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Integrating Molecular Biology into the Veterinary Curriculum

Article  in  Journal of Veterinary Medical Education · February 2007


DOI: 10.3138/jvme.34.5.658 · Source: PubMed

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Research and Education Reports

Integrating Molecular Biology into the


Veterinary Curriculum
Marion T. Ryan g Torres Sweeney

ABSTRACT
The modern discipline of molecular biology is gaining increasing relevance in the field of veterinary medicine. This trend
must be reflected in the curriculum if veterinarians are to capitalize on opportunities arising from this field and direct its
development toward their own goals as a profession. This review outlines current applications of molecular-based
technologies that are relevant to the veterinary profession. In addition, the current techniques and technologies employed
within the field of molecular biology are discussed. Difficulties associated with teaching a subject such as molecular biology
within a veterinary curriculum can be alleviated by effectively integrating molecular topics throughout the curriculum, pitching
the subject at an appropriate depth, and employing varied teaching methods throughout.

Key words: molecular biology; education; veterinary; genomics; CAL

INTRODUCTION of morphology—it must at the same time inquire


Molecular biology is the modern branch of biology that into genesis and function.1
explains biological factors, processes, and systems at a In parallel with the ever-widening body of knowledge
molecular level. It also encompasses biochemical and and concepts within the field of molecular biology there is
physical techniques used to investigate phenomena of also an extensive, constantly expanding and evolving
molecular origin. The field of molecular biology integrates ‘‘toolbox’’ of techniques.
a number of areas of biology, particularly genetics,
genomics, biochemistry, and cell biology (Figure 1), con- Numerous technologies associated with the exploration of
cerning itself with understanding the interactions between molecular biological phenomena have enabled researchers
the various systems of a cell. to explore veterinary-related research to new depths. In
addition, molecular-biology techniques are increasingly
The broad aim of scientists in this field include characteriz- being employed in veterinary diagnostics. The successful
ing genomes, understanding influences on gene expression and meaningful integration of molecular biology into the
and regulation, gaining insights into the molecular structure veterinary curriculum is a difficult but necessary challenge
and function of biologically important molecules, and for veterinary educators.
identifying how such factors and molecules interact. In
1961, Astbury described molecular biology as follows:
not so much a technique as an approach, an APPLICATIONS OF MOLECULAR BIOLOGY IN
approach from the viewpoint of the so-called VETERINARY MEDICINE
basic sciences with the leading idea of searching The veterinary curriculum is constantly evolving and
below the large-scale manifestations of classical adapting to the changing needs of society. Leighton2
biology for the corresponding molecular plan. makes the point that if veterinary medicine is to serve
It is concerned particularly with the forms of society, there must be enough veterinarians to supply the
biological molecules and . . . is predominantly demand for clinicians as well as ‘‘different’’ veterinarians
three-dimensional and structural—which does who can work in an area other than clinical practice. Only
not mean, however, that it is merely a refinement then can veterinarians provide a unique perspective in areas
such as bio-security, food safety, zoonotic diseases, ecology
of health and disease, and management of the environ-
ment.2, 3 If veterinarians are to adopt new roles, they must
Genomics Cell biology acquire the required new skills and knowledge. The way
molecular biology is being applied in veterinary practice
Molecular biology and in wider areas of veterinary endeavor needs to be
reflected in the curriculum. As Zemljit has written,
The veterinarians of the future must be armed
Genetics Biochemistry with new knowledge, which will be different
from that needed today.4
Figure 1: A schematic overview of the discipline of Molecular biology is relevant to a wide range of core
molecular biology. subjects within the veterinary curriculum and is applied in

658 JVME 34(5) ß 2007 AAVMC


Table 1: Important applications of molecular biology in veterinary medicine
Application Summary and Progress to Date

Detection of breed-specific Numerous diseases and traits in domestic species have a single-gene etiology.6, 7 Once a single-
single-gene disorders and traits gene disorder has been characterized, techniques such as PCR, RFLP, sequencing, and
microsatellite analyses can then be employed to determine whether an animal carries the
affected allele. Several of these single-gene traits/disorders have been identified in pedigree
animals, including dogs,8 cats,9 horses,10 cattle,11 pigs,12 and sheep.13 Owners can then take
this information into account when breeding. Various commercial companiesa–c offer services
that can detect the presence of a variety of single-gene disorders, including canine
leukocyte adhesion deficiency (CLAD), polycystic kidney disease, and severe combined
immunodeficiency (SCID).
Pathogen detection The routine use of molecular-based methods for human pathogen detection has been
established for a number of pathogens in the clinical setting.14 A process of optimization,
validation, and interpretation of results against clinical criteria is required when introducing a
molecular test into the microbiology laboratory.15 The development of new molecular-based
assays in the veterinary clinical setting has lagged behind human medicine, as companies have
traditionally focused on developing assays and products tailored to human pathogen diagnostics.
Increasingly, commercial companies are beginning to offer DNA-based pathogen detection as a
service.d, e In addition to this some commercial companies, are beginning to invest in molecular-
based products directed toward the veterinary diagnostic market. Artusf supplies optimized PCR
kits that can be used for detection and quantification of pathogen load; these are available
for Campylobacter, L. monocytogenes, M. paratuberculosis, and Salmonella. Qiageng is now
promoting a range of products targeted at molecular diagnostics and research in veterinary
medicine.
Parentage testing Several companies offer parentage testing as a service.d, h DNA testing is available for a number
of species, including horses,16 dogs,17 and cats.18 These DNA tests not only provide information
relating to parentage but also serve as a unique genetic profile that can be used for
identification purposes.
Epidemiology Typing of pathogens is carried out using a range of molecular-based techniques, including
plasmid analysis, pulse field gel electrophoresis, and multilocus allelic sequence-based typing.19
Epidemiologically related isolates share the same DNA profile, whereas unrelated isolates display
patterns that are different from one another. These techniques allow investigators to trace the
transmission of pathogens that have a negative effect on both production and animal welfare.
Molecular-based typing also has a role in food safety, giving a better understanding of the
mechanisms by which food-borne pathogens are transmitted. Monitoring antimicrobial-resistant
strains such as Methicillin-resistant Staphylococcus aureus 20 and other zoonotic diseases is
important from a public-health perspective.
Using animal models to under- A wide range of animal models is used to explore the etiology of human diseases.21–23 Animal
stand disease mechanisms models are a useful means of exploring disease states, as the ethical restrictions are less stringent
and therapeutics in human at a number of levels. Live-animal experiments, transgenic animals, breeding programs, and easy
medicine acquisition of tissue samples allow the completion of more comprehensive experiments than
could ever be contemplated in human medical research.
Animal cloning Of the 160 laboratories worldwide that carry out cloning experiments, 75% are working on
livestock, including cattle, pigs, goats, and sheep.24 Cloning technology’s current low success
rate, which is due to both technological and species-specific biological factors, 25 has hampered
its economic feasibility.26 It has potential uses in the many areas, including basic research,
generation of disease models, bioreactors, agricultural applications, and xenotransplantation.
Xenotransplantation Xenotransplantation has attracted considerable interest in recent years and is continually moving
closer to clinical application. This area of research is driven by the increasing demand for
replacement organs and the short supply of suitable donors. The prime candidate animal in this
field is the pig, which is most suitable in terms of breeding, rearing, cost, ethics, and anatomical
compatibility to humans as well as because of its physiological and biochemical characteristics.
The challenges in this area include graft rejection, acute inflammatory reactions, coagulation,
physiological incompatibilities, and the danger of zoonosis transmission. Genetic engineering of
donor animals has been seen as a way to overcome the challenges of acute rejection and
coagulation.27

(Continued )

