatives, acoustic surveys still
rely on empirically determined, frequency-specific TS L relation-
ships, which may be biased, particularly for physostomous fish.
Many experiments from which these relationships were derived
for herring were done in only one season and neglected important
biological effects such as feeding state and gonad development
(Ona, 1990; Ona et al., 2001) or the fact that dead or stunned
fish may have a different TS from live fish (e.g. McClatchie
et al., 1996). On the other hand, because for the same size and
species of fish the observed TS can cover a wide range, acoustic
surveys have relied on the expected values of the mean TS
(Simmonds and MacLennan, 2005; Foote, 1987). It is difficult to
quantify the effect of any one factor on TS by empirical obser-
vations, so theoretical TS models have been used to approximate
fish backscatter and investigate its dependence on various input
parameters (Clay and Horne, 1994; Foote and Francis, 2002;
Gorska and Ona, 2003). This theoretical modelling approach
Bayesian approach to estimating TS
Figure 3. Priors and posteriors of the backscattering model
parameters. The prior for the s.d. of the mean TS estimate (SDTS) is
uninformative.
offers the most promise if uncertainty arising from TS variability is
to be estimated and incorporated into the acoustic stock assess-
ments (Demer, 2004).
Total error in acoustic surveys is rarely quantified (Rivoirard
et al., 2000; Rose et al., 2000; Tjelmeland, 2002), despite its
obvious advantages and widespread use in fish-stock assessment.
Bayesian methods have been identified as a valuable tool to esti-
mate the total uncertainty in trawl surveys (Punt and Hilborn,
1997; MacAllister and Kirkwood, 1998; Meyer and Millar, 1999)
and could be used in acoustic surveys to quantify and incorporate
Page 5 of 8
Figure 4. Estimated herring abundances from the Scottish
component of the 2007 North Sea Herring Acoustic Survey using
various TS values: (i) dotted line: currently used TS sh length (L)
relationship TS = 20 log10(L) 2 71.2; (ii) solid-line histogram:
distribution of estimated mean TS for every sh length and 5-m
depth bin using a TS model and parameter estimates in a Bayesian
framework; (iii) dot-dashed line: depth-independent TS L
relationship estimated for the surface (10 20 m) backscattering
values obtained in (ii).
uncertainty into abundance and biomass estimates. The results of
these methods provide estimates of the random and systematic
components of measurement and sampling error, which are
important for managing fisheries. Uncertainty in the advice pro-
vided by fishery scientists can be based on a combination of data
from the stock in question and prior information on population
model parameters based on data from similar fish populations
(MacAllister and Kirkwood, 1998). Another advantage of
Bayesian methods is that the uncertainty associated with model
parameters can easily be propagated through the framework to
estimate distributions of model outcomes. Based on these proper-
ties, we have proposed a Bayesian system that allows unknown par-
ameters, or those that are difficult to estimate, to be incorporated
into a conventional fish-backscattering model to estimate distri-
butions of expected TS. This can then provide a basis for estimat-
ing the uncertainty associated with TS in an acoustic survey and
contribute to the overall estimate of uncertainty.
The herring abundance estimates obtained in this study indicate
that the uncertainty associated with this particular model is largely
systematic. Assuming that the ICES (2008) assessment of herring
abundance, which is based on the conventional TSL relationship,
is correct, the associated accuracy of the acoustic estimate based on
the Bayesian TS model is 40.6%. This error is in accordance with
previous upper estimates of the systematic error of the TS com-
ponent in acoustic estimates of absolute abundance, where values
of +26 to +41% (Tesler, 1989), 0 to +50% (Simmonds and
MacLennan, 2005), or 0 to +40% (Simmonds et al., 1992) have
been reported. Factors that may have affected the accuracy of the
abundance estimate in this study were the dataset used to derive
the model parameters and the TS model itself. It must be noted
that the herring TS data used in this study might not have been
entirely appropriate, being based on Norwegian spring-spawning
herring as opposed to North Sea herring. Although these stocks
are the same species and live in broadly similar environments, in
contrast, for example, to herring living in the Baltic Sea, which
have an entirely different salinity regime, the two herring stocks
may have a different morphology (Sinclair and Solemdal, 1988). It
is evident that the TSL relationship fitted to the whole Ona
(2003) dataset (TS = 20 log10(L) 2 67.3) is 3.9 dB higher than
that currently applied to estimate abundance of North Sea
herring. In terms of fish numbers, referring to the 2007 acoustic
survey, this difference would cause a 59.3% reduction in estimated
abundance of North Sea herring and ultimately bring the results
Page 6 of 8
Figure 5. Distribution of integrated acoustic energy (Nautical-Area-Scattering Coefcient, NASC) allocated to herring at various 5-m depth
bins in the Scottish component of the North Sea Herring Acoustic Survey 2007.
