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Yale University School of Medicine, Yale Center for Research On Reproductive Biology, New Haven,
CT 06511, USA and 1Mount Sinai School of Medicine, New York, NY 10029, USA
Received July 7, 2005; Revised August 15, 2005; Accepted October 10, 2005
*To whom correspondence should be addressed. Tel: +1 203 737 1218; Fax: +1 203 785 5294; Email: afadiel@yale.edu
The Author 2005. Published by Oxford University Press. All rights reserved.
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Nucleic Acids Research, 2005, Vol. 33, No. 19 6309
Table 1. General genomics and proteomics databases: comprised of resources for human, goat, mouse, deer, rat, and horse genomes
of similarity search programs, which operate on sequences, has The reproductive physiology of farm animals is more similar
helped in augmenting annotated databases, which house to humans than that of rodents because farm animals have
information about sequence domains. Information recording, longer gestation and pre-pubertal periods than mice (16). Spe-
retrieval and cataloging continue to advance further cific farm animal physiology, such as the digestive system of
database capabilities (13). Collaborating research centers, the pig is similar to that of humans (17). These attributes of
particularly in the US, Japan and the UK, have been actively farm animals reveal that they are an unparalleled resource for
collecting sequence data and making it accessible to public research replicating human physiological function (15).
(14) (Table 1). For decades, breeders have altered the genomes of farm
animals by first searching for desired phenotypic traits and
then selecting for superior animals to continue their lineage
Farm animals: an unexploited gold mine for biotech
into the next generation (18). This genomic work has already
Farm animals are quite valuable as resources, often notable as facilitated a reduction in genetic disorders in farm animals, as
models for pathology and physiology studies. Magnussen (15) many disease carriers are removed from breeding populations
discusses how a variety of these farm animal models are used. by purifying selection (19). By studying diverse phenotypes
6310 Nucleic Acids Research, 2005, Vol. 33, No. 19
over time, researchers can now monitor mutations that occur the economics of breeding, members of the commercial
as wild species become domesticated (20). Farm animal food pig industry are able to use this information to garner
safety will remain a concern for some time; however, advance- benefits (30).
ments such as the discovery of Escherichia coli resistant genes
in the pig (21) can mediate most of the problems. Moreover, The Bos taurus genome
resources devoted to investigating the genomes of farm anim- The mammalian order Cetartiodactyla (possessing B.taurus or
als can bring eventual economic benefits. For example, isola- cattle) is of great interest since it represents a group of
tion of DNA from animal tissue can be used as an inexpensive eutherian mammals phylogenetically distant from primates
method for tracking the origin of a meat sample, providing the (34,35). Working with the cow species, B.taurus, is significant
recipient with the quality assurance of that food (22). because the cow is such an economically important animal.
This form of livestock makes up the beef and milk production
2005: the year of the chicken industry, which is one of the largest industries in the United
There has been significant interest in the first complete ana- States. The identification of numerous single-nucleotide poly-
lysis of the draft genome sequence of the chicken. This morphisms (SNPs) makes it possible for geneticists to find
sequence has given rise to chicken genome array chips and associations between certain genes and cow traits that will
a number of web based mapping tools. The great importance of eventually lead to the production of superior-quality beef
the chicken as a scientific resource can be seen from the (36). After the completion of the September 2004 B.taurus
research on avian leucosis virus, which has led to develop- draft assembly (Table 4), this genome has functioned as a
vehicle for studies on non-primate and non-rodent genomes
hypothesis and showed the ability of the Fugu genome to aid www.genome.gov/Pages/Research/Sequencing/SeqProposals/
in the study of vertebrate functional non-coding sequences SticklebackSEQ.pdf). In general, fish genomic work is of
(41). Other studies of fish have focused on the stickleback, interest to the commercial fish farming community. Based
for its variable body shapes, ecology or its behavior (http:// on transgenic studies, antifreeze protein and fish
6312 Nucleic Acids Research, 2005, Vol. 33, No. 19
Table 3. Genomic and proteomic internet sites devoted to the study of the swine genome
growth hormone have been introduced into fish genomes, early stage. Currently available sources of information about
creating fish with greater cold tolerance and faster growth the rabbit on the web can be seen in Table 7.
rates (42).
