These
aquaculture-oriented commentaries reflect the opinions of GCL associates
-- usually Roger Doyle -- and are not the original abstracts of the
papers. Direct quotations from the papers or abstracts are
marked with inverted commas.
19. Competitive ability depends on the
neighbours
Group size and relative competitive ability: geometric progressions as
a conceptual tool. 2000. Humphries, S., G.D. Ruxton, and N. B. Metcalfe.
Behavioral Ecology and Sociobiology 47:113-118.
An ingenious mathematical and experimental study shows that there is no
simple, general relationship between the relative competitive ability of
two individual fish and the size of the group of fish they are living in.
Nevertheless, group size does make a tremendous difference in relative
competitive ability, and the optimal group size for each fish will depend
on its relative size. (It has to be said that this study might be enhanced
by a game-theoretic analysis of what is optimal.) In any case, for
aquaculture, it suggests that even more attention should be paid to
size-grading and tank size. For aquaculture genetics it gives additional
support to the idea that competitive interactions have totally fouled up
conventional heritability estimates and classically-designed selection
programmes based on related "independent errors" assumptions.
For biological conservation, it is one more thing to think about when
trying to design a hatchery programme for supplementing wild stocks that
won't domesticate the fish. s.humphries@bio.gla.ac.uk
18. Unlimited effective population size
Marker-Assisted Selection to Increase Effective Population Size by
Reducing Mendelian Segregation Variance. 2000. Wang, J., and W.G. Hill.
Genetics 154:475-489. Everyone knows that the effective population size of
a broodstock (the size that determines the rate of inbreeding and genetic
drift) is a "virtual" number. When you select an equal number of
offspring from every family the effective population size is approximately
twice the actual size. It had been supposed that this as good as you could
do to minimize inbreeding in genetic conservation, aquaculture broodstock
management etc. (Numerous mating schemes have been proposed which do
somewhat better than this, but at such tremendous rates of initial
inbreeding that they are impractical.)
The authors of this highly technical paper have now shown by theory and
simulation that the effective population size can be made indefinitely
large with the aid of marker-assisted selection. The procedure should work
best when the number of offspring is enormous, as in aquaculture.
Unfortunately, it is completely impractical at the present time. In 15
years or so, when marker maps have been filled in for the major species
and gene-chip technology is cheap, our inbreeding problems may be solved. jinliang.wang@ed.ac.uk
17. Better to know about house flies than to
know nothing at all
Experimental tests of minimum viable population size. 2000. Reed, D.H.,
and E.H. Bryant. Animal Conservation 3:7-14. Another experiment with house
flies. Some populations were maintained at a constant size, some put
through a severe bottleneck (down to 5 individuals) then allowed to expand
rapidly. The bottlenecked populations recovered quickly from their initial
inbreeding depression. "However, they were less adaptable to
environmental stress than constant size populations, suggesting that
populations founded with few numbers may do well within a single
environment but may do far less well if they are reintroduced to natural
environments or exposed to rapid environmental changes." This has
obvious consequences for genetic conservation and perhaps also for
aquaculture. Also, perhaps, for fisheries stock-enhancement programs.
16. Markers for the traits that matter
A logistic regression based extension of the TDT for continuous and
categorical traits. 1999. Waldman, I.D., B.F. Robinson, and D.C. Rowe.
Annals of Human Genetics 63:329-340. I've always argued that within-family
selection is the best way to go in aquaculture. These authors have come up
with a procedure for finding statistical associations between markers and
continuous traits such as growth and fecundity, a procedure that works
within families. (Other within-family procedures work only with
categorical yes-or-no traits.) I think it is about time research stations
or even farmers should start
collecting and storing fin-clips from fish within families. Just on a
speculative basis. When marker maps are adequate, these samples and their
associated size and rudimentary pedigree data might be extremely valuable 15. Inbreeding hits fitness first
A comparison of inbreeding depression in life-history and
morphological traits in animals. 1999. DeRose, M.A., and D.A. Roff.
Evolution 53:1288–1292.
Traits related to fitness (in aquaculture these would be growth, survival
and fertility) show greater inbreeding depression than other traits
(shape, colour etc.). This re-analysis of data from many published
research papers demonstrates large amounts of dominance variance in
fitness traits (see also #10), which is significant for broodstock
improvement. It is also another indication that inbreeding can cause big
problems in small populations. Perhaps by the time aquaculturists begin to
notice development (shape) abnormalities and deformities which they
ascribe to inbreeding, they are already losing much more production due to
mortality and slow growth than they realize.
