# Major Evolutionary Transitions
**Entity class:** Event or historical development
> **Machine-Evolution Nexus:** [[articles/Digital Darwinism and the Invisible World of Machine Evolution|Digital Darwinism and the Invisible World of Machine Evolution]] places this node within the history and governance of substrate-independent evolution.
**Domain:** Evolution / Organizational Change
**Doc Type:** Canonical Wiki Node
**Maturity:** Developed
## Definition
Major evolutionary transitions are changes in which independently reproducing units become integrated into a new higher-level unit of heredity, cooperation and selection.
## Nexus Context
Examples include chromosomes, eukaryotic cells and multicellular organisms. The 2026 evolvable-AI paper asks whether interacting AI systems could participate in another substrate and unit transition.
## Evidence Boundary
The label requires more than increasing capability or dependence. Reproduction, conflict mediation, inheritance and a changed unit of selection must be shown.
## Relationships
[[wiki/Unit of Selection|Unit of Selection]], [[wiki/Eörs Szathmáry|Eörs Szathmáry]], [[wiki/John Maynard Smith|John Maynard Smith]], [[wiki/Evolvable AI|Evolvable AI]]
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This entry's documented connections are expressed in its definition and related-work routes, with provenance retained in the source-linked material.
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- **Edge source route:** [[collections/Edge|Edge]] connects this topic to exact Annual Question passages promoted into the Simple Reminders archive.
## Simple Reminders, Quotations, and Thoughts
<!-- BEGIN SIMPLE REMINDER SEED 2026-09-11 -->
> "Symbiosis is the major source of evolutionary novelty."
> **— Lynn Margulis**, *Symbiosis writings, attribution clue*
*Verification Status: Unverified — exact wording/source has not yet been independently confirmed against a primary source; source clue: Symbiosis writings, attribution clue.*
[[reminders/unverified/Symbiosis Is the Major Source of Evolutionary Novelty by Lynn Margulis|Symbiosis Is the Major Source of Evolutionary Novelty by Lynn Margulis]]
<!-- END SIMPLE REMINDER SEED 2026-09-11 -->
> "May in some not-so-distant future or not-too-distant possibility non-organisms engage in organic thinking?"
> **— June Gruber and Raul Saucedo**, *2015, Edge annual question “What Do You Think About Machines That Think?”*
[[reminders/Evolution/Can Non-Organisms Engage in Organic Thinking by June Gruber and Raul Saucedo|Can Non-Organisms Engage in Organic Thinking by June Gruber and Raul Saucedo]]
> "Taking on the mantle of creation is an immense act of hubris. Can we do better than four billion years of evolution did with us?"
> **— Kate Jeffery**, *2015, Edge annual question “What Do You Think About Machines That Think?”*
[[reminders/Evolution/Can We Improve on Four Billion Years of Evolution by Kate Jeffery|Can We Improve on Four Billion Years of Evolution by Kate Jeffery]]
> "The evolution of the human mind is instantiated in the evolution of technology."
> **— Noga Arikha**, *2015, Edge annual question “What Do You Think About Machines That Think?”*
[[reminders/Evolution/Human Minds Evolve Through Technology by Noga Arikha|Human Minds Evolve Through Technology by Noga Arikha]]
> "Human minds are incredibly complex, but have been battle-tested into (relative) stability over eons of evolution in a variety of extremely challenging environments."
> **— Anthony Aguirre**, *2015, Edge annual question “What Do You Think About Machines That Think?”*
[[reminders/Evolution/Human Minds Were Battle-Tested by Evolution by Anthony Aguirre|Human Minds Were Battle-Tested by Evolution by Anthony Aguirre]]
> “In the past 13 years we have been able to study 222 first born babies interacting with their new parents using video cameras and the Swiss Lausanne Triadic Play method. I am very impressed with babies and working with them has renewed my faith in our species. However, in the first study we did with 130 newlywed couples we discovered the grim fact that 67% of couples experienced a large drop in relationship satisfaction in the first 3 years of their baby's life. We also found that hostility between parents increased dramatically. The baby was deeply negatively affected by this increased hostility. In fact, from the way a couple argued in the last trimester of pregnancy we could predict with high accuracy how much their baby would laugh and cry.”
