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.
211. Salmon Bacchanalia
in the River Dee
Spawning success in Atlantic
salmon (Salmo salar L.): a long-term DNA profiling-based study conducted
in a natural stream. 2001. Taggart, J.B., I.S. McLaren, D.W. Hay, J.H.
Webb, and A.F. Youngson. Molecular Ecology 10 (4):1047-1060.
The authors used minisatellite
markers to assign parentage to salmon progenies (eyed ova) collected from
an 8 km long stream in Scotland over three years. Below the surface of
this turbid Presbyterian tributary the behaviour of the salmon is
shockingly orgiastic.
"Multiple spawning was found to be prevalent. More than 50% of
anadromous spawners of both sexes contributed to more than one redd. Up to
six redds for a single female and seven for a single male were detected.
Both sexes ranged extensively. Distance between redds involving the same
parent varied from a few metres to > 5 km. Distances > 1 km were
common. Both males and females ranged to a similar extent. Range limit was
not correlated to fish size. Pairs were not monogamous, both males and
females mating with different partners at different sites. ... Redd
superimposition was found to be common . ... High levels of nonanadromous
mature parr mating success (40–50% of total progeny sampled) were
recorded, and these likely contribute greatly to the effective population
size."
The authors also note that parr
maturation is not detrimental to this stock. "...Mature parr appear
to exercise a viable life history strategy with high fitness which
contributes to population structure and persistence." j.b.taggart@stir.ac.uk
210. Confirming evidence for mutational
meltdown in small populations
Rapid mutational declines of
viability in Drosophila. 2001. Fry, J.D. Genetical Research 77:53-60.
"Mutational meltdown"
is the name given to one of the many bad things that can happen when a
population becomes too small. Deleterious mutations accumulate quickly and
the resulting decrease in fecundity and survival cause the population to
become even smaller. The positive feedback between declines in population
size and fitness can draw the population into an "extinction
vortex" (March-April #185, February #171, January #157). However,
doubts have been expressed concerning some of the key laboratory experiments
demonstrating the effect. Given their implications for genetic
conservation we would like to know whether the doubts are justified.
Apparently not. The author of
this paper writes "Here, using a [new method], I reanalyse the
previous mutation-accumulation (MA) experiments, and report the results of
a new one. I show that in each of four experiments, including Mukai's two
experiments, viability declines due to mildly deleterious mutations were
rapid. " jfry@mail.rochester.edu
209. The Neolithic birth of domestic breeds
Multiple maternal
origins and weak phylogeographic structure in domestic goats. 2001.
Luikart, G., L. Gielly, L. Excoffier, J.-D. Vigne, J. Bouvet, and Taberle.
P. Proceedings National Academy of Sciences (USA) 98 (10):5927-5932.
As well as being intrinsically
interesting, this is an unusually instructive paper because of the variety
of genetic and statistical techniques used to discover how, where and when
the domestic breeds of these useful animals evolved. The authors based
their analysis of the divergence of goat breeds on mitochondrial DNA
sequences from the rapidly-evolving control region of the mitochondrial
genome and calibrated the time scale with a molecular clock based on the
divergence of the sheep and goat mitochondrial cytochrome b gene.
Three main lineages of goat breeds
(80 breeds in total were studied) appear to have diverged 2 or 3 hundred
thousand years ago, i.e. long before the animals were first domesticated.
This evidence points strongly to the inference that goats were
domesticated from local wild animals at least three times, first about
10,000 years ago in southern Turkey/Euphrates valley, and then again
sometime between 3,000 -- 6,000 years ago farther to the east in the
fertile crescent (Iran) and in Asia (possibly northern India). The lineage
from the oldest domestication is much more genetically diverse than the
other two, as would be expected from this timing. The authors were also
able to estimate the time at which the initial domesticates underwent a
population expansion [for techniques see #202 below] and found that the
newer lineages expanded about 2,000 and 6,000 years ago relative to a
presumed 10,000 year date for the expansion of the first domesticate.