JVME 34(5) ß 2007 AAVMC 659


Table 1: Continued
Application Summary and Progress to Date

Tailored pharmacology Pharmacogenetics explores variations in individuals’ responses to drug therapy based on their
genotype (data from multiple genes). In veterinary medicine, a certain level of knowledge exists
in this field, mainly relating to polymorphisms in single genes concerned with drug metabolism.28
Another example is the well-characterized adverse reaction to halothane anesthesia that occurs
in pigs carrying a single-nucleotide polymorphism (SNP) in the ryanodine receptor.12 The
understanding of pharmacogenetics in human medicine is considerably more advanced, as
commercial interest is much greater. Current areas of interest in human medicine include cancer
therapy,29 treatment of mood disorders,30 responses to cholesterol-lowering drugs,31 and
variations in drug response between different ethnic groups.32
Quality-trait loci Quality-trait loci (QTL) are locations on the genome of genes or other genetic elements that
contribute to the expression of a quantitative trait.33 Quantitative traits show different degrees of
variation, depending on the trait, environmental factors, and the population in question.
Quantitative traits include production traits such as meat quality,34 wool quality,35 and milk
production;36 performance traits such as athletic performance;37 and non-production traits such
as body conformation.38 QTL relating to disease susceptibility, such as mastitis in cattle39 and
nematode resistance in sheep,40 are also of importance. All these traits are multifactorial, and
many can have low heritability, which has hindered any genetic gains resulting from conventional
selection and breeding.41 Efforts are underway to integrate QTL and phenotype data with
sequence and gene maps in a database form.42, i
Comparative genomics The comparison of genomes from different species, or comparative genomics, is now becoming
possible as the genomes of more and more species are uncovered. Comparative genomics will
give a greater insight into evolution, gene function, conserved regions, and non-coding DNA43
and is being employed as part of the functional annotation process.44

disciplines such as pathology, microbiology, endocrinology, Basic subjects


parasitology, immunology, animal husbandry, pharmacol-
ogy, embryology, and epidemiology (see Table 1 6–44). The Cell biology
Biochemistry
discipline of molecular biology has enriched our under- Gene structure and function
Principles of inheritance and genetic disease
standing of many of these subject areas by allowing
investigation of the underlying molecular processes within
them. In embryology, for example, molecular biology has
Techniques and Technologies
enabled developments that were formerly only described (See Table 1)
anatomically to be explained in terms of molecular
processes and regulating factors.5 Molecular-biology tech-
nologies have direct application to veterinary diagnostics, Applications of molecular biology in
veterinary medicine
veterinary epidemiology, and veterinary-related research.
Diagnostic testing for genetically based disease states and disease
susceptibility
Epidemiology of pathogens
Facilitating selective breeding programs for resistance to disease,
CHALLENGES OF DEVELOPING UNDERSTANDING enhanced production traits
IN BASIC MOLECULAR BIOLOGY THEORY Genetic changes in neoplastic diseases
Veterinary research
Molecular biology is a challenging subject in itself; Parentage testing
the subject matter is complex, and the scope wide. In the Pathogen identification and quantification
Pharmacogenomics
veterinary context, the material delivered needs to be
suitably detailed but not so extensive as to encourage
surface learning strategies.45, 46 When studying molecular Figure 2: An overview of molecular biology as a
biology, the student must commit a large amount of material discipline within the veterinary curriculum.
to memory, including definitions, terminology, names of
biologically active compounds, and biochemical processes
and pathways. Memorization can be used to master troublesome knowledge as that which is conceptually
unfamiliar terminology, as a first step toward understand- difficult, counterintuitive, inert, or alien.
ing.47 The student must possess a basic knowledge of Troublesome knowledge is believed to originate from the
genetics, cell biology, and biochemistry, as well as a new student’s inability to grasp more fundamental and unifying
lexicon for discussing these subjects, before progressing to concepts, termed ‘‘threshold concepts’’: conceptual gate-
the techniques and, finally, the applications of molecular
ways that are transformative, irreversible, and integrative
biology in veterinary medicine (Figure 2).
and lead to a previously inaccessible way of thinking about
Many areas of molecular biology can be considered the subject.49 Interestingly, these threshold concepts, once
‘‘troublesome’’ to students (Table 2). Perkins48 defines mastered, tend to give the learner a more privileged view of

660 JVME 34(5) ß 2007 AAVMC


Table 2: Selected troublesome areas in molecular biology, the nature of the difficulty, and related areas of
knowledge and understanding
Troublesome Area Nature of Difficulty Related Knowledge and Understanding

Higher organisms are com- Students may ‘‘know’’ this, but a true  Cell signaling
posed of specialized, hetero- appreciation of the concept requires an  Signal transduction
geneous populations of cells understanding of the areas listed in  Receptors
that communicate with each column 3.  Competence
other.  Cell differentiation
 Differential gene expression
Different cell types within an Students may ‘‘know’’ this, but a true  Transcription
organism have differential pat- appreciation of this concept requires an  Translation
terns of gene expression, understanding of some of the areas listed  Gene regulation
despite possessing the same in column 3.  Differentiation
genome.  Real-time PCR
 2D gel electrophoresis
 Microarrays
 Cell signaling
Protein metabolism, synthesis,  An appreciation of this concept requires  Amino acids
and secretion an understanding of the areas listed in  Codons
column 3.  Ribosomal, transfer, and messenger RNA
 The area contains a number of pathways  Protein metabolism
that have to be memorized.  Protein storage and secretion
 Students confuse digested protein with  Properties of proteins
proteins that are synthesized by a cell.  Post-translational modification
Hardy/Weinberg equilibrium Abstract  Alleles
 Phenotype/genotype
 Genetic fitness
 Epistatis
 Inbreeding depression
 Genetic drift
 Evolution
Genetic linkage Abstract  Genetic markers
 Co-inheritance of undesirable traits with
selected traits in animals
Mendelian principles of Abstract  Dominant/recessive alleles
inheritance  Penetrance
 Phenotype/genotype relationship
 Inheritance patterns of single-gene
diseases and traits
Multigenic inheritance Abstract  Additive effects of genes
 Heritability
 Quality-trait loci (QTL)
 Selection and breeding strategies
Polymerase chain reaction Dynamic  Chemical characteristics of DNA
(PCR)  DNA replication and synthesis
 Specificity of gene amplification
 Applications of PCR
DNA sequencing Dynamic  Chemical characteristics of DNA
 DNA replication and synthesis
 Applications of electrophoresis
Mitosis  Complex chain of events  Chemical characteristics of DNA
 Often represented in an unrealistic way  DNA replication and synthesis
 Students confuse chromosome number  Cell division for growth and replenishment
with the consequences of DNA replica-  Regulation of the cell cycle
tion at metaphase.  Neoplasia
Meiosis  Complex chain of events  Random segregation of genes
 Confused with mitosis  Aneuploidy
 A link may not be established between  Gametogenesis
the process and the consequences (see  Genetic variation
column 3).
the discipline. Threshold concepts have been identified in a of molecular and cellular processes within the cell. These
number of disciplines (e.g., the ideas of entropy in physics new technologies have become increasingly accessible, cost
and of pain in medicine).50 Taylor51 highlights the difficulty effective, and accurate. Developments and innovations
of characterizing true threshold concepts within the within the computer and engineering industries have
biological sciences; rather, the real threshold concept relates increased in response to the increased demand for the
to the complex and idiosyncratic nature of biological instrumentation that supports molecular-biology methodol-
sciences. Biological knowledge is troublesome because of ogies; numerous tools and technologies are now available
the high degree of interrelatedness of biological systems, that investigate cellular processes at a molecular level (see
the large number of processes, the constant need to Table 355–66). Many of the technologies employed are
reevaluate learned material in the light of new material, complex and are operated by experienced personnel
and the numerous layers of abstraction at which material who become specialists in that particular technology.
can be viewed. Other techniques are technically less complex and can be
completed by personnel with less experience. The challenge
Learning in biology usually requires the devel-
for educators will be to decide how these essential
opment of a gradually increasing awareness
technologies should be presented to veterinary students in
and constant re-evaluation of the topics being
a way that allows them to envisage molecular tools and
studied. Truisms taught at early stages are
technologies as something real as well as allowing them to
re-examined as further layers of complexity
appreciate both the potential of the technology and relevant
within the topic require a new set of truths to
applications of such technologies in veterinary medicine.
be constructed.51
Many of the recently developed molecular technologies
These points relate strongly to the field of molecular biology, are conceptually and technically complex (see Table 3).
which, infinitely complex, underpins so many biological Approaches to teaching molecular biology to veterinary
disciplines and can be viewed at numerous levels of students must ensure that the subject is integrated with
abstraction. other veterinary subjects and that focus on the details of
Some aspects of molecular biology require students to apply complex techniques and technologies is kept to a minimum.
alternative metaphors or cognitive paradigms when trying With reference to medical education, Jamieson supports this
to understand the same topic from a different viewpoint. point by stating that
A key example of this phenomenon relates to the under- non clinical teaching staff must accept that
standing of genes. At the highest level of abstraction, genes medical students do not necessarily need to
are best viewed as particles or units (e.g., Mendelian know the depth and detail required by science
inheritance and Hardy/Weinberg equilibrium).52, 53 This students. The emphasis should be on learning of
conception of genes must change, however, when learning underlying principles, paradigms and clinically
about transcription and translation: genes must be viewed relevant examples.67
as having unique properties of their own, composed of
promoter regions, introns/exons, and codons. The student In contrast, life-sciences degree students may require a more
must also be capable of viewing genes as a molecular in-depth and hands-on approach to molecular biology
compound (DNA) if they are to understand its molecular technologies, as these are a key aspect of their vocational
structure and chemical properties and before they can grasp training.
the concepts of technologies such as PCR, DNA sequencing, If veterinary students become too immersed in the complex-
and gel electrophoresis. ity of techniques, they may miss their relevance. Many
In addition, the real-world relevance of molecular biology textbooks illustrate landmark experiments; students often
and its discussion in the public domain may result in some do not distinguish between the conclusion and the methods
students’ having misconceptions about the discipline that used to reach the conclusion, instead learning the material
may not concur fully with the understanding educators in a disconnected manner.68
wish to instill.54 Entwistle and Smith47 suggest that the Technologies such as microarrays and sequencing are
target understanding of genetics courses should build on or technically complex and expensive, have long preparation
challenge ‘‘intuitive genetics’’ (i.e., that which is understood times, and are currently not established as routine veter-
from everyday experience and the media) so that scientific inary diagnostic tools. They are, however, important
meaning extends students’ existing understanding and components within the discipline of molecular biology
relates it to their everyday lives. Efforts are currently and are widely used in different areas of research. It is
underway to validate a series of questionnaires, collectively important that veterinary students know of the existence of
called the Biology Concept Inventory (BCI), aimed at these technologies, understand the underlying principles
accurately assessing students’ comprehension of concepts behind them, and know how and when they can be applied.
in introductory genetics, cell biology, and molecular
Learning about molecular-biological techniques in the
biology.54
veterinary context poses another subtle problem that may
not be appreciated by lecturers familiar with the techniques.
The small volumes, nano-scale concentrations, and manip-
CHALLENGES IN DEVELOPING UNDERSTANDING ulation of what are essentially large molecules can make the
OF MOLECULAR BIOLOGY TECHNIQUES AND subject seem inconceivable to many students. While this
TECHNOLOGIES may not necessarily affect students’ theoretical understand-
Over the last 15 years substantial technological develop- ing, it can affect their appreciation of the technology
ments have taken place, enabling the detailed exploration as something tangible and relevant.