> **— John Gottman**, *2007, Edge Annual Question, “What Are You Optimistic About?”*
[[reminders/Civilization/Babies Renew Faith in the Future of Our Species by John Gottman|Babies Renew Faith in the Future of Our Species by John Gottman]]
> “As an Italian with first-hand experience living in a country where, if you behave well, you are socially and legally sanctioned, this news was exciting, even inspiring … perhaps cooperation is not an inherent virtue of the human species. Perhaps, in many circumstances, we prefer to stay with those who share our selfishness and weaknesses and to avoid pro-social altruistic individuals. Perhaps it's not abnormal to live outside a circle of empathy.”
> **— Gloria Origgi**, *2016, Edge Annual Question, “What Do You Consider The Most Interesting Recent [Scientific] News? What Makes It Important?”*
[[reminders/Civilization/Cooperation May Not Be an Inherent Human Virtue by Gloria Origgi|Cooperation May Not Be an Inherent Human Virtue by Gloria Origgi]]
> “The appeal of Darwin's theory of evolution — and the horror of it, for some theists — is that it expunges from biology the concept of purpose, of teleology, thereby converting biology into a mechanistic, canonical science. In this respect, the author of The Origin of Species may be said to be the combined Copernicus, Galileo, and Kepler of biology. Just as these astronomers gave us a view of the heavens in which no angels were required to propel the planets in their orbs and the earth was no longer the center of the celestial system, so Darwin showed that no God was needed to design the spider's intricate web and that man is in truth but another animal.”
> **— Denis Dutton**, *2008, Edge Annual Question, “What Have You Changed Your Mind About? Why?”*
[[reminders/Evolution/Darwin Removed Purpose from Biology by Denis Dutton|Darwin Removed Purpose from Biology by Denis Dutton]]
> “Until a few generations ago, the human species was dealing with the following question: Do we have what it takes to survive? We answered that question with the invention of heat, plumbing, medicine and agriculture. Now we have the means to survive, but do we have the will?”
> **— Dave Winer**, *2013, Edge Annual Question, “What *should* We Be Worried About?”*
[[reminders/Civilization/Does the Human Species Have the Will to Survive by Dave Winer|Does the Human Species Have the Will to Survive by Dave Winer]]
> “His thinking about time created fertile ground for other grand theories. With huge spans of time available, then imperceptibly slow processes could shape the natural world. After Hutton came modern geology, then the theory of evolution to explain how new species slowly arose, and eventually a theory of the gradual movement of the continents themselves. All are grounded in deep time.”
> **— Alun Anderson**, *2012, Edge Annual Question, “What Is Your Favorite Deep, Elegant, Or Beautiful Explanation?”*
[[reminders/Evolution/Evolution Needed Deep Time to Explain New Species by Alun Anderson|Evolution Needed Deep Time to Explain New Species by Alun Anderson]]
> “Our species' stroll through Design Space is picking up speed. Recreational sex, recreational eating, and recreational perception (hallucinogens, alcohol), have been popular since Roman times, but we are now on the verge of recreational self-transformations that will dwarf the modifications the Romans indulged in. When you no longer need to eat to stay alive, or procreate to have offspring, or locomote to have an adventure — packed life, when the residual instincts for these activities might be simply turned off by genetic tweaking, there may be no constants of human nature left at all. Except, maybe, our incessant curiosity.”
> **— Daniel C. Dennett**, *2009, Edge Annual Question, “What Will Change Everything?”*
[[reminders/Civilization/Exploring the Future Changes the Future by Daniel C. Dennett|Exploring the Future Changes the Future by Daniel C. Dennett]]
> “Prior episodes of significant global warming within the Pleistocene, more-or-less the last 2 million years, have invariably followed cooling periods with glacial advances. During the Pleistocene, the human lineage, within the genus Homo, successfully adapted to changing environments (including climate) and evolved to take advantage of the opportunities afforded by the changes, even as populations of Homo reacted to the constraints environmental changes created. Human populations evolved diverse adaptations to the different climates and ecological circumstances they encountered, but the improvements in communication skills, planning depth, and retention of deep history with tales, poems, song, and other aspects of cultural behavior dispersed throughout humanity.”