more in line with estimates using the Bayesian model. It follows that,
because the accuracy of the Bayesian TS model is strongly dependent
on the input data, acoustic-survey analyses should be based on
reliable TS data. Further work will involve collection of appropriate
in situ TS data to tune the TS model, providing results more appli-
cable to North Sea herring.
Additionally, the TS model used may have been unsuitable for
the species in question, because it is based essentially on the coher-
ent addition of backscatter from simple approximations of the
herring swimbladder and fish-body shapes. This improves compu-
tational efficiency, but neglects structural details of the herring
anatomy. Moreover, the model does not take account of the
boundary conditions at the swimbladder wall, because it ignores
the fish-body components when calculating the swimbladder
backscatter. Nonetheless, coherent addition is applicable in the
present case, because the swimbladder is the dominant source of
backscatter (Ding and Ye, 1997; Gorska and Ona, 2003). In con-
trast to the low accuracy of the herring abundance estimate, it is
very precise (CV = 1.0%), so the random error associated with
the TS component is negligible. This high precision, however,
can be attributed to the central-limit theorem, because absolute
fish abundance was derived by averaging a large number of esti-
mates (Demer, 2004).
Another advantage of the Bayesian model is the ability to apply
a specific TS distribution to all herring schools encountered in the
survey. This is of particular importance when dealing with the
depth-dependence of herring TS. In the survey area, NASCs
were highest at depths between 100 and 150 m (Figure 5). At
these depths, swimbladder volumes are only 6 9% of those at
the surface, resulting in greatly reduced TS values (Fassler et al.,
2009). As a result, extending the TS dataset from values at the
sea surface (1020 m) to include values down to 300-m depth
resulted in an abundance estimate that was higher by about one
third (Figure 4).
There are disadvantages of the Bayesian approach linked with
the selection of prior probabilities. Unless carefully chosen, the
selection of prior probabilities may cause bias, and their construc-
tion can be tedious and require considerable effort (MacAllister
and Kirkwood, 1998). Prior distributions estimated for two of
the model parameters in this study were based on reliable
measurements on herring and are assumed valid. However, the
TS-model evaluation indicates a particular need for accurate
data on tilt-angle distributions. Fish behaviour, as reflected in
S. M. M. Fassler et al.
the distribution of tilt angles within an aggregation of fish, is the
most important factor influencing TS and hence the accuracy of
subsequent abundance estimates (Foote, 1980c; Blaxter and
Batty, 1990; Horne and Jech, 1999; Hazen and Horne, 2003).
Other potential limitations of the more complex Bayesian model
are the demand on computing resources, as well as posterior dis-
tributions that may sometimes be too complex to estimate.
Nonetheless, given appropriate input data, these methods can
provide an important step towards opportunistically incorporat-
ing the variability associated with factors affecting TS, to give an
overall estimate of the expected TS distribution. The resulting
probabilistic abundance, density, and biomass estimates could
become essential components in ecosystem studies and resource
management.
Acknowledgements
The authors are grateful to Dezhang Chu and Gareth Lawson from
the Woods Hole Oceanographic Institution, Woods Hole, MA, for
providing the MATLAB code for the fish-body model. We thank
Dave Borchers (CREEM, St Andrews University, UK), Liz Clarke
(FRS, Aberdeen, UK), and Colin Millar (CREEM and FRS) for
their expert advice and help with Bayesian statistical methods.
SMMF acknowledges the support received through an ORSAS
award of the British government, a studentship of the University
of St Andrews (Scotland), and an Ausbildungsbeitrag of the
Kanton Basel-Landschaft (Switzerland).
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doi:10.1093/icesjms/fsp008