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.
372. How to use frozen semen in a
live gene bank
The use of frozen
semen to minimize inbreeding in small populations. 2002. Sonesson, A.
K., M. E. Goddard and T. H. E. Meuwissen. Genetical Research 80:27-30.
This is a very useful
paper even though the authors suggest that their presentation may be
oversimplified. They suggest how one should use cryogenically preserved
semen to retain genetic diversity in a live gene bank or other small
population. It isn't enough just to fertilize the females in each
succeeding generation with sperm from the original founding males.
(Because of subsequent drift these should be the most genetically diverse
males in the pedigree.)
If you do that, all the founding female genomes will eventually be lost.
"We propose an
alternative scheme in which N sires from generation 1 (G1), as well as the
N sires from G0, have semen conserved, and the semen of G0 and G1 sires is
used for dams of odd and even generation numbers, respectively. With this
scheme, the ... genes of
founder dams are also conserved, because 50% of the genes of sires of G1
are derived from the founder dams. A computer simulation study shows that
this is the optimum design to minimize inbreeding, even if semen from
later generations is available."
Matings in the simulation
were random, i.e. were not arranged to minimize next-generation
inbreeding. This is a slick scheme that should be relatively easy to
manage (until the semen runs out). anna.sonesson@akvaforsk.nlh.no
371. Ranking the conservation value of
endangered breeds
Analysis of
genetic diversity for the management of conserved subdivided populations.
2002. Caballero, A. and M. A. Toro. Conservation Genetics 3 (3): 289-299.
The question addressed in
this paper is an important one: which breeds, or strains, or
subpopulations of an endangered species should be preserved in live gene
banks, given that one cannot preserve them all? The authors acknowledge
that unique adaptations, allele richness, evolutionary potential, and even
cultural and historical factors should be taken into account in deciding
this question -- when these factors are identified and measurable. What is
more commonly available to decision makers, though, is molecular marker
data and/or pedigree records. The availability of this kind of information
usually leads to a narrower objective, namely, to preserve as much genetic
diversity as possible in the species or metapopulation as a whole.
If that is accepted as an
expedient objective, which particular subpopulations should be conserved
and which allowed to go extinct? One commonly accepted view is that the
subpopulations which are most valuable are those which are
"distinctive" by some measure of genetic distance. By giving
priority to such subpopulations, a conservation strategy will (in essence)
maximise the total branch length of the phylogenetic tree of the surviving
metapopulation. In this calculus the contribution of each subpopulation to
the global diversity is proportional to its contribution to total tree
length.
The authors of this paper
show, with real as well as with simulated data, that the above strategy
can lead to exactly the wrong choice for conservation. This can
happen when animals within a "distinctive" population have a
high level of coancestry, which of course will often be the case. The
relative sizes of the subpopulations or breeds matters, too.
Their argument
requires a demonstration that maximisation of genetic diversity (as
expected heterozygosity) follows from the minimisation of coancestry in a
subdivided population. The paper includes this demonstration, along with a
clear analysis of the relationship between total genetic diversity
(heterozygosity), Wright's F statistics, Nei's genetic distance, and
Malecot's coancestry coefficient. (See also Nov 2001 #261, Jul 2001 #212.)
Fortunately, coancestry is now easy to estimate from marker data (e.g. Apr
2002 #320). The suggestions in this paper appear to be eminently
practical. armando@uvigo.es ; this
and other recent papers by Caballero can be downloaded from http://www.uvigo.es/webs/c03/webc03/XENETICA/XB2/Caballero.htm
370. Increased susceptibility to disease
in small salmon populations
Resistance to
three pathogens in the endangered winter-run Chinook salmon (Oncorhynchus
tshawytscha): effects of inbreeding and major histocompatibility
complex genotypes. 2002. Arkush, K. D., A. R. Giese, H. L. Mendonca,
A. M. McBride, G. D. Marty and P. W. Hedrick. Canadian Journal of
Fisheries and Aquatic Science 59:966-975.
The authors distinguish
two genetic consequences of small population size which might reduce
disease resistance: (a) increased homozygosity due to inbreeding and (b)
loss of genetic variation, specifically at the MHC locus. In their
experiment matings were arranged such that both full-sib inbred progeny
and outbreds could be created from the same female. "In addition, a
number of parents were selected so that their progeny would segregate for
heterozygotes and homozygotes at... [a class II MHC locus ]."
