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.
96. Shrimp: the host vs. pathogen evolutionary arms race
Evolution of parasite virulence against qualitative or quantitative
host resistance. 2000. Gandon, S., and Y. Michalakis. Proceedings of
the Royal Society (UK) Series B 267:985-990.
The authors distinguish between qualitative or absolute host resistance to a pathogen, in which
there is no infection whatsoever, and quantitative or partial host resistance
in which the spread of a pathogen within the host occurs but at a reduced
rate. They show that "a qualitative form of resistance reduces
parasite virulence, while a quantitative form of resistance generally
selects for higher virulence". This is because after
successfully infecting the host, the variable pathogen
genotypes will compete to overcome the partial resistance, and pathogen evolution by natural selection
will take place.
The paper is one of the few that
studies the linked coevolution of a host and its pathogen. Most studies
are concerned with the evolution of only one of the pair while the other
is assumed to remain unchanged. In shrimp aquaculture, this is not a
reasonable assumption.
The analysis is consistent with
the work reported in July list #84, where a somewhat different distinction
is made between host resistance (both quantitative and qualitative) and
host tolerance. In the case of increasing host tolerance the virulence
of a pathogen does not evolve by natural selection. This is because the
relative fitness of pathogen genotypes will not be influenced by
improved tolerance.
If we consider these two papers together, it would seem
that host tolerance, for a pathogen which is always present in the
population, is evolutionarily stable, and may be a desirable target for
selective breeding. Absolute, qualitative resistance is also a desirable
trait to select if there are any such genes available in the host
population. However, selecting
for a quantitative increase in resistance appears to invite a
coevolutionary arms race with the pathogen. Or in the case of an artificially
applied quantitative resistance -- for instance an antibiotic dip that
is less than 100% effective -- we should have a coevolutionary arms race
between the pathogen and the pharmaceutical industry.
sgandon@snv.jussieu.fr
95. The problem for fish conservation is entropy,
not extirpation
Homogenization of fish faunas across the United States. 2000.
Rahel, F.J. Science 288:854-856.
European settlers introduced
fish into North America for sport and for food. "Fish faunas across
the continental United States have become more similar through time. ...
On average, pairs of states have 15.4 more species in common now than
before European settlement of North America. The 89 pairs of states that
formerly had no species in common now share an average of 25.2 species.
Introductions have played a larger role than extirpations [of local
endemic species] in homogenizing fish faunas." frahel@uwyo.edu
94. Does ex situ conservation work? Loss
of fish genes
Effective population size and maintenance of genetic diversity in
captive-bred populations of a Lake Victoria cichlid. 2000. Fiumera ,
A.C., P.G. Parker, and P.A. Fuerst. Conservation Biology 14:886-892.
The endangered cichlid species
had been maintained in captive subpopulations for five generations as part
of a Lake Victoria cichlid species survival plan (the major threats being over fishing
and introduced Nile perch). A study of four microsatellite
loci (24 alleles) showed that the effective population size was only 1% to
10% of the observed census size in the gene bank. Not only that, around 20% of the alleles
were lost in the first 4 generations. Expected diversity declined by 6% -
12% per generation. The subpopulations are being maintained in different
places, in facilities that include one to several tanks of 10 - 30 adults.
None of this sounds very good,
either for genetic conservation by ex situ methods (i.e. where fish
are maintained in "zoos" outside their natural habitat) or for tilapia aquaculture (to the extent
that the conclusions can be generalized). The authors make several
recommendations for reducing genetic loss including "periodically
removing dominant males, which will encourage reproduction by additional
males". The authors did not attempt to calculate the rate of
accumulation of inbreeding in the populations. Readers should think about the
way inbreeding in small populations amplifies male-male competition and
reduces genetic variance (July #82, June #60, May #48). fuerst.1@osu.edu
93. All-male tilapia and opportunities for
Frankensex
Sexually dimorphic expression of two types of DM (Doublesex/Mab-3)-domain
genes in a teleost fish, the tilapia. 2000. Guan, G. , T. Kobayashi,
and Y. Nagahama. Biochemical and Biophysical Research Communications
272:662-666.
The authors have identified two
sex-determining genes in tilapia that appear to be different versions of
the "doublesex" gene first identified in Drosophila. One version is already known to be found in vertebrates
(including zebra fish) and is transcribed mainly in the testis. The authors
found that tilapia also has a very different variant of this
same gene which is expressed in the ovary and
is structurally somewhat like the female version of the Drosophila
doublesex gene. The expression of the two tilapia genes is mutually
exclusive in any particular fish (which must become either male or female
during development), and so is presumably under the control of additional
genes and environmental factors.
This important discovery of two doublesex genes may ultimately, perhaps, help explain the spooky sex
ratios that often result when tilapia species and even populations are
crossed. It might also become of practical interest to aquaculture if some
of the truly bizarre transgenic experiments in Drosophila could ever be
replicated in fish for the purpose of controlling sexual differentiation.
