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.
137. Outbred fish (population hybrids) are more resistant to ich
Differences in initial and
acquired resistance to Ichthyophthirius multifiliis between
populations of rainbowfish. 2000. Gleeson, D.J., H.I. Mccallum, and I.
P.F. Owens. Journal of Fish Biology 57 (2):466-475.
Two pure species of rainbowfish
(Melanotaenia) from three locations in Australia were
experimentally infected with the parasite Ichthyophthirius multifiliis,
a.k.a. "ich".
One of the pure species, M. eachamensis, was much more susceptible
to the parasite than the other species, M. splendida. Splendida served
as a control for a follow-up hybridization experiment which involved
crossing eachamensis from the original population with another
population of the same species located some distance away. The population
hybrids had significantly higher resistance than the single-population
fish. "It is tentatively suggested that there may be a link between
the heterozygosity of populations of rainbowfish and their initial ability
to resist infection by Ichthyophthirius multifiliis." dgleeson@zoology.uq.edu.au
136. Recombination distorts our
understanding of the evolution of a viral epidemic
Consequences of recombination on
traditional phylogenetic analysis. 2000. Schierup, M.H., and J. Hein.
Genetics 156: 879-891.
When sequence data are analysed
to generate a phylogenetic tree, the assumption is usually made that
recombination within the sequence does not occur. Yet we know that this
assumption is definitely wrong for viruses and for nuclear DNA in higher
organisms, and possibly for mitochondrial DNA too.
(Viruses can recombine when a
host is simultaneously infected with more than one strain. It has been
suggested that Asian integrated aquaculture of ducks, fish and pigs will
soon bring an end to us all when avian and human influenza viruses
recombine in the pig. The pig can catch both strains and pass the
recombinant on to the farmer.)
Recombination means that
different parts of a sequence will have different evolutionary histories.
The ground shifts: we are no longer trying to discover the one
"true" tree for the sequence. The sequence is the outcome of a
number of correlated trees.
The authors of this paper used
the same phylogenetic software most people use (PHYLIP) to investigate the
effects of recombination. "With recombination present, the length of
terminal branches and the total branch length are larger, and the time to
the most recent common ancestor smaller, than for a tree reconstructed
from sequences evolving with no recombination."
In the context of this GCL
website -- aquaculture and aquatic genetic conservation -- the paper may
be most relevant to viral disease. The authors found that "The
results for the [HIV and foot-and-mouth disease] viral data sets show
strong evidence for recombination with very large estimated values. ...
For HIV, [the analysis] also provides evidence for exponential growth,
which agrees with the rapid spread of this virus. ... the rates appear
here to be so high that phylogenetic analysis may be of very limited
value."
The authors conclude from their
analysis of four data sets that "recombination may be sufficiently
high to invalidate the use of phylogenetic trees in many population
studies." Plainly, recombination needs to be taken into account if
sequence data are used to draw epidemiological conclusions about the
spread of aquacultural diseases, as suggested in October list #111. The
computer program for calculating the various statistics is available at http://www.bioinf.au.dk/~mheide . The contact e-mail address is mikkel.schierup@biology.au.dk
135. Virus attacks a protected species
Canine virus blamed in
Caspian seal deaths. 2000. Stone, R. Science 289 (5487): 2017-2018.
The Caspian seal, listed as
vulnerable by the World Conservation Union, may recently have lost as many
as 20,000 individuals to an infection identified as canine distemper virus
(CDV), i.e. a dog virus. The identification was made by a research group
in Belfast working in collaboration with scientists from the former
bio-weapons lab VECTOR, which is located upwind of Novosibirsk. VECTOR
believes that the situation is even more serious, and that the deaths are
due to simultaneous infection by CDV and an influenza virus in the seals
[where the viruses recombine? -- just kidding]. For more on this
development see the September-October publication from the Belfast group
led by Seamus Kennedy at http://www.cdc.gov/ncidod/eid/vol6no6/kennedy.htm
E-mail seamus.kennedy@dardni.gov.uk
134. Hybridization of locally-adapted
finches promotes adaptive radiation
The mitochondrial and
nuclear genetic homogeneity of the phenotypically diverse Darwin's
groundfinches. 1999. Freeland, J.R., and Boag. P.T. Evolution 53
(5):1553-1563.
