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.
85. US Patent application for the first Frankenfish -- a salmon
FDA, researchers consider first transgenic fish. 2000.
Anonymous. Nature Biotechnology 18:143.
The US Food and Drug
Administration is now evaluating a patent application from A/F Protein for
transgenic salmon -- apparently the first transgenic fish to be patented
in the US if the application succeeds. The A/FP technique is to
attach the salmon growth hormone DNA sequence to a foreign promoter
sequence (from the ocean pout) which causes the gene to produce growth
hormone in the liver. Unlike the normal gene which is expressed in
the pituitary, the transgene is continuously switched on by its promoter.
The salmon reach a size of 8 pounds in 1.5 years. The article
briefly and fairly reviews a number of recent studies on the possible
ecological risks of transgenic fish including the so-called "Trojan
gene" identified in Medaka. The article acknowledges that
medaka are not salmon and that the transgenic salmon are sterilized by
triploidization. It also notes that sterilization is not 100%
effective..... As Maureen Dowd said, "I have seen the future,
and it is backing out of the room". [No e-mail address
available]
84. Genetics may explain why shrimp diseases are
spreading as stocks recover
Evolutionary dynamics of pathogen resistance and tolerance. 2000. Roy,
B.A., and J.W. Kirchner. Evolution 154:51-63.
The authors of this mainly
theoretical paper discuss the evolution of disease resistance (defense
mechanisms which block infection, kill the pathogen or limit its spread
within the host) and disease tolerance (ability to survive despite being
infected). The selective advantage of resistant host
individuals becomes less as they become more numerous, because the
incidence of disease declines and selection intensity decreases in a
negative feed-back loop. Therefore, "... genes conferring
complete resistance cannot become fixed (i.e., universal) by selection in
a host population, and diseases cannot be eliminated solely by natural
selection for host resistance." Disease resistance -- including
the immune response -- is predicted always to be variable and incomplete.
This is because the extra physiological costs of resistance genes in the
host causes them to be selected against in regions where the disease
challenge is rare.
Disease tolerance is a
different story. As more host individuals become tolerant the
incidence of disease increases, thus increasing the selective advantage of
the tolerance gene. Tolerance genes in the host increase the fitness
of the host and the pathogen as well in a positive feedback loop.
Tolerance genes should rapidly evolve to a frequency of 100% and the
disease should become universally present.
The point for aquaculture is
that the evolution of host resistance causes the prevalence of a pathogen
to decrease while tolerance causes it to increase. Does this
prediction allow us to guess what is happening during the successive waves
of diseases in shrimp aquaculture? Anecdotal evidence may suggest
that tolerance, rather than resistance is evolving in shrimp aquacultural
systems, and that diseases are becoming more prevalent while shrimp
production recovers in infected regions. The usual explanations for
the spread of shrimp disease -- human greed, criminality & fear, etc.
-- explain the spread but not the correlation. The time scale is
right for evolution especially if the underlying genetic mechanisms
involve dominant or co-dominant genes. Does anyone know what the
possible tolerance mechanisms in shrimp might be?
Incidentally, the pathogen also
evolves, so the shrimp and the disease are in what is called a "coevolutionary
arms race". More about this in the August posting. The
conclusion is that tolerance, but not resistance, is an evolutionary
stable defense strategy and may also be the best tactic for aquaculture
geneticists. roy@geobot.umnw.ethz.ch .
83. Atlantic salmon sneak into paradise
Evidence of natural reproduction of aquaculture-escaped Atlantic salmon in
a coastal British Columbia river. 2000. Volpe, J.P., E.B. Taylor, D.W.
Rimmer, and B.W. Glickman. Conservation Biology 14:889-903.
The authors captured 12
juvenile salmon which they identified as Atlantics in the Tsitika River,
British Columbia. Species identity was confirmed by analysis of rDNA
and mtDNA. Morphological characteristics and condition lead the authors to
believe that juveniles were the offspring of escaped farm salmon and were
themselves successfully maturing. This is the first direct evidence
that self-sustaining Salmo salar populations may be developing in BC, with
many possible negative implications for the survival of the effete native
salmonids. jvolpe@uvic.ca
82. More competition problems for inbred males
A comparative study of ejaculate traits in three endangered ungulates with
different levels of inbreeding: fluctuating asymmetry as an indicator of
reproductive and genetic stress. 2000. Gomendio, M., J. Cassinello,
and E.R.S. Roldan. Proceedings Royal Society Ser. B. 267:875-882.