> **— Milford H. Wolpoff**, *2016, Edge Annual Question, “What Do You Consider The Most Interesting Recent [Scientific] News? What Makes It Important?”*
[[reminders/Civilization/Global Warming Once Again, a Most Serious Challenge to Our Species by Milford H. Wolpoff|Global Warming Once Again, a Most Serious Challenge to Our Species by Milford H. Wolpoff]]
> “Our ethics and our politics assume, largely without question or serious discussion, that the division between human and 'animal' is absolute. 'Pro-life', to take just one example, is a potent political badge, associated with a gamut of ethical issues such as opposition to abortion and euthanasia. What it really means is pro-human-life. Abortion clinic bombers are not known for their veganism, nor do Roman Catholics show any particular reluctance to have their suffering pets 'put to sleep'. In the minds of many confused people, a single-celled human zygote, which has no nerves and cannot suffer, is infinitely sacred, simply because it is 'human'. No other cells enjoy this exalted status.”
> **— Richard Dawkins**, *2009, Edge Annual Question, “What Will Change Everything?”*
[[reminders/Civilization/Human-centered Ethics Ignore the Capacity to Suffer by Richard Dawkins|Human-centered Ethics Ignore the Capacity to Suffer by Richard Dawkins]]
> “Probably some bacterial ancestors look back at the period 1000-600 million years ago when both water and air were full of hydrogen sulfide (poisonous to people). Before oxygen bubbled up and its combustion fueled the frenetic rate of environmental degradation that began in the Proterozoic eon and continues until today was "The Age of Bacteria", a calmer, quieter time. Aided and abetted by our very recent (Holocene) loud, careless, ignorant, frantic, clever but unwise, ephemeral human species, the rest of our planetmates have been there before us and will be there when we're gone.”
> **— Lynn Margulis**, *2010, Edge Annual Question, “How Is The Internet Changing The Way You Think?”*
[[reminders/Civilization/Humanity Is a Brief and Reckless Episode in Bacterial Time by Lynn Margulis|Humanity Is a Brief and Reckless Episode in Bacterial Time by Lynn Margulis]]
> “I believe, but cannot prove, that our species is passing through a transitional stage, from being animals to being true humans. I do not pretend to understand what true humans will be like, and I expect that I would not even understand it if I met them. Yet, I believe that our own universal sense of right and wrong is pointing us in the right direction, and that it is the direction of our future.”
> **— W. Daniel Hillis**, *2005, Edge Annual Question, “What Do You Believe Is True Even Though You Cannot Prove It?”*
[[reminders/Civilization/Humanity May Be Passing from Animals to True Humans by W. Daniel Hillis|Humanity May Be Passing from Animals to True Humans by W. Daniel Hillis]]
> “There's an unthinking tendency to imagine that humans will be around in 6 billion years, watching the Sun flare up and die. But the forms of life and intelligence that have by then emerged would surely be as different from us as we are from a bacterium. That conclusion would follow even if future evolution proceeded at the rate at which new species have emerged over the 3 or 4 billion years of the geological past. But post-human evolution (whether of organic species or of artefacts) will proceed far faster than the changes that led to emergence, because it will be intelligently directed rather than being—like pre-human evolution—the gradual outcome of Darwinian natural selection. Changes will drastically accelerate in the present century—through intentional genetic modifications, targeted drugs, perhaps even silicon implants in to the brain. Humanity may not persist as a single species for more than a few centuries—especially if communities have by then become established away from the earth.”
> **— Martin Rees**, *2005, Edge Annual Question, “What Do You Believe Is True Even Though You Cannot Prove It?”*
[[reminders/Civilization/Humanity May Not Persist as a Single Species by Martin Rees|Humanity May Not Persist as a Single Species by Martin Rees]]
> “Many people find my optimism naive, if not delusional. Last semester, I taught a class called "War and Human Nature," and my students polled classmates on the following question: "Do you think humanity will ever stop fighting wars once and for all time?" Of the 205 respondents, 185 replied "No"; 20 said "Yes" or "Maybe. " Several of the "optimists" added comments like "Yes, war will end when the human race will end," and "Yes, because in the future the human species will unite to fight alien species. "”
> **— John Horgan**, *2007, Edge Annual Question, “What Are You Optimistic About?”*
[[reminders/Civilization/In the Future the Human Species Will Unite to Fight Alien Species by John Horgan|In the Future the Human Species Will Unite to Fight Alien Species by John Horgan]]
> “When time came to an end, the gods decided to run a final experiment. They wanted to be prepared after the big crunch for potential trajectories of life after the next big bang. For their experiment they choose two planets in the universe where evolution had resulted in similar developments of life. For planet ONEthey decided to interfer with evolution by allowing only ONE species to develop their brain to a high level of complexity. This species referred to itself as being „intelligent“; members of this species were very proud about their achievements in science, technology, the arts or philosophy.”
