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.
321. A gene for salinity tolerance in tilapia:
microsatellites are directly selected!
Microsatellite variation associated with prolactin expression and
growth of salt-challenged tilapia. 2002. Streelman, J.T, and T.D.
Kocher. Physiological Genomics 9 (1):1-4.
Tilapia geneticists and
many population geneticists should be
interested in this paper. The relative growth of individual O.
mossambicus X niloticus hybrids in saline water is influenced
by variation in the relative expression of two prolactin genes. (Prolactin
is a peptide growth hormone which also affects osmoregulation.)
The variability in expression is caused in part by variation in the length
of a microsatellite repeat embedded in the control region of the prolactin
genes.
This is a very good story: not only is the microsatellite marker
decidedly non-neutral, it may be under selection in an environment that
has considerable importance in aquaculture and, perhaps, conservation (such
as feral mossambicus taking over the seas). The effect of the microsatellite
is apparently not caused by linkage, i.e. because the microsatellite is close
to the prolactin gene on the chormosome (association overdominance). It is
a functional effect. The authors speculate that "repeats of varying length can induce
promoter conformations that differ in their ability to bind
transcriptional regulators". Fish homozygous for the long
microsatellite were only half as big, in half-strength seawater, as
fish homozygous or heterozygous for the short microsatellite. There was
strong genotype-environment interaction. "Each genotype grew best at
a different salinity treatment".
The authors conclude, "From an evolutionary perspective, our results
run counter to the textbook interpretation that dinucleotide
microsatellite variation lacks functional consequences. ...
[microsatellites] provide a cellular means to alter the amount of gene
product among individuals without changing amino acid sequence. The
association of microsatellites with a long list of transcription factors
and environmentally regulated genes suggests an under-appreciated role in
the fine-scale modulation of gene expression." Streelman e-mail: jts3@hopper.unh.edu .
320. A new relatedness estimator for use
when pedigrees aren't available
An estimator for pairwise relatedness using molecular markers. 2002. Wang, J. Genetics 160:1203-1215.
When true pedigree data are lacking, microsatellites
or other markers may possibly be used instead for practical purposes like
estimating heritabilities in the wild, managing fish farms or increasing
founder diversity in captive populations (Nov 2001 #261; Jan 2002 #283).
This new estimator can be used to estimate inbreeding of individuals as
well as the relatedness of pairs of individuals. It is unbiased and
appears to offer advantages over previous estimators in its relative
insensitivity to problem data. "The new estimator is also robust for
small sample sizes and for unknown relatives being included in samples for
estimating allele frequencies. Compared to previous estimators, the new
one is generally advantageous, especially for highly polymorphic loci
and/or small sample sizes." For a recent review of other relatedness estimators
see Sep 2001 #227. jinliang.wang@ioz.ac.uk
319. Genetic biomarkers for disease and toxic stress in oysters
Potential indicators of stress response identified by expressed
sequence tag analysis of hemocytes and embryos from the American oyster,
Crassostrea virginica. 2002. Jenny, M.J., A.H. Ringwood, E.R. Lacy, A.J.
Lewitus, J.W. Kempton, P.S. Gross, G.W. Warr, and R.W. Chapman. Marine
Biotechnology 4:81-93.
C. virginica is an important commercial crop and is also a useful
indicator of environmental quality in the coastal waters of eastern North
America. The animal is susceptible to a variety of water-borne diseases,
pollutants and toxic chemicals in low concentrations. This paper describes
a search for oyster genes that are expressed -- turned on and
manufacturing their products -- in the presence of these harmful agents.
(See Sep 2001 #233 for a bit more information about expressed sequence
tags.) The idea is to find biomarkers for environmental stress and, more
generally, to learn more about what molluscs pretend to be their immune
systems. "Several potential biomarkers identified include an
antimicrobial peptide [see May 2000 #55], recognition molecules (lectin
receptors), proteinases and proteinase inhibitors, and a novel
metallothionein."
The authors conclude that " ... the
observation that most of the messages (ca. 70%) encode proteins that are
either completely novel or are related to genes of unknown function may be
the most interesting finding. If taken at face value, it suggests that a
large proportion of metabolic energy is utilized, in oyster embryos and
hemocytes, to generate proteins whose functions are unknown." chapmanr@mrd.dnr.state.sc.us
318. Fluctuating populations may be larger than we
thought
The effective size of fluctuating populations. 2001. Iizuka,
Masaru. Theoretical Population Biology 59:281-286.
