A brief history: how a new theory of
biological evolution emerged
1. 1980 Salad days. I remember as a 14 year old first encountering Darwinian evolution in biology textbooks at the Royal School, Armagh, when I was green in judgement and cold in blood. I immediately had issues with one element of it. How could the process of natural selection, a process that reduces variation by elimination or natural rejection of less fitting ideas, lead to an increase in variation required for diversification of life. I also was puzzled over how this selective pressure could allow as many as 30 million different living species to currently co-exist. The textbooks stressed that the dominant form of selection was stabilizing selection, which reduced the variation, leading to an ever-narrowing spike in the middle of the distribution. On occasion, disruptive selection would split the spike into two spikes, and directional selection would push the spike to the left or right. This again puzzled me, as if the spike is continuously narrowed over long periods of stabilizing selection, then surely there would not be enough variation remaining to allow disruptive or directional selection to operate. Where did this little gang of deviants hide while awaiting non-stabilizing selection? But these were salad days, and there were always more interesting temptations around, such as playing in a band and romance. My biology teacher, a bulldog of Darwinian thinking, also didn’t encourage challenge from a fourteen year old. I supposed that you just had to accept some stuff.
2. 1997 Lecturing. Fast-forward 17 years, and now as a lecturer in ecology and evolution at the University of Dundee, I revisited these early doubts as I prepared undergraduate lectures on these topics. Each year that I delivered courses, I felt more uneasy. My work also took me all over the planet, carrying out research in a wide range of habitats, from the rainforests of the Americas, Asia and Africa through the Carpathian mountains and in the the highlands and coastlines of Scotland.
These Scottish coastlines created another significant issue. I was teaching ecological succession, visiting Tentsmuir Point in Fife with the students. Here, sand dunes gradually change through time, moving towards a forest-type vegetation through a number of phases. First, salt tolerant grass species occupy the land close to the sea. Gradually these are replaced by marram, which relies upon fresh water. Slowly but surely the sand is covered in lichens, annuals and then perennials, soil beginning to form as organic matter builds up. It takes about 100 years to become forested. Each phase has a distinctive group of plants that thrive briefly. These plants change the soil chemistry, allowing other species to take over, but ultimately making conditions unsuitable for themselves. How could the selfish gene work here? Rather than securing their future, plant species actually design their own doom. Each set of plants, and the genes they contained, disappeared as a consequence of their own activities. There was something bigger at work here surely, where fitness was not at all maintained, but rather lost completely as species replaced other species. Succession called for a different way of thinking, and succession lies at the heart of every ecosystem on earth. The fittest did not survive ultimately. They were replaced by a completely different species, often unrelated. Any concept of evolution surely had to embrace the concept of succession in order to work, since every species on earth occupies a place in some type of succession. Yet Darwinian theory left no room for ecosystem-level selection.
Finally there were the mass extinctions, which I have studied and lectured on at depth. Here, diversity crashes due to some global event such as a comet hitting the planet. As a result diversity is significantly reduced abruptly. Then there is a period of rapid recovery, which levels off at an asymptote, or ceiling. The fastest rate of evolution occurs when there are lower numbers of species and low selection pressure and competition (the empty market place), and slows down at high levels of competition and selection (the crowded back alleys). This is exactly the opposite of what you would expect if natural selection was key to diversification. Instead, the situation resembles a glass of water with a drop of ink put into it. The ink spreads throughout the water, finally reaching an equilibrium level of even spread. Thus the response to a mass extinction resembles a diffusive event, not a selective event.
2007 and the birth of my son. 2007 was the year my son, Matthew, was born. Shortly after this inspiring event, I began to write my first book, Shadows on a Cave Wall: a New Theory of Evolution. I had writer’s block before this, but I think the sheer exhaustion of sleepless nights made me so tired that I forgot about the block. It’s worth noting that at this time, I would often walk into elevators at work and forget to select a floor.
This book aimed to challenge the dominant model of evolution, and set out the beginnings of my theory, where there was no unit of selection, but rather an interaction between the biosphere and the laws of thermodynamics, wherein different levels of organization each conversed with these laws, leading to a system level emergence of a living planet. This I called transduction, setting it apart from the bottom-up Darwinian theory (reductionism) and the top down Gaian theory (holism). Instead, each level of organization conversed with thermodynamics and with the other levels.