Within populations of three
species of endangered gazelles the individual coefficient of inbreeding is
correlated with ejaculate quality only when the overall level of
inbreeding is high. However fluctuating asymmetry (FA, see June list
#70) is correlated with inbreeding in all populations. "Thus,
FA appears in individuals whose levels of inbreeding are still not high
enough to affect male reproductive potential and should therefore be
considered a sensitive indicator of genetic stress. ... Finally, FA is
also a reliable indicator of male reproductive stress since it is related
to individual semen quality in all the species studied". [FA is
easy to measure in shrimp and fish. It may also be of practical
interest to aquaculturists that recent studies of what human beings find
attractive in the opposite sex can be summarized in one word:
"symmetry".] mcnc125@mncn.csic.es .
81. Frankensalmon have more muscle cells and need
more air
Coho salmon (Oncorhynchus kisutch) transgenic for a growth hormone gene
construct exhibit increased rates of muscle hyperplasia and detectable
levels of differential gene expression. 2000. Hill, J.A., A. Kiessling,
and R.H. Devlin. Can. J. Fish Aquat. Sci. 57:939-950.
The authors studied a variety
of traits in coho salmon (Oncorhynchus kisutch) which were made transgenic
(as heterozygotes) for growth hormone. "Transgenic fish were found to
have significantly higher numbers of small-diameter muscle fibres [In
somite muscles. The alternative possibility is to have larger fibres.].
... Higher levels of activity were found for phosphofructokinase and
cytochrome oxidase in white muscle of the transgenic fish. This
difference indicates a higher glycolytic and aerobic requirement in the
muscle of transgenic fish." An unusual and progressive aspect
of this study was the use of subtractive RNA hybridization to identify
other genes that are being switched on in rapidly growing transgenic fish.
Sure enough, the authors found
(after constructing the differential cDNA library and amplifying and
sequencing some of the cloned fragments) that many of the additional
messenger RNAs in the transgenic fish were specifying myosin, consistent
with " high level of expression in the early stages of muscle fibre
construction."
[in the Journal of
Fish Biology Vol. 56, pp. 191-195. 2000. E. D. Stevens, & R. H.
Devlin. report that the two-fold increase in growth rate of transgenic
coho was paralleled by a two-fold increase in the surface area of the
intestine.] devlinr@pac.dfo-mpo.gc.ca .
80. Microsatellite mutation models used by
population geneticists are probably OK
A phylogenetic perspective on sequence evolution in microsatellite loci.2000.
Zhu, Y., D.C. Queller, and J.E. Strassmann. Journal of Molecular Evolution
50:324-338.
The authors studied 140 million
years of evolution of microsatellite repeat regions in a 158-species
subfamily of wasps. They found that the number of repeats evolved
more rapidly than other features such as "imperfections" but
that imperfections are phylogenetically informative. "Overall,
phylogenetic variation in repeat regions can be explained by adding
neutral evolution to what is already known about the mutation process.
The life cycle of microsatellites appears to reflect a balance between
growth by slippage and degradation by an essentially irreversible
accumulation of imperfections."
This paper will be of interest
to people who use measures of genetic differentiation in their work on
fisheries population genetics and conservation genetics. Many such
measures, including Rst and some procedures used to detect bottlenecks,
outbreeding and introgression are dependent on assumptions concerning the
underlying mutational process in microsatellite repeat polymorphisms.
This paper on the whole supports the validity of these assumptions. strassm@rice.edu .