The drab-looking finches of the
Galapagos islands, which turned Charles Darwin towards the discovery of
natural selection, are among the best-studied and best-understood examples
of this evolutionary process. In this paper the authors describe the first
successful attempt to resolve the evolutionary history and genetic
relationships of the birds. It is not surprising that the paper is
relevant to issues in genetic conservation and even aquaculture.
The authors found that
"The differentiation of the ground finch species based on
morphological data is not reflected in either mitochondrial or nuclear DNA
sequence phylogenies." Not only that, inferred genealogies based on
mitochondrial and nuclear markers are not even concordant with each other.
The authors suggest that "the absence of species-specific [DNA
sequence] lineages can be attributed to ongoing hybridization involving
all six species of Geospiza." Hybrids have an advantage in
some years so there is no selection against hybridization even though
there is strong, ongoing selection for morphology (e.g. the size of the
beak in relation to the size of the seeds available for food).
"Hybridization has apparently played a role in the adaptive radiation
of Darwin’s finches."
There is a lot of evidence for
local adaptation of migratory salmonid populations (e.g. October #115),
but those who argue that this will result ipso facto in outbreeding
depression might ponder Darwin's finches. We need to have direct evidence
for outbreeding depression when salmonids hybridize between spawning
areas. Then we can know for sure whether hybridization is an evolutionary
problem, rather than an evolutionary solution, for salmonids at risk of
extinction. j.r.freeland@reading.ac.uk
133. The genetics (?) of acquired (?)
tolerance to shrimp WSSV
Quasi-immune response of
Penaeus japonicus to penaeid rod-shaped DNA virus. 2000. Venegas, C.A.
, L. Nonaka, K. Mushiake, T. Nishizawa, and K. Murog. Diseases of Aquatic
Organisms 42 (2):83-89.
The authors experimentally
infected Kuruma prawns with white spot syndrome virus (WSSV). The shrimp
that survived this experimental challenge, as well as the survivors of a
natural outbreak of WSSV, demonstrated high levels of survival to
subsequent experimental challenges by injection of the virus. PCR testing
showed that many of the survivors continued to carry the virus even though
they were apparently healthy. The paper refers to recent work in Thailand
by Flegel and others which also may show induction of tolerance to WSSV
and yellowhead virus YHV within a single generation of shrimp following
massive mortalities.
The authors of this
carefully-designed experiment conclude that the "high survival rates
suggested that survivors (natural or experimental) were able to resist
infection and that the resistance was not due to selection of naturally
resistant [or tolerant?] shrimp ... but due to enhancement of an
immune-like system (quasi-immune response) after exposure to [the
virus]."
Whether to tag this phenomenon
as "tolerance" or "resistance" is not a mere quibble.
The two processes have very different implication for the long-term
evolution of a host/pathogen system. See for example September #105,
August #96, July #84. My highly non-expert interpretation of this
experiment is that surviving animals carry the virus but aren't sick,
therefore both components survive indefinitely in the aquaculture system,
which means tolerance in the host. From the long-term genetic perspective
the evolutionary implications of this may be good news for the industry.
Also from the genetics
perspective, one is tempted to speculate about the variation in the
proposed quasi-immune response. Is the partial survival at the first
(inducing) challenge due to variation in the strength of the response? Or
in the threshold for induction or enhancement? Or is there any selectable
genetic variation at all in the shrimp (the details of the experimental
design suggest that the environment is involved in a complicated way)? The
answers to such questions will profoundly influence the design of genetic
improvement programmes. fpath@hiroshima-u.ac.jp
132. Inbreeding depresses fitness in the
wild
Inbreeding depression
influences lifetime breeding success in a wild population of red deer
(Cervus elaphus). 2000. Slate, J., L.E.B. Kruuk, T.C. Marshall, J.M.
Pemberton, and T.H. Clutton-Brock. Proceedings of the Royal Society (B)
267 (1453):1657-1662.