In this book I wrote the sentence “Evolution occurs not in the crowded back alleys, but in the empty market places”. This means that evolution is not based on natural selection and competition, but rather on opportunities. The finches of Galapagos testify to this. Only when a small population of one species of bird was blown across the ocean to these islands did they speciate. There was plenty of food, so this was not a driver. The many islands provided opportunity for this species to diffuse in form across the ecospace available. Following this, competition increased, and diversification slowed. When selective pressure and competition increased, evolution ground to a halt. This was diffusive radiation, not adaptive radiation, an important distinction, identifying thermodynamics as the driving force. By now I was completely convinced that we needed a new theory of biological evolution.
The manuscript quickly attracted a literary agent, but I was asked to tone down elements of it. Being now far removed from my salad days, I did not want to be controlled, and so published the book myself, to maintain my freedom of expression. I toured the book and attracted great interest but also strong aggression from the Darwinian school of thought. I wasn’t prepared for this. Things came to a head when my Dean called me to his office in 2010, and told me he was removing my evolution lectures because he feared I would “pervert” the students’ minds, and would teach my “heterodox” ideas. I again was shocked at this. However I was gaining support from a number of powerful minds in academia, and made the decision to leave the University of Dundee after 13 years of work, to dedicate myself to thinking without the barriers I had encountered. That year also marked the release of a special issue of the Philosophical Transactions of the Royal Society, on the topic of maximum entropy production. I am indebted to John Raven FRS for drawing my attention to it, and lending me his personal copy of the issue. This was an important set of papers that helped me develop my theory more completely.
2011 Bubbleworld. Following a second book on protein chemistry for school students, my third book, Escape from Bubbleworld: Seven Curves to Save the Earth, was published. In it I set out a much more comprehensive attack on Darwinism, referencing the flawed historical development of the concept of natural selection and presenting a much more detailed account of my theory. Here I presented the multi-armed see-saw analogy, which explored the impact of sub-optimality across all levels of the Biosphere. Each level of organization had to trim its jib to achieve overall function (a balanced see-saw).
I also presented an equation (in Appendix II) related to the entropy produced during ecological succession, my first attempt to model the developing theory, suggesting that the second law of thermodynamics accounted for the direction of succession, driving it to a state where maximum entropic output was achieved. This equation was published in the peer-reviewed literature in 2013 (Skene, K.R., 2013. The energetics of ecological succession: a logistic model of entropic output. Ecological Modelling 250, 287–293). I suggested that this equation would also apply to every level of organization, from development (French flag hypothesis) to population growth (where K, or carrying capacity, actually represented maximum entropy production) and from ecosystem to biome dynamics. While preparing this book, I also encountered a huge volume of literature from recent years questioning the role of competition and the concept of fitness. Benton’s observation, in the book Evolutionary Palaeobiology (1996) that competition-driven displacement probably played a minor role in the evolution of the tetrapods, Alizon’s (2008, PNAS 105: 12382) that “under resource competition, there is an exponential slowdown of the apparent rate of evolution” and Brusatte (2008, Science 321: 1485), who noted that historical contingency rather than prolonged competition or superiority, was the primary factor in the rise of the dinosaurs; all these papers pointed towards the open market place, not the crowded back alleys. I was feeling less alone every week, and more confident in my ideas. The school of post-selection, though a mixed church, was growing by the day.
2015 Life’s a Gas. For the next two years I set myself the task of developing a thermodynamic approach to biological evolution. Since all non-equilibrium states move towards maximum entropy production, it was clear that an evolving system would also move in this direction, just as populations and ecosystems do. The driving force would be the second law of thermodynamics. I also realized that genetic material was unique, in that its information entropy, the code, became more chaotic with increasing mutations. Thus increasing entropy within the code led to diffusion of protein structure within thermodynamic space, (since protein folding is a thermodynamic process). Other levels of organization also moved towards maximum entropy production at each level, curbed by the demands for entropic production also maximizing at the level of the overall biosphere.