The authors note that a similar
pattern exists in cattle, sheep, and pigs. However, "Goat populations
are surprisingly less genetically structured than cattle populations. In
goats only 10% of the mtDNA variation is partitioned among continents. In
cattle the amount is 50%. This weak structuring suggests extensive
intercontinental transportation of goats and has intriguing implications
about the importance of goats in historical human migrations and
commerce." The authors suspect that goat genes flow travel rapidly
because goats are more transportable and also because they represent
smaller and more durable units of wealth than cattle. gordon.luikart@ujf-grenoble.fr.
208. Complicated behavioural evolution in
domesticated salmon
Behavioural and heart
rate responses to predation risk in wild and domesticated Atlantic salmon. 2001. Johnsson, J.I., J. Höjesjö, and I.A. Fleming. Can. Jour. Fisheries
and Aquatic Sciences 58:788-794.
In this study
seventh-generation farmed salmon were compared with wild salmon from the
same founder population. One-year old wild fish (1+) had a stronger heart
rate and flight response from a simulated predator than the domesticated
fish, but the differences were weaker or reversed in two-year-old fish.
The authors suggest that if there is a genetic domestication response it
is not a simple one. It involves the "reaction norm", i.e. the
parameters of the functional interaction between current environmental
stress, prior environmental history, and the state of development.
"This should be considered when attempting to predict the
consequences of release or escape of domesticated animals in the
wild." jorgen.johnsson@zool.gu.se
207. Do population bottlenecks reduce the
ability to evolve?
Effects of bottlenecks
on quantitative genetic variation in the butterfly Bicyclus anyana. 2001. Saccheri, I.J., R.A. Nichols, and P.M. Brakefield. Genetical
Research 77:167-181.
How does a bottleneck, a brief
period of very low numbers, affect the ability of a population to meet new
selective challenges and evolve new adaptations? This is an important
question for aquaculture broodstock management as well as for genetic
conservation.
We know that only a random
sample of genes (alleles) will pass through the bottleneck while other
genes are lost by chance during the period when few matings take place. In
an extreme case, with only one pair of breeders passing through the
bottleneck, only 4 alleles per locus will be available for population
re-growth and evolutionary adaptation. Some of those alleles may be
identical. Either of two things can happen to evolutionary capacity in a
bottleneck, depending on how the surviving genes combined with each other
to influence the phenotype in the pre-bottleneck environment:
(1) The effects of
individual genes simply add together. Genetic variance will decrease when
genes are lost by chance, and the rate at which the population can evolve
will therefore decrease because of the bottleneck.
(2) Pairs of genes
interact non-additively to produce their effects on the phenotype, either
in combination at a single locus (called dominance) or at different loci
(called epistasis). Non-additive genetic variance does not respond in a
simple way to selection in sexual species even though it contributes to
phenotypic variation among individuals. Generally speaking, only additive
genetic variance allows an evolutionary response to selection. Theory has
shown, however, what when some alleles are lost by chance from a
non-additive genetic system the surviving genes will sometimes make an
increased contribution to additive genetic variance. In this case the
capacity to evolve will be enhanced by the bottleneck. (See also
October 2000 #116, #112)
So much for theory. What
actually happens? (See February list #165 for an instance in which genetic
diversity did not reduce neutral microsatellite diversity in sockeye
salmon.) This interesting experimental paper makes a strong case for
believing that additive variance will increase in traits closely related
to fitness, like fecundity and survival. Quantitative traits that are
under weak selection, on the other hand, will lose additive genetic
variance in the bottleneck, just like neutral marker alleles. The
explanation is that traits related to fitness have an especially high
proportion of dominance variance. Recent papers pointing out this
difference between the genetic architectures of fitness and non-fitness
traits have already been noted on this website (February 2000 #15 and
#10).
The present experiment
describes what happens when butterfly populations are bottlenecked to
different extents in the laboratory. Additive genetic variance of wing
size (a neutral trait) decreases in bottlenecks according to theoretical
expectation while additive variance and heritability of egg hatching rate
(a component of fitness) increases. The authors use genetic markers in an
interesting way to calculate inbreeding in the bottleneck (see May list
#193), which allows them to show that the change of wing-size variance,
but not hatching, follows random sampling expectation exactly. saccheri@liverpool.ac.uk
206. No evolution of crossover resistance
to whirling disease
Salmonids resistant to
Ceratomyxa shasta are susceptible to experimentally induced infections
with Myxobolus cerebralis. 2001. Hedrick, R.P., T.S. McDowell, K.
