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.
385. News flash: Effective population size five orders of magnitude smaller than
census size in a pelagic species
Loss of
microsatellite diversity and low effective population size in an
overexploited population of New Zealand snapper (Pagrus auratus). 2002. Hauser, L., G. J. Adcock, P. J. Smith, J. H. B. Ramírez and G. R.
Carvalho. Proceedings of the National Academy of Sciences (US)
99:11742-11747.
This study is based on a
50-year sequence of scale samples collected from a population of New
Zealand snapper in Tasman Bay. The sample covers most of the period of
commercial exploitation, which by now has reduced the census size by about
75%.
Microsatellite DNA analysis of the preserved scales and recent
samples allowed comparison between the effective population number Ne and
contemporaneous estimates of population size (census number) which were
made using standard fisheries methods. Amazingly, although the census
number was several millions the long-term effective population number was
somewhere between 46 and 186 individuals!
This is a
much lower ratio of effective to census number than previously guessed or
inferred for other species. The New Zealand snapper resembles many other
exploited marine fish (e.g. cod) in having a long life, indeterminate body
size, size-dependent fecundity, iteroparity and overlapping generations.
The authors make the following important points: (1) Fish stocks of
several million individuals can rapidly lose genetic diversity. (2) The low ratio of effective to
census number may explain the poor correspondence between stock sizes and
recruitment which bedevils fisheries managers. And finally, (3) "The
recruitment processes suggested by genetic data also open the exciting
possibility of identifying the demographic section of the population most
likely to reproduce successfully and to protect these individuals from
excessive exploitation". lhauser@u.washington.edu
384. Nine more genes on the tilapia
linkage map
Fishing in silico :
searching for tilapia genes using sequences of microsatellite DNA markers. 2002. Cnaani, A., M. Ron, G. Hulata and E. Seroussi. Animal Genetics
33:474-476.
The linkage map of the
tilapia genome is coming along nicely due to the efforts of several
research groups, but until now the map included only 14 known genes. The
other tilapia markers in the map -- of which there are hundreds -- consist
mainly of "neutral" microsatellite and AFLP markers and some
allozymes. The authors of this paper have added nine more genes without
doing any lab work.
They found unique flanking sequences of 312 tilapia
microsatellites in the GenBank database, and entered them into the
appropriate search-and-match engines for comparison with the human and Fugu genomes. Sixteen tilapia sequences with high homology to known genes were
identified. Nine of these microsatellites had already been assigned to
tilapia linkage groups, so now the genes which flank them are mapped too.
The genes are a mixed bag of transporter and structural proteins, enzymes
etc. seroussi@agri.huji.ac.il
383. Domesticated salmon have more
active growth hormone systems
Effects of
domestication on growth physiology and endocrinology of Atlantic salmon
(Salmo salar). 2002. Fleming, I. A., T. Agustsson, B. Finstad, J. I.
Johnsson and B. T. Björnsson. Canadian Journal of Fisheries and Aquatic
Science 59:1323-1330.
Seventh- generation,
domesticated Atlantic salmon from the Norwegian National Breeding Program
were compared with their principal wild founder population with respect to
growth rate, plasma growth hormone (GH), pituitary GH and insulin-like
growth factor (IGF-I). Artificial selection or, more probably, several
decades of domestication have brought about a three-fold increase in the
growth rate of this strain (now called the AquaGen). "Pituitary GH
content was positively correlated with growth rate and correspondingly was
significantly higher in the faster growing domesticated fish than in the
wild fish. Plasma GH levels were also significantly higher in the
domesticated fish, whereas IGF-I levels did not differ."
The paper includes an interesting discussion of
"accidental" selection in domesticated salmon; for example, there is indirect
selection for high GH activity, and therefore for rapid growth, because
there is little "selective payoff from cautious feeding behaviour [in
the absence of predators]". The authors also note that their results
may explain why insertion of transgenic GH genes doesn't increase the
growth of a strain that has already been selected for rapid growth (see
Feb 2001 #174). ian.fleming@hmsc.orst.edu
382. New formula for effective
population number
Effective size of
populations with unequal sex ratio and variation in mating success. 2002.
Nomura, T. Journal of Animal Breeding and Genetics 119:297-310.
This paper derives a
useful formula for calculating effective population size when mating
success varies in both sexes. Non-overlapping generations are assumed.