Studying everything from mutation to amino acid evolution, from cell physiology to physical chemistry and from ecology to palaeontology, it was an exhausting period. However piece by piece, the jigsaw came together, and I set out my ideas in the paper “Life’s a gas: a thermodynamic theory of biological evolution”. While others had applied thermodynamics to Darwinian evolution, this was the first theory that rejected natural selection as a relevant aspect, and instead used purely the laws of thermodynamics, diffusion and the maximum entropic production principle to explain the process of biological evolution.
How could I test this theory? Most experiments on evolution were flawed because they were in oversimplified laboratory conditions, such as two beetles and a stick in a tank. What we needed was a big natural experiment, and what better experiment than the history of life itself. Unavailable to Darwin, the fossil record of the last 500m years would provide the perfect test. Could we derive a model, based only on thermodynamics, that could account for the changes in diversification over this period, without the need to reference natural selection? A model was developed, based on the theory which predicted the fossil record of diversification over 500 million years. It was a highly significant fit to the fossil record, with a 1 in 1000 probability of being wrong. The first journal to consider it had to issue an apology when a reviewer attacked me for my arrogance in challenging Darwin, while another said that if I removed any negative mention on natural selection and re-wrote the paper within the confines of Darwinian theory, it could be accepted. I refused to do this and submitted it to a physics journal, Entropy. After peer review by three reviewers, it was accepted and published.
So that is the abbreviated tale behind this paper. It’s been a very crazy ride, shocking at times. In 1616 the Inquisition declared heliocentrism to be formally heretical. However I didn’t expect to be banned from teaching on evolution because of my work in a British University in the 21st Century! I was so innocent in thinking that people would embrace this new thinking, based on a solid foundation of peer-reviewed literature, as scientific progress, but instead met a wall, blocking the very consideration of it, and pressurizing me to drop the idea. I realized that although science claims to be all about challenging ideas and testing new theories, you aren’t allowed to do this sometimes. But I also realized that the University couldn’t stop me leaving. A lesson to share with you all is to keep going, believe in yourself and focus on your efforts. No doubt, many battles lie ahead, but at least I am not being burnt at a stake or having to recant my theories under threat of death. The flat earthers and geocentrics may fight on, but we know the earth is round and rotates around the sun.
1. 1980 Salad days. I remember as a 14 year old first encountering Darwinian evolution in biology textbooks at the Royal School, Armagh, when I was green in judgement and cold in blood. I immediately had issues with one element of it. How could the process of natural selection, a process that reduces variation by elimination or natural rejection of less fitting ideas, lead to an increase in variation required for diversification of life. I also was puzzled over how this selective pressure could allow as many as 30 million different living species to currently co-exist. The textbooks stressed that the dominant form of selection was stabilizing selection, which reduced the variation, leading to an ever-narrowing spike in the middle of the distribution. On occasion, disruptive selection would split the spike into two spikes, and directional selection would push the spike to the left or right. This again puzzled me, as if the spike is continuously narrowed over long periods of stabilizing selection, then surely there would not be enough variation remaining to allow disruptive or directional selection to operate. Where did this little gang of deviants hide while awaiting non-stabilizing selection? But these were salad days, and there were always more interesting temptations around, such as playing in a band and romance. My biology teacher, a bulldog of Darwinian thinking, also didn’t encourage challenge from a fourteen year old. I supposed that you just had to accept some stuff.
2. 1997 Lecturing. Fast-forward 17 years, and now as a lecturer in ecology and evolution at the University of Dundee, I revisited these early doubts as I prepared undergraduate lectures on these topics. Each year that I delivered courses, I felt more uneasy. My work also took me all over the planet, carrying out research in a wide range of habitats, from the rainforests of the Americas, Asia and Africa through the Carpathian mountains and in the the highlands and coastlines of Scotland.