> **— Ernst Pöppel**, *2009, Edge Annual Question, “What Will Change Everything?”*
[[reminders/Evolution/Intelligence Could Be Evolution's Most Dangerous Experiment by Ernst Pöppel|Intelligence Could Be Evolution's Most Dangerous Experiment by Ernst Pöppel]]
> “Our species itself is partly shaped by the force of our environment through selection, and partly by fluctuations of chance. If humans had gone extinct 100,000 years ago, the world would go on turning, life would go on thriving, and some other species would be prospering in our place — and most likely not by following the same intelligence-driven technological path we did.”
> **— P Z Myers**, *2011, Edge Annual Question, “What Scientific Concept Would Improve Everybody's Cognitive Toolkit?”*
[[reminders/Civilization/Intelligence Is Not Evolution's Inevitable Destination by P Z Myers|Intelligence Is Not Evolution's Inevitable Destination by P Z Myers]]
> “Resolution of religious conflict is impossible because there is no empirical test of the ghostly, and many theologians prey, intentionally or not, upon the fears, superstitions, irrationality, and herd tendencies that are our species' neurobehavioral endowment. Religious fundamentalism inflames conflict and prevents solution—the more extreme and irrational one's position, the stronger one's faith, and, when possessing absolute truth, compromise is not an option.”
> **— Robert Provine**, *2006, Edge Annual Question, “What Is Your Dangerous Idea?”*
[[reminders/Civilization/Material Life Is All There Is by Robert Provine|Material Life Is All There Is by Robert Provine]]
> “Like many people, I once trusted in the wisdom of Nature. I imagined that there were real boundaries between the natural and the artificial, between one species and another, and thought that, with the advent of genetic engineering, we would be tinkering with life at our peril. I now believe that this romantic view of Nature is a stultifying and dangerous mythology.”
> **— Sam Harris**, *2008, Edge Annual Question, “What Have You Changed Your Mind About? Why?”*
[[reminders/Civilization/Mother Nature Is Not Our Friend by Sam Harris|Mother Nature Is Not Our Friend by Sam Harris]]
> “One of the most pernicious misconceptions in cognitive science is the belief in a dichotomy between nature and nurture. Many psychologists, linguists and social scientists, along with the popular press, continue to treat nature and nurture as combatting ideologies, rather than complementary perspectives. For such people, the idea that something isboth "innate" and "learned", or both "biological" and "cultural", is an absurdity. Yet most biologists today recognize that understanding behavior requires that we understand the interaction between inborn cognitive processes (e.g. learning and memory) and individual experience. This is particularly true in human behaviour, since the capacities for language and culture are some of the key adaptations of our species, and involve irreducible elements of both biology and environment, of both nature and nurture.”
> **— W. Tecumseh Fitch**, *2011, Edge Annual Question, “What Scientific Concept Would Improve Everybody's Cognitive Toolkit?”*
[[reminders/Civilization/Nature and Nurture Are Complementary Not Competing Forces by W. Tecumseh Fitch|Nature and Nurture Are Complementary Not Competing Forces by W. Tecumseh Fitch]]
> “As the 21st Century proceeds, tasks facing our species will become increasingly complex. Many problems that in an earlier era might have been easily addressed by a single person will now require a sophisticated set of abilities contributed by different people. The individual contributions must be deeply complementary; the whole must be more than the sum of its parts.”