The authors of this paper make it clear that there are likely to be many targets for
genetic intervention in the teleost sex-determining system.
A note on Drosophila.
Drosophila has only one doublesex gene, not two like tilapia, and produces
male and female versions by RNA splicing after transcription. Sexual differentiation in Drosophila is controlled by a short cascade of
regulatory genes, the expression pattern of which determines all aspects
of maleness and femaleness, including complex behaviors displayed by males
and females. Doublesex is found near the end of the cascade. Other genes in this pathway have been given names like
"transformer" and "dissatisfaction".
Mutational and
transgenic manipulation of certain genes in the cascade have produced
flies that are genetic (XX) females but develop and behave as males. This
genetic sex-reversal can be blocked by transforming another gene in the
cascade. But if the blocking gene is coupled with a suitable promoter the
blocking action can be stopped by applying a heat shock during
development. Then, although the transformed female flies develop as males
and display vigorous male courtship, they are still producing female
pheromones and are themselves attractive to males. Their chronically
amorous condition (as males) may be caused by self-stimulation from their
own female pheromones. Whew! At this point we must draw a veil
across their behaviour but note again that the analogous
tilapia doublesex system could provide transgenic opportunities useful in
aquaculture. nagahama@nibb.ac.jp
92. Does ex situ genetic conservation
work? Loss of plant genes
Genetic diversity of barley landrace accessions (Hordeum vulgare
ssp. vulgare) conserved for different lengths of time in ex situ gene
banks. 2000. Parzies, H.K., W. Spoor, and R.A. Ennos. Heredity
84:476-486. There are large numbers of samples of seeds of crop plants
stored in gene banks throughout the world. These seeds must be planted on
a regular basis and replaced by offspring seed. In the course of this
rejuvenation of the collection there is an opportunity for gene frequency
changes and erosion of genetic diversity due to "bottlenecking"
and genetic drift.
"In order to determine
whether these undesirable effects occur, genetic diversity levels were
assessed for morphological and isozyme markers within gene bank accessions
of two barley landraces from Syria that had been stored for 10, 40 and 72
years. These were compared with genetic diversity levels for the same
markers in barley landraces collected recently at locations in Syria where
they are still under cultivation. Average gene diversity, alleles per
locus and percentage polymorphic loci all showed very
significant declines with length of time in storage, and genetic
differentiation among accessions increased over time.... [It] was estimated that the
effective population size Ne of rejuvenation populations over their period
in storage was only 4.7."
The authors also note that the
"consistently high diversity of all recently collected barley
landraces found in this study is a clear justification for the use of in
situ conservation approaches. ("In situ" conservation
means conserving organisms in their natural location and habitat, like a
national park.) rennos@ed.ac.uk
91. How fish evolve genetic adaptation to thermal
stress
Adaptive variation in lactate dehydrogenase-B gene expression: Role
of a stress-responsive regulatory element. 2000. Schulte, P.M., H. C.
Glémet, A.A. Fiebig, and D.A. Powers. Proceedings of the National Academy
of Sciences (USA)
97:6597-6602.
Northern and southern
populations of the fish Fundulus heteroclitus have different levels of
expression of the lactate dehydrogenase-B gene (Ldh-B). The northern, e.g.
Newfoundland, fish are superior at lower temperatures while the southern,
e.g. Florida, fish are superior at higher temperatures. The differences
are known to be genetic. This exceptionally interesting study is one of
the first to elucidate how adaptive gene regulation
can evolve. The experimental details are too complicated to be easily
summarized here, but they include temporary transgenesis of the
regulatory sequences into the livers of
experimental fish; deletion studies to identify the approximate location
within the regulatory sequence where the adaptive changes in the
transcript occurred; stress tests of live fish to see which alleles (northern or
southern) drive the transcription of the gene.
The authors found that a
difference of only one base pair in the regulatory sequence accounts for
the adaptive difference between the northern and southern
populations. Marker mappers and QTL selectors should
take heart, but also note the experiment reported in June #65 where
selection for temperature stress tolerance did not work. pschulte@uwaterloo.ca
90. Endangered horsemeat sushi
Predicted decline of protected whales based on molecular genetic
monitoring of Japanese and Korean markets. 2000. Baker, C.S., G.M.
Lento, F. Cipriano, and S.R. Palumbi. Proceedings of the Royal Society (U.K)
Series B 267:1191-1199.
The authors used mtDNA data to
identify the species of products sold as whale meat (which turned out to
include sheep and horse as well as many species of whale, porpoise and
dolphin) and sub-populations (which included a greater than expected
number of samples from protected minke whales in the Sea of Japan).