79. Sneaky parr help genetic conservation
Multiple paternity increases effective size of southern Atlantic salmon
populations. 2000. Martinez, J.L., P. Moran, J. Perez, B. De Gaudemar,
E. Beall, and E. Garcia-Vazquez. Molecular Ecology 9:293-298.
This parentage-determination
study was done on small, endangered Atlantic salmon populations in rivers
running into the Bay of Biscay. A large proportion of the broods
sampled in the redds had been fertilized by more than one precociously
mature parr. The authors show that "multiple paternity derived
from mature parr is crucial for the conservation of genetic variability in
small populations of Atlantic salmon. egv@sauron.quimica.uniovi.es .
78. New procedures for estimating hybridization
and gene flow
Estimating multilocus linkage disequilibria. 2000. Barton, N.H.
Heredity 84:373-389.
When loci are situated close
together on the same chromosome, the frequencies of the genotypes will, in
populations that are less than infinitely large and infinitely old, often
be non-random. (That is, they will not occur in the frequencies
predicted by simple multivariate combination of the component allele
frequencies.) This linkage disequilibrium occurs because of the
physical connection between alleles at different loci on one long strand
of DNA. Statistical information concerning the joint distributions
of physically linked alleles is very useful for mapping loci on
chromosomes when the genotypes originate from a specific cross or
hybridisation.
Even when loci are located on
different chromosomes, or are so far apart that the DNA strand connecting
them is effectively broken and randomized during replication, linkage
disequilibrium will still often be observed in real populations. In
this case the statistical disequilibrium ("non-randomness")
carries information about the mixing of genetically different populations,
or non-random mating among individuals, or drastic changes in population
size, or other phenomena that are of great practical interest both in
aquaculture and genetic conservation. The question is how to develop
a meaningful story out of the frequency data.
A number of different
approaches which have been used in the past are reviewed in this paper.
Barton is here particularly concerned with hybrid and mixed populations.
His work extends previous work by providing a framework for analysing data
from several loci at a time and for using "non-diagnostic" loci
-- that is, loci at which all populations carry all the alleles, but at
different frequencies.
Biologists who have taken less
than an infinite level of indefinitely well-taught statistics classes may
find this interesting paper hard sledding. Nonetheless, it "
... sets out methods for estimating multilocus genotype frequencies which
are appropriate for unlinked neutral loci, and for populations that are
ultimately derived by mixing of two source populations. ...Two methods
...are described: a simple method based on multivariate moments [variance,
skewness etc.], and a maximum likelihood procedure, which uses the
Metropolis algorithm. Both methods perform well when tested against
simulations with two or four loci. ... "
The procedures "can be
used to infer quantities such as rates of gene flow and degree of
assortative mating. ... [and] likelihood that offspring were sired by
sampled individuals of known genotype, rather than by some father from the
unsampled population. These methods make possible a variety of
statistical analyses of hybrid populations" n.barton@ed.ac.uk .
77. Genetic signal traces the progressive
development of WSSV in individual shrimp
Long-term presence of white spot syndrome virus (WSSV) in a cultivated
shrimp population without disease outbreaks. 1999. Tsai, M.F., G.H.
Kou, H.C. Liu, K.F. Liu, C.F. Chang, S.E. Peng, H.C. Hsu, C.H. Wang, and
C.F. Lo. Diseases of Aquatic Organisms 38:107-114.
Polymerase Chain Reaction (PCR)
is a procedure for amplifying DNA (in this case associated with WSSV)
until there is enough to work with. When the initial quantity of
target (template) DNA is very small PCR amplification may have to be done
in several stages, each of which involves many rounds of self-catalyzing
amplification.
The shrimp PLs in this
experiment came from a single WSSV-carrying brooder. At 7 months of
grow-out 2-stage PCR was required to detect the WSSV signal in the heart,
gill and other tissues in some of the offspring shrimp tested. Prior
to harvest at 13 months, most tested shrimp were scoring positive by the
2-stage procedure. The first deaths attributable to WSSV occurred at
13 months when, for the first time, the 1-stage PCR began picking up the
WSSV signal. "Although superficially healthy, 10 % of the
surviving adults had tiny white spots on their carapace, and in situ
hybridization analysis revealed WSSV-positive cells in 40 % of the
specimens examined."