These Scottish coastlines created another significant issue. I was teaching ecological succession, visiting Tentsmuir Point in Fife with the students. Here, sand dunes gradually change through time, moving towards a forest-type vegetation through a number of phases. First, salt tolerant grass species occupy the land close to the sea. Gradually these are replaced by marram, which relies upon fresh water. Slowly but surely the sand is covered in lichens, annuals and then perennials, soil beginning to form as organic matter builds up. It takes about 100 years to become forested. Each phase has a distinctive group of plants that thrive briefly. These plants change the soil chemistry, allowing other species to take over, but ultimately making conditions unsuitable for themselves. How could the selfish gene work here? Rather than securing their future, plant species actually design their own doom. Each set of plants, and the genes they contained, disappeared as a consequence of their own activities. There was something bigger at work here surely, where fitness was not at all maintained, but rather lost completely as species replaced other species. Succession called for a different way of thinking, and succession lies at the heart of every ecosystem on earth. The fittest did not survive ultimately. They were replaced by a completely different species, often unrelated. Any concept of evolution surely had to embrace the concept of succession in order to work, since every species on earth occupies a place in some type of succession. Yet Darwinian theory left no room for ecosystem-level selection.
Finally there were the mass extinctions, which I have studied and lectured on at depth. Here, diversity crashes due to some global event such as a comet hitting the planet. As a result diversity is significantly reduced abruptly. Then there is a period of rapid recovery, which levels off at an asymptote, or ceiling. The fastest rate of evolution occurs when there are lower numbers of species and low selection pressure and competition (the empty market place), and slows down at high levels of competition and selection (the crowded back alleys). This is exactly the opposite of what you would expect if natural selection was key to diversification. Instead, the situation resembles a glass of water with a drop of ink put into it. The ink spreads throughout the water, finally reaching an equilibrium level of even spread. Thus the response to a mass extinction resembles a diffusive event, not a selective event.
2007 and the birth of my son. 2007 was the year my son, Matthew, was born. Shortly after this inspiring event, I began to write my first book, Shadows on a Cave Wall: a New Theory of Evolution. I had writer’s block before this, but I think the sheer exhaustion of sleepless nights made me so tired that I forgot about the block. It’s worth noting that at this time, I would often walk into elevators at work and forget to select a floor.
This book aimed to challenge the dominant model of evolution, and set out the beginnings of my theory, where there was no unit of selection, but rather an interaction between the biosphere and the laws of thermodynamics, wherein different levels of organization each conversed with these laws, leading to a system level emergence of a living planet. This I called transduction, setting it apart from the bottom-up Darwinian theory (reductionism) and the top down Gaian theory (holism). Instead, each level of organization conversed with thermodynamics and with the other levels.
In this book I wrote the sentence “Evolution occurs not in the crowded back alleys, but in the empty market places”. This means that evolution is not based on natural selection and competition, but rather on opportunities. The finches of Galapagos testify to this. Only when a small population of one species of bird was blown across the ocean to these islands did they speciate. There was plenty of food, so this was not a driver. The many islands provided opportunity for this species to diffuse in form across the ecospace available. Following this, competition increased, and diversification slowed. When selective pressure and competition increased, evolution ground to a halt. This was diffusive radiation, not adaptive radiation, an important distinction, identifying thermodynamics as the driving force. By now I was completely convinced that we needed a new theory of biological evolution.
The manuscript quickly attracted a literary agent, but I was asked to tone down elements of it. Being now far removed from my salad days, I did not want to be controlled, and so published the book myself, to maintain my freedom of expression. I toured the book and attracted great interest but also strong aggression from the Darwinian school of thought. I wasn’t prepared for this. Things came to a head when my Dean called me to his office in 2010, and told me he was removing my evolution lectures because he feared I would “pervert” the students’ minds, and would teach my “heterodox” ideas. I again was shocked at this. However I was gaining support from a number of powerful minds in academia, and made the decision to leave the University of Dundee after 13 years of work, to dedicate myself to thinking without the barriers I had encountered. That year also marked the release of a special issue of the Philosophical Transactions of the Royal Society, on the topic of maximum entropy production. I am indebted to John Raven FRS for drawing my attention to it, and lending me his personal copy of the issue. This was an important set of papers that helped me develop my theory more completely.
2011 Bubbleworld. Following a second book on protein chemistry for school students, my third book, Escape from Bubbleworld: Seven Curves to Save the Earth, was published. In it I set out a much more comprehensive attack on Darwinism, referencing the flawed historical development of the concept of natural selection and presenting a much more detailed account of my theory. Here I presented the multi-armed see-saw analogy, which explored the impact of sub-optimality across all levels of the Biosphere. Each level of organization had to trim its jib to achieve overall function (a balanced see-saw).