Mukkatira, and M.P. Georgiadis. Journal of Aquatic Animal Health 13:35-42.
Whirling disease is a nasty,
cartilage-destroying condition of salmonids caused by the myxosporean
pathogen Myxobolus cerebralis. Strains of rainbow trout with known
differences in susceptibility to another myxosporean, Ceratomyxa shasta,
were exposed in the laboratory to waterborne infectious stages of whirling
disease. "In general [possible exception of one strain] the
mechanisms of resistance that have developed among certain salmonid
populations to one myxosporean, C. shasta, do not extend [or extend
minimally] to protection or resistance to a second myxosporean, the
causative agent of salmonid whirling disease". The mechanisms by
which some rainbow trout resist infection by C. shasta is unknown, but
breeding experiments suggest that its genetic architecture is polygenic
and at least partly additive. rphedrick@ucdavis.edu
205. Useful software for genetic analysis
TOOLS: Phylogenetics
Crunch. 2001. Anonymous. Science 292 (5518):815.
"Wayne and David Maddison
at the University of Arizona are working on Mesquite, a modular system for
evolutionary analysis. Their goal is to build a Java-based system that
makes it possible to integrate disparate packages, such as the popular
PAUP program and MacClade, which the same duo also developed. Download a
beta version of Mesquite at: http://mesquite.biosci.arizona.edu/mesquite/mesquite.html "
The website has additional
information, "Modules written include ones for basic character
analysis (parsimony and likelihood), comparative biology, molecular
evolution, population genetics and morphometrics. The Mesquite
documentation has a summary of plans as to what it will do. ... Also, we
plan to increase Mesquite's connectedness in the world outside the local
hard disk via distributed processing and interaction with Internet
databases such as the Tree of Life . ... we hope to have it link to other
programming efforts including Drummond and Strimmer's PAL." wmaddisn@u.arizona.edu.
204. Pacific salmon Y chromosome
Identification of the Y chromosome in
chinook salmon (Oncorhynchus tshawytscha). 2001. Stein, J., R.B.
Phillips, and R.H. Devlin. Cytogenet Cell Genet 92 (1-2):108-110.
"The Y chromosome in
chinook salmon, Oncorhynchus tshawytscha, was identified using
fluorescence in situ hybridization (FISH) with a probe to a male-specific
repetitive sequence isolated from this species. The probe highlights the
distal end of the short arm of an acrocentric chromosome with a DAPI-bright
interstitial band of variable size. " This will be a useful technique
for ploidy manipulation and other types of genetic study including extreme
linkage analyses that make use of androgenesis. jstein@csd.umw.edu
203. Recognizing evolutionary adaptation to
local environments
Nonclinality of
molecular variation implicates selection in maintaining a morphological
cline of Drosophila melanogaster. 2001. Gockel, J., W.J. Kennington,
A. Hoffmann, D.B. Goldstein, and L. Partridge. Genetics 158:319-323.
When populations differ
genetically in a trait that appears to affect fitness, it may be that the
populations are adapted to different environments. On the other hand, the
genetic differences may have arisen merely by accidents of population
dynamics and dispersal and not reflect adaptive differences at all. These
two alternative explanations have very different implications for genetic
conservation (since neutral variation is probably less worth conserving
than adaptive variation) , for the definition of evolutionarily
significant units and, in the case of fisheries, for choosing broodstock
for aquaculture, enhancement and rehabilitation of endangered wild
populations.
In this paper the authors have
compared the body size of Drosophila with neutral variation at 16
microsatellite loci to see whether variation in body size is adaptive over
a 2000 km north-south transect in Australia. Body size was several
standard deviations smaller at the extreme north than the extreme south.
