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    Mirrored from http://bioinformatics.org/faq/  thanks to the open publication licence adopted by Damian Counsell for this document

    Bioinformatics Frequently Asked Questions

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    Latest changes

    • added Georgia State University's courses---thanks to Eric VanWieren
    • added FH Weihenstephan in Freising---thanks to Tobias Kailich
    • added Johns Hopkins' courses---thanks to Tim Young
    • added three new bioinformatics courses from Germany---thanks to Sebastian Kurscheid
    • added courses at University of Illinois---thanks to Amit Sabnis

    $Revision: 1.207 $ $Date: 2005/04/05 13:06:07 $

    Introduction

    Mail your questions to me, Damian Counsell, and I'll try to bring you answers. Alternatively, if you have your own answers, mail them to me and I'll incorporate them. The practical section in particular is full of gaps so your contributions to that are particularly welcome; I am slowly completing and extending the entries when I have the time.

    Although I am happy to tackle questions of general interest to all visitors to the site, please note that:

    • I cannot answer queries specific to you alone,
    • I am not a careers adviser,
    • I try not to offer opinions on the relative merits of bioinformatics courses,
    • I won't answer your essay questions, assignments, or homework,
    • I won't provide you with a list of companies for you to market your bioinformatics product to,
    • I won't suggest a project for your Master's/PhD,
    • I have not devised a bioinformatic cure for cancer---and neither have you, and
    • This FAQ is perpetually under construction.
    I hope, however, that the information here helps with your studies, career and work.

    I acknowledge the help of many other individuals in creating this part of the Bioinformatics.Org site. If you have contributed and I have forgotten to credit you, please email me and I will correct my oversight immediately.

    Bioinformatics is, I believe, a special kind of engineering discipline---it certainly isn't a "pure" science. It has been enormously successful in its short existence and I think its successes have been the result of a practical and rigorous approach which I hope to encourage in anyone interested in entering the field.

    This document is not a scientific paper or textbook (yet). You will find blunt opinions here. If you disagree with me about any of the following please tell me. I hope to learn a lot from your inevitable and welcome criticisms.

    There is certainly one sense in which I consider myself a pure scientist: I'm open to rational persuasion.

    I write this resource and hold the copyright for the purposes of protecting its content from intellectual property pirates. By that I mean I want to keep this out of the hands of people who steal the work of others for commercial gain, and those who abuse and extend the powers of IP law at the expense of the disadvantaged---rather than those who would like to copy or mirror this resource for educational reasons. (This may sound overdramatic, but the FAQ has already been pirated for doubtful purposes.)

    Overview

    Contents

    Definitions: What is Bioinformatics?

    Definition of Bioinformatics: What is bioinformatics?

    Roughly, bioinformatics describes any use of computers to handle biological information.

    In practice, the definition used by most people is narrower; bioinformatics to them is a synonym for "computational molecular biology"---the use of computers to characterize the molecular components of living things.

    What is Bioinformatics?---The Tight Definition

    "Classical" bioinformatics

    Most biologists talk about "doing bioinformatics" when they use computers to store, retrieve, analyze or predict the composition or the structure of biomolecules. As computers become more powerful you could probably add simulate to this list of bioinformatics verbs. "Biomolecules" include your genetic material---nucleic acids---and the products of your genes: proteins. These are the concerns of "classical" bioinformatics, dealing primarily with sequence analysis.

    Fredj Tekaia at the Institut Pasteur offers this definition of bioinformatics:

    "The mathematical, statistical and computing methods that aim to solve biological problems using DNA and amino acid sequences and related information."

    It is a mathematically interesting property of most large biological molecules that they are polymers; ordered chains of simpler molecular modules called monomers. Think of the monomers as beads or building blocks which, despite having different colours and shapes, all have the same thickness and the same way of connecting to one another.

    Monomers that can combine in a chain are of the same general class, but each kind of monomer in that class has its own well-defined set of characteristics.

    Many monomer molecules can be joined together to form a single, far larger, macromolecule. Macromolecules can have exquisitely specific informational content and/or chemical properties.

    According to this scheme, the monomers in a given macromolecule of DNA or protein can be treated computationally as letters of an alphabet, put together in pre-programmed arrangements to carry messages or do work in a cell.

    "New" bioinformatics

    The greatest achievement of bioinformatics methods, the Human Genome Project, is currently being completed. Because of this the nature and priorities of bioinformatics research and applications are changing. People often talk portentously of our living in the " post-genomic" era. My personal view is that this will affect bioinformatics in several ways:

    • Now we possess multiple whole genomes we can look for differences and similarities between all the genes of multiple species. From such studies we can draw particular conclusions about species and general ones about evolution. This kind of science is often referred to as comparative genomics.
    • There are now technologies designed to measure the relative number of copies of a genetic message (levels of gene expression) at different stages in development or disease or in different tissues. Such technologies, such as DNA microarrays will grow in importance.
    • Other, more direct, large-scale ways of identifying gene functions and associations (for example yeast two-hybrid methods) will grow in significance and with them the accompanying bioinformatics of functional genomics.
    • There will be a general shift in emphasis (of sequence analysis especially) from genes themselves to gene products. This will lead to:
      • attempts to catalogue the activities and characterize interactions between all gene products (in humans): proteomics ).
      • attempts to crystallize and or predict the structures of all proteins (in humans): structural genomics.
      • fewer DNA double-helices in bad sci-fi movies.
    • What some people refer to as research or medical informatics, the management of all biomedical experimental data associated with particular molecules or patients---from mass spectroscopy, to in vitro assays to clinical side-effects---will move from the concern of those working in drug company and hospital I.T. (information technology) into the mainstream of cell and molecular biology and migrate from the commercial and clinical to academic sectors.
    This FAQ concentrates on classical bioinformatics, but will, I hope, grow to cover more of the "post-genomic" aspects of the field. It is worth noting that all of the above non-classical areas of research depend upon established sequence analysis techniques.

