Brian Goodwin
Encyclopedia
Brian Carey Goodwin was a Canadian mathematician
and biologist
, a Professor Emeritus at the Open University
and a key founder of the field of theoretical biology.He made key contributions to the foundations of biomathematics, complex systems and generative models in developmental biology. He was one of the prominent scientists who suggested that a reductionist view of nature will fail to explain complex features. He was also a visible member of the Third Culture
movement.
, Canada
in 1931. He studied biology at McGill University
and then emigrated to the UK
, under a Rhodes Scholarship
for studying mathematics at Oxford. He got his PhD at the University of Edinburgh
under the supervision of Conrad Hal Waddington
. He then moved to Sussex University until 1983 when he became a full professor
at the Open University
in Milton Keynes
until retirement in 1992. He became a major figure in the early development of mathematical biology
, along with other researchers. He was one of the attendants to the famous meetings that took place between 1965 and 1968 in Villa Serbelloni, hosted by the Rockefeller Foundation
, under the topic "Towards a theoretical Biology". The workshop involved, among other key scientists, Conrad Waddington, Jack Cowan
, Michael Conrad
, Christopher Zeeman, Richard Lewontin
, Robert Rosen, Stuart Kauffman
, John Maynard Smith
, Rene Thom
and Lewis Wolpert
. As a result of the conference talks and discussions, a four-volumes came out, becoming at the time a major reference in the area.
and Jacques Monod
developed their first model of gene regulation, Goodwin proposed the first model of a genetic oscillator, showing that regulatory interactions among genes allowed periodic fluctuations to occur. Shortly after this model became published, he also formulated a general theory of complex gene regulatory networks using statistical mechanics
.
In its simplest form, Goodwin's oscillator involves a single gene that represses itself. Goodwin equations were originally formulated in terms of conservative (Hamiltonian) systems, thus not taking into account dissipative effects that are required in a realistic approach to regulatory phenomena in biology. Many versions have been developed since then. The simplest (but realistic) formulation considers three variables, X, Y and Z indicating the concentrations of RNA
, protein
and end product which generates the negative feedback
loop.The equations are
and closed oscillations can occur for n>8 and behave limit cycles: after a perturbation of the system's state, it returns to its previous attractor. A simple modification of this model, adding other terms introducing additional steps in the transcription machinery allows to find oscillations for smaller n values. Goodwin's model and
its extensions have been widely used over the years as the basic skeleton for other models of oscillatory behavior, including circadian clocks, cell division
or physiological
control systems.
) to multicellular organisms, including early development in Drosophila
. One of his key contributions was to link morphogenetic fields, defined in terms of spatial distributions of chemical
signals (morphogenes), and the shape of the system experiencing morphogenetic changes. In this wasy, geometry and development were linked through a mathematical formalism. Along with his colleague Lynn Trainor, Goodwin developed a set of mathematical equations describing the changes of both physical boundaries in the porganism and chemical gradients.
By considering the mechanochemical behaviour of the cortical cytoplasm (or cytogel) of plant cells, a viscoelastic material mainly composed of actin microfilaments and reinforced by a microtubules network, Goodwin & Trainor (1985) showed how to couple calcium and the mechanical properties of the cytoplasm. The cytogel is treated as a continuous viscoelastic medium in which calcium ions can diffuse and interact with the cytoskeleton. The model consists in two non-linear partial differential equations which describe the evolution of the mechanical strain field and .of the calcium distribution in the cytogel.
It has been shown (Trainor & Goodwin, 1986) that, in a range of parameter values, instabilities may occur and develop in this system, leading to intracellular patterns of strain and calcium concentration. The equations read, in their general form:
These equations describe the spatiotemporal dynamics of the displacement from the reference state and the calcium concentration, respectively. Here
x and t are the space and time coordinates, respectively. These equations
can be applied to many different scenarios and the different functions P(x) introduce the specific mechanical properties of the medium. These equations
are very rich in terms of the static and dynamic patterns that can generate, including both complex geometrical motifs to oscillations
and chaos (Briere 1994).
of biological systems. In that sense, he became one of the strongest defenders of the systems view against reductionism
. Among other contributions, he suggested that nonlinear phenomena and the fundamental laws defining their behavior were essential in order to understand biology and its evolutionary paths. His position within evolutionary biology can be defined as a structuralist
one. To Goodwin, many patterns that we observe in nature are a byproduct of constraints imposed by complexity. The limited repertoire of motifs observed in the spatial organization of plants and animals (at some scales) would be, in Goodwin's opinion, a fingerprint of the role played by such constraints. The role of selection would be secondary.