Instead of the usual formula Ne=4NmNf/(Nm + Nf), the author recommends
Ne=4NmNf/(2Nm+Nf) for populations with a harem mating system (very
frequent in aquaculture). "The effective population sizes of several
wild, experimental and domestic animals are estimated by applying the
derived equations to the published demographic and ecological data." nomurat@cc.kyoto-su.ac.jp
381. MHC diversity doesn't require
overdominance
MHC heterozygosity
confers a selective advantage against multiple-strain infections. 2002. Penn, D. J., K. Damjanovich and W. K. Potts. Proceedings National
Academy of Sciences (U.S.) 99:11260-11264.
Interest in the genetics
of the fish immune system is running high these days. Atlantic salmon --
and other fish -- may be choosing mates (in nature)
to maximise the genetic diversity of their offspring at the major
histocompatibility complex (Dec 2001 #272). The preferred explanation for
this "disassortative mating", in which fish choose mates which
are genetically unlike themselves, is that MHC heterozygotes are
intrinsically more fit than homozygotes (overdominant selection at
MHC loci; see for example Dec 2002 #370). The thought is that
heterozygosity at MHC loci may enhance a host animal's resistance to
pathogens by increasing both the diversity of peptide antigens it presents
to T cells and the diversity of the T cells themselves.
But there is also
evidence (Mar 2002 #302) that particular MHC alleles may also be directionally selected (i.e. towards homozygosity), possibly on a lake-specific basis.
Heterozygotes would not be more fit than MHC homozygotes and
disassortative mating should not be selected in such lakes. And in an
experiment in an aquaculture-like environment where exposure to specific
pathogen strains was controlled, particular MHC alleles appeared to have a
selective advantage but heterozygosity did not (Dec 2002 #369).
So, we would like to know what the best genetic management strategy is:
selection for homozygosity of particular MHC alleles, or selection for MHC diversity per
se. The answer is important both to conservationists and to
geneticists who hope to profit from the development of proprietary
"super breeds" for aquaculture.
The experiment reported here was done on mice, which have an MHC system
similar to that of fish although more complex. Mice strains were crossed
to produce a variety of MHC genotypes which were released into a house
that had been specially designed to harbour mice. There they were exposed
simultaneously to a defined mix of pathogen strains plus whatever natural
pathogens and parasites had been left behind by an aboriginal mousy
population. Surprisingly, the experimenters found that "contrary to
what is widely assumed, the benefits of heterozygosity were due to
resistance being dominant rather than overdominant, i.e., heterozygotes
were more resistant than the average of parental homozygotes, but they
were not more resistant than both."
In other words the MHC
heterozygotes were never more pathogen resistant than the most resistant
homozygote. The authors point out, however, that heterozygotes may be more
fit than homozygotes on average, thus maintaining polymorphism in the
population, even if specific pathogen resistance is merely a
dominant trait. (Hedrick has reached the same conclusion on theoretical
grounds; Evolution 56:1902-8, 2002). They add that "The fact that MHC
heterozygotes were more resistant to infection and had higher fitness than
homozygotes provides a functional explanation for MHC-disassortative
mating preferences".
It appears to reconcile the observations that specific particular alleles
can have selective advantage in laboratory (and some aquaculture?)
conditions and that heterozygosity may be an advantage in nature. It
does not necessarily help one decide whether to artificially select for
homozygosity or heterozygosity in managed broodstocks. This would depend
on the variety and timing of challenges anticipated from pathogens. The
stakes could be very high. Optimal selection strategies for
extensive and "biosecure" aquaculture systems may be even more
different than we thought. Dustin Penn's address: Konrad Lorenz Institut,
Austrian Academy of Sciences, Savoyenstrasse 1a, A-116 D Wien, Austria.
380. Danish trout populations maintain
their genetic identity for a long time
Long-term
effective population sizes, temporal stability of genetic composition and
potential for local adaptation in anadromous brown trout ( Salmo trutta )
populations. 2002. Hansen, M. M., D. E. Ruzzante, E. E. Nielsen, D.
Bekkevold and K.-L. D. Mensberg. Molecular Ecology 11:1003-1015.
DNA from an 80-year
sequence of scale samples was analysed to estimate the effective
population number from the 1910s to the 1990s. Calculations were based on
temporal fluctuations in allele frequencies at eight microsatellite loci.
The authors conclude that some Danish rivers maintained their
genetic identities reasonably well over this period despite intense
stocking with hatchery trout and (which is more or less the same
conclusion) the long-term effective population numbers were 300 to more
than 500.