    Definitions of Fields Related to Bioinformatics

    What is Biophysics?

    Molecular biology itself grew out of biophysics.The British Biophysical Society defines biophysics as:

    "an interdisciplinary field which applies techniques from the physical sciences to understanding biological structure and function"
    More information about the various facets of the discipline can be found at the society's site hosted at Birkbeck College, London.

    Mike Goodrich wrote to ask what the status of biophysics was given the definition of computational biology submitted by Paul Schulte (below). A recent article in The Scientist [free registration required] dealt with this question---thanks to Jo Wixon (Managing Editor of Comparative and Functional Genomics) for the reference.

    What is Computational Biology?

    Computational biologists might object (please do), but, I find that people use "computational biology" when discussing that subset of bioinformatics (in the broadest sense) closest to the field of classical general biology.

    Computational biologists interest themselves more with evolutionary, population and theoretical biology rather than cell and molecular biomedicine. It is inevitable that molecular biology is profoundly important in computational biology, but it is certainly not what computational biology is all about (see next paragraph). In these areas of computational biology it seems that computational biologists have tended to prefer statistical models for biological phenomena over physico-chemical ones. This is often wise...

    One computational biologist (Paul J Schulte) did object to the above and makes the entirely valid point that this definition derives from a popular use of the term, rather than a correct one. Paul works on water flow in plant cells. He points out that biological fluid dynamics is a field of computational biology in itself. He argues that this, and any application of computing to biology, can be described as "computational biology" (see also the "loose" definition of bioinformatics below). Where we disagree, perhaps, is in the conclusion he draws from this---which I reproduce in full:

    "Computational biology is not a "field", but an "approach" involving the use of computers to study biological processes and hence it is an area as diverse as biology itself."

    Richard Durbin, Head of Informatics at the Wellcome Trust Sanger Institute, expressed an interesting opinion on this distinction in an interview:

    "I do not think all biological computing is bioinformatics, e.g. mathematical modelling is not bioinformatics, even when connected with biology-related problems. In my opinion, bioinformatics has to do with management and the subsequent use of biological information, particular genetic information."

    What is Medical Informatics?

    The Medical Informatics FAQ (no relation) provides the following definition:

    "Biomedical Informatics is an emerging discipline that has been defined as the study, invention, and implementation of structures and algorithms to improve communication, understanding and management of medical information."

    That FAQ also points here

    Aamir Zakaria, the author of the FAQ, emphasises that medical informatics is more concerned with structures and algorithms for the manipulation of medical data, rather than with the data itself.

    This suggests that one difference between bioinformatics and medical informatics as disciplines lies with their approaches to the data; there are bioinformaticians interested in the theory behind the manipulation of that data and there are bioinformatics scientists concerned with the data itself and its biological implications. (I believe that a good bioinformatics researcher should be interested in both of these aspects of the field.)

    Medical informatics, for practical reasons, is more likely to deal with data obtained at "grosser" biological levels---that is information from super-cellular systems, right up to the population level---while most bioinformatics is concerned with information about cellular and biomolecular structures and systems.

    On both of these points I'd be happy for any medical informatics specialists to correct me.

    What is Cheminformatics?

    The Web advertisement for Cambridge Healthtech Institute's Sixth Annual Cheminformatics conference describes the field thus:

    "the combination of chemical synthesis, biological screening, and data-mining approaches used to guide drug discovery and development"

    but this, again, sounds more like a field being identified by some of its most popular (and lucrative) activities, rather than by including all the diverse studies that come under its general heading.

    The story of one of the most successful drugs of all time, penicillin, seems bizarre, but the way we discover and develop drugs even now has similarities, being the result of chance, observation and a lot of slow, intensive chemistry. Until recently, drug design always seemed doomed to continue to be a labour-intensive, trial-and-error process. The possibility of using information technology, to plan intelligently and to automate processes related to the chemical synthesis of possible therapeutic compounds is very exciting for chemists and biochemists. The rewards for bringing a drug to market more rapidly are huge, so naturally this is what a lot of cheminformatics works is about.

    Here is a page with a commercial slant which links to some interesting discussions of the term "cheminformatics", what it means, whether or not it exists as a distinct discipline, and even whether it should be replaced by "chemoinformatics".

    The span of academic cheminformatics is wide and is exemplified by the interests of the cheminiformatics groups at the Centre for Molecular and Biomolecular Informatics at the University of Nijmegen in the Netherlands. These interests include:

    • Synthesis Planning
    • Reaction and Structure Retrieval
    • 3-D Structure Retrieval
    • Modelling
    • Computational Chemistry
    • Visualisation Tools and Utilities

    Trinity University's Cheminformatics Web page, for another example, concerns itself with cheminformatics as the use of the Internet in chemistry.