The rates at which a population accumulates inbreeding and loses genetic
diversity by random drift are often expressed in terms of the
"effective number" of the population. The effective number, Ne,
is a theoretical number which is usually smaller than the actual number of
animals. It is smaller mainly because of unequal sex ratio, the failure of
many animals to breed, non-random mating, and the exaggerated success of a
few breeders. These problems lower the effective number in hatcheries and in nature.
When population size fluctuates from generation to generation the
effective population number over the long term is a lot smaller than the
ordinary average value. It is usually considered to be
the harmonic mean (the reciprocal of the average of the reciprocals) of
the individual values. Therefore, when a population occasionally crashes and
recovers, the long-term effective population number (the harmonic mean) is actually much
closer to the smallest size reached during the crash. Or at least
that is what people think. This leads to concern about the genetic health
of endangered natural populations and aquacultural populations, which are
often reduced to a few breeders whenever a new broodstock is started.
This theoretical paper by Iizuka turns the usual thinking upside down.
He demonstrates (in his theorem #5) that the long-term effective number for inbreeding
and drift is actually very close to the largest value reached by a
fluctuating population if the sequence of values is highly, and
positively, autocorrelated. (Population size is autocorrelated when a
"good" year tends to be followed by another relatively good
year, a poor year is likely to be followed by another poor year etc.)
How
important is this new insight concerning effective population number? At
the very least, it would be interesting to try out the effect of autocorrelation
using a sequence of real data. Population sizes of fish
generally are positively autocorrelated. Even if they aren't, overlapping
generations should induce positive autocorrelation. No e-mail available.
The address is: Division of Mathematics, Kyushu Dental College, 2-6-1 Manazuru,
Kokurakita-ku, Kitakyushu 803-8580, Japan.
317. Alien invaders mate with locals in
Wisconsin
Implications of hybridization between introduced and resident Orconectes crayfishes. 2001. Perry, W.L., J.L. Feder, and D.M. Lodge.
Conservation Biology 15 (6):1656-1666.
There are several hundred
species of crayfish in North America (80% of the world total) and almost
one-third of these species are listed as threatened or endangered. One of
the threats to species survival is introduction of non-native species,
which not only homogenize species distributions across the continent, i.e.
increases within-site diversity while reducing between-site diversity,
but can cause local extinction as well. (For an analogous effect on the
diversity of N.A. fish see Aug 2000 #95). The authors found that Orconectes rusticus,
native to northern Ohio but introduced widely elsewhere in the U.S., is
displacing O. virilis and O. propinquus in northern Wisconsin. This is a purely
ecological displacement as far as O. virilis is concerned; The
authors detected no hybrids of this species.
The problem facing O.
propinquus is more genetic. "Over 6% of the crayfish at one
sympatric site were putative F1 hybrids [rusticus X propinquus],
4% were putative F2 individuals ( hybrid × hybrid origin), and 13% were
putative backcrosses ( product of hybrid × parental matings). ... Our
results suggest that genetic mechanisms play a role in the extirpation of O.
propinquus by O. rusticus and are consistent with observations
of other researchers suggesting that hybridization with non-native species
is common among crayfishes at many other locations. High rates of endemism
and widespread introductions of non-native crayfish suggest that invasions
and hybridization are a major threat to crayfish biodiversity." wlperry@ilstu.edu
316. Markers may be a poor substitute for pedigrees
The use of marker-based relationship information to estimate the
heritability of body weight in a natural population: a cautionary tale. 2002.
Thomas, S.C., D.W. Coltman, and J.M. Pemberton. Journal of Evolutionary
Biology 15 (1):92.
It may sometimes be possible to use genetic relationships inferred from
microsatellite data as a substitute for pedigree records. Many papers
mentioned on this website have explored the idea, including an earlier
paper by these same authors (Aug 2000 #88). Apparently it doesn't work very
well with the feral Soay sheep of St. Kilda, for which a good pedigree
record exists. Several methods of estimating heritabilities from
microsatellite-inferred relationships gave unreliable
estimates of body weight heritability.