The authors have shown that
with sufficiently sensitive procedures WSSV can be detected in a shrimp
population a long time before massive mortality takes place. They
hypothesize that "disease outbreaks do not occur if shrimp defense
mechanisms manage to contain low-intensity viral infections under
low-stress culture conditions. Conversely, outbreaks may occur under
stressful conditions". This hypothesis is not new but
multi-stage PCR makes it testable. Interesting tool for studying
tolerance vs. resistance in shrimp. gracelow@ccms.ntu.edu.tw .
76. Possible negative effects of supplementation
on fitness
Microsatellite analysis of hatchery stocks and natural populations of
Arctic charr, Salvelinus alpinus, from the Nordic region: Implications for
conservation. 1999. Primmer, C.R, T. Aho, J.; Piironen, A. Estoup,
J.M. Cornuet, and E. Ranta. Hereditas 130:277-289.
This is a study of
microsatellite variation in Finnish populations of land-locked Arctic
charr. Population differentiation was high even among nearby lakes.
The hatchery stocks were similar to the wild in allele number and average
heterozygosity but they showed higher levels of single- and multi-locus
genotypic disequilibrium. The population in a lake that was
completely reliant on hatchery supplementation showed particularly low
levels of genetic variation. "Although the hatchery stocks of
this population suffer from increased egg and alevin mortality and disease
susceptibility, it remains to be determined if this is due directly to a
lack of genetic variation as some abundant unstocked natural populations
possessed similarly low levels of microsatellite variability." craig.primmer@helsinki.fi .
75. Inbreeding may reduce population fitness for
a long time
Does inbreeding affect the extinction risk of small populations?:
predictions from Drosophila. 2000. Bijlsma, R., J. Bundgaard, and A.C.
Boerema. Journal of Evolutionary Biology 13:502-514.
The authors studied the
extinction rates of experimental Drosophila populations that had been
inbred to varying degrees before being transferred into optimal and
stressful experimental environments. "We show that inbred
populations have a significantly higher short-term probability of
extinction than non-inbred populations, even for low levels of
inbreeding.... the impact of environmental stress becomes significantly
greater for higher inbreeding levels, demonstrating explicitly that
inbreeding and environmental stress are not independent but can act
synergistically. These effects seem long lasting as the impact of
prior inbreeding was still qualitatively the same after the inbred
populations had been expanded to appreciable numbers and maintained as
such for approximately 50 generations." r.bijlsma@biol.rug.nl .
74. Tracing parentage in a natural fish
population
Genetic parentage in large half-sib clutches: theoretical estimates and
empirical appraisals. 2000. DeWoody, J.A., D.E. Walker, and J.C.
Avise. Genetics 154:1907-1912.
Many people would like to
assign parentage to fish in natural or aquacultural populations.
Applications range all the way from genetic variance estimation, to
controlling inbreeding and genetic drift in endangered populations, to
saving aquaculture escapees from paternity suits, to protecting investment
in new breeds. This is an example of a successful assignment study
on the sand goby Pomatoschistus minutus. In addition to the
observational work reported in this paper "a general dilocus matrix
procedure is suggested for organizing and interpreting otherwise
cumbersome data sets when extremely large numbers of full-sib and half-sib
embryos ... are genotyped at two or more hypervariable loci. " dewoody@arches.uga.edu .
73. Sterilized trout are just as much fun
Relative return to creel of triploid and diploid rainbow trout stocked in
eighteen Idaho streams. 2000. Dillon, J.C., D.J. Schill, and D.M.
Teuscher. North American Journal of Fisheries Management 20:1–9, 2000
20:1-9.
Triploid salmonids are almost
entirely sterile and thus pose no genetic risks to native stocks if they
are used for supplementation. The authors released diploid and
triploid fish that had been reared to hatchable size and found no
significant differences in the rate of returns to anglers. They
conclude that " triploid rainbow trout can provide stream angling
opportunity equal to that provided by fertile diploid fish. Although
there are other concerns regarding the stocking of hatchery trout in
streams containing native trout, we suggest that using triploid rainbow
trout in stream-stocking programs can help balance the demands for both
consumptive fishing opportunity and conservation of native stocks." dschill@micron.net
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