Population sizes of this order are large enough so that small,
but distinguishing, gene frequencies can persist for many generations
despite drift. However, the population sizes are nevertheless small enough that weak
directional selection (weak local adaptation) should to be overwhelmed by
drift. There is no reason in principle to expect local adaptation to be
weak; it can be and sometimes is very strong. But since the environment of
these Danish rivers appears more or less similar the authors don't think
that strong, divergent, local selection is likely to have caused the persistent gene
frequency differences. mmh@dfu.min.dk
379. Baltic salmon stocks not losing
diversity too quickly
Maintenance of
genetic diversity of Atlantic salmon (Salmo salar) by captive breeding
programmes and the geographic distribution of microsatellite variation. 2002.
Koljonen, M.-L., J. Tähtinen, M. Säisä and J. Koskiniemi. Aquaculture
212:69-92.
According to the authors
over 90% of Atlantic salmon smolts from Baltic gene pools are now reared
in hatcheries. In Finland, where this work was done, only two out of an
original 18 rivers have any naturally reproducing salmon at all. The
survival of Baltic stocks is now entirely dependent on costly maintenance
in hatcheries. Therefore it is interesting to see how well hatcheries are
doing from a genetic conservation perspective.
On the whole, the record is
not so bad in the hatchery and natural populations studied here. "In
short-term breeding programmes [where there are some wild breeders every
generation], the average rate of loss of heterozygosity was 1.4% per
generation and the average observed rate of loss of alleles was 4.7% per
generation. The estimated Ne for the broodstock were 32 and 238. The
average Ne/Nc ratio was 0.81. Changes in present-day broodstocks were not
alarming and the Ne/Nc ratios [effective number / census number] were
higher than in wild populations in general." marja-liisa.koljonen@rktl.fi
378. Genetic declines follow population
crashes in sea mammals
(a) Loss of
genetic diversity in sea otters (Enhydra lutris) associated with the fur
trade of the 18th and 19th centuries. 2002. Larson, S., R. Jameson, M.
Etnier, M. Fleming and B. Bentzen. Molecular Ecology 11:1899-1903.
(b) Impact of a
population bottleneck on symmetry and genetic diversity in the northern
elephant seal. 2002. Hoelzel, A. R., R. C. Fleischer, C. Campagna, B.
J. Le Boeuf and G. Alvord. Journal of Evolutionary Biology 15:567-575.
These before-and-after studies of the impact of human activities are
similar in that both are based on DNA extracted from samples preserved for
more than one hundred years.
The fur trade in the Pacific Northwest was
bad for the genetic well-being of sea otters. "While mtDNA sequence
variability was low within both modern and extinct populations, analysis
of microsatellite allelic data revealed that the prefur trade population
had significantly more variation than all the extant sea otter
populations."
Similarly for elephant seals: "The northern
elephant seal (NES) suffered a severe population bottleneck towards the
end of the nineteenth century. ... Measures of genetic diversity [before
and after the bottleneck] show a loss of variation consistent with
expectations and suggest a strong disruption in the pattern of allele
frequencies following the bottleneck. Measures of bilateral characters
show an increase in fluctuating asymmetry." shawn.larson@ci.seattle.wa.us
; a.r.hoelzel@durham.ac.uk
377. More anti-pathogen genes found in
Penaeid shrimp
Identification of
immune-related genes in hemocytes of black tiger shrimp (Penaeus monodon). 2002. Supungul, P., S. Klinbunga, R. Pichyangkura, S. Jitrapakdee, I.
Hirono, T. Aoki and A. Tassanakajon. Marine Biotechnology 4:487-494.
Hemocytes are probably the most important component of the
anti-pathogen defense system in shrimp. These cells are known to engage in
pathogen recognition, phagocytosis and release of anti-microbial peptides
as well as performing other immunological services. Some of the genes that
are involved in the immunological functioning of hemocytes have previously
been identified in Litopenaeus (or Penaeus) vannemai using the EST
technique (see Sep 2001 #233). The authors of this new paper have done
rather similar work on P. monodon, a species that makes up about 50% of
commercial shrimp production world wide. Homologies were found with known
defense functions including heat-shock proteins, the clotting system,
anti-oxidative enzymes, anti-bacterial and anti-fungal peptides etc. anchalee.k@chula.ac.th
376. Bottlenecking can cause
developmental instability in an introduced population
Increased
frequency of scale anomalies and loss of genetic variation in serially
bottlenecked populations of the dice snake, Natrix tessellata. 2002. Gautschi, B., A. Widmer, J. Joshi and J. C. Koella. Conservation
Genetics 3:235-245.
