These aquaculture-
and conservation-oriented
commentaries are not abstracts written by the original authors.
They reflect the opinions of someone else -- usually Roger Doyle. Direct quotations from the papers or
abstracts are marked with inverted commas.
345. Fish evolve to defeat the fishermen
Sustaining
fisheries yields over evolutionary time scales. 2002. Conover, D.O.,
and S.B. Munch. Science 297 (5578):94-96.
This is a direct
experimental proof that when you selectively harvest the larger fish in a
population, the population will evolve in a way that reduces your catch.
The experimental fishery consisted of a number of lab populations of the
Atlantic silverside (Menidia menidia) which were selectively fished for
four years.
In one pair of
populations all but the smallest 10% were harvested (mimicking a heavily
exploited fishery), in another pair all but the largest 10% were
harvested, and in the third pair the harvest was random with respect to
size. This is, of course, just a selection experiment and the fish
responded in the expected way: when only the small fish were left to breed
the harvested "yield to the fishery" grew less and less as the
generations passed. When the large fish were left to breed the yield
increased.
The idea that
size-selective fishery (the usual practice) will reduce yields has been
around for decades and indeed was included in Palumbi's "cost of
evolution" estimate (Dec 2001 #262). Analogous situations are
notorious in aquaculture -- in Chinese carps and Macrobrachium, for
instance. However, this is the first time it has been proved to happen in
an experimental model of a wild fishery. dconover@notes.cc.sunysb.edu
344. Pacific salmon require special Ne
calculations
Effective size of
fluctuating salmon populations. 2002. Waples, R.S. Genetics
161:783-791.
The salmon species
referred to are those which only breed once as individuals
but mature in several different years as a cohort, so that generations
overlap even though each year brings out an entirely new set of spawners.
That includes chinook and chum, but not coho. Since there is a lot of variation among years in the total number of
spawning adults (from several overlapping generations), how should we
calculate the long-term effective population number, Ne?
The author explores this
problem analytically and by simulation, and finds that variation in the
number of breeders among years within a cohort enters the calculation as
the harmonic mean, just like variation among generations in the well known
non-overlapping generation model. The result is that Ne can be smaller
even than the usual harmonic-mean estimate, and much smaller than
the average census count. This is useful to know when evaluating the
genetic status of endangered populations of Pacific salmon.
The detailed calculations
presented here may be about as far as one can go in estimating Ne in
Pacific salmon solely from demographic data. However, it is likely that
DNA marker data will soon be adequate to permit some sort of direct
pedigree determination, or pedigree sampling, in non-experimental
populations of salmon and other fish. When that happens, inbreeding and
heterozygosity-by-descent can be calculated exactly from the pedigree and
the loss of QTL diversity estimated, in even the most complicated
situations, by gene-drop simulation. Who then will care about Ne? [To this
facetious question the author has replied: "I think Ne will still have a lot of
general interest as the "currency" of evolution, so that you
make predictions about processes in unstudied systems"]. robin.waples@noaa.gov
343. Growth rate and disease resistance
negatively correlated
Genetic variation
for growth rate, feed conversion efficiency, and disease resistance exists
within a farmed population of rainbow trout. 2002. Henryon, M., A.
Jokumsen , P. Berg, I. Lund, P.B. Pedersen, N.J. Olesen, and W.J.
Slierendrecht. Aquaculture 209 (1-4):59-76.
An excellent experimental
design for aquaculture genetic studies was used here -- an incomplete
factorial in which each male was mated with two females and each female
with two males. Each full-sib family was grown out separately, with
periodic random culling to reduce the biomass density in the tanks.
Feeding was restricted, with the ration determined by the anticipated
growth rate.
The results of the
genetic (animal model) analysis of size-at-age and feed conversion
efficiency were somewhat odd, in that the genetic variance and covariance
components differed between the successive stages of growout, but the
differences didn't show any easily interpreted pattern. Growth and feed
conversion efficiency were observed to be positively correlated, which the
authors ascribe to the dual effect of the same genes on both traits.
Correlation would have been expected anyway, however, because of the way
in which feeding rates were fixed. The authors discuss this problem. There
were also problems with the genetic variance-covariance matrix which may
(in my opinion) have been the result of competition among individuals in a
tank. The statistically independent (individual) effects are not really
independent at all. This problem would be exacerbated by restricted
feeding but it may turn out to be insoluble with current genetic models
and estimation procedures even when fish are fed ad libitum. Which is not
to say that trout will fail to show a realized response to selection for
increased growth.