These opinions were highly controversial and
Goodwin came into conflict with many prominent Darwinian evolutionists, whereas many physicists found some of his view natural. Physicist Murray Gell-Mann
for example acknowledged that "when biological evolution — based on largely random variation in genetic material and on natural selection — operates on the structure of actual organisms, it does so subject to the laws of physical science, which place crucial limitations on how living things can be constructed.". Richard Dawkins
, the former professor for public understanding of science at Oxford University and a well known Darwinian evolutionist, conceded: "I don't think there's much good evidence to support [his thesis], but it's important that somebody like Brian Goodwin is saying that kind of thing, because it provides the other extreme, and the truth probably lies somewhere between." Dawkins also agreed that "It's a genuinely interesting possibility that the underlying laws of morphology allow only a certain limited range of shapes.". For his part, Goodwin did not reject basic Darwinism, only its excesses.
in Devon
, UK
, where he was instrumental in starting the college's MSc in Holistic Science. He was made a Founding Fellow of Schumacher College shortly before his death. Goodwin also had a research position at MIT and was a long time visitor of several institutions including the UNAM
in Mexico City. He was a founding member of the Santa Fe Institute
in New Mexico
where he also served as a member of the science board for several years.
Brian Goodwin died in hospital in 2009, after surgery resulting from a fall from his bike.. Goodwin is survived by his third wife, Christel, and his daughter, Lynn.
Selected Scientific papers
Essays
Mathematician
A mathematician is a person whose primary area of study is the field of mathematics. Mathematicians are concerned with quantity, structure, space, and change....
and biologist
Biologist
A biologist is a scientist devoted to and producing results in biology through the study of life. Typically biologists study organisms and their relationship to their environment. Biologists involved in basic research attempt to discover underlying mechanisms that govern how organisms work...
, a Professor Emeritus at the Open University
Open University
The Open University is a distance learning and research university founded by Royal Charter in the United Kingdom...
and a key founder of the field of theoretical biology.He made key contributions to the foundations of biomathematics, complex systems and generative models in developmental biology. He was one of the prominent scientists who suggested that a reductionist view of nature will fail to explain complex features. He was also a visible member of the Third Culture
Third culture
Third culture can refer to:* Third Culture Kid, a term for children who have lived a significant portion of their lives in a country that is not their passport country.* The Third Culture, a book by John Brockman....
movement.
Biography
Brian Goodwin was born in MontrealMontreal
Montreal is a city in Canada. It is the largest city in the province of Quebec, the second-largest city in Canada and the seventh largest in North America...
, Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...
in 1931. He studied biology at McGill University
McGill University
Mohammed Fathy is a public research university located in Montreal, Quebec, Canada. The university bears the name of James McGill, a prominent Montreal merchant from Glasgow, Scotland, whose bequest formed the beginning of the university...
and then emigrated to the UK
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...
, under a Rhodes Scholarship
Rhodes Scholarship
The Rhodes Scholarship, named after Cecil Rhodes, is an international postgraduate award for study at the University of Oxford. It was the first large-scale programme of international scholarships, and is widely considered the "world's most prestigious scholarship" by many public sources such as...
for studying mathematics at Oxford. He got his PhD at the University of Edinburgh
University of Edinburgh
The University of Edinburgh, founded in 1583, is a public research university located in Edinburgh, the capital of Scotland, and a UNESCO World Heritage Site. The university is deeply embedded in the fabric of the city, with many of the buildings in the historic Old Town belonging to the university...
under the supervision of Conrad Hal Waddington
Conrad Hal Waddington
Conrad Hal Waddington CBE FRS FRSE was a developmental biologist, paleontologist, geneticist, embryologist and philosopher who laid the foundations for systems biology...