The authors used microsatellite
heterozygosity as an indicator of individual inbreeding coefficients among
unmanaged deer on the island of Rum, Scotland. They found that
heterozygosity is correlated with lifetime breeding success (total
offspring) in both males and females. They believe that this is the first
observation of such a correlation between inbreeding and something close
to overall fitness, in both sexes. Similar correlations have
previously been found between heterozygosity/inbreeding and major
components of fitness such as mating success (July #82). Lifetime breeding
success is not quite identical with fitness measures such as r or R0 which might be calculated from a life table or a Leslie matrix. Both total
production and the schedule of the production of offspring (as a function
of overall population growth rate) are important to fitness. j.pemberton@ed.ac.uk
131. Inbreeding depression in the wild no
worse than in the lab
Equivalent inbreeding
depression under laboratory and field conditions in a tree-hole-breeding
mosquito. 2000. Armbruster, P., R.A. Hutchinson, and T. Linvell.
Proceedings of the Royal Society (Biology) 267 (1456):1939-1945.
The authors expected to find
that inbreeding had a more profound depressive effect on fitness in the
natural environment than in an optimized laboratory environment where
animals are well taken care of. This is a reasonable expectation based on
theory as well as many recent observations. However, in this case, they
found that while the fitness of the mosquito populations did decline with
inbreeding, the decline was the same in natural tree-holes as under
favourable laboratory conditions. A 10% increase in inbreeding coefficient
reduced fitness, as measured by R0 an index which is
essentially the absolute rate of population growth calculated from the
life table (see #132, above) , by 12%-15%. Thus we can not be totally
confident in predicting that inbred small and endangered populations (e.g.
of fish, by analogy) must inevitably be in worse shape than predicted by
inbreeding experiments in the laboratory, aquarium or farm.
The authors suggest that a
composite fitness index may be a more appropriate measure of inbreeding
depression than individual component of fitness such as fecundity or
survival. If trade-offs between fitness components occur this may well be
true. Note September list #101 in which inbreeding depression was found to
be specific to each trait, not a generalized phenomenon. parmbrus@zoo.uvm.edu
130. Escaped farmed salmon are unfit in
Norway -- and their offspring too
Lifetime success and
interactions of farm salmon invading a native population. 2000.
Fleming, I.A., K. Hindar, I.B. Mjølnerød, B. Jonsson, T. Balstad, and A.
Lamberg. Proceedings Royal Society (UK), Ser. B 267 (1452):1517-1523.
The authors report the result
of an experiment in which mature, farmed salmon were released into the
Imsa river in Norway. Radio tagging and genetic analysis revealed that
"The farm fishes were competitively and reproductively inferior,
achieving less than one-third the breeding success of the native
fishes." Most of the gene flow involved native males mating with
farmed females -- farmed males were relatively uncompetitive. "There
were also indications of selection against farm genotypes during early
survival but not thereafter. ... Ultimately, the lifetime reproductive
success (adult to adult) of the farm fishes was 16% that of the native
salmon." The authors note, however, that the invasion of the Imsa by
farmed fish reduced the productivity of the native population by more than
30% through resource competition and competitive displacement. ian.fleming@ninatrd.ninaniku.no
129. The right way to estimate quantitative
genetic parameters in nature
Estimating variance
components and heritabilities in the wild: a case study using the ‘animal
model’ approach. 2000. Milner, J.M., J.M. Pemberton, S. Brotherstone,
and S.D. Albon. Journal of Evolutionary Biology 13 (5):804-813.
The authors of this paper have
published a fine example of a type of study that a lot of people are
attempting -- estimation of quantitative genetic parameters in wild
populations. In this case the population was the Soay sheep living on the
island of St. Kilda, and the techniques included markers to estimate
first-order genetic relationships (the relationship matrix) which were
then used for the modern animal model technique for parameter estimation.
The trait that experienced the strongest selection was body weight. This
trait also had both the highest additive variation and the lowest
heritability. jos.milner@ic.ac.uk
128. Prawn markers and biogeography in the
Pacific
Population structure of
the giant tiger prawn Penaeus monodon in Australian waters, determined
using microsatellite markers. 2000. Brooker, A.L., J.A.H. Benzie, D.
Blair, and J.-J. Versini. Marine Biology 136 (1):149-157.