The progenies were
challenged with three major pathogens: a bacterium (vibrio), a virus (IHNV)
and a parasitic protozoan (whirling disease).
Inbreeding increased the
probability and severity of the protozoan infection but had no significant
effect on the viral or bacterial pathogen. MHC heterozygosity increased
the resistance to IHNV challenge but did not affect resistance to vibrio.
"Overall, we have shown that the endangered winter-run Chinook salmon
appears to be genetically susceptible to the detrimental effects of
pathogens. If there were further losses of genetic variation because of
continued small population size in winter-run, then one would predict both
a higher level of homozygosity at MHC genes and higher levels of
inbreeding. As a result, the pathogens that we have examined, and
potentially others that may infect winter-run, may cause in a decline in
the number of winter-run Chinook salmon and an additional extinction
threat to this critically endangered species." Other papers on
salmonid MHC diversity are Mar 2002 #302, Dec 2001 #272, Oct 2001 #240,
Jul 2001 #221 and #369, below. philip.hedrick@asu.edu
369. Experimental evidence for allele-specific
MHC resistance to infection
Experimental evidence
for major histocompatibility complex-allele-specific resistance to a
bacterial infection. 2002. Lohm, J., M. Grahn, Å. Langefors, Ø.
Andersen, A. Storset and T. von Schantz. Proceedings Royal Society U.K.
(B) 269:2029-2033.
The authors of this
important paper report on the resistance to furunculosis in Atlantic
salmon originating from the Akvaforsk strain and reared by AquaGen AS in
Norway. Families of salmon were mated on the basis of their MHC genotypes
so as to generate mixtures of homozygous and heterozygous individuals
within the same full-sib families, thus controlling the genetic background
against which specific MHC alleles were expressed. A lot of research
activity is currently focused on the importance of MHC diversity in
natural populations, including the possibility that in natural populations
mates are chosen so as to produce MHC heterozygous offspring (Mar 2002
#302, Dec 2001 #272, Oct 2001 #240, Jul 2001 #221 and #370, above.
Surprisingly, in this controlled
breeding experiment MHC heterozygosity per se did not
improve resistance to the furunculosis challenge test. It was
particular Class II MHC alleles that conferred resistance. "We show
that a pathogen has the potential to cause very intense selection pressure
on particular MHC alleles; the relative fitness difference between
individuals carrying different MHC alleles was as high as 0.5."
The authors also write,
"This study clearly shows a strong survival advantage for individuals
carrying a high-resistance allele when exposed to a bacterial infection.
… directional selection acting on the MHC despite its high polymorphism
stresses the importance of renewal of genetic variation at these kinds of
loci, either from mutation, recombination or immigration from other
populations, when combating new or coevolving virulent pathogens." jakob.lohm@zooekol.lu.se
368. All European carp breeds have a
common Asian origin
mtDNA sequence data
supports an Asian ancestry and single introduction of the common carp into
the Danube Basin. 2002. Froufe, E., I. Magyary, I. Lehoczky and S.
Weiss. Journal of Fish Biology 61:301-304.
Common carp have been
cultivated in Europe since late Roman times and in Asia for several
thousand years. It is not known where the diverse, current breeds of
domesticated carp in Europe originally came from; possibilities include a
Pleistocene glacial refuge in the Danube basin, or western Asia (Caspian
sea), or eastern Asia. The authors of this paper found no variation in
mitochondrial DNA haplotypes among European carps and found the same
haplotype in Japanese koi. Individuals from the Amur river, on the other
hand, showed a lot of sequence variation in the mitochondrial control
region.
"The lack of control
region variation in European common carp (as well as Japanese koi)
supports a recent common ancestry of all individuals sampled. Thus, the
observed phenotypic diversity is assumed to have developed through intense
selection after a relatively recent bottleneck event."
No fish from the Caspian
sea were studied but the high mitochondrial diversity of fish from the Amur
river makes eastern
Asia look like an evolutionary center of origin, and the authors conclude
that Caspian fish are more likely to have immigrated from eastern Asia
rather than the other way around. Therefore the authors rule out the
Caspian as well as the Danube as the source of all modern European carp
breeds. steven_weis@hotmail.com
367. Sex-specific DNA sequence in tilapia (O.
niloticus)
Molecular-cytogenetic
analysis reveals sequence differences between the sex chromosomes of Oreochromis
niloticus: evidence for an early stage of sex-chromosome
differentiation. 2002. Harvey, S. C., J. Masabanda, L. A. Carrasco, N.