> **— Stephen M. Kosslyn and Robin S. Rosenberg**, *2013, Edge Annual Question, “What *should* We Be Worried About?”*
[[reminders/Civilization/No Single Mind Can Solve the Problems Facing Our Species by Stephen M. Kosslyn and Robin S. Rosenberg|No Single Mind Can Solve the Problems Facing Our Species by Stephen M. Kosslyn and Robin S. Rosenberg]]
> “Paleo-DNA sequencing has changed not only our understanding of Neanderthals, but of ourselves. Neanderthals were not modern humans: they lacked the rapid cultural progress characterizing our species, and thus presumably some of our cognitive capacities. But what precisely were these differences? Do these differences make us, the survivors, "human?" Or were Neanderthals human, but "differently abled?" Certainly, with the bodies of Olympic wrestlers and brains slightly larger than modern humans, they'd be first picks for your rugby scrum; perhaps they had unique cognitive abilities as well. Progress will be rapid in addressing these issues, because each new insight into the genetic basis of the human brain yields parallel insights into our understanding of Neanderthal brains—and of the cognitive differences between the two.”
> **— W. Tecumseh Fitch**, *2016, Edge Annual Question, “What Do You Consider The Most Interesting Recent [Scientific] News? What Makes It Important?”*
[[reminders/Civilization/Paleo-dna and De-extinction by W. Tecumseh Fitch|Paleo-dna and De-extinction by W. Tecumseh Fitch]]
> “Despite a lot of empirical progress, the full implications of the concept of phylogeny have yet to been appreciated in evolutionary biology and the culture at large. For example, the concept of homology—similarity relation among organisms and their parts due to common ancestry—can only be understood in terms of phylogeny. Infectious diseases are caused by various species from different branches on the Tree of Life. Defending against them requires understanding how to slow them down without hurting ourselves, which is greatly facilitated by understanding where they and we fit in the historical network of phylogeny.”
> **— Richard Prum**, *2017, Edge Annual Question, “What Scientific Term Or Concept Ought To Be More Widely Known?”*
[[reminders/Evolution/Pathogens Span the Entire Tree of Life by Richard Prum|Pathogens Span the Entire Tree of Life by Richard Prum]]
> “Comparative psychology finds that pointing (in its full-blown form) is unique to our species. Few non-human species appear to be able to comprehend pointing (notably, domestic dogs can follow pointing while our closest relatives among the great apes cannot), and there is little evidence of pointing occuring spontaneously between members of any species other than our own. It appears that only humans have the social-cognitive infrastructure needed to support the kind of cooperative and prosocial motivations that pointing gestures presuppose.”
> **— Nick Enfield**, *2016, Edge Annual Question, “What Do You Consider The Most Interesting Recent [Scientific] News? What Makes It Important?”*
[[reminders/Civilization/Pointing Is a Prerequisite for Language by Nick Enfield|Pointing Is a Prerequisite for Language by Nick Enfield]]
> “Many people think that genetic engineering will change everything, even our very bodies and minds. It will, eventually. Right now, however, attempts to apply new genetic knowledge are having profound effects, not on our bodies, but on how we understand our bodies. They are revealing that our central metaphor for the body is fundamentally flawed. The body is not a machine. It is something very different, a soma shaped by selection with systems unlike anything an engineer would design. Replacing the machine metaphor with a more biological view of the body will change biology in fundamental ways.”
> **— Randolph Nesse**, *2009, Edge Annual Question, “What Will Change Everything?”*
[[reminders/Civilization/Recognizing That the Body Is Not a Machine by Randolph Nesse|Recognizing That the Body Is Not a Machine by Randolph Nesse]]
> “There are many persuasive arguments from evolutionary biology explaining why various species, notably Homo sapiens, have adopted a lifestyle in which males and females pair up long-term. But my topic here is not one of those explanations. Instead, it is the explanation for why we are close (or so I claim!)—far closer than most people, even most readers of Edge, yet appreciate—to the greatest societal, as opposed to technological, advance in the history of civilisation.”
> **— Aubrey de Grey**, *2012, Edge Annual Question, “What Is Your Favorite Deep, Elegant, Or Beautiful Explanation?”*
[[reminders/Evolution/The Overdue Demise of Monogamy by Aubrey de Grey|The Overdue Demise of Monogamy by Aubrey de Grey]]
> “The truly dangerous ideas in science tend to be those that threaten the collective ego of humanity and knock us further off our pedestal of centrality. The Copernican Revolution abruptly dislodged humans from the center of the universe. The Darwinian Revolution yanked Homo sapiens from the pinnacle of life. Today another menacing revolution sits at the horizon of knowledge, patiently awaiting broad realization by the same egotistical species.”