    What is Genomics?

    Genomics is a field which existed before the completion of the sequences of genomes, but in the crudest of forms, for example the oft-re-referenced estimate of 100 000 genes in the human genome derived from a(n) (in)famous piece of "back of an envelope" genomics, guessing the weight of chromosomes and the density of the genes they bear. Genomics is any attempt to analyze or compare the entire genetic complement of a species or species (plural). It is, of course possible to compare genomes by comparing more-or-less representative subsets of genes within genomes.

    What is Mathematical Biology?

    Mathematical biology is easier to distinguish from bioinformatics than computational biology. Mathematical biology also tackles biological problems, but the methods it uses to tackle them need not be numerical and need not be implemented in software or hardware. Indeed, such methods need not "solve" anything; in mathematical biology it would be considered reasonable to publish a result which merely establishes that a biological problem belongs to a particular general class.

    The distinction between bioinformatics and mathematical biology was illuminated by an email I received from Alex Kasman at the College of Charleston. According to his working definition, he distinguished bioinformatics which (under the tight definition at least)...

    "...seems to focus almost exclusively on specific algorithms that can be applied to large molecular biological data sets..."

    ...from mathematical biology which...

    "...includes things of theoretical interest which are not necessarily algorithmic, not necessarily molecular in nature, and are not necessarily useful in analyzing collected data."

    What is Proteomics?

    A recent review on proteomics in the journal Nature defined the field this way:

    "The term proteome was first coined to describe the set of proteins encoded by the genome1. The study of the proteome, called proteomics, now evokes not only all the proteins in any given cell, but also the set of all protein isoforms and modifications, the interactions between them, the structural description of proteins and their higher-order complexes, and for that matter almost everything 'post-genomic'."

    Michael J.Dunn, the Editor-in-Chief of Proteomics defines the "proteome" as:

    "the PROTEin complement of the genOME"

    and proteomics to be concerned with:

    "qualitative and quantitative studies of gene expression at the level of the functional proteins themselves"

    that is:

    "an interface between protein biochemistry and molecular biology"

    Characterizing the many tens of thousands of proteins expressed in a given cell type at a given time---whether measuring their molecular weights or isoelectric points, identifying their ligands or determining their structures---involves the storage and comparison of vast numbers of data. Inevitably this requires bioinformatics. Here is a constructively skeptical review by Lukas Huber.

    What is Pharmacogenomics?

    Pharmacogenomics is the application of genomic approaches and technologies to the identification of drug targets. Examples include trawling entire genomes for potential receptors by bioinformatics means, or by investigating patterns of gene expression in both pathogens and hosts during infection, or by examining the characteristic expression patterns found in tumours or patients samples for diagnostic purposes (possibly in the pursuit of potential cancer therapy targets).

    The term "pharmacogenomics" is used for the more "trivial"---but arguably more useful---application of bioinformatics approaches to the cataloguing and processing of information relating to pharmacology and genetics, for example the accumulation of information in databases like this one. (Thanks to Ivanovi.)

    What is Pharmacogenetics?

    All individuals respond differently to drug treatments; some positively, others with little obvious change in their conditions and yet others with side effects or allergic reactions. Much of this variation is known to have a genetic basis. Pharmacogenetics is a subset of pharmacogenomics which uses genomic/bioinformatic methods to identify genomic correlates, for example SNPs (Single Nucleotide Polymorphisms), characteristic of particular patient response profiles and use those markers to inform the administration and development of therapies. Strikingly, such approaches have been used to "resurrect" drugs thought previously to be ineffective, but subsequently found to work with in subset of patients. They can also be used for optimizing the doses of chemotherapy for particular patients.

    Overview of most common bioinformatics programs

    Everyday bioinformatics is done with sequence search programs like BLAST, sequence analysis programs, like the EMBOSS and Staden packages, structure prediction programs like THREADER or PHD or molecular imaging/modelling programs like RasMol and WHATIF.

    Overview of most common bioinformatics technology

    Currently, a lot of bioinformatics work is concerned with the technology of databases (Thanks again to Ivanovi.) These databases include both "public" repositories of gene data like GenBank or the Protein DataBank (the PDB), and private databases, like those used by research groups involved in gene mapping projects or those held by biotech companies. Making such databases accessible via open standards is very important. Consumers of bioinformatics data use a range of computer platforms: from the more powerful and forbidding UNIX boxes favoured by the developers and curators to the far friendlier Macs often found populating the labs of computer-wary biologists.

    Databases of existing sequencing data can be used to identify homologues of new molecules that have been amplified and sequenced in the lab. The property of sharing a common ancestor, homology, can be a very powerful indicator in bioinformatics (see below).

    Acquisition of sequence data

    Bioinformatics tools can be used to obtain sequences of genes or proteins of interest, either from material obtained, labelled, prepared and examined in electric fields by individual researchers/groups or from repositories of sequences from previously investigated material.

    Analysis of data

    Both types of sequence can then be analysed in many ways with bioinformatics tools.

