Do abiotic mechanisms determine interannual
variability in length-at-age of juvenile Arcto-
Norwegian cod?
Geir Ottersen, Kristin Helle, and Bjarte Bogstad
Abstract: For the large Arcto-Norwegian stock of cod (Gadus morhua L.) in the Barents Sea, year-to-year variability
in growth is well documented. Here three hypotheses for the observed inverse relation between abundance and the
mean length-at-age of juveniles (ages 1–4) are suggested and evaluated. Based on comprehensive data, we conclude
that year-to-year differences in length-at-age are mainly determined by density-independent mechanisms during the
pelagic first half year of the fishes’ life. Enhanced inflow from the southwest leads to an abundant cohort at the 0-
group stage being distributed farther east into colder water masses, causing lower postsettlement growth rates. We can
not reject density-dependent growth effects related to variability in food rations, but our data do not suggest this to be
the main mechanism. Another hypothesis suggests that lower growth rates during periods of high abundance are a
result of density-dependent mechanisms causing the geographic range of juveniles to extend eastwards into colder
water masses. This is rejected mainly because year-to-year differences in mean length are established by age 2, which
is too early for movements over large distances.
Résumé : La variabilité inter-annuelle de la croissance chez l’important stock arcto-norvégien de Morues franches
(Gadus morhua L.) de la mer de Barents est bien connue. On trouvera ici la proposition et l’évaluation de trois
hypothèses pour expliquer la relation inverse observée chez les juvéniles (âges 1–4) entre la densité et la longueur
moyenne à un âge donné. L’étude de données détaillées nous ont amenés à conclure que les différences inter-annuelles
des longueurs à un âge donné sont déterminées principalement par des mécanismes indépendants de la densité durant
la première demi-année pélagique de vie des poissons. Des influx accrus du sud-ouest entraînent le déplacement à l’âge
0 d’une cohorte importante vers l’est dans des masses d’eau plus froide, ce qui diminue les taux de croissance après
l’établissement au fond. Il ne nous est pas possible de nier les effets dépendants de la densité de la variabilité des
quantités de nourriture sur la croissance, mais nos données nous laissent croire que ce n’est pas là le mécanisme
principal. Une autre hypothèse veut que les taux réduits de croissance durant les périodes de haute densité soient le
résultat de mécanismes reliés à la densité qui amènent les juvéniles à étendre leur aire géographique vers les masses
d’eau plus froide vers l’est. Cette hypothèse n’est pas retenue, car les différences inter-annuelles des longueurs
moyennes sont déjà en place dès l’âge 2, âge trop précoce pour permettre des déplacements sur de grandes distances.
[Traduit par la Rédaction] Ottersen et al. 65
Introduction
Controlled experimental evidence has established that
growth of fish is often inversely related to stock density. A
change in the population size of a predator is assumed to af-
fect intraspecific competition for food and thereby consump-
tion and growth. An inverse relationship between growth and
stock abundance is therefore indicative of density-dependent
growth. Density-dependent growth in fish populations has
been reported since early in the 20th century and further evi-
dence has accumulated as stocks have become overexploited
(e.g., Raitt 1939; Marshall and Frank 1999). However, other
studies have found no significant correlation between
growth and stock abundance (e.g., Daan 1974). It should
also be noted that even when such a correlation is observed
in field data it could be caused by a third variable, e.g., tem-
perature acting on both stock abundance and growth
(Jørgensen 1992).
Individual growth in fish depends on density-dependent
factors, such as the availability of prey. However, growth is
also the integration of a series of processes (feeding, assimi-
lation, metabolism, transformation, and excretion), the rates
of which are all controlled by temperature (Michalsen et al.
1998). Brander (1995) examined 17 North Atlantic cod stocks
and found that most of the variability in growth among stocks
could be attributed to variation in temperature.
The Arcto-Norwegian (or Northeast Arctic) cod spawn in
March–May in patchy areas off northern and mid-Norway,
with spawning mainly taking place in the Lofoten region
(Fig. 1). Spawning occurs at the same sites each year, but
spawning depth varies considerably, following the thermocline
between the cold water of the Norwegian coastal current and
Can. J. Fish. Aquat. Sci. 59: 57–65 (2002) DOI: 10.1139/F01-197 © 2002 NRC Canada
57
Received 20 March 2001. Accepted 22 October 2001.
Published on the NRC Research Press Web site at
http://cjfas.nrc.ca on 14 January 2002.
J16271
G. Ottersen,
1,2
K. Helle, and B. Bogstad. Institute of
Marine Research, P.O. Box 1870, Nordnes, N-5817 Bergen,
Norway.
1
Corresponding author (e-mail: geir.ottersen@bio.uio.no).
2
Present address: University of Oslo, Department of Biology,
Zoology Division, P.O. Box 1050, Blindern, N-0316 Oslo,
Norway.