662 JVME 34(5) ß 2007 AAVMC


JVME 34(5) ß 2007 AAVMC

Table 3: Selected techniques employed in molecular biology


Technique Purpose Clinical and Research Applications Complexity Form of Data Generated

Polymerase chain Amplification of specific gene targets Used clinically and in research. This techni- Low Image of an ethidium bromide–stained gel,
reaction (PCR)55 que can be used to detect a specific DNA visualized under UV light
template or to provide starting materials for
further DNA analysis.
Restriction fragment Cutting DNA at specific sequence Used routinely in research; also used Low Image of an ethidium bromide–stained gel,
length polymorphism locations using restriction enzymes clinically. Can be used for rapid detectiong visualized under UV light
analysis of SNPs without DNA sequencing.
Agarose gel Separation of DNA or RNA on the This is one of the few ways that DNA and Low Image of an ethidium bromide–stained gel,
electrophoresis basis of size RNA can be visualized and, so it is used in visualized under UV light
all areas of molecular biology.
Real-time PCR56 Relative and absolute quantification of Applied in research as means of Low–Moderate Image of amplification plot Numerical
specific gene targets originating from exploring functional genomics and clinically data in the form of CT values
RNA or DNA in the determination of viral load, pathogen
detection, and therapy monitoring
Microsatelite analysis Determination of size of DNA Parentage testing Epidemiology Allele Moderate Image of fluorescently tagged PCR product
(GeneScanj) fragment (accurate to 1 base pair determination and co-migrating size marker Numerical
difference) by high-resolution data relating to size
polyacrylamide gel electrophoresis
Pulse field gel Separation of large DNA fragments Bacterial typing Moderate Image of an ethidium bromide–stained gel,
electrophoresis57 visualized under UV lightNumerical data
Bioinformatics58 Broadly encompasses the use of Analysis of sequence data, genome Variable (depends Sequence and numerical data
computational methods in the analysis data, macromolecular structure, and on complexity of
of biological data. The tools used to analyze gene-expression data. data and analysis)
different types of data range from relatively
simple algorithms to more complex
multivariate analysis techniques. Expression
data are analyzed using multivariate
statistical methods and have extended to
mathematical modeling, neural networks,
and genetic algorithms.59

(Continued )
663
664

Table 3: Continued
Technique Purpose Clinical and Research Applications Complexity Form of Data Generated

Automated DNA Determination of DNA sequence Used for research purposes; it can also be High Image of chromatogram
sequencing60 employed in diagnostics (i.e., to detect specific Sequence data
mutations)
Microarrays61 Detection of differentially expressed Generally only used in research at present, but High Image of Array
genes clinical applications are being explored Numerical data
Two-dimensional Separation of proteins based Generally only used in research at present, but High Image of gel
polyacrylamide gel on both iso-electric point and clinical applications are being explored
electrophoresis62 molecular weight
Matrix-assisted Provides accurate analysis of SNP genotyping Microsatellite typing Protein High Spectral data
laser desorption/ the molecular weight and structure characterization
ionisation of bio-molecules
time-of-flight
(MALDI TOF)63
Flow cytometry64 Separation and characterization of cells Has both clinical and research applications High Numerical data
65
Pyrosequencing Typing of SNPs and scoring haplotypes Has both clinical and research applications High Sequence data
RNA Selective disruption of distinct mRNA Defines gene function in vivo by knocking out High Phenotypic analysis of cell or
interference66 transcripts or reducing gene expression at the organism function
post-transcriptional level; has applications
in reverse genetics and therapeutics
JVME 34(5) ß 2007 AAVMC
GENETIC DATABASES AND BIO-INFORMATICS NCBI’s repertoire of databases reflects the increasing
Sequence information is now integrated with related interest in domestic species. The availability of whole
information, including gene and protein structure, con- genome sequences for many different species is, and will
served domains, and publications relevant to that continue to be, an important resource contributing to
sequence.69 Bio-informatics databases such as PubMedk comparative mapping initiatives, informing human medi-
are now the main source of molecular-biological informa- cine, and improving animal health and production.71
tion for graduate students.70
The automation of sequencing means that sequence data are APPROACHES TO TEACHING
being generated at an exponential rate. As of April 2006, It is reasonable to assume that veterinary educators will
there were over 130 billion bases in GenBank and RefSeq.k differ in their approach to basic science education depend-
Databases such as those curated by the National Centre for ing on whether they themselves work predominantly in a
Biotechnology (NCBI) and the European Molecular Biology clinical or in a basic science environment. With regard to
Laboratory (EMBL) provide platforms for placing sequen- medical education, basic scientists may emphasize concep-
cing and genome data in the public domain (see Table 4). tual coherence over clinical application and favor teaching
the basic sciences in more depth;78 it is felt that basic
Over the last decade the major sequencing projects (see scientists do not feel fully confident in deciding the level of
Table 4) have focused primarily on the sequencing of human knowledge medical—or, indeed, veterinary—students need
genomes and those of model organisms such as mouse, at graduation and may not tailor their teaching to
E. coli, Drosophila melanogaster, and zebra fish. Subsequent emphasize clinical relevance, as they may not appreciate it
genome projects have followed in the footsteps of the fully themselves.78 Clinicians who are involved exclusively
Human Genome Project, employing the most successful in clinical practice may understandably feel unconnected
strategies and operating on a fraction of the budget.71 Large- with the basic sciences; hence, despite its relevance to their
scale sequencing projects are now extending their interest to area, they will be less likely to discuss a subject such as
a number of domestic species, including cattle,72 horses,73 molecular biology in great depth. The fact that molecular
pigs,74 cats,75 dogs,76 and sheep.77 The recent addition of biology is a rapidly evolving and growing discipline will
Online Mendelian Inheritance in Animals (OMIA)k to the increasingly exacerbate this problem.