    They can be assembled. Note that this is one of the occasions when the meaning of a biological term differs markedly from a computational one (see the amusing confusion over the issue at Web-based geek forum Slashdot). Computer scientists, banish from your mind any thought of assembly language. Sequencing can only be performed for relatively short stretches of a biomolecule and finished sequences are therefore prepared by arranging overlapping "reads" of monomers (single beads on a molecular chain) into a single continuous passage of "code". This is the bioinformatic sense of assembly.

    They can be mapped---that is, their sequences can be parsed to find sites where so-called "restriction enzymes" will cut them.

    They can be compared, usually by aligning corresponding segments and looking for matching and mismatching letters in their sequences. Genes or proteins that are sufficiently similar are likely to be related and are therefore said to be "homologous" to each other---the whole truth is rather more complicated than this. Such cousins are called "homologues".

    If a homologue (a related molecule) exists, then a newly discovered protein may be modelled---that is the three dimensional structure of the gene product can be predicted without doing laboratory experiments.

    Bioinformatics is used in primer design. Primers are short sequences needed to make many copies of (amplify) a piece of DNA as used in PCR (the Polymerase Chain Reaction).

    Bioinformatics is used to attempt to predict the function of actual gene products.

    Information about the similarity, and, by implication, the relatedness of proteins is used to trace the "family trees" of different molecules through evolutionary time.

    There are various other applications of computer analysis to sequence data, but, with so much raw data being generated by the Human Genome Project and other initiatives in biology, computers are presently essential for many biologists just to manage their day-to-day results

    Molecular modelling / structural biology is a growing field which can be considered part of bioinformatics. There are, for example, tools which allow you (often via the Net) to make pretty good predictions of the secondary structure of proteins arising from a given amino acid sequence, often based on known "solved" structures and other sequenced molecules acquired by structural biologists.

    Structural biologists use "bioinformatics" to handle the vast and complex data from X-ray crystallography, nuclear magnetic resonance (NMR) and electron microscopy investigations and create the 3-D models of molecules that seem to be everywhere in the media.

    note

    Unfortunately the word "map" is used in several different ways in biology/genetics/bioinformatics. The definition given above is the one most frequently used in this context, but a gene can be said to be "mapped" when its parent chromosome has been identified, when its physical or genetic distance from other genes is established and---less frequently---when the structure and locations of its various coding components (its "exons") are established.

    What is Bioinformatics?---The Loose definition

    There are other fields---for example medical imaging / image analysis which might be considered part of bioinformatics. There is also a whole other discipline of biologically-inspired computation; genetic algorithms, AI, neural networks. Often these areas interact in strange ways. Neural networks, inspired by crude models of the functioning of nerve cells in the brain, are used in a program called PHD to predict, surprisingly accurately, the secondary structures of proteins from their primary sequences.

    What almost all bioinformatics has in common is the processing of large amounts of biologically-derived information, whether DNA sequences or breast X-rays.

    How old is the discipline?

    "How old is bioinformatics?" The answer to this one depends on which source you choose to read.

    From T K Attwood and D J Parry-Smith's "Introduction to Bioinformatics", Prentice-Hall 1999 [Longman Higher Education; ISBN 0582327881]:

    "The term bioinformatics is used to encompass almost all computer applications in biological sciences, but was originally coined in the mid-1980s for the analysis of biological sequence data."

    From Mark S. Boguski's article in the "Trends Guide to Bioinformatics" Elsevier, Trends Supplement 1998 p1:

    "The term "bioinformatics" is a relatively recent invention, not appearing in the literature until 1991 and then only in the context of the emergence of electronic publishing...

    "...However, some of my role models when I was a graduate student (Margaret O. Dayhoff, Russell F. Doolittle, Walter M. Fitch and Andrew D. McLachlan) had been building databases, developing algorithms and making biological discoveries by sequence analysis since the 1960s---long before anyone thought to label this activity with a special term (if anything it was called `molecular evolution'). Even a relatively new kid on the block, the National Center for Biotechnology Information (NCBI), is celebrating its 10th anniversary this year, having been written into existence by US Congressman Claude Pepper and President Ronald Reagan in 1988. So bioinformatics has, in fact, been in existence for more than 30 years and is now middle-aged."

    Books: Can you recommend any bioinformatics books?

    It's notoriously difficult to find any books on bioinformatics itself that cater well for all of those coming from computing, from mathematics and from biology backgrounds. The few textbooks available in the field tend to be eyewateringly expensive as well. I've divided suggested reading into books of general interest, those best suited to people coming from a computational/mathematical background and books for biologists interested in bioinformatics. Where a book is also listed in Bioinformatics.Org's books section I have linked the title to the relevant entry there. Links to other lists of bioinformatics books follow this section of suggested reading.

    General introductions

    Many people are curious about the Human Genome (Project). The completion of the first draft probably represents bioinformatics' coming of age as a discipline. The first couple of books are aimed at the intelligent layperson.

    A gossipy and insightful account of the race to sequence the genome can be found in "The Sequence" by Kevin Davies [Weidenfeld; ISBN 0297646982]. Matt Ridley's "Genome" [Fourth Estate; ISBN 185702835X] is both an interesting layperson's introduction to the issues raised by the bioinformatic revolution and an overview of its biology and enormous scope. If I remember rightly, Ridley's book received a slightly snooty review from Walter Bodmer. This is understandable, since his and Robin McKie's excellent "pre-genomic" guide to the Human Genome Mapping Project, "The Book of Life" [Oxford Paperbacks; ISBN 0195114876] was undeservedly in a remainders bin when I bought my copy a couple of years ago.