Does this represent adaptation? The authors found that regression of body
size on latitude explained approximately 80% of the variance, i.e. there
is a strong correlation. Regression of their measure of supposedly neutral
gene frequency on latitude explained only about 20% of the variation on
average, although one of the 16 loci had 76% of its variance explained by
latitude. The conclusion of the paper is that body size is indeed under
selection. Some of the loci may be under selection or closely linked to
quantitative trait loci (QTL) that are under selection.
One of the innovative features
of this paper which makes it interesting to fisheries is that the authors
have used regression on an extrinsic variable to compare phenotypic with
neutral variation. This allows them to study an explicit environmental
variable, namely latitude. Presumably adaptation of morphological and
behavioral traits to temperature, photoperiod, salinity or other
continuous but not necessarily clinal variables could be studied in the
same way. It might even be a fast way to identify candidate markers for
QTL. ucbhlop@ucl.ac.uk [Linda
Partridge]
202. Use DNA markers to infer historical
changes in population size
A power analysis of
microsatellite-based statistics for inferring past population growth.
2000. King, J.P., M. Kimmel, and R. Chakraborty. Molecular Biology and
Evolution 17 (12):1859-1868.
The authors evaluate the
relative power of several different statistical procedures that use
microsatellite data to detect past population expansion. They find that
the statistical test based on the imbalance between allele size variance
and heterozygosity is better for detecting population growth than are
methods based on between-locus variability or allele size distributions.
The underlying concepts are presented with admirable clarity in this
paper. rc@hgc9.sph.uth.tmc.edu
201. Differentiating European carp strains
Genetic variability in
reared stocks of common carp (Cyprinus carpio L.) based on allozymes and
microsatellites. 2001. Desvignes, J.F., J. Laroche, J.D. Durand, and
Y. Bouvet. Aquaculture 194:291-301.
The authors looked at allozyme
and microsatellite gene frequencies in a number of cultured carp strains
in France and the Czech Republic. Rather surprisingly, heterozygosity was
lower in the French strains but the number of alleles was higher. The
difference between these two measures of genetic diversity may be the key
to sorting out what has happened under the different cultivation and
selection regimes in the two regions of Europe. Generally speaking the two
marker systems told the same story, but the authors judge that
microsatellites are better at discriminating strains within and between
countries. laroche@cismsun.univ-lyon1.fr
200. A crustacean evolves, loses and
re-evolves anti-predation behaviour
Rapid, local adaptation
of zooplankton behavior to changes in predation pressure in the absence of
neutral genetic changes. 2001. Cousyn, C., L. De Meester, J.K.
Colbourne, L. Brendonck, D. Verschuren, and F. Volckaert. Proceeding of
the National Academy of Sciences (USA) 98:6256-6260.
The waterflea Daphnia magna
regularly produces resting eggs that can survive for decades in mud until
conditions are right for emerging. The authors of this article hatched out
resting eggs collected at various depths in the sediment of a 30-year-old
man-made pond in Belgium. Daphnia and fish have coexisted in the pond over
its entire history, but the density of fish, and therefore the intensity
of fish predation in the environment, has varied considerably. The
question asked by the authors was whether the waterfleas adapted
genetically to the changing predation. It is known that Daphnia which
live in ponds characterized by strong predation pressure tend to avoid the
light in the presence of chemical substances (kairomones) released by
fish. This was the possibly adaptive trait, a change in phototactic
behavior in the presence of "fishwater". Three microsatellite
loci provided a presumably neutral genetic trait with which the behavioral
trait was compared.
The genetical logic, although not the
statistical procedures, is the same in this paper as in #203
above. Much of the genetic variance in the behavior of clones developed
from the buried eggs was explained by historical variation in fish
predation, which constituted the independent environmental variable. The
neutral microsatellite frequencies varied little over the 30 year period.
Therefore the observed genetic change in phototaxis was considered to be
an adaptive evolutionary response to changes in the local selective
regime.
Whether the behaviour of crustaceans such as Penaeid shrimp would
evolve similarly in the presence/absence of fish predation remains to be
discovered. If it does it might have interesting genotype-environment
interaction effects when broodstocks are selected for extensive
aquaculture and/or "fishwater free" (?) intensive aquaculture. Luc.DeMeester@bio.kuleuven.ac.be.
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