R. Bromage, D. J. Penman and D. K. Griffin. Cytogenetics Genome Research
97:76-80.
Sophisticated
microscopical and molecular techniques were used to identify a
sex-specific region in the suspected XX/XY chromosome pair of O.
niloticus (see Mar 2002 #308 for similar procedures applied to O.
aureus.) This paper reports the development of in-situ hybridization
probes for a critical region in the long arm of
chromosome 1. The authors write, "... analysis of the
comparative hybridization of X and Y chromosome-derived probes to
different genotypes provides the first demonstration that sequence
differences exist between the sex chromosomes of O. niloticus."
The binding difference
between the probe sequences from X and Y chromosomes is small, which does
not surprise the authors. "Only limited sequence divergence between
the X and Y chromosomes would be expected as YY individuals can develop
into males or, if hormone treated, females that are both viable and
fertile, although growth and survival rates are somewhat lower in YY than
XY males [see Jun 2002 #332].... This suggests that only a very limited
loss of function can have occurred in Y-linked genes and that sequence
differences between the X and Y chromosomes are largely confined to
non-coding regions." d.j.penman@stir.ac.uk
366. Importance of starting an
aquaculture program with the best strain
Genetic
differences in growth among wild populations of the yabby, Cherax
destructor (Clark). 2002. Jerry, D. R., I. W. Purvis and L. R.
Piper. Aquaculture Research 33:917-923.
Cultivation of this
tasty, tolerant, freshwater crayfish is expanding in Australia. "As a
precursor to a breeding programme ... five geographically isolated
populations were evaluated for the traits weight at age, abdomen length
and abdomen width over a period of 9 months." Total weight and the
relative lengths of the head and abdomen (the edible part) varied
significantly among test animals derived from the different source
populations. The fastest growing were almost twice as heavy as the slowest
growing at the end of the experiment. "This study emphasizes that
rapid genetic gains can be made simply by starting a breeding programme
based on faster growing populations." Very true. dean.jerry@li.csiro.au
365. Sex determination in mussels
Genetics of
mother-dependent sex ratio in blue mussels (Mytilus spp.) and
implications for doubly uniparental inheritance of mitochondrial DNA.
2002. Kenchington, E., B. MacDonald, L. Cao, D. Tsagkarakis and E. Zouros.
Genetics 161:1579-1588.
According to this paper
the mechanism of sex determination is not known for any bivalve,
nor have sex chromosomes been identified. Here the authors present new
breeding data that test the recent hypothesis (due to them and others)
that the sex of mussels is directly determined by the genotype of
their mothers, rather than the nuclear genes they receive from their mothers
and/or fathers. There may be a nuclear gene expressed in females which
when homozygous produces either all-male or no-male broods and when
heterozygous produces both males and females.
The sex ratio of
offspring from heterozygous females varies widely depending on dominance
at this locus and other factors. The principal other factor may be the mitochondrial
genome of the father. Mussels, and perhaps many or most other bivalves,
inherit mitochondria from both parents, a phenomenon which is very unusual
in the animal kingdom. kenchingtone@mar.dfo-mpo.gc.ca
364. Adaptive natural selection beats genetic
drift in grayling
Contemporary
fisherian life-history evolution in small salmonid populations. 2002.
Koskinen, M. T., T. O. Haugen and C. R. Primmer. Nature 419:826-830.
Theory and some
experimental evidence suggest that when populations are very small,
genetic drift becomes the dominant evolutionary force. Accidents of allele
segregation and reproductive success allow unfavorable mutatations to
accumulate and beneficial genes to be lost, rapidly, by chance. The
outcome can be descent into an "extinction
vortex" in which populations lose fitness because they are small and
consequently become smaller still (Sep 2001 #230 and #370, above). Natural
selection is supposed to counteract these chance processes (Fisherian
evolution) but unless it is very strong selection should be relatively
ineffective in very small populations. But we really know hardly
anything about the relative power of drift and selection in real-world
situations.
This is another paper on
the evolution of isolated populations of Norwegian grayling (Apr 2002
#315, Jan 2001 #162). It is known that the founding parents were very few
because 90 years ago a fisherman carried them in a bucket for five hours
uphill along a mountain trail. Previously published quantitative genetic
analyses of life history traits in these populations is here combined with
population genetic analysis of neutral microsatellite markers.