Table 4: Selected sequence and mapping databases, genome centers, and bioinformatics institutes
Online Resource Web Address Comments

National Centre for <www.ncbi.nih.gov> Established in 1988 as a national resource for molecular biology
Biotechnology Information information, NCBI creates public databases, conducts research in
(NCBI) computational biology, and develops software tools for analyzing
genome data.
European Molecular Biology <www.embl.org> EMBL’s mission is to conduct basic research in molecular biology; to
Laboratory (EMBL) provide essential services to scientists in its member states; to
provide high-level training to its staff, students, and visitors; to
develop new instrumentation for biological research; and ensure
technology transfer. These core functions are combined with
significant outreach activities in the areas of science and society
and training for science teachers.
DNA Databank of Japan (DDBJ) <www.ddbj.nig.ac.jp> DDBJ has been functioning as the international nucleotide sequence
database in collaboration with EBI/EMBL and NCBI/GenBank.
The Wellcome Trust Sanger <www.sanger.ac.uk> The Sanger Institute is a genome research institute primarily funded
Institute (WTSI) by the Wellcome Trust. Their purpose is to further the knowledge of
genomes, particularly through large-scale sequencing and analysis.
The Institute for Genomic <www.tigr.org> The Institute for Genomic Research (TIGR) is a non-profit center
Research (TIGR) dedicated to deciphering and analyzing genomes.
(J. Craig Venter Institute)
European Bioinformatics <www.ebi.ac.uk> The EBI is a center for research and services in bioinformatics. The
Institute (EBI) institute manages databases of biological data, including nucleic
acid, protein sequences, and macromolecular structures.
Ensembl <www.ensembl.org> Ensembl is a joint project between EMBL, the EBI, and the WTSI to
develop a software system that produces and maintains automatic
annotation on selected eukaryotic genomes.
Expert Protein Analysis <www.expasy.ch> The ExPASy proteomics server from the Swiss Institute of
System (ExPASy) Bioinformatics (SIB) is dedicated to molecular biology with an
emphasis on data relevant to proteins.

JVME 34(5) ß 2007 AAVMC 665


The involvement that many veterinary clinicians have in Computer-aided learning (CAL) includes a wide range of
research containing a biomedical or molecular component tools, from online resources available in browser format to
may help to bridge the gap between clinical and basic structured tutorials, simulations, animations, and multi-
sciences. This will not only increase awareness and knowl- media presentations.91 CAL can offer students an alternative
edge of molecular biology among clinicians but also insight into areas that may otherwise be difficult to
promote communication between teaching staff in clinical conceptualize.92 CAL can be particularly useful when
and basic sciences. Collaboration through research should applied to molecular biology education. Animations are
give both parties a more holistic view of molecular biology available to illustrate dynamic topics such as PCR93 and
as applied to veterinary medicine, which should have a automated sequencing (see Table 5), in addition to concep-
positive impact on both teaching content and practice. tually difficult areas such as Mendelian inheritance85 and
In the broader context, teaching style is influenced by the population genetics.94 With reference to learning styles,
lecturer’s perception of the students’ needs and prior learners with a strong preference for visual information as
knowledge, organizational support structures, timetabling opposed to verbal information89, 90 may find CAL tools
constraints, and class sizes. Other influences include the helpful (see Figure 4).
lecturer’s own openness to varied teaching methods as well
as his or her personal teaching and work responsibilities.79 16
Academics possessing a more holistic view of their subject
adopt more student-focused approaches to teaching,80 14
including teaching methods such as tutorials, practicals,
12
problem-based learning, and group discussions. Generally

Number of students
these teaching methods are considered superior to more- 10
teaching focused approaches, such as lectures, but require
greater effort and resources.81 8

Problem-based learning (PBL)82, 83 is a student-focused 6


approach that aims to encourage students to construct
4
knowledge and uncover concepts while developing analy-
tical and critical thinking skills through a process of inquiry 2
and reflection. It can be used as a means of integrating 0
molecular biology with clinical scenarios at all stages of the
0 1 2 3 4 5 6 7 8 9 10
curriculum; this approach has the potential to give students
Score (Global)
a more applied view of molecular biology. PBL can also
motivate students to explore the many online resources and
Figure 3: The distribution of scores from first-year
genetic databases outlined above.
veterinary students (N ¼ 75) for the global learning
Electives and intercalated degrees in a molecular or biome- style on the global/sequential dimension, as deter-
dical subject afford veterinary students the opportunity mined by Felder’s Learning Styles questionnaire.l
to explore a career in research or diagnostics. This is Students scoring above 5 show a preference for a
important if students wish to deepen their existing under- global learning style.
standing of molecular biology. Many molecular-biology
experiments take a number of days or weeks to complete.
Electives allow students to appreciate the inconsistencies, 14
quality-control issues, and troubleshooting that take place
continually in research and veterinary practice, as well as 12
gaining a deeper insight into molecular and research
concepts. 10
Number of students

8
ALTERNATIVE REPRESENTATIONS OF MOLECULAR
6
BIOLOGY INFORMATION
In view of the conceptual challenges the discipline of 4
molecular biology presents to students, a significant amount
of literature advocates the use of alternative representations 2
in teaching this subject.84–86 Molecular biology is a broad
0
and interconnected subject, and many processes within it
2 3 4 5 6 7 8 9 10 11
are dynamic. Students need to appreciate the interconnect-
edness of molecular biology as well as knowing all its parts. Score (Visual)
This process takes time and will be more difficult for those
studying at undergraduate level, who have not yet formed a
Figure 4: The distribution of scores from first-year
global picture in their minds. Providing students with visual veterinary students (N ¼ 75) for the Visual learning
overviews and concept maps87, 88 can help them visualize style on the visual/verbal dimension, as determined
the interrelatedness of molecular biological topics; this is by Felder’s Learning Styles questionnaire.l Students
particularly useful for students that have a strongly scoring above 5 show a preference for a visual
Wholist89 or Global90 learning style (see Figure 3). learning style.

666 JVME 34(5) ß 2007 AAVMC


Table 5: A selection of freeware that can be used as educational tools to support education in molecular biology
Source Type of Application Web Address

The Roche Genetics An interactive CD-ROM developed to promote <http://www.roche.com/home/science/


Education Program basic awareness of genetics in the general public sci_gengen/sci_gengen_cdrom.htm>
and to offer an interactive tool for learning basic
principles of genetics
Howard Hughes Medical Fully interactive biomedical laboratory simula- <http://www.hhmi.org/biointeractive/>
Institute (HHMI) tions, animations, lectures, and video clips relat-
ing to different aspects of biomedical science,
medicine, genetics, and evolution
Dolan DNA Learning Animations for automated sequencing, PCR, DNA <http://www.dnalc.org/ddnalc/resources/
Centre restriction, DNA transformation, DNA arrays animations.html>
Davidson College Animation for microarrays <http://www.bio.davidson.edu/Courses/
genomics/chip/chip.html>
Life Sciences Learning Interactive animations for PCR, electrophoresis, <http://lifesciences.envmed.rochester.edu/
Centre genetic disorders, transcription/translation animation.html>
North Harris College Tutorials and multimedia animations in cell <http://science.nhmccd.edu/biol/
biology, genetics, molecular biology, and popula- bio1.html#regulation>
tion genetics

The World Wide Web is a useful educational recourse for DNA-extraction practical. The format of the scrapie prac-
molecular biology and genetics topics.95 Numerous free- tical places less emphasis on technique and more on the
ware applications available online can be used to illustrate application of molecular biology to clinical practice, and
molecular biological techniques, including PCR, sequen- thus could prove much more useful for conducting a
cing, and sub-cellular processes such as transcription and molecular biology–based practical class in the veterinary
translation (see Table 5). The breadth of information that the curriculum.
Web gives students is extensive; this resource should be
used with caution, as it may encourage unproductive
browsing and students may encounter material that is VERTICAL INTEGRATION OF MOLECULAR BIOLOGY IN
beyond the scope of what is being taught at the time.96 THE VETERINARY CURRICULUM
Accreditation bodiesm–o have ensured that cell biology and
molecular biology are included in the veterinary curriculum
PRACTICALS as a core subject at accredited institutions. The content and
Practicals are widely employed in science education to delivery of courses is at the discretion of the institution
illustrate theoretical information.97 Practicals in molecular itself, and molecular biology is mainly taught as a discrete
biology can be difficult to conduct in any setting, as many of module or unit in the pre-clinical curriculum. Considering
the techniques involved are too complex, time consuming, the relevance of molecular biology to so many areas of the
and expensive. Simpler techniques such as DNA extrac- veterinary curriculum, relatively little guidance is given on
tions, PCR, and RFLP can be incorporated into a practical how to effectively incorporate this non-clinical subject into
setting, but, like many molecular biological experiments, the veterinary curriculum.
they may not be very visual and will have long preparation McCrorie100 comments that vertical integration in medical
and incubation times.98 Many veterinary students may find curricula is largely unidirectional. Clinical topics are
the technical aspects of molecular biology uninteresting and integrated in the early years, when the basic sciences are
irrelevant to their vocational training. Allowing students to principally taught, but efforts to reintroduce the basic
interpret simplified outputs (see Table 3) from various sciences later in the curriculum are less actively driven as
technologies based on experiments relevant to veterinary the course content acquires a more clinical slant.
practice is an inexpensive and relatively easy way of Coordinated vertical integration of molecular-biology
familiarizing them with the techniques and allowing them topics is essential if veterinary students are to maximize
to appreciate their relevance. To date, little research has been their understanding of molecular biology and realize its
carried out to confirm the educational merit of practicals; applicability to clinical veterinary medicine. Nickerson101
however, research undertaken on a cohort of veterinary describes ‘‘understanding’’ as an active process that
medical students has illustrated that deep learning correlated connects factual information, relates newly acquired infor-
with high ratings for practical classes as a teaching mation, and weaves this knowledge into a cohesive whole.
method.46 Further to this, two practicals—(1) DNA extrac- This process takes time. Reintroducing molecular biology at
tion and visualization and (2) scrapie genotyping—were relevant points during the curriculum will not only
evaluated98 in the context of students’ approaches to reinforce the information already learned but also give
study.99 The scrapie-genotyping practical was rated higher veterinary students time to reflect on previously studied
and had a stronger association with deep learning than the material while reevaluating it in the context of greater