    If you are a non-biological scientist (or a non-scientist) and are hooked by these, why not go back to the "real beginning" of the race and read James Watson's entertaining and indiscreet memoir of his and Francis Crick's determination of the structure of DNA, "The Double Helix" [Penguin; ISBN 0140268774]---now updated with an introduction by media don Steve Jones.

    Nigel Barber at Peterborough Regional College in the UK recommends Gary Zweiger's "Transducing the Genome" [McGraw-Hill Professional Publishing: ISBN 0071369805]. The summary at Amazon makes it sound a tad pretentious, but all the reviews seem pretty positive so it might be worth a read.

    If you are a quantitative scientist and would like a deeper knowledge of contemporary (molecular) biology, but you want to acquire it as painlessly as possible you could try the following:

    • Donna Rae Siegfried's Biology for Dummies [Wiley; ISBN 0-7645-5326-7] is fun, well thought out and a lot more informative than the title might suggest. If only all biology textbooks were this entertaining and unpretentious.
    • If you already have some biological knowledge and would like to get a grip on modern biomolecular science then Richard J. Epstein's Human Molecular Biology is an elegant, colourful and detailed guide.

    There are two classic competing texts in cell and molecular biology which Maximilian Haeussler reminds me to include: Alberts et al's Molecular Biology of the Cell [Garland Science: ISBN 0815340729] and Molecular Biology of the Gene [Benjamin Cummings: ISBN 0321248643].

    Computational/Mathematical aspects

    If you are a hardcore maths/computing person Michael Waterman's "Introduction to Computational Biology" [Chapman & Hall/CRC Statistics and Mathematics; ISBN 0412993910] and Pavel Pevzner's "Computational Molecular Biology - An Algorithmic Approach" [The MIT Press (A Bradford Book); ISBN 0262161974] will give you all the discrete maths you can shake a stick at, but perfunctory introductions to the biology.

    Bioinformatics.Org's very own Jeff Bizzaro recommends Dan Gusfield's "Algorithms on Strings, Trees and Sequences" [Cambridge, 1997 ISBN 0-52158-519-8], Richard Durbin, S. Eddy, A. Krogh, G. Mitchison "Biological Sequence Analysis: Probabilistic Models of Proteins and Nucleic Acids" [Cambridge, 1997 ISBN 0-52162-971-3] (which I think is one of the clearest and most comprehensive guides to alignment algorithms) and---for that full "computers-to-biology conversion"--- Geoffrey M. Cooper "The Cell: A Molecular Approach" [ASM Press, 1996 ISBN 0-87893-119-8]. Jeff Ames writes that a second edition of this book is now available [Sinauer Associates, Incorporated, 2000 ISBN 0-87893-106-6] and that this version---if you can find it in the shops---comes with a CD.

    Applying bioinformatics to biological research

    One outstanding general text for the biologist is David W. Mount's "Bioinformatics" [Cold Spring Harbor Press; ISBN 0879696087]. It's not cheap, but it's the best I've seen if you are studying bioinformatics itself.

    Bioinformatics has been dismissed by some as "the science of BLAST searches". The best collection of advice so far on doing BLAST searches is O'Reilly's BLAST book by Ian Korf, Mark Yandell and Joseph Bedell [O'Reilly ISBN 0-596-00299-8]. I reviewed it enthusiastically, but not uncritically, for the UK UNIX Users' Group magazine. I'd go as far as to say that all biologists thinking of using BLAST in their research should read the relevant sections before they even go near a computer.

    If you wish to use general bioinformatics tools, especially if you are a little wary of computers, my new "best" book is "Bioinformatics for Dummies" [John Wiley and Sons ISBN 0764516965]. It is (obviously) aimed at people who are beginners, who are happier using the Web rather than typing commands, and who are more interested in learning than in impressing people---the writing is friendly clear and unpretentious. However, like several of my other tips (below) it concentrates on Web-based resources so it will, inevitably, date. (This is partially compensated for by there being a companion Website.)

    Also, if you're coming to the subject as a computer user with a biological background, looking to exploit the many tools available, you might want to try Terry Attwood and David Parry-Smith's "Introduction to Bioinformatics" [Longman Higher Education; ISBN 0582327881], or Des Higgins and Willie Taylor's "Bioinformatics: Sequence Structure and Databanks" [Oxford University Press; ISBN 0199637903]. Another excellent practical introduction is Andreas Baxevanis and Francis Oulette's "Bioinformatics: A Practical Guide to the Analysis of Genes and Proteins" [Wiley-Interscience; ISBN 0471383910], now in its new and improved second edition. Bax teaches bioinformatics all over Canada and the experience shows. Arthur Lesk has also produced an excellent teaching book particularly for protein bioinformatics in his Introduction to Bioinformatics

    Bioinformatics.Org also recommends Cynthia Gibas and Per Jambeck's "Developing Bioinformatics Skills" [O'Reilly, 2001 ISBN 1-56592-664-1].

