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Elemental Composition as a Natural Tag
To the extent that populations or stocks of fish inhabit different
environments, the otolith elemental composition should serve as a
proxy for population identity. Two assumptions underlie the use of the
otolith elemental composition as a stock discriminator: 1) material
deposited on the otolith is metabolically inert after deposition, and is
not susceptible to resorption; and 2) the physical and chemical
environment influences the rate of trace element incorporation into the
growing otolith surface. Elements under strong physiological
regulation (eg- Na, K, S, P, Cl) probably do not meet the second
assumption, and thus are of limited value for stock identification
studies. However, both assumptions appear to be met with respect to
elements such as Sr, Ba, Mn, Fe and Pb (and perhaps Li, Mg, Zn, Cu
and Ni), in which both ambient element:Ca concentrations and
temperature produce significant effects on otolith composition.
Environmental responses such as these, recorded permanently in the otolith, imply that the otolith concentrations of selected elements and
isotopes can be used as a biological tag to discriminate among groups
of fish which have spent at least part of their lives in different
environments. Discriminatory power is increased greatly by treating
the selected elements as a group, using multivariate statistics, rather
than individually; hence the term "elemental fingerprint". An
appealing feature of this application is that the elemental fingerprint
need not be linked to potential sources or locations in the environment.
Rather, the presence of significant differences in the fingerprints of two
or more groups of fish implies that the groups cannot all be random
samples from the same population. This deduction holds even if
physiological effects have influenced elemental composition, since
random samples from the same population would have experienced the
same mean physiological effects. Of course, the presence of different
fingerprints cannot be used to infer the length of time that the groups of
fish remained separate, since even occasional residency in a different
environment has the potential to introduce a detectable difference in the
elemental composition. By corollary, the absence of differences does
not necessarily imply that the groups of fish are of common origin. As
a result, it is fair to categorize otolith elemental fingerprints as
powerful discriminators when differences exist, but of negligible value
when differences cannot be detected.
It is probably inappropriate to refer to the use of elemental fingerprints
as stock discriminators, since genetic differences are not implied and
spatial heterogeneity in the stock environment can result in different
fingerprints for different stock components. Perhaps more importantly,
ontogenetic effects and age-related differences in exposure history can
result in very different fingerprints for fish of different size classes
from the same population. Nevertheless, the presence of different
fingerprints among matched groups of fish necessarily implies different
environmental histories. Accordingly, the elemental fingerprint would
appear to be an excellent biological tracer of groups of fish, and it is
this application of elemental fingerprints which has met with success in
both freshwater and saltwater.
Two forms of elemental fingerprinting are in general use: one based
on whole dissolved otoliths, and the other based on analysis of the
otolith core. Advantages of whole-otolith assays include ease of
preparation, absence of error associated with sampling or identifying
growth increments, and the availability of accurate and precise assay
protocols. The major disadvantage is associated with the inability to
take advantage of the chronological growth sequence recorded in the otolith. Atomic absorption spectrometry
(AAS), inductively-coupled plasma atomic emission spectroscopy (ICP-AES), neutron
activation analysis, and inductively-coupled plasma mass
spectrometry (ICPMS) are among the techniques which have been
used to analyze otoliths. However, it is ICPMS which has emerged
as the instrument of choice for such assays, due largely to its
capability for rapid and accurate isotopic and elemental assays over
a wide range of elements and concentrations. Isotope dilution
ICPMS (ID-ICPMS), a variant of ICPMS often used to certify
reference materials, is the most accurate of the otolith analytical
techniques currently available. Sample sizes required for most of
the above assays are on the order of 5-10 mg of otolith material,
although ICPMS units outfitted with high efficiency nebulizers are
capable of handling otolith weights as low as 0.3 mg. See the Methods
page for detailed information on preparing otoliths for elemental analysis.
With the realization that elemental fingerprints can be used very effectively to
separate mixtures of fish coming from different sources, there is increasing
demand for the maximum-likelihood based software to separate the group
mixtures. Discriminant analysis is not a good option here, since the
'priors' parameter is unknown. In the Methods
section of this web site, we've released a working copy of the Integrated Stock
Mixture Analysis (ISMA) program (written for the S-Plus environment) for use in
separating stock mixtures based on elemental fingerprints or other continuous or
categorical variables.
Our laboratory has devoted much effort to the study and application
of otolith elemental fingerprints, both with respect to stock
identification and to factors influencing trace element uptake into
the otolith. This work can be reviewed in Thorrold et al.
(1998),
Farrell and Campana (1996), Campana and Gagné
(1995), Campana
et al. (1995), and Fowler et al. (1995a,b). A comprehensive review of the field
of otolith composition and chemistry is provided in Campana
(1999). An in-depth examination
of the value of elemental fingerprints as natural tags of fish is presented
in Campana et al. (2000).
Recent applications of otolith elemental fingerprints as natural tags are
available in Campana
et al. (2007) for redfish and Jonsdottir
et al. (2006) for cod around Iceland.
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