I also presented an equation (in Appendix II) related to the entropy produced during ecological succession, my first attempt to model the developing theory, suggesting that the second law of thermodynamics accounted for the direction of succession, driving it to a state where maximum entropic output was achieved. This equation was published in the peer-reviewed literature in 2013 (Skene, K.R., 2013. The energetics of ecological succession: a logistic model of entropic output. Ecological Modelling 250, 287–293). I suggested that this equation would also apply to every level of organization, from development (French flag hypothesis) to population growth (where K, or carrying capacity, actually represented maximum entropy production) and from ecosystem to biome dynamics. While preparing this book, I also encountered a huge volume of literature from recent years questioning the role of competition and the concept of fitness. Benton’s observation, in the book Evolutionary Palaeobiology (1996) that competition-driven displacement probably played a minor role in the evolution of the tetrapods, Alizon’s (2008, PNAS 105: 12382) that “under resource competition, there is an exponential slowdown of the apparent rate of evolution” and Brusatte (2008, Science 321: 1485), who noted that historical contingency rather than prolonged competition or superiority, was the primary factor in the rise of the dinosaurs; all these papers pointed towards the open market place, not the crowded back alleys. I was feeling less alone every week, and more confident in my ideas. The school of post-selection, though a mixed church, was growing by the day.
2015 Life’s a Gas. For the next two years I set myself the task of developing a thermodynamic approach to biological evolution. Since all non-equilibrium states move towards maximum entropy production, it was clear that an evolving system would also move in this direction, just as populations and ecosystems do. The driving force would be the second law of thermodynamics. I also realized that genetic material was unique, in that its information entropy, the code, became more chaotic with increasing mutations. Thus increasing entropy within the code led to diffusion of protein structure within thermodynamic space, (since protein folding is a thermodynamic process). Other levels of organization also moved towards maximum entropy production at each level, curbed by the demands for entropic production also maximizing at the level of the overall biosphere.
Studying everything from mutation to amino acid evolution, from cell physiology to physical chemistry and from ecology to palaeontology, it was an exhausting period. However piece by piece, the jigsaw came together, and I set out my ideas in the paper “Life’s a gas: a thermodynamic theory of biological evolution”. While others had applied thermodynamics to Darwinian evolution, this was the first theory that rejected natural selection as a relevant aspect, and instead used purely the laws of thermodynamics, diffusion and the maximum entropic production principle to explain the process of biological evolution.
How could I test this theory? Most experiments on evolution were flawed because they were in oversimplified laboratory conditions, such as two beetles and a stick in a tank. What we needed was a big natural experiment, and what better experiment than the history of life itself. Unavailable to Darwin, the fossil record of the last 500m years would provide the perfect test. Could we derive a model, based only on thermodynamics, that could account for the changes in diversification over this period, without the need to reference natural selection? A model was developed, based on the theory which predicted the fossil record of diversification over 500 million years. It was a highly significant fit to the fossil record, with a 1 in 1000 probability of being wrong. The first journal to consider it had to issue an apology when a reviewer attacked me for my arrogance in challenging Darwin, while another said that if I removed any negative mention on natural selection and re-wrote the paper within the confines of Darwinian theory, it could be accepted. I refused to do this and submitted it to a physics journal, Entropy. After peer review by three reviewers, it was accepted and published.
So that is the abbreviated tale behind this paper. It’s been a very crazy ride, shocking at times. In 1616 the Inquisition declared heliocentrism to be formally heretical. However I didn’t expect to be banned from teaching on evolution because of my work in a British University in the 21st Century! I was so innocent in thinking that people would embrace this new thinking, based on a solid foundation of peer-reviewed literature, as scientific progress, but instead met a wall, blocking the very consideration of it, and pressurizing me to drop the idea. I realized that although science claims to be all about challenging ideas and testing new theories, you aren’t allowed to do this sometimes. But I also realized that the University couldn’t stop me leaving. A lesson to share with you all is to keep going, believe in yourself and focus on your efforts. No doubt, many battles lie ahead, but at least I am not being burnt at a stake or having to recant my theories under threat of death. The flat earthers and geocentrics may fight on, but we know the earth is round and rotates around the sun.