Even if this turns out to
be a general problem with variance component estimation it need not
obviate the use of relatedness estimators for other purposes. Note
that is a close correspondence between inferred relatedness and that
determined by actual pedigrees in thoroughbred race horses (Mar 2002
#303). stuart.thomas@ed.ac.uk
315. Graylings may have evolved in response to size-selective fishing
A century of life-history evolution in grayling. 2001. Haugen, T.O.
and L.A. Vøllestad. Genetica 112-113:475-491.
This is a study of several populations of graylings (Thymallus
thymallus, a sort of trout or salmon) that have been isolated from
each other and from a common source for 9 - 22 generations in Norway. The
source population still flourishes in one of the lakes studied.
Experiments demonstrating that adaptation to new temperature regimes has
evolved in these fish have already been published (Jan 2001 #162). The
authors have now extended the work to include changes in other phenotypic
traits, including size and age at maturity, size-specific fecundity,
growth rate, size at swim-up etc. For some of the traits they used
preserved scale samples to look at the changing phenotype over the years
in the source lake. For other traits they grew fish from different lakes
at a single location ("common garden" experiments). For some
adult traits they collected samples from the lakes and measured
contemporary variation in the phenotype.
They estimate the selection intensity caused by size-selective fishing
(gill-netting) on the basis of a life-history model which maximizes
lifetime contribution to the growth of a population with a stable age
structure. The evolutionary conclusions in the paper therefore derive from
direct genetic observations, i.e. from the common garden experiments, and
also from inferences about phenotypic measurements that span a range of
time, space and real and hypothetical selection regimes.
The paper supports the conclusion that size-selective fishing in the
lakes has caused rapid evolution towards earlier maturation. An
evolutionary change in the growth pattern (faster early growth and slower
late growth) and increased size-specific fecundity has also been caused by
selective fishing. All these changes are in the direction predicted by
theory. The speed and magnitude of the changes show that evolutionary
considerations may be important for fisheries management on a decadal time
scale. throndh@bio.uio.no
314. Swimbladder inflation in bass is not heritable:
a useful negative result
Heritability of swimbladder inflation in striped bass. 2002.
Harrell, R.M., W.V. Heukelem, J.M. Jacobs, J.R. Schutz, J.U. Takacs, and
D. Jacobs. North American Journal of Aquaculture 64:117-121.
Striped bass (Morone saxatilis) that don't inflate their swim
bladders in a timely way have serious skeletal and other developmental
abnormalities, if they survive at all. The authors of this paper conducted
a factorial mating experiment to see if the problem is heritable. They found
that it isn't. Their estimate of half-sib heritability was essentially
zero. The full-sib estimate was a robust 0.35, however, and the authors
conclude that "swim bladder inflation is predominantly influenced by
dominance, epistasis, or environmental factors".
They draw the
conclusion that there may be genotypic variation for the trait but it
isn't additive and will not respond to selection. The authors should be
praised for publishing this useful, negative result, which is likely to
save people a lot of time and effort. (For a more aggressive approach to
the marketing of full-sib data see Feb 2002 #295.) rh116@umail.emd.edu
313. Useful additions to the "effective population size"
concept
Relationship of effective to census size in fluctuating populations. 2002. Kalinowski, S.T., and R.S. Waples. Conservation Biology 16
(1):129-136.
Population biologists and hatchery managers
often like to talk about the effective population number even if they are not geneticists.
It seems like a simple idea -- to the extent there is such thing in population
genetics -- which evidently relates to the size of some sort of
population. The effective population is a conceptual one that has tidy
properties like random mating and 1:1 sex ratio, and which loses genetic
diversity and/or accumulates inbreeding at the same rate as the real one
with which it is being compared.
The ratio of effective number to the real
census number, Ne/N is an intuitive measure of how well a population can
protect its genetic quality against random degradation. Rather like
comparing the mechanical advantage of an ideal "frictionless pulley"
to a real pulley to see how well a pulley is made. The effective
population size and the frictionless pulley are highly practical,
theoretical concepts.
This paper by Kalinowski and Waples extends the intuitive appeal of the
concept by defining several variables for use when the census size of a
population fluctuates from generation to generation. These variables are
related to the ratio Ne/N and give it some intuitive depth when numbers
keep changing from census to census. For example,
the meaning given to the variable beta is "a description of how much
each real individual contributed to the effective population size"
during a period of fluctuation. Their examples and discussions help
clarify the meaning of the effective population number in real-world situations. The definitions are
algebraic and not tied to any particular genetic model, so there is hope
they will continue to apply even in complex situations such as #318,
above. steven.kalinowski@noaa.gov
312. Heterozygosity is probably the best way to detect inbreeding using
markers
Heterosis, marker mutational processes and population inbreeding
history. 2001. Tsitrone, A., F. Rousset, and P. David. Genetics
159:1845-1859.