The dice snake is a
non-venomous Eurasian snake which spends much of its life in or near
water. Several populations introduced into Swiss lakes are known to have
been severely bottlenecked one or more times. As would be expected,
individuals in populations which have been bottlenecked more often have
fewer alleles per locus and less individual heterozygosity (8
microsatellite loci) than those in populations bottlenecked only once, and
much less than those in natural populations.
What is interesting in this
study is the finding that the frequency of scale anomalies (thought to be
the result reduced developmental stability caused by inbreeding) follow
the same pattern as the loss of genetic diversity. The population sizes are small and the species is red
listed in Switzerland and Germany. The authors point out that even if the
snake populations grow because of habitat protection etc. their genetic health will be
rather poor [for a while?] because of the bottlenecking. babagaut@uwinst.unizh.ch
375. Mass-spawned families may stay
together in plankton
Genetic and
demographic variation in new recruits of Naso unicornis. 2002. Planes,
S., G. Lecaillon, P. Lenfant and M. Meekan. Journal of Fish Biology
61:1033-1049.
"Demographic data
showed that larvae of Naso unicornis colonizing the reefs of Moorea,
French Polynesia, on the same night within a restricted area originated
from several spawning events that occurred 67 to 94 days previously." Age
was determined from daily growth rings, and genetic variables (Fst, Fis,
relatedness etc.) were calculated from allozyme data. Larvae which were
the same age (spawned on the same night) sometimes showed significant
genetic relatedness (Queller & Goodnight's relatedness measure)
despite the fact that they had spent an average of about seven weeks in
the plankton. Only some age groups (i.e. nights) showed this relatedness, and not all individuals which had been born on those nights were
related.
The authors conclude from their analyses that despite the mass
spawning behaviour and larval dispersal of this species, the accidents of
survival and dispersal are such that effective population size is greatly reduced. Reduced, in fact, to the point where there is large stochastic
fluctuation between the genetic composition of larval groups and between
larval and later stages, which become progressively more homogenous in the
lagoon as time passes, presumably because of physical dispersal and
mixing. (Why families would want to stay together in a
"sweepstakes" or "elm-oyster" (see Nov 2001 #259)
reproduction and selection regime is a mystery to me.). planes@univ-perp.fr
374. Shame! Shame!
Paper trail
reveals references go unread by citing authors. 2002. Ball, P. Nature
420:594.
It is shocking to learn
that authors don't bother to
read 80% of the papers they cite [or 80% of authors don't read any
papers?]. This conclusion was reached by two
physicists, M.V. Simkin and V.P. Chowdery, who studied the propagation of
errors in published citation lists. "The pair concluded that four out
of five authors had not done their homework." Most people
(physicists, anyway) just copy other people's reference lists. The
original paper can be downloaded from
http://xxx.lanl.gov/ftp/cond-mat/papers/0212/0212043.pdf . It is interesting not only for its embarrassing conclusions but for its
statistical treatment (did you know about the Zipf distribution, which has
obvious genetic applications?). It ends with the following threat: "We conclude
that misprints in scientific citations should not be discarded as mere
happenstance, but, similar to Freudian slips, – analyzed". An e-mail source for the Nature paper is not available.
373. Useful recent Excel add-ins for
analysis of genetic data
(a) PopTools looks like a useful and fun set of Excel add-ins for doing things which
can usually only be done in expensive, complicated programs like
Mathematica or S-plus. "PopTools is a versatile add-in for PC
versions of Microsoft Excel (97, 2000 or XP) that facilitates analysis of
matrix population models and simulation of stochastic processes. It was
originally written to analyse ecological models, but has much broader
application. It has been used for studies of population dynamics,
financial modeling, calculation of bootstrap and resampling statistics,
and can be used for preparing spreadsheet templates for teaching
statistics."
"Also included in PopTools are routines for iterating
spreadsheets. These make it possible to run Monte Carlo simulations,
conduct randomisation tests (including the Mantel test) and calculate
bootstrap statistics. Some facilities are available for function
minimisation and parameter estimation using maximum likelihood techniques,
and there are a number of auditing and other tools that the author finds
useful in his everyday work."
"PopTools allows the construction, analysis, and simulation of
quite complex models in a simple spreadsheet format. The only programming
required is the ability to construct formulas in Excel...." PopTools and descriptive files can be downloaded from http://www.cse.csiro.au/client_serv/software/poptools/index.htm
(b) DadShare is
an Excel macro designed to help interpret patterns of shared paternity
among groups of offspring. The program and explanatory documentation are
available at http://www.zoo.cam.ac.uk/zoostaff/amos/index.html .
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