Additive genetic
variation for survival was low in these experiments but overall survival
was high except in the disease challenge experiments.
The analysis of the viral
haemorrhagic septicaemia (VHS) challenge experiments is an admirable model
for aquacultural geneticists. The "Survival Kit" software of
Ducrocq and Sölkner http://www.boku.ac.at/nuwi/software/softskit.htm
was used to estimate individual breeding values and population-level
genetic variances and covariances for "frailty", a technical
term which can be interpreted intuitively as the probability that an
animal will die from the disease. Additive genetic variance for VHS
resistance did exist and the estimated heritability was 0.13. This could
be important in practical selection programs. "Only 63% of the fish
from the sire and dam with high resistance were predicted to die after 21
days following challenge with VHS. By contrast ... all of the fish from
the sire and dam with low resistance were predicted to die after only 12
days following challenge."
Most interestingly,
genetic variation for resistance to VHS tended to be negatively correlated
with genetic variation for growth. This is potentially very important for
the design of broodstock improvement programs, as the authors point out.
The cautions expressed above about the growth-rate variance estimates also
apply to the estimates of covariation between growth and VHS resistance,
however. Mark.Henryon@agrsci.dk
342. Initial advantage, terminal
disadvantage to immigrant genes
Heterosis and
outbreeding depression in descendants of natural immigrants to an inbred
population of song sparrows (Melospiza melodia). 2002. Marr, A.B., L.F.
Keller, and P. Arcese. Evolution 56 (1):131-142.
Does immigration help a
small population by reducing inbreeding depression, or does it do harm by
bringing in genes that are poorly adapted to local conditions? We hardly
ever know the answer to this question in the real world. Therefore this
new information on the sparrows living on the tiny island of Mendarte is
very interesting. The resident population includes several hundred birds
of each sex. The annual immigration rate is less than 5%.
The effect of an
occasional immigrant on genetic diversity and inbreeding depression of the
population has been published earlier (Sep 2001 #235). Now it appears that
the F1 hybrid offspring of residents and immigrants are somewhat more fit
than the local residents in the same generation, a fact which the authors
attribute to heterosis. The F2 generation, however, was much less fit than
would be predicted merely from the breakdown of F1 hybrid vigour due to
segregation of loci. The authors found "large performance differences
between F1s and F2s, which suggested that either heterosis was associated
with epistasis in F1s, that F2s experienced outbreeding depression, or
that both phenomena occurred". The authors prefer outbreeding
depression as an explanation – i.e. the breakdown of locally adapted
gene complexes. amarr@interchange.ubc.ca
341. Rapid immigration of genes caused by
hybrid vigour
A selective
advantage to immigrant genes in a Daphnia metapopulation. 2002. Ebert,
D., C. Haag, M. Kirkpatrick, M. Riek, J.W. Hottinger, and I. Ilmari
Pajunen. Science 295 (5554):485-488.
Should tiny, endangered
populations of West Coast salmonids be kept pure and uncontaminated by
stock from elsewhere? More generally, is immigration good or bad for small
populations from a genetic point of view? Good, presumably, if the
population is suffering from inbreeding depression and has drifted away
from its fitness optimum. Presumably bad if the population is uniquely
well adapted to its local environment. Complicated, if inbreeding
depression purges the population of deleterious recessive genes and
immigration then brings them back (e.g. Sep 2001 #235).
This paper reports on a
genetic study of Daphnia magna populations in small shoreline pools in
Scandinavia. Every now and then one of the pools goes extinct and is
recolonized from some other pool(s). Sometimes, when pools are recolonized
by only one individual, there is a period of clonal reproduction before
sexual reproduction takes place between identical, but not homozygous,
individuals. This is a classic metapulation in which inbreeding and loss
of genetic diversity are major evolutionary considerations.
The authors
experimentally manipulated immigration rates both in natural pools and in
laboratory populations. They found that the rate of introgression of
immigrant genes was enhanced by hybrid vigor of the offspring in almost
all cases. That is, fitness increased, first at the individual and then
the population level. Since different immigrant clones were used in each
experiment the authors conclude the effect was due to hybrid vigour not to
the selective superiority of particular alleles.