The authors argue that,
"natural selection was the dominant diversifying agent in the
evolution of the quantitative [life history] traits. However, the
populations were founded by a small number of individuals, exhibit very
low microsatellite-based effective sizes and show genetic imprints of
severe 'bottlenecks'; which are conditions often suggested to constrain
selection and favour drift. This study demonstrates a very clear case of
fisherian evolution in small natural populations across a contemporary
timescale. mtkoskin@cc.helsinki.fi
363. When genetic distance doesn't mean anything
Is a multivariate
consensus representation of genetic relationships among populations always
meaningful? 2002. Moazami-Goudarzi, K. and D. Laloë. Genetics
162:473-484.
This is an interesting
critique of the reliability of multivariate analysis of gene
frequencies, such as those used to differentiate populations for
conservation and other purposes. The meaning of "reliability"
is, "do these methods say anything useful?" It turns out that
unless all or most of the individual marker loci tell the same story about
the genetic separation among populations, that is unless the individual
patterns are congruent, then "the compromise structure is not meaningful".
Even worse, "Our analysis suggests that for closely related
populations, it is not always possible to accept the hypothesis that an
increase in the number of markers will increase the reliability of the
typology analysis."
The statistical structure analysed in this paper
are the dimensions produced by principal component analysis (PCA).
The authors say or imply that their conclusions apply to other methods as well,
including multidimensional scaling and phylogenetic trees. They
reach their conclusions by correlating, or failing to correlate, the
different patterns of population differentiation inferred from marker loci
taken one at a time.
They also suggest that another
useful and simple indication that inferred genetic distances may not mean
much is the degree to
which genetic variance is distributed among the principal components. If
the first couple of dimensions don't carry most of the variance, then the
compromise pattern of population differentiation is dubious. In the case of
phylogenetic trees, the warning signal is
lack of strong bootstrap support for the consensus phylogeny despite
significant population differentiation at all or most loci. There is some good news though: "The lack of congruence among structures is not a
disadvantage for assignment studies, where independence of discriminating
factors is a great asset." ugendla@dga2.jouy.inra.fr
362. Poor correspondence between true
and inferred coancestries in a captive population
Estimation of
coancestry in Iberian pigs using molecular markers. 2002. Toro, M., C.
Barragán, C. Óvilo, J. Rodrigañez, C. Rodriguez and L. Silió.
Conservation Genetics 3:309-320.
Several of the same
authors published a paper last year (Feb 2001 #173) which was an excellent
example of the use of pedigree information in genetic conservation. In
this new paper coancestry calculations using the same pedigrees,
which extend over more than 50 years, were compared with coancestries
inferred from microsatellite data. Four molecular similarity measures and
no fewer than eight molecular coancestry estimators, based on 49
microsatellites, were correlated with the true pedigrees.
Unfortunately the
correlation within breeds (populations) was not good, usually less than
0.5. "All the correlations were similar to those obtained when using
simple coefficients of molecular resemblance such as molecular coancestry
or similarity indexes.... It is concluded that lack of information on the
allele frequencies in the base population may explain the bias of these
estimators in populations with complex pedigrees." Compare, however,
Mar 2002 #303 which reported good correspondence between inferred and true
pedigrees in race horses.
This is an important
topic for the genetic conservation of small populations which have
been brought under human protection recently, and where relevant pedigree
data from earlier wild matings do not exist (January 2002 #283). toro@inia.es
361. Frankenfish review
Engineered fish:
friend or foe of the environment? 2002. Stokstad, E. Science
297:1797-1799.
This is an entertaining review
of the current state of play in Frankenculture. Recent developments in salmon, trout,
tilapia, carp and other species are outlined, as are the real and
rhetorical environmental issues. Two useful letters to the editor follow
the main news article, one from an ornamental fish specialist and the
other from an environmentalist. E-mail address not available.
360. Website for Frankeninfo
AGBIOS is basically a
consulting company in policy and educational aspects of agricultural
biotechnology. Their website is a gold mine of free news items, research
reports, web links, definitions etc. relating to genetic engineering and
the environment. Although the focus is plants much of their
material is relevant to aquaculture. I noticed several entries on the
acceptability of bioengineered ingredients in animal feeds. http://www.agbios.com/main.php
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