. He then moved to Sussex University until 1983 when he became a full professor
Professor
A professor is a scholarly teacher; the precise meaning of the term varies by country. Literally, professor derives from Latin as a "person who professes" being usually an expert in arts or sciences; a teacher of high rank...
at the Open University
Open University
The Open University is a distance learning and research university founded by Royal Charter in the United Kingdom...
in Milton Keynes
Milton Keynes
Milton Keynes , sometimes abbreviated MK, is a large town in Buckinghamshire, in the south east of England, about north-west of London. It is the administrative centre of the Borough of Milton Keynes...
until retirement in 1992. He became a major figure in the early development of mathematical biology
Mathematical biology
Mathematical and theoretical biology is an interdisciplinary scientific research field with a range of applications in biology, medicine and biotechnology...
, along with other researchers. He was one of the attendants to the famous meetings that took place between 1965 and 1968 in Villa Serbelloni, hosted by the Rockefeller Foundation
Rockefeller Foundation
The Rockefeller Foundation is a prominent philanthropic organization and private foundation based at 420 Fifth Avenue, New York City. The preeminent institution established by the six-generation Rockefeller family, it was founded by John D. Rockefeller , along with his son John D. Rockefeller, Jr...
, under the topic "Towards a theoretical Biology". The workshop involved, among other key scientists, Conrad Waddington, Jack Cowan
Jack Cowan
Jack Cowan was a Canadian association football player, who played for Scottish club Dundee from 1949 to 1954. He was inducted into the Canadian Soccer Hall of Fame as a player in 2000.-External links:**...
, Michael Conrad
Michael Conrad
Michael Conrad was an American television actor.-Career:Conrad had a long acting career in television from the 1950s to the 1980s...
, Christopher Zeeman, Richard Lewontin
Richard Lewontin
Richard Charles "Dick" Lewontin is an American evolutionary biologist, geneticist and social commentator. A leader in developing the mathematical basis of population genetics and evolutionary theory, he pioneered the notion of using techniques from molecular biology such as gel electrophoresis to...
, Robert Rosen, Stuart Kauffman
Stuart Kauffman
Stuart Alan Kauffman is an American theoretical biologist and complex systems researcher concerning the origin of life on Earth...
, John Maynard Smith
John Maynard Smith
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....
, Rene Thom
René Thom
René Frédéric Thom was a French mathematician. He made his reputation as a topologist, moving on to aspects of what would be called singularity theory; he became world-famous among the wider academic community and the educated general public for one aspect of this latter interest, his work as...
and Lewis Wolpert
Lewis Wolpert
Lewis Wolpert CBE FRS FRSL is a developmental biologist, author, and broadcaster.-Career:Wolpert was educated at the University of Witwatersrand , at Imperial College London, and at King's College London...
. As a result of the conference talks and discussions, a four-volumes came out, becoming at the time a major reference in the area.
Gene networks and development
Shortly after François JacobFrançois Jacob
François Jacob is a French biologist who, together with Jacques Monod, originated the idea that control of enzyme levels in all cells occurs through feedback on transcription. He shared the 1965 Nobel Prize in Medicine with Jacques Monod and André Lwoff.-Childhood and education:François Jacob is...
and Jacques Monod
Jacques Monod
Jacques Lucien Monod was a French biologist who was awarded a Nobel Prize in Physiology or Medicine in 1965, sharing it with François Jacob and Andre Lwoff "for their discoveries concerning genetic control of enzyme and virus synthesis"...
developed their first model of gene regulation, Goodwin proposed the first model of a genetic oscillator, showing that regulatory interactions among genes allowed periodic fluctuations to occur. Shortly after this model became published, he also formulated a general theory of complex gene regulatory networks using statistical mechanics
Statistical mechanics
Statistical mechanics or statistical thermodynamicsThe terms statistical mechanics and statistical thermodynamics are used interchangeably...
.
In its simplest form, Goodwin's oscillator involves a single gene that represses itself. Goodwin equations were originally formulated in terms of conservative (Hamiltonian) systems, thus not taking into account dissipative effects that are required in a realistic approach to regulatory phenomena in biology. Many versions have been developed since then. The simplest (but realistic) formulation considers three variables, X, Y and Z indicating the concentrations of RNA
RNA
Ribonucleic acid , or RNA, is one of the three major macromolecules that are essential for all known forms of life....
, protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
and end product which generates the negative feedback
Feedback
Feedback describes the situation when output from an event or phenomenon in the past will influence an occurrence or occurrences of the same Feedback describes the situation when output from (or information about the result of) an event or phenomenon in the past will influence an occurrence or...
loop.The equations are
and closed oscillations can occur for n>8 and behave limit cycles: after a perturbation of the system's state, it returns to its previous attractor. A simple modification of this model, adding other terms introducing additional steps in the transcription machinery allows to find oscillations for smaller n values. Goodwin's model and
its extensions have been widely used over the years as the basic skeleton for other models of oscillatory behavior, including circadian clocks, cell division
Cell division
Cell division is the process by which a parent cell divides into two or more daughter cells . Cell division is usually a small segment of a larger cell cycle. This type of cell division in eukaryotes is known as mitosis, and leaves the daughter cell capable of dividing again. The corresponding sort...
or physiological
control systems.
Models in developmental biology
Later on he explored the problem of self-organization in pattern formation, using different case studies, from single-cell organisms (as AcetabulariaAcetabularia
Acetabularia is a genus of green algae, specifically of the Polyphysaceae family, Typically found in subtropical waters, Acetabularia is a single-celled organism, but gigantic in size and complex in form, making it an excellent model organism for studying cell biology...
) to multicellular organisms, including early development in Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
. One of his key contributions was to link morphogenetic fields, defined in terms of spatial distributions of chemical
signals (morphogenes), and the shape of the system experiencing morphogenetic changes. In this wasy, geometry and development were linked through a mathematical formalism. Along with his colleague Lynn Trainor, Goodwin developed a set of mathematical equations describing the changes of both physical boundaries in the porganism and chemical gradients.
By considering the mechanochemical behaviour of the cortical cytoplasm (or cytogel) of plant cells, a viscoelastic material mainly composed of actin microfilaments and reinforced by a microtubules network, Goodwin & Trainor (1985) showed how to couple calcium and the mechanical properties of the cytoplasm. The cytogel is treated as a continuous viscoelastic medium in which calcium ions can diffuse and interact with the cytoskeleton. The model consists in two non-linear partial differential equations which describe the evolution of the mechanical strain field and .of the calcium distribution in the cytogel.
It has been shown (Trainor & Goodwin, 1986) that, in a range of parameter values, instabilities may occur and develop in this system, leading to intracellular patterns of strain and calcium concentration. The equations read, in their general form:
These equations describe the spatiotemporal dynamics of the displacement from the reference state and the calcium concentration, respectively. Here
x and t are the space and time coordinates, respectively. These equations
can be applied to many different scenarios and the different functions P(x) introduce the specific mechanical properties of the medium. These equations
are very rich in terms of the static and dynamic patterns that can generate, including both complex geometrical motifs to oscillations
and chaos (Briere 1994).
Structuralism
He was also a strong advocate of the view that genes cannot fully explain the complexityComplexity
In general usage, complexity tends to be used to characterize something with many parts in intricate arrangement. The study of these complex linkages is the main goal of complex systems theory. In science there are at this time a number of approaches to characterizing complexity, many of which are...
of biological systems. In that sense, he became one of the strongest defenders of the systems view against reductionism
Reductionism
Reductionism can mean either an approach to understanding the nature of complex things by reducing them to the interactions of their parts, or to simpler or more fundamental things or a philosophical position that a complex system is nothing but the sum of its parts, and that an account of it can...
. Among other contributions, he suggested that nonlinear phenomena and the fundamental laws defining their behavior were essential in order to understand biology and its evolutionary paths. His position within evolutionary biology can be defined as a structuralist
Structuralism (biology)
Biological or process structuralism is a school of biological thought that deals with the law-like behaviour of the structure of organisms and how it can change....
one. To Goodwin, many patterns that we observe in nature are a byproduct of constraints imposed by complexity. The limited repertoire of motifs observed in the spatial organization of plants and animals (at some scales) would be, in Goodwin's opinion, a fingerprint of the role played by such constraints. The role of selection would be secondary.
These opinions were highly controversial and
Goodwin came into conflict with many prominent Darwinian evolutionists, whereas many physicists found some of his view natural. Physicist Murray Gell-Mann
Murray Gell-Mann
Murray Gell-Mann is an American physicist and linguist who received the 1969 Nobel Prize in physics for his work on the theory of elementary particles...
for example acknowledged that "when biological evolution — based on largely random variation in genetic material and on natural selection — operates on the structure of actual organisms, it does so subject to the laws of physical science, which place crucial limitations on how living things can be constructed.". Richard Dawkins
Richard Dawkins
Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...