JVME 34(5) ß 2007 AAVMC 667


clinical understanding. It is anticipated that this approach genes on a chip-based format.113 The ongoing identification
will deepen understanding, improve retention, and directly of QTLs and SNPs relating to animal health will become
highlight the relevance of molecular biology to significant drivers for selective breeding in the future.38
veterinarians. Veterinarians will play a vital role in using this new
information to advise breeders on animal welfare and
Gamulin, writing about medical education, supports this
health in production animals. It is reasonable to postulate
argument:
that by 2025 the bio-informatics revolution may even have
The molecular medicine topics should be endowed veterinarians with tools that predict the final
included into various subjects of undergraduate phenotypic outcome based on large quantities of genotype
curricula and vertically integrated rather than information, predicting the conformation and biological
treated as a separate subject in preclinical or characteristics of a ‘‘virtual animal.’’ Breeding for health and
clinical courses.102 welfare will be the ultimate step in preventive medicine.
Koens et al. make an important point in favor of teaching Veterinarians have important roles to play in comparative
basic sciences in the clinical context: medicine and biomedical research.105 Veterinarians and
Clinical context increases commitment to learn those researching animal diseases will also play a vital
biomedical knowledge because it increases the role in informing human medicine, especially on issues
willingness to invest more effort in the learning relating to animal models, comparative genomics, and
task.103 pharmacogenetics.

Achieving vertical integration of a non-clinical subject such Veterinarians’ current role in food safety and the traceability
as molecular biology demands a coordinated effort from of zoonotic disease will be better facilitated by tools that
teaching staff in relevant disciplines in all years. To permit the rapid identification and characterization of
determine how and at what level molecular biology learning pathogens ‘‘in the field.’’ Rapid surveillance will be vital if
materials should be incorporated at each stage of the antibiotic use is to be reduced.114
curriculum requires coordination and communication Transgenic technologies, which have the potential to confer
between educators in the pre-clinical and clinical years. new characteristics on existing species and even result in the
This goal can be achieved only through a conscious creation of new species or breeds, may increase as a result of
commitment. consumer demand. Veterinarians will play a role in advising
on the health and welfare of these animals as well as
treating them. Cloning technologies, if improved, could
HOW MIGHT THE VETERINARIAN OF 2025 BE enable the conservation of rare,115 commercially valuable,24
APPLYING MOLECULAR BIOLOGY? or even extinct or endangered species,115 a scenario that will
The veterinarians of 2025 could have a new range of also generate unique clinical challenges for veterinarians.
molecular-based diagnostic tools at their disposal and, as a New areas of understanding (e.g., small interfering RNA,66
profession, may have new roles beyond the ones they copy-number variation,116 and epigenetic variation117) are
currently play. However, concerns exist about the small continually being uncovered in the field of molecular
numbers of veterinarians attracted to biomedical biology. This new understanding may also significantly
research,104, 105 as well as about the lack of non-clinical affect diagnostics, treatment, and breeding strategies in
veterinary role models teaching in current veterinary some as yet unforeseen way.
program.3 The profession’s ability to capitalize fully on
these new technologies will depend as much on how the
profession broadens its own horizons as on the existence of
SUMMARY
the technologies in the wider arena.
Molecular biology is increasingly affecting veterinary
If veterinary diagnostics follows current trends in medical medicine, a trend that needs to be reflected on the veterinary
diagnostics, techniques such as PCR and real-time PCR will curriculum. Effective incorporation of a non-clinical subject
increasingly be employed as routine methods of pathogen such as molecular biology onto the veterinary curriculum
detection,106, 107 SNP analysis,108 cancer diagnostics,109 and needs to be addressed in a way that maximizes under-
quantification of viral load.110 This technology will improve standing of the discipline without overloading students
the speed of both diagnosis and treatment. Microarray with unnecessary detail. The central concepts within
technologies will not only help to characterize disease states molecular biology and its applications to veterinary med-
but also provide information on the optimal course of icine should be emphasized. Effective vertical integration
treatment.111 The applications of molecular-based imaging will facilitate the teaching of molecular biology topics in the
techniques will also evolve in the clinical arena, giving the light of students’ understanding at the time, building on
veterinarian a greater depth of clinical information beyond and reinforcing their existing knowledge and illustrating its
that currently available.112 applications in context. Effective communication among
those contributing to molecular-biology education at differ-
One area that should significantly affect veterinarians will
ent stages of the curriculum is essential if a global learning
be the availability of data relating to the genome, proteome,
outcome is to be defined and implemented.
and transcriptome of most species. By 2025, these data
should be better annotated, and clearer phenotypic relation- Educators also need to be sympathetic to the learning
ships established. Veterinarians may increasingly need to challenges that molecular biology presents to veterinary
advise clients on genetic screening tests, which, in the students. In light of these particular challenges, a wide
future, could possibly involve the screening of multiple range of teaching methods and alternative representations