Farm animal genomics: current statistics
The forgotten rabbit
As researchers delve into the composition of farm animal
Little genomics attention is paid to rabbits in comparison genome sequences, new functional and biological data
with other animal models, such as the mouse, rat and fruit emerge. Table 8 is compiled from the Ensembl database
fly in the pre- and post-genomic era. However, experimental (http://www.ensembl.org/) and illustrates a summary of cur-
models, such as the Alba or the ‘mighty lighting rabbit’, were rent analysis on coding regions within genomes for selected
developed as glowing mutants that shine under special light farm animals. For example, in the cow genome, 239 000
for commercial reasons. French scientists created Alba using exon regions and just over 29 000 transcripts have been tallied
a process called zygote microinjection. In this process, the from among the 565 million bp found in the genome.
scientists plucked a fluorescent protein from the fluorescent One of the major sources of information on farm animal
jellyfish Aequorea victoria. Then, they modified the gene to genomics is the ArkDB (43), which is available through the
make its glowing properties twice as powerful. This gene, Roslin Institute (UK) and Texas A&M University (USA). The
called EGFG (enhanced green fluorescent gene, was then ArkDB provides detailed genomic mapping data on sheep,
inserted into a fertilized rabbit egg cell that eventually grew chicken, cow and pig genomes (44), including data on PCR
into Alba (Amanda onion). Debates about the project itself primers, genetic linkage map assignments of specific loci and
and about the practice of manipulating genes in animals markers, and cytogenetic map assignments. Information about
for research have quickly arisen in the research community. farm animals in this database is displayed in Table 9 and
The French National Institute of Agronomic Research provides a snapshot of current resources available for each
hesitated to release these rabbits owing to protests over its genome. Quantitative functional information, such as the num-
development. ber of clones, microsatellites and associated mapping assign-
Chinese scientists have placed rabbit genes in cotton plants, ments, can lend insight into the complexity of the models
producing cotton fibers as bright and soft as rabbit hair but available. For example, the number of primers tabulated in
stronger and warmer. This indicates not only the rise of the the swine genome currently far exceeds that known for the
rabbit in experimental models for genetics engineering, but cow, indicating more varieties of genes are available for study
also for future medical experimentation models. Despite the in the swine. Much of this information remains at an early
controversy that the ‘shining rabbit’ and the ‘cotton rabbit’ stage and with increasing experimentation, compilation and
raised, it is clear that this type of genetic engineering is at an analysis, will be refined.
Nucleic Acids Research, 2005, Vol. 33, No. 19 6313
Table 8. Ensemble (gene build July 2005) and NCBI statistics of available farm animal genomesa
Ensembl (e)
1 eGene predictions 17 784 22 013 NA NA 22 877
2 Genscan predictions 77 600 103 597 NA NA 49 697
3 egene exons 185 326 239 889 NA NA 231 799
4 egene transcripts 28 491 29 363 NA NA 32 143
5 Base pairs 1 054 180 845 565 382 643 NA NA 1 688 467 974
NCBI (entrez 2005 records)
1 Nucleotide 923 899 1 961 869 628 750 28 716 912 447
2 Protein 29 634 51 726 8835 3160 47 458
3 UniSTS 2936 13 320 7851 2408 27 616
4 Gene 18 449 39 815 2388 625 18 429
5 HomoloGene 14 433 12 909 10 795 1016 11 026
6 EST 554 084 624 652 461 891 10 960 651 991
a
The ESTs in this table are based on the dbEST release 082605-August 26, 2005 (http://www.ncbi.nlm.nih.gov/dbEST/dbEST_summary.html) BLAST alignment
data, associated Human UniGene and RH maps (EBI GenomeMap’99 and NCBI GeneMap’99), pig–human comparative map information and mapped pig genes
in the pig genome database (PigBase).