, the former professor for public understanding of science at Oxford University and a well known Darwinian evolutionist, conceded: "I don't think there's much good evidence to support [his thesis], but it's important that somebody like Brian Goodwin is saying that kind of thing, because it provides the other extreme, and the truth probably lies somewhere between." Dawkins also agreed that "It's a genuinely interesting possibility that the underlying laws of morphology allow only a certain limited range of shapes.". For his part, Goodwin did not reject basic Darwinism, only its excesses.
Academic life
Thereafter, he taught at the Schumacher CollegeSchumacher College
Schumacher College was founded in 1991 in Dartington, Totnes, Devon, UK by Satish Kumar, John Lane and others. It was named after E.F. Schumacher. It is an international centre offering transformative learning for sustainable living, and runs holistic education courses...
in Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...
, UK
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...
, where he was instrumental in starting the college's MSc in Holistic Science. He was made a Founding Fellow of Schumacher College shortly before his death. Goodwin also had a research position at MIT and was a long time visitor of several institutions including the UNAM
Unam
UNAM or UNaM may refer to:* National University of Misiones, a National University in Posadas, Argentina*National Autonomous University of Mexico , the large public autonomous university based in Mexico City...
in Mexico City. He was a founding member of the Santa Fe Institute
Santa Fe Institute
The Santa Fe Institute is an independent, nonprofit theoretical research institute located in Santa Fe and dedicated to the multidisciplinary study of the fundamental principles of complex adaptive systems, including physical, computational, biological, and social systems.The Institute houses a...
in New Mexico
New Mexico
New Mexico is a state located in the southwest and western regions of the United States. New Mexico is also usually considered one of the Mountain States. With a population density of 16 per square mile, New Mexico is the sixth-most sparsely inhabited U.S...
where he also served as a member of the science board for several years.
Brian Goodwin died in hospital in 2009, after surgery resulting from a fall from his bike.. Goodwin is survived by his third wife, Christel, and his daughter, Lynn.
Publications
Books- 1989. Theoretical Biology: Epigenetic and Evolutionary Order for Complex Systems with Peter Saunders, Edinburgh University Press, 1989, ISBN 0852246005
- 1994. Mechanical Engineering of the Cytoskeleton in Developmental Biology (International Review of Cytology), with Kwang W. Jeon and Richard J. Gordon, Academic Press, London 1994, ISBN 0123645530
- 1996. Form and Transformation: Generative and Relational Principles in Biology, Cambridge Univ Press, 1996.
- 1997. How the Leopard Changed its Spots: The Evolution of Complexity, Scribner, 1994, ISBN 0025447106 (German: Der Leopard, der seine Flecken verliert, Piper, München 1997, ISBN 3492038735)
- 2001. Signs of Life: How Complexity Pervades Biology, with Ricard V. Sole, Basic Books, 2001, ISBN 0465019277
- 2007. Nature's Due: Healing Our Fragmanted Culture, Floris Books, 2007, ISBN 0863155960
Selected Scientific papers
- 1997, "Temporal organization and disorganization in organisms". in: Chronobiology International 14 (5): 531-536 1997
- 2000, "The life of form. Emergent patterns of morphological transformation". in: Comptes rendud de la Academie des Science III 323 (1): 15-21 JAN 2000
- Miramontes O, Solé RV, Goodwin BC (2001). Neural networks as sources of chaotic motor activity in ants and how complexity develops at the social scale. International Journal of bifurcation and chaos 11 (6): 1655-1664.
- Goodwin BC (2000). The life of form. Emergent patterns of morphological transformation. Comptes rendus de l'academie des sciencies III - Sciences de la vie-life sciences 323 (1): 15-21
- Goodwin BC. (1997) Temporal organization and disorganization in organisms. Chronobiology international 14 (5): 531-536
- Solé, R., O. Miramontes y Goodwin BC. (1993)Collective Oscillations and Chaos in the Dynamics of Ant Societies. J. Theor. Biol. 161: 343
- Miramontes, O., R. Solé y BC Goodwin (1993), Collective Behaviour of Random-Activated Mobile Cellular Automata. Physica D 63: 145-160
Essays
- 2002, "In the Shadow of Culture". in: "The Next Fifty Years: Science in the First Half of the Twenty-First Century" Edited by John Brockman, Vintage Books, MAY 2002, ISBN 0-375-71342-5