668 JVME 34(5) ß 2007 AAVMC


should be employed to promote understanding of molecular <http://www.vgl.ucdavis.edu/service/parentage/
biology. These learning challenges relate not only to the process.html>.
subject itself but to wider issues such as the necessary depth
i Bovine QTL Viewer, Texas A&M University
and breadth of material required by veterinary students.
<http://bovineqtl.tamu.edu>.
Haga95 clarifies this point further, stating that the goal of
enhancing basic genetics education is to provide a founda- j Applied BioSystems, Foster City, CA 94404
tion of knowledge that will allow learners to understand <http://www.appliedbiosystems.com>.
genetics concepts, applications, and ethical issues and k National Centre for Biotechnology Information
become informed users of genetics technologies and their (NCBI), Bethesda, MD 20894
applications. <http://www.ncbi.nih.gov>.
We believe that the core competencies that veterinary l Felder R, Index of Learning Styles (ILS)
students should obtain in molecular biology include the <http://www2.ncsu.edu/unity/lockers/users/f/
following: felder/public/ILSpage.html>. Accessed 10/15/07.
. an understanding of the background facts and m American Veterinary Medical Association,
concepts of molecular biology Schaumburg, IL 60173-4360 <http://www.avma.org>.
. knowledge of terminology and language specific to n European Association of Establishments for Veterinary
the subject Education <http://www.eaeve.org>.
. knowledge of the technology and techniques used to o Australasian Veterinary Boards Council
study and investigate molecular processes <http://www.avbc.asn.au/>.
. an understanding of the potential applications of
molecular biology to veterinary medicine
REFERENCES
. an understanding of the wider implications of 1 Astbury WT. Molecular biology or ultrastructural
employing molecular-based technologies in veterin- biology. Nature 190: 1124, 1961 p1124.
ary medicine and to society
2 Leighton FA. Veterinary medicine and the lifeboat test:
The means by which these learning goals are achieved a perspective on the social relevance of the veterinary
within any individual veterinary institution is still a matter profession in the twenty-first century. J Vet Med Educ 31:
of choice. Clear learning objectives and varied teaching 329–333, 2004.
strategies combined with good communication between
educators and coordinated vertical integration should 3 Barthold SW. Musings of a Connecticut Yankee in King
encourage the production of a comprehensive, clinically Arthur’s court: antemortem analysis of the veterinary
relevant, and constructively aligned molecular biology profession. J Vet Med Educ 32: 306–313, 2005.
course that spans the entire veterinary curriculum. 4 Zemljic B. Standards of veterinary education for
the future: preparing the profession for the new century.
J Vet Med Educ 31: 13–14, 2004. p13.
ACKNOWLEDGMENTS
We would like to thank Professors Boyd Jones and 5 McGeady TA, Quinn PJ, Fitzpatrick EP, Ryan MT.
P.J. Quinn for their extremely helpful editorial comments Veterinary Embryology. Oxford: Blackwell, 2006.
during the writing of this paper. 6 Anderson L, Georges M. Domestic-animal
genomics: deciphering the genetics of complex traits.
Nat Rev Genet 5: 202–212, 2004.
NOTES 7 Bannasch DL, Hughes AM. Recent advances in small
a Animal DNA Services, Fairfield, OH 45014 animal genetics. Vet Clin North Am Small Anim Pract 36:
<http://www.vetdnacenter.com/>. 461–474, 2006.
b Optigen, Ithaca, NY 14850 <www.optigen.com>. 8 Ostrander EA, Gailbert F, Patterson DF. Canine genetics
comes of age. Trends Genet 16(3):117–124, 2000.
c VetGen, Ann Arbor, MI 48108
<http://www.vetgen.com/>. 9 Helps CR, Tasker S, Barr FJ, Wills SJ, Lyons LA,
Gruffydd-Jones TJ. Detection of Feline autosomal-dominant
d VITA-TECH, Markham, ON L3R 5V2
polycystic kidney disease (AD-PKD) by real-time PCR
<http://www.vita-tech.com/canineparentage.cfm>.
genotyping. J Vet Intern Med 19: 949, 2005.
e HealthGene, Toronto, ON M6M 3Z4
10 Perryman LE, Torbeck RE. Combined immunodefi-
<www.healthgene.com/vet>.
ciency of Arabian horses: confirmation of autosomal
f QIAGEN Pathogen real-time PCR detection kits, recessive mode of inheritance. J Am Vet Med Assoc 76:
QIAGEN <http://www1.qiagen.com/products/ 1250–1251, 1980.
assays/pathogendetection/?r¼777>.
11 Nagahata H. Bovine leukocyte adhesion deficiency
g QIAGEN <http://www1.qiagen.com/>. (BLAD): a review. J Vet Med Sci 66: 1475–1482, 2004.
h Parentage Testing service, Veterinary Genetics 12 Fujii J, Otsu K, Zorzato F, De Leon S, Khanna VK,
Laboratory, UC Davis, CA 95616 Weiler JE, O’Brien PJ, MacLennan DH. Identification of a

JVME 34(5) ß 2007 AAVMC 669


mutation in porcine ryanodine receptor associated with 28 Shah R. Pharmacogenetics in drug regulation: promise,
malignant hyperthermia. Science 253: 448–451, 1991. potential and pitfalls. Phil Trans R Soc B 360: 1617–1638,
2005.
13 Beever JE, Smit MA, Meyers SN, Hadfield TS,
Bottema C, Albretsen J, Cockett NE. A single-base change 29 Yong WP, Innocenti F, Ratain MJ. The role of
in the tyrosine kinase II domain of ovine FGFR3 causes pharmacogenetics in cancer therapeutics. Brit J Clin
hereditary chondrodysplasia in sheep. Anim Genet 37: 66–71, Pharmacol 62: 35–46, 2006.
2006.
30 Nnadi CU, Goldberg JF, Malhotra AK.
14 Gunson RN, Collins TC, Carman WF. Practical experi- Pharmacogenetics in mood disorder. Curr Opin Psychiatr 18:
ence of high throughput real time PCR in the routine 33–39, 2005.
diagnostic virology setting. J Clin Virol 35: 355–367, 2006.
31 Chasman DI, Posada D, Subrahmanyan L, Cook NR,
15 Forbes BA. Introducing a molecular test into the clinical Stanton VP Jr, Ridker PM. Pharmacogenetic study of
microbiology laboratory. Arch Pathol Lab Med 127: 1106–1111, statin therapy and cholesterol reduction. J Am Med Assoc
2003. 291: 2821–2827, 2004.
16 Bowling AT, Eggleston-Stott ML, Byrns G, Clark RS, 32 Massart F. Human races and pharmacogenomics of
Dileanis S, Wictum E. Validation of microsatellite markers effective bone treatments. Gynecol Endocrinol 20(1):36–44,
for routine horse parentage testing. Anim Genet 28: 247–252, 2005.
1997.
33 Nicholas FW. Introduction to Veterinary Genetics. Oxford:
17 Binns MM, Holmes NG, Marti E, Bowen N. Dog Oxford University Press, 1996.
parentage testing using canine microsatellites. J Small Anim
34 Thomsen H, Lee HK, Rothschild MF, Malek M,
Pract 36: 493–497, 1995.
Dekkers JCM. Characterization of quantitative trait loci for
18 Menotti-Raymond M, David VA, Lyons LA, growth and meat quality in a cross between commercial
Schaffer AA, Tomlin JF, Hutton MK, O’Brien SJ. A genetic breeds of swine. J Anim Sci 82: 2213–2228, 2004.
linkage map of microsatellites in the domestic cat.
35 Purvis IW, Franklin IR. Major genes and QTL
(Felis catus). Genomics 57: 9–23, 1999.
influencing wool production and quality: a review. Genet Sel
19 Pfaller MA. Molecular approaches to diagnosing and Evol 37: S97–S107, 2005.
managing infectious diseases: practicality and costs.
36 Kucerova J, Lund MS, Sorensen P, Sahana G,
Emerging Infect Dis 7: 312–318, 2001.
Guldbrandtsen B, Nielsen VH, Thomsen B, Bendixen C.
20 O’Mahony R, Abbott Y, Leonard FC, Markey BK, Multitrait quantitative trait loci mapping for milk
Quinn PJ, Pollock PJ, Fanning S, Rossney AS. production traits in Danish Holstein cattle. J Dairy Sci 89:
Methicillin-resistant Staphylococcus aureus (MRSA) isolated 2245–2256, 2006.
from animals and veterinary personnel in Ireland.
37 Chowdhary BP, Bailey E. Equine genomics:
Vet Microbiol 109: 285–296, 2005.
galloping to new frontiers. Cytogenet Genome Res 102:
21 Rivera B, Ahrar K, Kangasniemi MM, Hazle JD, 184–188, 2003.
Price RE. Canine transmissible venereal tumor: a
38 Hiendleder S, Thomsen H, Reinsch N, Bennewitz J,
large-animal transplantable tumor model. Comp Med 55:
Leyhe-Horn B, Looft C, Xu N, Medjugorac I, Russ I, Kühn C,
335–343, 2005.
Brockmann GA, Blümel J, Brenig B, Reinhardt F, Reents R,
22 Olsen AK, Keiding S, Munk OL. Effect of hypercapnia Averdunk G, Schwerin M, Förster M, Kalm E, Erhardt G.
on cerebral blood flow and blood volume in pigs studied by Mapping of QTL for body conformation and behavior in
positron emission tomography. Comp Med 56: 416–420, 2006. cattle. J Hered 94: 496–506, 2003.
23 Tiefenbacher S, Fahey MA, Rowlett JK, Meyer JS, 39 Rupp R, Boichard D. Genetics of genetics to mastitis in
Pouliot AL, Jones BM, Novak MA. The efficacy of diazepam diary cattle. Vet Res 34: 671–688, 2003.
treatment for the management of acute
40 Raadsma HW, Gray GD, Woolaston RR. Breeding for
wounding episodes in captive rhesus macaques. Comp Med
disease resistance in Merino sheep in Australia. Rev Sci Tech
55: 387–392, 2005.
17: 315–328, 1998.
24 Oback B, Wells DN. Cloning cattle: the methods in the
41 Hiendleder S, Bauersachs S, Boulesteix A, Blum H,
madness. Adv Exp Med Biol 591: 30–57, 2007.
Arnold GJ, Fröhlich T, Wolf E. Functional genomics: tools for
25 Rhind SM, Taylor JE, De Sousa PA, King TJ, improving farm animal health and welfare. Rev Sci Tech Off
McGarry M, Wilmut I. Human cloning: can it be made safe? Int Epiz 24: 354–377, 2005.
Nat Rev Genet4: 855–864, .
42 Polineni P, Aragonda P, Suresh XR, Furuta R,
26 Vajta G, Gjerris M. Science and technology of Adelson DL. The Bovine QTL viewer: a Web accessible
farm cloning: state of the art. Anim Reprod Sci 92: 211–230, database of bovine quantitative trait loci. BMC Bioinformatics
2006. 7: 283, 2006.
27 Bucher P, Morel P, Bühler LH. Xenotransplantation: an 43 Windsor AJ, Mitchell-Olds T. Comparative genomics
update on recent progress and future perspectives. Transpl as a tool for gene discovery. Curr Opin Biotechnol 17: 161–167,
Int 18: 894–901, 2005. 2006.