The authors used microsatellite
polymorphisms to discover that the monodon populations in western
Australia are genetically quite different (and less variable) than those
on the north and east. "Reduced variability is consistent with a
recent population bottleneck or colonization by a small founding
population from the east when sea links between Indonesia, New Guinea and
Australia were re-established following the last ice age." Good
microsatellite markers have been rather hard to find in shrimp, for
technical reasons, and these look like winners: 34, 35 and 85
alleles! Amanda.Brooker@jcu.edu.au
127. BLUP selection increases inbreeding
Prediction of rates of
inbreeding in populations selected on best linear unbiased prediction of
breeding value. 2000. Bijma, P., and J.A. Woolliams. Genetics
156:361-373.
The best linear unbiased
predictor (BLUP) combines information from relatives to maximise the rate
of genetic gain from selection. BLUP is now being used for breed
improvement in aquaculture and, with the advent of marker-based
reconstruction of pedigrees, related techniques are used to estimate
quantitative genetic parameters in natural populations. "Predictions
for the rate of inbreeding (F) in populations with discrete generations
undergoing selection ... were developed. ... Expected contributions of
individuals were predicted using a linear model, ui(x) = + ßsi, where si
denotes the selective advantage as a deviation from the contemporaries,
which was the sum of the breeding values of the individual and the
breeding values of its mates." .... [Simulation showed] that,
contrary to random selection, F less than halved when the number of
parents was doubled and that in specific cases F may increase with the
number of dams."
The authors provide a useful
rule of thumb for dealing with this situation, which may have considerable
importance in aquaculture and, possibly, in genetic conservation of fish,
where artificial selection might be performed to reduce domestication
effects in the hatchery. "The cost of raising offspring from parents
that are destined not to contribute [genes] in the long term is a hidden
cost of the high rates of inbreeding associated with BLUP selection." piter.bijma@alg.vf.wau.nl
126. PCR shrimp for virus without killing
them
A non-destructive
method based on the polymerase chain reaction for detection of
hepatopancreatic parvovirus (HPV) of penaeid shrimp. 2000. Pantoja,
C.R., and D.V. Lightner. Diseases of Aquatic Organisms 39:177-182.
"Current methods to detect
hepatopancreatic parvovirus (HPV) infection of penaeid shrimp depend on
invasive techniques that require dissecting the organs infected by this
virus. ... A method was developed for HPV detection by applying a
polymerase chain reaction (PCR) assay to fecal samples collected from live
HPV-infected shrimp Penaeus chinensis. ...Analysis of fecal samples by PCR
may prove useful for non-lethal screening of valuable shrimp [e.g.
broodstock shrimp] of unknown HPV status."
Presumably the techniques could
also be useful for distinguishing and then selecting resistant as opposed
to tolerant animals. cpantoja@u.arizona.edu
125. How to measure growth in genetic
experiments
Statistical models for
estimating the genetic basis of repeated measures and other
function-valued traits. 2000. Jaffrézic, F., and S.D. Pletcher.
Genetics 156:913-922.
Growth rate, one of the prime
targets for genetic improvement in aquaculture as well as for QTL mapping,
is not easy to measure in a meaningful way. In fact the definition of
growth gets more complex -- mushy -- the more you worry about it. Similar
problems arise with other important traits that are measured repeatedly as
a function of another variable: fecundity vs. age, mating success vs.
size, size vs. temperature, survival vs. salinity, etc.
There is a choice of three
mathematical models that are sometimes used for this purpose: regression,
approximation of the data by orthogonal polynomials, and character process
models. The discussion of the contrasting features of the 3 approaches is
admirably clear in this paper. The authors elaborate on the third approach
(which they find to be the best) and evaluate all of them using both
simulated and real data. All models were estimated from data by maximum
likelihood (REML). The data consisted of a genetic design (sire half-sib
families) that is standard in experimental quantitative genetics and
achievable in some natural populations using genealogies reconstructed
from DNA markers.
"The simplicity of the
character process model allows quantitative statements about the
predominant attributes of the genetic covariance function. Genetic
variance for Drosophila mortality declines significantly with age, while
genetic variance is constant at all ages for reproductive output."
The authors also analysed growth in beef cattle. They are making the
software available for free. s.pletcher@ucl.ac.uk |