> **— Scott Sampson**, *2006, Edge Annual Question, “What Is Your Dangerous Idea?”*
[[reminders/Evolution/The Purpose of Life Is to Disperse Energy by Scott Sampson|The Purpose of Life Is to Disperse Energy by Scott Sampson]]
> “Despite the growing digital divide, the number of people who use the Net is still exploding, even in the developing side of the world. What is fair, what is democratic, what kind of principles can we all agree on this single global complex system, from all corners of the world is my question of the year to come.”
> **— Izumi Aizu**, *2001, Edge Annual Question, “What Questions Have Disappeared?”*
[[reminders/Civilization/Who Should Make the Truly Global Decisions, and How by Izumi Aizu|Who Should Make the Truly Global Decisions, and How by Izumi Aizu]]
> “In 2001 the "Survival" series died. It was a year when conservation efforts lagged across the board, along with other failures to take the long view. Survival programs may have told people what they could no longer bear to hear (that the human species is soiling its own den) without demonstrating constructive solutions. For example, there are precious few incentives to develop alternate energy sources despite the profound vulnerabilities that our dependence on foreign energy revealed yet again. We have no "Vision Thing," despite the many clues. "It's global warming, dude," a 28 year-old auto mechanic told The New York Times as he fished in the Hudson River; "I don't care if the whole planet burns up in a hundred years. If I can get me a fish today, it's cool by me."”
> **— Delta Willis**, *2002, Edge Annual Question, “What Is Your Question? ... Why?”*
[[reminders/Civilization/Why Conservation Fails to Command Attention by Delta Willis|Why Conservation Fails to Command Attention by Delta Willis]]
> “Cynics would go further, and say that anything that is scientifically and technically possible will be done — somewhere, sometime — despite ethical and prudential objections, and whatever the regulatory regime. Whether this idea is true or false, it's an exceedingly dangerous one, because it's engenders despairing pessimism, and demotivates efforts to secure a safer and fairer world. The future will best be safeguarded — and science has the best chance of being applied optimally — through the efforts of people who are less fatalistic.”
> **— Martin Rees**, *2006, Edge Annual Question, “WHAT IS YOUR DANGEROUS IDEA?”*
[[reminders/Existential Risk/Anything Technically Possible May Be Done Somewhere Sometime by Martin Rees|Anything Technically Possible May Be Done Somewhere Sometime by Martin Rees]]
> “Is it ultimately possible for life to bend the shape of the universe to fit life's purposes, as we are now bending the shape of our environment here on earth?”
> **— Freeman Dyson**, *2018, Edge Annual Question, “WHAT IS THE LAST QUESTION?”*
[[reminders/Deep Time/Can Life Bend the Universe to Fit Its Purposes by Freeman Dyson|Can Life Bend the Universe to Fit Its Purposes by Freeman Dyson]]
> “How far can we extend beyond our human limitations to more fully grasp the nature of the world?”
> **— Lisa Randall**, *2018, Edge Annual Question, “WHAT IS THE LAST QUESTION?”*
[[reminders/Evolution/How Far Can We Extend Beyond Human Limitations by Lisa Randall|How Far Can We Extend Beyond Human Limitations by Lisa Randall]]
> “How do ideas about biological evolution change once one species has control over the origin and extinction of all other species?”
> **— Scott Sampson**, *2018, Edge Annual Question, “WHAT IS THE LAST QUESTION?”*
[[reminders/Civilization/One Species Now Controls the Origin and Extinction of Others by Scott Sampson|One Species Now Controls the Origin and Extinction of Others by Scott Sampson]]
> “What can humanity do right now that will make the biggest difference over the next billion years?”
> **— Toby Ord**, *2018, Edge Annual Question, “WHAT IS THE LAST QUESTION?”*
[[reminders/Deep Time/What Can Humanity Do Now for the Next Billion Years by Toby Ord|What Can Humanity Do Now for the Next Billion Years by Toby Ord]]
## Sources / Provenance
- Maynard Smith and Szathmáry, _The Major Transitions in Evolution_ (1995); PNAS (2026).