670 JVME 34(5) ß 2007 AAVMC


44 Miller W, Makova KD, Nekrutenko A, Hardison RC. 60 Smith LM, Sanders JZ, Kaiser RJ, Hughes P, Dodd C,
Comparative genomics. Ann Rev Genomics Hum Genet 5: Connell CR, Heiner C, Kent SBH, Hood LE. Fluorescence
15–56, 2004. detection in automated DNA sequence analysis. Nature 321:
674–679, 1986.
45 Kember D, Ng s, Tse H, Wong ETT. An examination of
the interrelationships between workload, study time, 61 Fathallah-Shaykh HM. Microarrays: applications and
learning approaches and academic outcomes. Stud High pitfalls. Arch Neurol 62: 1669–1672, 2005.
Educ 21: 347–358, 1996.
62 O’Farrell PH. High resolution two-dimensional
46 Ryan MT, Irwin JA, Bannon, FJ, Mulholland CW, electrophoresis of proteins. J Biol Chem 250: 4007–4021, 1975.
Baird AW. Observations of veterinary medicine students’
63 Karas M, Bachman D, Hillenkamp F. Influence of the
approaches to study in preclinical years. J Vet Med Educ 31:
wavelength in high-radiance ultraviolet laser desorption
242–254, 2004.
mass spectrometry of organic molecules. Anal Chem 57:
47 Entwistle N, Smith C. Personal understanding and 2935–2939, 1985.
target understanding: mapping influences on the outcomes
64 Giaretti W. Flow cytometry and applications in
of learning. Brit J Educ Psychol 72: 321–342, 2002.
oncology. J Clin Pathol 50: 275–277, 1997.
48 Perkins D. The many faces of constructivism.
65 Diggle MA, Clarke SC. Pyrosequencing: sequence
Educ Leadership 57(3):28–35, 1999.
typing at the speed of light. Mol Biotechnol 28(2):129–137,
49 Meyer JHF, Land R. Threshold concepts and 2004.
troublesome knowledge (2): epistemological considerations
66 Lenz G. The RNA interference revolution. Braz J Med
and a conceptual framework for teaching and learning.
Biol Res 38: 1749–1757, 2005.
High Educ 49: 373–388, 2005.
67 Jamieson S. Cell and molecular biology in the medical
50 Meyer J, Land R. Threshold Concepts and Troublesome
curriculum. Med Educ 35: 85–86, 2001 p86.
Knowledge: Linkages to Ways of Thinking and Practising
within the Disciplines. Occasional Report 4, Enhancing 68 Pukklia PJ. Genetics education: innovations in
Teaching–Learning Environments in Undergraduate teaching and learning genetics—introducing student
Courses, May 2003. inquiry in large introductory genetic classes. Genetics 166:
11–18, 2004.
51 Taylor CE. Threshold concepts in biology—do they fit
the definition?. In Meyer J, Land R, eds. Overcoming 69 Benson DA, Karsch-Mizrachi I, Lipman DJ, Ostell J,
Barriers to Student Understanding: Threshold Wheeler DL. GenBank: update. Nucleic Acids Res 32:
Concepts and Troublesome Knowledge. London: Routledge, D23–D26, 2004.
2006:87–99 p89. 70 Brown C. Where do molecular graduate students find
52 Venville GJ, Treagust DF. Exploring conceptual information? Sci Tech Libr 25(3):89–104, 2005.
change in genetics using a mulitdimensional interpretive 71 Womack JE. Advances in livestock genomics: opening
framework. J Res Sci Teach 35: 1031–1055, 1998. the barn door. Genome Res 15: 1699–1705, 2005.
53 Dupré J. Understanding contemporary genomics. 72 Everts-van der Wind A, Larkin DM, Green CA,
Perspectives on Science 12: 320–338, 2004. Elliott JS, Olmstead CA, Chiu R, Schein JE, Marra MA,
54 Klymkowsky MW, Garvin-Doxas K, Zeilik M. Womack JE, Lewin HA. A high-resolution whole-genome
Bioliteracy and teaching efficacy: what biologists can learn cattle–human comparative map reveals details of mamma-
from physicists. Cell Biol Educ 2: 155–161, 2003. lian chromosome evolution. Proc Natl Acad Sci 102: 18526–
18531, 2005.
55 Mullis K, Faloona F, Scharf S, Saiki R, Horn G, Erlich H.
Specific enzymatic amplification of DNA in vitro: the 73 Chowdhary BP, Raudsepp T, Kata SR, Goh G,
polymerase chain reaction. Cold Spring Harb Symp Quant Biol Millon LV, Allan V, Piumi F, Guerin G, Swinburne J,
51(1):263–273, 1986. Binns M, Lear TL, Mickelson J, Murray J, Antczak DF,
Womack JE, Skow LC. The first-generation whole-genome
56 Gibson UE, Heid CA, Williams PM. A novel method
radiation hybrid map in the horse identifies conserved
for real time quantitative RT-PCR. Genome Res 6: 995–1001,
segments in human and mouse genomes. Genome Res 13:
1996.
742–751, 2003.
57 Olson MV. Separation of large DNA molecules by
74 Wernersson R, Schierup MH, Jorgensen FG, Gorodkin J,
pulsed-field gel electrophoresis: a review of the basic
Panitz F, Staerfeldt HH, Christensen OF, Mailund T,
phenomenology. J Chromatogr 470: 377–383, 1989.
Hornshoj H, Klein A, Wang J, Liu B, Hu S, Dong W, Li W,
58 Luscombe NM, Greenbaum D, Gerstein M. What is Wong GK, Yu J, Wang J, Bendixen C, Fredholm M, Brunak S,
bioinformatics? A proposed definition and overview of the Yang H, Bolund L. Pigs in sequence space: a 0.66X coverage
field. Method Inform Med 4: 346–358, 2001. pig genome survey based on shotgun sequencing. BMC
Genomics 6(1):70, 2005.
59 Kapetanovic IM, Rosenfeld S, Izmirlian G.
Overview of commonly used bioinformatics 75 Menotti-Raymond M, David VA, Agarwala R,
methods and their applications. Ann NY Acad Sci 1020: 10– Schäffer AA, Stephens R, O’Brien SJ, Murphy WJ.
21, 2004. Radiation hybrid mapping of 304 novel microsatellites in