In practical aquaculture and conservation one often wants to know whether
a particular population, in a particular situation, is suffering the
effects of inbreeding depression. There are various ways inbreeding can
arise and it may be very important to know what is happening. In
aquaculture, for example, inbreeding can be caused by systematic mating
between relatives in a mis-managed breeding program, accidental mating
between relatives in a broodstock that is chronically too small, a
too-small founding broodstock, etc. Analogous things happen to natural
populations.
At any given time there will be variation
in the
inbreeding levels of individuals in the population. If inbreeding
depression is causing problems for the population as a whole, then there
usually will be some degree of correlation between individual fitness (size,
fecundity, parasite load etc.) and individual inbreeding. It is easy to
check for this correlation if pedigree records exist. If they don't exist,
DNA markers might provide an alternative. But how should DNA markers be used
for this purpose?
One solution is to estimate an inbreeding coefficient for individuals
using markers to reconstruct, or deduce, the absent pedigree records (e.g. Dec 2000
#142), and then look for a correlation between fitness and the inbreeding
coefficient. The second solution is to look for a correlation
between the marker heterozygosity (as an inverse indicator of inbreeding) and
the fitness of individuals. Such correlations are often found, and the three
or four most likely explanations have been debated by geneticists for
about 20 years. A third solution is to construct a genotypic index which
depends on the mutational dynamics of the marker locus, rather than its
heterozygosity, as an indicator of inbreeding. One such index is d2 (Mar
2002 #303) which is the square of the difference in length of the two
microsatellite alleles at a marker locus in an individual.
This paper compares the power of the heterozygosity and the d2 methods
to detect inbreeding, using simulations based on the two prevalent models
of the mutation process and a variety of population scenarios. In almost every scenario examined, inbreeding
was more highly
correlated with marker heterozygosity than with marker d2. This is useful
to know. It would be useful to compare heterozygosity and d2 in some
of the more pathological breeding scenarios one encounters in hatcheries,
to see if it holds true there as well. tsitrone@cefe.cnrs-mop.fr
311. Beautiful visual display of a good genetic
story
African pastoralism: genetic imprints of origins and migrations. 2002. Hanotte, O., D.G. Bradley, J.W. Ochieng, Y. Verjee, E.W. Hill, and
J.E.O Rege. Science 296:336-339.
This paper about the domestication of cattle in Africa should interest all
conservation geneticists, for two reasons. The first is the way the
microsatellite data are displayed visually. The authors extracted the
first three principal components (PCs) from microsatellite data on more
than 50 breeds of indigenous cattle scattered throughout Africa, and
plotted the PCs as a "synthetic map" superimposed on an ordinary
map of Africa. This sort of thing has been done
before but the result here is absolutely stunning. You can understand
10,000 years of genetic history at a glance. The visual display renders
the tale told by the authors instantly convincing.
The second reason is the tale itself. Apparently
cattle were first domesticated locally, in the north-east of Africa, and
the original breed of Bos taurus was walked southwards all the way
to the Cape by pastoralists. This process may have begun about 8,000 years
ago. The animals are pictured in pre-historic rock paintings and both they
and their living descendents look very strange to my Canadian eyes. The
southward diffusion is shown beautifully in the map of the second
principal component (PC2).
Most of the cattle now in Africa are also
descended from another type of cattle (B. indicus) which was
domesticated in the Near East, and which PC1 shows to have entered
East Africa and the Horn, probably by sea trade. Pictorial representation
of these animals, which unlike B. taurus are hump-backed, appear in
Egyptian tomb paintings after about 2,000 BC. The
site of introduction of hump-back cattle and the spread of their paternal
genes over the continent is obvious in the map of PC1. The third PC shows
that there has also been more recent introduction of B. taurus genes from Europe and the Near East. An elegant paper. The maps with
superimposed principal components were drawn with the GIS program ArcView
(Spatial Analyst Extension) which is available, for a price, at http://www.esri.com/software/arcview.
The program runs on LINUX and MSWindows. o.hanotte@cgiar.org
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