"The source of these products was assumed to be undocumented
'incidental takes' from fisheries' by-catch, although we cannot exclude
the possibility of illegal hunting or smuggling. The demographic impact of
this undocumented exploitation was evaluated using the model of population
dynamics adopted by the Scientific Committee of the International Whaling
Commission. For the range of exploitation consistent with the market
sample, this protected stock [Sea of Japan] was predicted to decline
towards extinction over the next few decades. These results confirmed the
power of molecular methods in monitoring retail markets and pointed to the
inadequacy of the current moratorium for ensuring the recovery of
protected species." csbaker@auckland.ac.nz
89. Selection against hatchery trout that go to
sea
Microsatellite and mitochondrial DNA polymorphism reveals
life-history dependent interbreeding between hatchery and wild brown trout
(Salmo trutta L.). 2000. Hansen, M.M., D.E. Ruzzante, E.E. Nielsen,
and K-L.D. Mensberg. Molecular Ecology 9:583-594.
Microsatellite and mtDNA
markers were examined in wild trout from stocked and unstocked sections of
a Danish river. The authors concluded that genetic admixture from the
hatchery trout used for stocking was much higher in the resident,
non-migratory wild trout than in the migratory "sea trout" that
spawn in the river but also spend time in the ocean. It appeared that more
hatchery males than hatchery females are the culprits in these genetic
transactions. However, this sex bias may be due to natural selection
rather than sexual selection. "We suggest that stronger selection
acts against stocked hatchery trout that become anadromous compared to
hatchery trout that become resident. As most resident trout are males this
could also explain why gene flow from hatchery to wild trout appeared to
be male biased." mmh@dfu.min.dk
88. Estimating heritabilities in aquacultural and
captive populations
Estimating variance components in natural populations using
inferred relationships. 2000. Thomas, S.C., J.M. Pemberton, and W.G.
Hill. Heredity 84:427-436.
Microsatellite and other marker
loci have been used for several years to establish pedigrees (or at least
full- and half-sib relationships) which can be used to estimate quantitative genetic parameters such as
heritability of growth in natural populations. To my knowledge no one has worked out
the effect of pedigree error on the accuracy of the parameter estimates.
Instead, several authors have tried to estimate heritabilities and other
quantitative genetic variance components from micro-satellite based
relatedness calculations that do not depend on exact pedigrees. The authors
carry this approach farther and evaluate the various procedures which have
been developed to date.
In this paper, two techniques have been developed
which "use marker information to estimate heritabilities without
reference to the exact nature of the relationships: a regression-based
estimator that regresses phenotypic similarity for a pair of individuals
against an estimate of their relationship and a likelihood-based estimator
that maximizes the probability of the genotypic and phenotypic data given
a known population structure." The could be very useful for
conducting genetics experiments cheaply in commercial aquaculture
environments sthomas@srv0.bio.ed.ac.uk
87. High genetic diversity in introduced oysters
Genetic variation of wild and hatchery populations of the Pacific
oyster, Crassostrea gigas (Thunberg), in Australia. 2000. English, L.J.,
G.B. Maguire, and R.D. Ward. Aquaculture 187:283-298.
"Pacific oysters were introduced
to Tasmania about 50 years ago from Japan; fears had been expressed that
they would have lost genetic variation during their subsequent
naturalisation. Using 17 allozyme loci [the authors determined that].
...The introduced oysters appear to have retained most of the genetic
variation present in the Japanese populations." The overall level of
genetic variation is high but the authors acknowledge that microsatellite
loci, which typically have more rare alleles, might have been more
powerful for detecting genetic erosion. bob.ward@marine.csiro.au
86. Improving the theoretical basis of
conservation genetic predictions
Derivation of the relationship between neutral mutation and
fixation solely from the definition of selective neutrality. 2000.
Tomizawa, J-I. Proceedings of the National Academy of Sciences (USA)
97:7372-7375.
Anyone who has taken any
population genetics at all -- including the pop section of a general
genetic course -- believes that the relative probabilities of fixation of
neutral alleles are equal to their current relative frequencies. This
commonsense prediction is actually due to M. Kimura, who used a
mathematical argument that includes some assumptions and which has not
been experimentally verified. Now this elegant note by Tomizawa should allow countless former genetics
students who have worried about this soft spot in their educations for years and years to sleep more
easily. The paper contains no formal mathematics.
"[Tomizawa's hypothetico-deductive] approach shows that the
equality of the alleles during the fixation process is equivalent to the
equality of probability of their ultimate fixation in a steady state. Both
are manifestations of the definition of selective neutrality. Then, solely
from this dual nature of the definition, the equality between u and p is
derived directly." ("u" is the probability
of fixation and "p" is the current allele frequency.) You only thought you knew this already. The
relevance to aquaculture and genetic conservation is the additional
theoretical support which is given to predictions and models. tomizawa@lab.nig.ac.jp |