JVME 34(5) ß 2007 AAVMC 671


the domestic cat genome. Cytogenet Genome Res 102: 272–276, 87 Novek JD. The promise of new ideas and new
2003. technology for improving teaching and learning. Cell Biol
Educ 2: 122–132, 2003.
76 Lindblad-Toh K, Wade CM, Mikkelsen TS, Karlsson EK,
Jaffe DB, Kamal M, Clamp M, Chang JL, Kulbokas EJ, 88 Pinto AJ, Zeist HJ. Concept mapping: a strategy for
Zody MC, Mauceli E, Xie X, Breen M, Wayne RK, promoting meaningful learning in medical education. Med
Ostrander EA, Ponting CP, Galibert F, Smith DR, deJong PJ, Teach 19(2):114–121, 1997.
Kirkness E, Alvarez P, Biagi T, Brockman W, Butler J,
89 Riding R, Rayner SG. Cognitive Styles and Learning
Chin C, Cook A, Cuff J, Daly MJ, DeCaprio D, Gnerre S,
Strategies. London: David Fulton, 1998.
Grabherr M, Kellis M, Kleber M, Bardeleben C, Goodstadt L,
Heger A, Hitte C, Kim L, Koepfli K, Parker HG, Pollinger JP, 90 Felder RM. Matters of style. ASEE Prism 6(4):18–23,
Searle SMJ, Sutter NB, Thomas R, Webber C, Lander ES. 1996.
Genome sequence, comparative analysis and haplotype
91 Sweeters W. Multimedia electronic tools for learning.
structure of the domestic dog. Nature 438: 803–819, 2005.
Educ Technol 34: 47–52, 1994.
77 Maddox JF, Davies KP, Crawford AM, Hulme DJ,
92 Ryan M, Mulholland CW, Gilmore WS. Applications
Vaiman, D, Cribiu EP, Freking BA, Beh KJ, Cockett NE,
of computer aided learning in biomedical sciences:
Kang N, Riffkin CD, Drinkwater R, Moore SS, Dodds KG,
considerations in design and evaluation. Brit J Biomed Sci 57:
Lumsden JM, van Stijn TC, Phua SH, Adelson DL, Burkin,
28–34, 2000.
HR, Broom JE, Buitkamp J, Cambridge L, Cushwa WT,
Gerard E, Galloway, SM, Harrison B, Hawken RJ, 93 Jenkins RO, Scott NW. DNA to PCR: computer-aided
Hiendleder S, Henry HM, Medrano JF, Paterson KA, learning software for DNA sequencing and
Schibler L, Stone RT, van Hest B. An enhanced linkage map amplification techniques of molecular biology. Biochem Educ
of the sheep genome comprising more than 1000 loci. 27: 166–168, 1999.
Genome Res 11: 1275–1289, 2001.
94 Peng B, Kimmel M. simuPOP: a forward-time
78 Koens F, Custers EJFM, Ten Cate OTJ. Clinical and basic population genetics simulation environment. Bioinformatics
science teachers’ opinions about the required depth of 21: 3686–3687, 2005.
biomedical knowledge for medical students. Med Teach 28:
95 Haga SB. Teaching resources for genetics. Nat Rev Genet
234–238, 2006.
7: 223–229, 2006.
79 Center for Science, Mathematics, and Engineering
96 Smith JB, Weiss SF. Hypertext. Comm Assoc Comp Mach
Education. Science Teaching Reconsidered: A Handbook
31: 816–819, 1988.
<http://www.nap.edu/catalog.php?record_id¼5287#toc>.
Accessed 10/15/07. Washington, DC: National Academies 97 Boud D, Dunn J, Hegarty-Hazel E. Teaching in
Press, 1997. Laboratories. Guildford, UK: Society for Research into Higher
Education and NFER-NELSON, 1986.
80 Prosser M, Martin E, Trigwell K, Ramsden P,
Luekenhausen G. Academics’ experiences of understanding 98 Ryan MT, Sweeney T. Genetic susceptibility to scrapie
of their subject matter and the relationship of this to their in sheep: a clinically relevant theme in veterinary medical
experiences of teaching and learning. Instruct Sci 33: education. J Vet Med Educ 32: 544–550, 2005.
137–157, 2005.
99 Entwistle N, Tait H, McCune V. Patterns of response to
81 O’Neill G, McMahon T. Student-centred learning: what approaches to studying inventory across contrasting groups
does it mean for students and lecturers? <http:// and contexts. Eur J Psychol Educ 15: 33–48, 2000.
www.aishe.org/readings/2005-1/oneill-mcmahon-
100 McCrorie P. The place of the basic sciences on the
Tues_19th_Oct_SCL.pdf>. Accessed 10/15/07. In O’Neill G,
medical curricula. Med Educ 34: 594–595, 2000.
Moore S, McMullin B, eds. Emerging Issues in the Practice of
University Learning and Teaching. Dublin: All Ireland Society 101 Nickerson RS. Understanding understanding.
for Higher Education (AISHE), 2005. Am J Educ 93: 201–239, 1985.
82 Barrows HS. Problem-based learning in medicine and 102 Gamulin S. Impact of molecular medicine on patho-
beyond: a brief overview. New Dir Teach Learn 68: 3–11, 1996. physiology, medical practice, and medical education. Croat
Med J 44: 374–385, 2003 p382.
83 Newman MJ. Problem based learning: an introduction
and overview of the key features of the approach. J Vet Med 103 Koens F, Mann KV, Custers EJFM, Ten Cate OTJT.
Educ 32: 12–20, 2005. Analysing the concept of context in medical education. Med
Educ 39: 1243–1249, 2005 p1245.
84 Arwood L. Teaching cell biology to nonscience majors
through forensics, or how to design a killer course. Cell Biol 104 Halliwell R. Whither veterinary education—have we
Educ 3: 131–138, 2004. lost our direction? J Vet Med Educ 33: 309–316, 2006.
85 Tsui C, Treagust DF. Genetics reasoning with multiple 105 Fox JG, Obernier J. Veterinarians in biomedical
external representations. Res Sci Educ 33: 111–135, 2003. research: a perilous future? J Vet Med Educ 32: 301–305, 2005.
86 Venville GJ, Treagust, DF. Exploring conceptual 106 Kaltenboeck B, Wang C. Advances in real-time PCR:
change in genetics using a multidimensional interpretive application to clinical laboratory diagnostics. Adv Clin Chem
framework. J Res Sci Teach 35: 1031–1105, 1998. 40: 219–259, 2005.

672 JVME 34(5) ß 2007 AAVMC


107 Schmitt B, Henderson L. Diagnostic tools for animal Cho EK, Dallaire S, Freeman JL, Gonzalez JR, Gratacos M,
diseases. Rev Sci Tech 24: 243–250, 2005. Huang J, Kalaitzopoulos D, Komura D, MacDonald JR,
Marshall CR, Mei R, Montgomery L, Nishimura K,
108 Gibson NJ. The use of real-time PCR methods in
Okamura K, Shen F, Somerville MJ, Tchinda J, Valsesia A,
DNA sequence variation analysis. Clin Chim Acta 363: 32–47,
Woodwark C, Yang F, Zhang J, Zerjal T, Zhang J,
2006.
Armengol L, Conrad DF, Estivill X, Tyler-Smith C,
109 Winegarden N. Microarrays in cancer: moving from Carter NP, Aburatani H, Lee C, Jones KW, Scherer SW,
hype to clinical reality. Lancet 362: 1428, 2003. Hurles ME. Global variation in copy number in the human
genome. Nature 444: 444–454, 2006.
110 Piiparinen H, Höckerstedt K, Lappalainen M, Suni J,
Lautenschlager I. Monitoring of viral load by quantitative 117 Morgan HD, Santos F, Green K, Dean W, Reik W.
plasma PCR during active cytomegalovirus infection of Epigenetic reprogramming in mammals. Hum Mol Genet
individual liver transplant patients. J Clin Microbiol 40: 14(Suppl 1):R47–R58, 2005.
2945–2952, 2002.
111 Statnikov A, Aliferis CF, Tsamardinos I, Hardin D,
Levy S. A comprehensive evaluation of multicategory
AUTHOR INFORMATION
classification methods for microarray gene expression Marion T Ryan, MSc, D.Med.Sci., is a Senior Technical Officer
cancer diagnosis. Bioinformatics 21: 631–643, 2005. in the Molecular Biology Facility, College of Life Sciences,
School of Agriculture, Food Science and Veterinary Medicine,
112 Jaffer FA, Weissleder R. molecular imaging in the
University College Dublin, Belfield, Dublin 4, Republic of
clinical arena. J Am Med Assoc 293: 855–862, 2005.
Ireland. E-mail: marion.ryan@ucd.ie. She has published in a
113 Sauer S, Lange BM, Gobom J, Nyarsik L, Seitz H, number of areas including distance education, computer-aided
Lehrach H. Miniaturization in functional genomics and learning, learning approaches, veterinary embryology, and
proteomics. Nat Rev Genet 6: 465–476, 2005. molecular genetics.
114 Ungemach FR, Muller-Bahrdt D, Abraham G. Torres Sweeney, PhD, is a Senior Lecturer in Cell and
Guidelines for prudent use of antimicrobials and their Molecular Biology in the College of Life Sciences, School of
implications on antibiotic usage in veterinary medicine. Agriculture, Food Science and Veterinary Medicine and
Int J Med Microbiol 41: 33–38, 2006. Conway Institute of Biomolecular and Biomedical Research,
115 Lanza RP, Cibelli JB, Diaz F, Moraes CT, Farin PW, University College Dublin, Belfield, Dublin 4, Republic of
Farin CE, Hammer CJ, West MD, Damiani P. Cloning of an Ireland. E-mail: torres.sweeney@ucd.ie. Her major research
endangered species (Bos gaurus) using interspecies nuclear interests are: genetic polymorphisms in commercially impor-
transfer. Cloning 2(2):79–90, 2000. tant traits in bovine and sheep, gene expression profiling of
meat-quality traits, and immunogenomics of disease resis-
116 Redon R, Ishikawa S, Fitch KR, Feuk L, Perry GH, tance in livestock animals.
Andrews TD, Fiegler H, Shapero MH, Carson AR, Chen W,

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