14. Hatchery salmon contaminating
wild salmon
Genetic changes in Atlantic salmon (Salmo salar) populations of northwest Irish
rivers resulting from escapes of adult farm salmon. 1998. Clifford, S.L., P.
Mcginnity, and A. Ferguson. Can. Jour. Fish. Aquatic Sci. 55:358-363.
Escaped farm salmon did interbreed with wild salmon. The proportion of juveniles
of maternal farm parentage in two rivers averaged 7% and reached a high of 70%
in an individual sample. "Only a small proportion of 29 000 adult farm
salmon that escaped in spring 1992 appear to have bred successfully in the
rivers studied." Note that the authors found fish whose mothers came from
the farm.
13. Someone really smart should
count the dead fish in advance
Effect of Selection Against Deleterious Mutations on the Decline in
Heterozygosity at Neutral Loci in Closely Inbreeding Populations. 1999. Wang,
J., and W.G. Hill. Genetics 153:1475-1489.
Highly technical, but unusually realistic, computer simulation. Shows that
natural selection does in fact "purge" a population of genes that
depress fitness when the population is randomly and partially inbred, and that
this can be observed at marker loci. The simulation is realistic enough to
resemble aquaculture in that not all animals in the same generation have the
same inbreeding. The question for aquaculture is, what is the economic cost of
all the dead animals that are purged out before they're harvested? Should be
possible to estimate and/or simulate this too.
12. Local adaptation made impossible
Gene flow and local adaptation in a sunfish-salamander system. 1999. Storfer, A.
Behavioral Ecology and Sociobiology 46:273-279
The adaptation of salamanders in local populations to fish predators (a genetic
predisposition to hide) is overcome by gene flow from nearby salamander
populations where there are no predators. Analogous to the situation in which huge
numbers of salmon escaping from aquaculture may (might perhaps possibly
maybe sometimes) reduce the fitness of natural populations.
11. Good environments accumulate bad
genes?
Fitness Decline under Relaxed Selection in Captive Populations. 1998. Bryant, E.
H., and D.H. Reed. Conservation Biology 13:665-669.
A neat laboratory study on house flies in which the authors relaxed selection on
genes that affect fitness late in life by a simple expedient -- killing the
flies young. Late-life fecundity declined drastically after a few generations of
this treatment. This would seem to be relevant to aquaculture. If natural
selection is reduced by improving the environment or equalizing the size of
families produced by aquaculture broodstock, there may be a rapid reduction in
the ability of these populations to exist under natural conditions. The authors
note that this "domestication" effect is independent of population
size. It is a genetic problem that occurs because new, bad mutations are allowed
to hang around, and is somewhat independent of the deleterious effects of
inbreeding and drift.
10. Plenty of selectable genetic
variance for fitness
Genetic architecture of fitness and nonfitness traits. 1999. Merila, J., and B.
C. Sheldon. Heredity 83:103-109.
It used to be thought that traits related to fitness (survival, fecundity etc.)
would not respond to selection because additive genetic variance would be low
(mutations with good or bad effects already having been selected to frequencies
of one or zero, respectively). Professor Rom Moav used this idea 20 years ago to
explain why his carp selection experiment made no progress. It seemed plausible
to many people at the time. However, recent studies - none in aquaculture - have
completely reversed this view. Fitness traits have more additive genetic
variance than other traits, probably because there are a lot more genes
involved. However, they also have a lot more dominance variance as well, which
reduces heritability.
9. If you're inbred you're going to
die of parasites in hard times
Parasite-mediated selection against inbred Soay sheep in a free-living island
population. 1999. Coltman, D.W., J.G. Pilkington, J.A. Smith, and J.M.
Pemberton. Evolution 53 (4):1259–1267
Small population of wild sheep on a remote North Atlantic island. The individual
sheep that are inbred are more susceptible to parasitism by gastrointestinal
nematodes at high population density. During periods of high overwinter
mortality highly parasitised individuals were less likely to survive. More
inbred individuals were also less likely to survive, which is due to their
increased susceptibility to parasitism. Dave Coltman's paper is causing a
general stir.
Extension to aquaculture would be that the effect of inbreeding
should show up under stress, in those individuals that are most inbred, and
would be manifested in mortality from parasites and disease?. Point is, this can
be investigated on working fish farms by a properly interpreted assay of
microsatellite heterozygosity. For future reference fish farmers should start storing
tiny bits of tissue from their
morts and healthy fish in 95% alcohol. The depressing effects of
inbreeding can be identified. |