    Stuart Brown recommends his own book "Bioinformatics: A Biologist's Guide to Biocomputing and the Internet" [Eaton Pub Co; ISBN: 188129918X]. If he sends me a review copy I might recommend it too ;-) .

    Fiction books

    "Darwin's Radio" by Greg Bear [Ballantine Books, ISBN: 0345435249] is a wonderful hard SF thriller which stretches ideas derived from genome discoveries to their breaking point. It's gripping and humane.

    Leonard Crane, the author of Ninth Day of Creation kindly sent me a copy for review. So far it's an excellent read. I haven't finished it yet, not because it isn't a rattling good story, but because, like "Darwin's Radio", it is very long and because I am very busy. If you'd like to read a well-researched, but speculative, novel containing actual scenes of practising bioinformatics then try it.

    Ken Allen contributed the following reviews:

    "Frameshift [Tor Books, ISBN: 0812571088] by Robert J. Sawyer---based around the HGP---reasonable read, but poor / confused ending."

    Calculating God [Tor Books, ISBN: 0812580354]by the same author---has a subtler bio connection and is a much better read. Near the start an alien spacecraft lands, the alien emerges and says 'take me to your paleontologist'

    Further suggestions for this section are welcome.

    Other lists of bioinformatics books

    See also compbiology.org's list, Steve Brenner's list, and Aik Choon Tan's collection of books.

    Centres of Bioinformatics Activity: Where is bioinformatics done?

    The biggest and best source of bioinformatics links I have encountered is the Genome Web at the Rosalind Franklin Centre for Genomics Research at the Genome Campus near Cambridge, UK. Most of the links below come from that resource. My list is necessarily limited by comparison.

    Research centres

    Sequencing centres

    [XXXX INSERT DETAILS OF MORE SEQUENCING CENTRES HERE]

    Standards centres

    [XXXX INSERT DETAILS OF STANDARDS CENTRES HERE]

    What virtual centres (for example consortia and communities) for bioinformatics activity are there?

    [XXXX INSERT MORE DETAILS OF VIRTUAL BIOINFORMATICS CENTRES HERE]

    Online Resources: What bioinformatics Websites are there?

    'Blogs

    The front page of Bioinformatics.Org itself is a bioinformatics 'Blog.

    The Bio-Web links to resources online for molecular and cell biologists and covers current news in various biological/computational fields.

    Genehack is the first bioinformatics 'Blog I ever encountered.

    Information

    The Australian National Genomic Information Service (ANGIS) is operated by the Australian Genomic Information Centre (currently at The University of Sydney) to offer software, databases, documentation, training and support for biologists

    "The University of Maryland AgNIC gateway is a guide to quality agricultural biotechnology information on the Internet."

    Directories

    Christy Hightower, Engineering Librarian at the Science and Engineering Library, University of California Santa Cruz has already done this better than me. Visit her excellent article about bioinformatics Net resources in Issues in Science and Technology Librarianship.

    Societies

    Humberto Ortiz Zuazaga kindly introduced me to The International Society for Computational Biology which he points out "has links to programs of study and online courses in computational biology and to job postings".

    Collections of Tools

    You can start right here at Bioinformatics.Org if you are looking for a bioinformatics toolbox.

    I cannot recommend strongly enough the Rosalind Franklin Centre's "GenomeWeb".

    Of historical interest only now, I guess, is the legendary " Pedro's Molecular Biology Search and Analysis Tools".

    Portals

    Bioinformatics.Org is an international organization which promotes freedom and openness in the field of bioinformatics and is the root domain of a damned fine Website :-) .

    CCP11 (Collaborative Computational Project 11) is another product of the UK's Genome Campus. To quote their Web site, it was...

    "...established to foster the broad bioinformatics community and the UK research community in particular. Its purpose is to facilitate the transfer of knowledge and expertise through conferences, workshops, a newsletter and the use of the world wide web. CCP11 is funded by the BBSRC and is hosted at the MRC Rosalind Franklin Centre for Genomics Research RFCGR located on the Wellcome Trust Genome Campus, Cambridge."

    Jennifer Steinbachs runs compbiology.org which is a general computational biology site as well as being a portal to her own work.

    BioPlanet is well worth visiting. It describes itself as "a not-for-profit site, funded with our resources, for [its users'] benefit"

    ColorBasePair is a densely packed portal with lots of bioinformatics links

    Nick Yates runs his own informative bioinformatics site, unsurprisingly called nick-yates.com. He doesn't aim to make money from it, but it may have paid-for ads. Check out the glossaries---they are better than mine.

    Tutorials

    A great place to start, whether you come from a biological, physical or computational background is at Martin Vingron's superb online bioinformatics tutorial. (Begin by choosing a section from the left-hand-side menu bar.)

    Tom Smith and Don Emmeluth have produced a nice little exploration of bioinformatics using NCBI resources and tools.

    I recently stumbled upon a promising set of online lecture notes currently under construction by B. Steipe at the Genzentrum (Gene Center) at the Ludwig-Maximilians-Universität München (University of Munich).

    Chemistry for all

    A defiantly frames-free chemistry tutorial site.

    Mathematics for biologists

    First of all, an almost completely painless introduction to the horrors of the quadratic equation by Peter Whalen, James Walker, and Drew Marticorena.

    C. J. Schwarz of the Department of Statistics and Acturial Science, Simon Fraser University has produced a course in statistics which is accompanied by set of sound, online PDF handouts.