Expression sequence tags gene identification and positional mapping within a genome
(http://www.ncbi.nlm.nih.gov/About/primer/est.html). The
Expression sequence tags (ESTs) are small pieces of cDNA NCBI provides the most comprehensive EST database for
sequence (usually 200–500 nt long), which are useful as mark- many farm animals. Tabulation of chicken EST information,
ers for a desired portion of RNA and DNA that can be used for including quantities of ESTs and their tissue sources can
Nucleic Acids Research, 2005, Vol. 33, No. 19 6315
Table 9. Current status summary provided by the ArkDB 410 loci with 1327 homologue loci on 129 species, 21 poly-
morphisms, 314 genes and their physical location on chromo-
No. Element Chicken Pig Cow Sheep
somes. Other information about primers, probes and enzymes
Markers are also included. Genes in the RabbitMap database are hyper-
1 Loci 2530 4081 2725 2030 linked to a separate page that provides locus information.
2 Designated genes 765 1588 746 543 Physical mapping of the zebrafish is available through the
3 5 3 Sine-PCR 47 18 146 3
4 Microsatellite 1277 1673 1219 2257
Ensemble database at http://www.ensembl.org/Danio_rerio/
5 RFLP 255 258 250 index.html. The Ensemble database also provides a compre-
6 Clones 316 602 1 206 hensive map viewer for zebrafish and the chicken (http://www.
Map assignments ensembl.org/Gallus_gallus/index.html). High-resolution maps
1 Linkage 3400 5141 2209 14 909 for sheep chromosomes are available at http://www.ncbi.nlm.
2 Cytogenetic 307 1927 659 843
3 Total 3707 7068 2868 15 752 nih.gov/genome/guide/sheep/index.html.
be found in the BBSRC ChickEST Database (http://chick. One of the primary challenges in modern biology is the under-
standing of the genetic basis of phenomic diversity within and
umist.ac.uk/) (45,46). ChickEST provides access to
among species (20). The foundation for this diversity lies in
339 000 Gallus ESTs, generated from 21 different embry-
genetic governance of both how traits are expressed and the
onic and adult tissues. Cow EST information is available on
ethical consideration. Although a number of gene-based The potential of farm animal genomics
technological advances, such as the prediction of illnesses
Farm animal genomic studies continue to attract audiences
or adverse drug response are beneficial for commercial
excited by the multitude of applications (62). The meat indus-
interests (54), many look to protect how genetic information
try can now use cow and chicken genomic data to confirm the
is used. The corollary in human genomics is the recent
quality of meat products. For example, meat producers can
Genetic Information Nondiscrimination Act of 2005 (http://
now confirm the parentage of an Angus cattle breed by per-
www.genome.gov/PolicyEthics/). This bill, which is pending
forming a genetic blood test or attempt to identify the SNPs
in the US Congress, prohibits among other things ‘a health
associated with high-quality beef (57). Other companies are
insurance issuer from: adjusting premiums on the basis of
using genomic information to determine disease-resistant
genetic information’ (http://thomas.loc.gov/cgi-bin/bdquery/
genes in shrimp and then are selectively mating the shrimp
z?d109:SN00306:@@@L&summ2=m&). Concerns over
that carry them in order to create disease resistant strains (57).
such actions reveal the understanding by the US government
In the healthcare arena, farm animal genomic work will aid in
that the use of human genomic data must be closely regulated
enterprises such as xenotransplantation (the transfer of animal
in order to hinder any commercial abuses. Although genetic
tissues or organs into humans). Though animal organs may be
discrimination and health insurance might not be issues rel-
used someday to satiate organ donor shortage, genomic work
evant to farm animal genomics, a great need exists to establish
in this area is still in its early stage (63). Many of the
a similar legal framework for farm animals in order to ensure
immediate practical applications of farm animal genomics
the proper use of farm animal genomic information.
show potential for growth in this field.
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