"One key effect of
hybrid vigor is an increase in effective gene flow ...[in these
experiments] the effective rate of gene flow is about 35 times larger than
would be predicted by the number of immigrants alone. ... An important
effect of hybrid vigor is the "genetic rescue" of populations
from extinction .... In this light, gene flow is an essential component
for the persistence of metapopulations. Thus, our study gives clear
empirical support for the need to maintain gene flow in the management and
conservation of subdivided populations." dieter.ebert@unifr.ch
340. Inbreeding is bad even in good
environments
Relationship
between coefficient of inbreeding and parasite burden in endangered
gazelles. 2001. Cassinello, J., M. Gomendio, and E.R.S. Roldan.
Conservation Biology 15 (4):1171-1174.
Pedigree records on
captive populations of these three species of gazelle enabled the authors
to calculate inbreeding levels exactly, rather than estimating them
indirectly from heterozygosities (as for example in Apr-May 2002 #320,
June 2001 #207, Mar 2000 #32). The species with the highest inbreeding
also had the highest intensity of nematode infection. Within that species
(but not the other two) there was a positive correlation between an
individual's parasite burden and its inbreeding coefficient. The authors
conclude that when inbreeding is high, it makes individuals more
susceptible to parasites even in a relatively benign artificial
environment. cass@mncn.csic.es
339. Fast growing Frankenloaches
Accelerated growth
performance and stable germ-line transmission in androgenetically derived
homozygous transgenic mud loach, Misgurnus mizolepis. 2002. Nam, Y.K.,
Y.S. Cho, H.J. Cho, and D.S. Kim. Aquaculture 209 (1-4):257-270.
These fish are truly
bioengineered. First a growth-promoting transgene (mud-loach growth
hormone fused to mud-loach h-actin regulatory region) was introduced into
the loach genome, then sperm from the transgenic loach were used to
fertilize eggs of a related fish (common carp) in which the female genome
had been destroyed by UV irradiation. Finally, a carefully-timed heat
shock was used to double the male (loach) haploid genome. "One
homozygous transgenic line displayed consistently faster growth (up to
1.6-fold) than its heterozygous transgenic counterpart. Conversely, growth
acceleration in the other two homozygous transgenic lines was limited to
the first 3 months of age only, and subsequently grew far slower than
their heterozygous transgenic, especially when some physiological and
morphological abnormalities began to appear." dongskim@nuri.net
338. Selection of laid-back trout
Differences in
behaviour between rainbow trout selected for high- and low-stress
responsiveness. 2002. Overli, O., T.G. Pottinger, T.R. Carrick, E.
Overli, and S. Winberg. Journal of Experimental Biology 205 (3):391-395.
Is it possible to speed
the domestication of farmed trout by selecting a line which is indifferent
to stress? Trout which showed either high or low plasma cortisol levels
after a standardized stress were selected as breeders. The offspring of
these divergent lines also differed in their mean post-stress cortisol,
showing that the endrocrinological response to stress was heritable in the
parental population. Offspring in the high-cortisol line (sensitive to
stress) were more active than the low-cortisol line, but only when an
intruding fish was present. "A significantly higher incidence of feed
intake was seen in [low cortisol] trout when held in observation tanks (40
% versus 0 % of the fish took food when in isolation), suggesting that
these fish acclimated more successfully to the experimental conditions
than [high cortisol] fish did." So it appears that one-generation of
selection for reduced physiological stress response produced trout that
were less excitable and fed more than fish selected in the opposite
direction. ooverli@usd.edu
337. Fluctuating asymmetry important, but
maybe not genetically important
Fluctuating asymmetry is
non-genetically related to mating success in Drosophila buzzatii. 2001. Santos, M. Evolution 55 (11):2248-2256.
Fluctuating asymmetry
(FA) is the occurrence of random differences between the left and right
side of the body. Since a body usually has the same genotype on both
sides, FA reflects random failures of developmental stability. It is known
that stability can be decreased, and FA increased, by the non-heritable genetic stress of inbreeding, and by environmental stress (Jun 2000
#70). Because FA is easy to measure it is potentially useful as an
indicator of poor genetic quality in aquaculture and genetic conservation.
But in addition to inbreeding, is it also a signal of heritable
differences in the fitness of animals in particular environments, e.g. in
their reproductive fitness?
In these experiments with
laboratory and wild Drosophila populations, males with higher FA did
indeed have lower reproductive success, but "the higher FA in single
males is most likely due to a poorer average phenotypic condition because
there was no evidence of a genetic basis for this trait".