    Here is a great guide to a whole array of statistical learning/teaching resources prepared by Juha Puranen of the University of Helsinki (English).

    Computers for biologists

    Programming for biologists

    General introduction to biology for computer scientists

    Estrella Mountain Community College in the States offers this excellent short introduction to biology (actually "The Nature of Science and Biology". It's a great place for keyboard jockeys to start their journey to enlightenment. Thanks to Alex O'Neill for pointing out the broken link.

    Genetics

    The Dolan DNA Learning Center at Cold Spring Harbor has an outstanding interactive tutorial introducing genetics. To take full advantage of the multimedia elements you should download the Flash and Real players.

    Molecular biology for computer scientists

    The Institute of Arable Crop Research Beginner's Guide to Molecular Biology

    Protein chemistry for computer scientists

    Unilever Education Advanced Series tutorial on proteins.

    Cell biology for computer scientists

    The University of Arizona has made available a high-quality tutorial in cell biology. Not only does it cover the facts, but it also attempts to introduce some of the philosophy of the field---recommended. Even better, it's also available en Español and in Italiano.

    Once you've worked your way through that you might like to see some scanning electron microscope images of some of the structures you've read about taken by members of John Heuser's lab.

    Evolution for computer scientists

    Bob Patterson maintains his "Darwiniana" with amazing diligence.

    Practical bioinformatics

    Other lists of bioinformatics tutorials

    Education: Where can I study Bioinformatics...

    jump straight to introduction to education section

  • ...in Europe?
  • ...remotely?
  • This section is not complete, but contributions to broaden its coverage are welcome. Please do not direct questions about eligibility, course quality or admissions policy to me, but to ask the individual institutions directly. Use the links to obtain contact details. If an institution doesn't provide telephone numbers/email addresses or snailmail details on its Web site it doesn't deserve your patronage.

    This resource focuses on complete, full-time degree programmes rather than on individual study modules. Curating a list of the latter would be a full-time job. You can go to other places, however, if you are looking for short courses. Thanks to various contributors, including Wentian Li who pointed me to this list at Rockefeller which is mirrored at various other sites. And to Humberto Ortiz Zuazaga for mailing me a link to the ICSB, where you can find this list.

    If you are interested in U.S. programmes, here's a list from Curtin and here's a list from Stanford. Thanks to Amelie Stein who also supplied some of the individual entries in this section.

    Those wanting to find programmes in the Asia Pacific region could have a look at this resource maintained by the Asia Pacific Bioinformatics Network APBioNet. Thanks to Sentausa.

    In the UK The Bioinformatics Resource (part of the BBSRC's CCP11 project) project maintains (among many other resources) lists of (mainly) British Masters and PhDs in bioinformatics. If you have any suggestions or updates please contact me with them. You can publicize your course and offer a public service at the same time.

    Africa

    Rhodes University, Grahamstown, South Africa offers an MSc. in Bioinformatics and Computational Molecular Biology. Thanks to Natalie Twine.

    Cathal Seoighe wrote a while back about the South African National Bioinformatics Institute (SANBI). Ruediger Braeuning has since written to point out that bioinformatics training in South Africa has been radically reorganized. He says:

    "A new institute, the National Bioinformatics Nework (NBN), has been created. We have nodes at Universities all over the country (UWC, UCT, SUN, RU, UKZN, UP, WITS). Our main tasks are to:

    • develop capacity in Bioinformatics
    • perform world-class research
    • support local Biotechnology initiatives

    "We do offer courses on various topics in Bioinformatics ranging in length from 3 days to several weeks. We also train Bioinformaticists on MSc, PhD and post doc level. Undergraduate programs are currently being developed. Bursaries are available. For more information visit our Website."

    South African National Bioinformatics Institute (SANBI) Honours Bioinformatics Course at the University of the Western Cape. Next year the same institute will be offering a Master's in bioinformatics---thanks to Cathal Seoighe.

    If you know of any other bioinformatics courses on the African continent please feel free to mail me about them.

    The Americas

    Canada

    Thanks to Jordan Patterson for the information that the University of Alberta offers four-year Biology or Computer Science degrees with a specialization in bioinformatics. The Faculty of Computer Science there offers Master's and PhD training in bioinformatics.

    Benjamin Horsman wrote to tell me that Simon Fraser University and the University of British Columbia are collaborating on a new Bioinformatics training program with the British Columbia Cancer Agency. The program offers post-graduate diploma, Master's, and PhD training in Bioinformatics. Now Simon Fraser University also offers a joint major programme in Molecular Biology and Biochemistry (MBB) and Computer Science in Bioinformatics. Thanks to Brittany Nielsen for the info.

    Thanks to Olga Likhodi for the information that Seneca College, Toronto offers a post-graduate diploma in Bioinformatics.

    Peter Kublik informs me that from 2003/2004 the University of Calgary will offer a bioinformatics programme. He's part of the first intake.

    The University of Waterloo, Department of Computer Science offers undergraduate and graduate courses in bioinformatics. More information is here.

    California

    The Keck Graduate Institute claims that computational biology is a core element of the curriculum in its Master of Bioscience degree.

    Stanford University offers academic and professional (distance-learning) MSs in Biomedical Bioinformatics as well as its PhD programme. Thanks to Betty Cheng.

    Thanks to Momchil Georgiev for the information that the University of California at San Diego offers a Bioinformatics graduate programme and to Dana Brehm that there is now a new bachelor's program, to quote her:

    "[This is an] undergraduate, interdisciplinary program for undergraduates leading to a B.S. degree. The new Bioinformatics major is offered by the Division of Biology, and the departments of Chemistry/Biochemistry, Computer Science and Engineering, and Bioengineering. A student may choose to major in Bioinformatics in any one of the four departments or division. The Division of Biology currently offers two Bioinformatics courses, and with the advent of the cross-disicplinary major, even more courses are going to be taught 2002-03 and 2003-04."
    .

    University of California, Irvine Informatics in Biology and Medicine

    David Delong wrote to me to point out that the College of Natural and Agricultural Sciences at the University of California, Riverside is developing a "Center in Genomics and Bioinformatics" which will offer a PhD curriculum in genomics and bioinformatics from academic year 2001-2002 onwards.

    Catherine Velazquez says that The University of California, Santa Cruz offers a new undergraduate BS course in bioinformatics. They have a Frequently Asked Questions. Now they also offer an MS/PhD in Bioinformatics. Thanks to Kevin Karplus for the update.

    Connecticut

    Javier Rojas Balderrama emailed me to point out thatYale University offers a Bioinformatics and Computational Biology track as part of its combined Biological and Biomedical Sciences graduate programme.

    Georgia

    Georgia Institute of Technology Masters of Science in Bioinformatics

    According to Eric VanWieren Georgia State University offers a Master's and PhD in Computer Science with a focus on bioinformatics. The university's Bachelor of Science in Computer Science also offers a "Fundamentals of Bioinformatics" course.

    Illinois

    The University of Illinois at Chicago offers graduate programmes covering Bioengineering Bioinformatics through its Bioengineering department as well as an undergraduate course track. Thanks to Amit Sabnis.

    Indiana

    IUPUI offers an MS programme in Bioinformatics.

    Indiana University also offers an MS programme in Bioinformatics.

    Iowa

    Iowa State University offers an Interdisciplinary Ph.D. Program in Bioinformatics and Computational Biology (BCB).

    Maine

    The Jackson Lab, a World centre of mouse genome informatics offers a graduate training program.

    Maryland

    Tim Young wrote to say that Johns Hopkins University in Maryland offers an MS in Bioinformatics through the Zanvyl Krieger School of Arts and Sciences Advanced Academic Programs and Whiting School of Engineering Engineering and Applied Science Programs for Professionals. They are also offering a Bioinfomatics concentration with their MS in Biotechnology program.

    Massachusetts

    Boston University offers a graduate programme and so does its partner North Eastern University. North Eastern also offers a Graduate Certificate in the subject.

    Brandeis University offers both a Master of Science in Bioinformatics and a Graduate Certificate in Bioinformatics. Thanks to Matt Foster.

    The Department of Computer Science at UMass Lowell offers various degrees from Bachelor's through to PhD. level in Computer Science with Bioinformatics options.

    Mexico

    At the National Autonomous University of Mexico a doctoral program in biomedical sciences is available. Their Computational Molecular Biology Group is here.

    Minnesota

    The University of Minnesota offers a graduate programme in bioinformatics.

    Thanks to Anu Haniharan for drawing my attention to mixing up the Minnesota and New Jersey paragraphs.

    Nebraska

    The University of Nebraska Lincoln offers an Interdisciplinary Bioinformatics Specialization.

    The Graduate Program of the Pathology-Microbiology Department at the University of Nebraska Medical Center (University of Nebraska at Omaha) offers a specialty track in bioinformatics.

    NewJersey

    Rama Penta wrote to say that Stevens Institute of Technology offers a Master's programme in Bioinformatics.

    The message also states that the University of Medicine and Dentistry New Jersey (UMDNJ) offers a programme in biomedical informatics.

    Thanks to Anu Haniharan for drawing my attention to mixing up the Minnesota and New Jersey paragraphs.

    Moustafa wrote to say that Ramapo College in New Jersey is the only school in New Jersey offering a Bachelor's degree in bioinformatics.

    New York State

    The University at Buffalo has been involved in establishing a "Center of Excellence in Bioinformatics". It used to a range of courses in bioinformatics and related subjects, but all the course links seem to be dead now. Thanks to Jeff Ligas for the original notification.

    Canisius College---also in Buffalo, NY---has had a state-approved B.S. in Bioinformatics since 2001. Thanks to Deb Burhans.

    Cornell and Rockefeller Universities, together with the Sloan-Kettering Research Institute offer a "Tri-institutional program in Computational Biology and Medicine". Thanks to Brant Inman.

    Rensselaer Polytechnic Institute offers both undergraduate and graduate programmes in bioinformatics

    Rochester Institute of Technology offers BS MS and BS/MS programmes in Bioinformatics. Thanks to Brandon H.

    According to Maureen Downey, the College of Staten Island, part of the City University of New York also offers a challenging program in bioinformatics.

    If you know of any other bioinformatics courses on the American continent please feel free to mail me about them.

    North Carolina

    Duke University's Center for Bioinformatics and Computational Biology offers var