There is as yet no
consensus on the usefulness of FA as a general indicator of genetic
quality: Jun 2000 #70 FA not related to components of fitness in D.
melanogaster; Jul 2000 #82 FA related to inbreeding and fitness in
ungulates; Feb 2001 #167 FA not heritable in chinook salmon; Feb 2002 #292
not related to heterozygosity (inbreeding?) in partridge. This paper makes
a useful methodological contribution to estimating FA when the animals
also exhibit non-random variation in symmetry ("handedness"). mauro.santos@uab.es
336. Selecting temperature-insensitive sex
determination in bass
Temperature
effects and genotype-temperature interactions on sex determination in the
European sea bass (Dicentrarchus labrax L.). 2002. Saillant, E., A.
Fostier, P. Haffray, B. Menu, J. Thimonier, and B. Chatain. Journal of
Experimental Zoology 292 (5):494-505.
The European sea bass is
an important aquacultural species. Females grow faster than males and are
favoured for cultivation, but unfortunately the sex ratio is biased
towards males. In the work reported here 27 sea bass families were
produced according in a factorial mating design. Microsatellites were used
to sort out the parentage of animals reared together. The sex ratio was
affected by temperature, with more males developing when the fish were
grown at lower temperatures. (The temperature effect on tilapia sex ratio
goes the other way.) There was an interaction between the temperature
effect and the developmental stage at which the treatment was applied.
" In groups reared at high temperature, both parents had a
significant additive effect on the percentage of females, and the
interaction between sire and dam was not significant. Genotype temperature
interactions were also detected and their existence suggests the
interesting possibility of selecting nonsensitive genotypes in breeding
programs." esaillant@wfscgate.tamu.edu
335. Procedure for reducing inbreeding
during artificial selection
Non-random mating
for selection with restricted rates of inbreeding and overlapping
generations. 2002. Sonesson, A.K., and T.H.E. Meuwissen. Genetics
Selection Evolution 34:23-39.
The authors of this paper
have applied (in theory and simulation) the concept of minimal kinship
selection to the problem of artificially selecting a trait such as growth.
Minimum kinship (= minimum coancestry mating; Dec 2001 #270, Nov 2001
#261) is generally used to reduce long-term inbreeding and genetic drift
in a broodstock. One of the problems afflicting aquaculture is the high
rate of inbreeding that results from various culture procedures which are
hard or expensive to change. Fish in hatcheries conspire to inbreed even
when they are not artificially selected. When they are selected the
inbreeding problem gets a lot worse.
This paper shows that when
animals are selected on their merits (e.g. by phenotypic selection or BLUP)
and the matings among them are arranged to minimize kinship, not only is
the accumulation of inbreeding reduced but the rate of genetic gain is
increased (consult paper for details). None of this would have been
relevant to aquaculture 5 years ago because pedigree records are required
to use these procedures. However, we can now use DNA markers instead of
pedigrees to minimize kinship (e.g. January 2002 #283) and to estimate
breeding values when animals are reared in common pools, as in aquaculture
(Jun 2002 #330).
Breeder populations of
moderate size (a few hundreds), having one or few selected offspring per
mating, are optimal for combining minimal coancestry with selection,
according to this paper. Good news, because it is also the aquacultural
setup which would lead to very high selection intensities and great
savings in hatchery infrastructure. anna.sonesson@akvaforsk.nlh.no
334. Evolution of spawning date in captive
Pacific salmon
Artificial
selection and environmental change: countervailing factors affecting the
timing of spawning by coho and chinook salmon. 2002. Quinn, T.P., J.A.
Peterson, V.F. Gallucci, W.K. Hershberger, and E.L. Brannon. Transactions
of the American Fisheries Society 131 (4):591-598.
Water temperatures in
salmon hatcheries in the U.S. state of Washington have been getting warmer
over the years. This is an environmental factor that would normally
produce a later spawning date in the two species of Pacific salmon
studied here. However, breeding and spawning procedures in the hatchery
since the 1950s for chinook and 1960s for coho have been selecting for earlier spawning. Given that spawning date is known to be highly heritable, which
way has it been moving? Records kept in three hatcheries including one run
by Washington State University show that spawning is occurring earlier and
earlier as the years go on. "Thus, inadvertent selection at all three
hatcheries appears to have resulted in progressively earlier spawning,
overcoming selection from countervailing temperature trends". tquinn@u.washington.edu
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