[(The Major Transitions in Evolution)] [Author: John Maynard Smith] published on

by John Maynard Smith

Over the history of life there have been several major changes in the way genetic information is organized and transmitted from one generation to the next. These transitions include the origin of life itself, the first eukaryotic cells, reproduction by sexual means, the appearance of multicellular plants and animals, the emergence of cooperation and of animal societies, and the unique language ability of humans. This ambitious book provides the first unified discussion of the full range of these transitions. The authors highlight the similarities between different transitions--between the union of replicating molecules to form chromosomes and of cells to form multicellular organisms, for example--and show how understanding one transition sheds light on others. They trace a common theme throughout the history of after a major transition some entities lose the ability to replicate independently, becoming able to reproduce only as part of a larger whole. The authors investigate this pattern and why selection between entities at a lower level does not disrupt selection at more complex levels. Their explanation encompasses a compelling theory of the evolution of cooperation at all levels of complexity. Engagingly written and filled with numerous illustrations, this book can be read with enjoyment by anyone with an undergraduate training in biology. It is ideal for advanced discussion groups on evolution and includes accessible discussions of a wide range of topics, from molecular biology and linguistics to insect societies.

Genres: Science, Evolution, Biology, Nonfiction, History, Philosophy, Education

Paperback

First published January 1, 1995

About the author

John Maynard Smith FRS was a British theoretical and mathematical 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. Haldane

Reviews

[Jon Richfield · Rating 5 out of 5]

Though not a formal textbook, this is not a popular book; it is a get-down-to-business book, and very edifying. If you feel frustrated because you find it difficult to follow, read the simplified version, published "The Origins of Life". If you cannot deal with that one either, then don't get into any arguments about evolution; you are not equipped for it.
The book is written in pleasantly competent English (well, with Maynard Smith as the senior author, what would one expect?) but much of it takes slow and patient reading; no speed reading please -- the material is rich and takes proper digestion.
There are certain basic principles in science, very, very important principles, that are so simple that most people encountering them for the first time think that the subject also must be simple and accordingly that they understand that subject. Probability theory is one example, and natural selection is another. Such fields are snares for the unwary, and this book deals with certain classes of the snares clearly and systematically, presenting a powerful overall view that should enrich the insights of most biologists and many evolutionists as well. JMS left a legacy of thoughtful, sound, edifying writings without pretension or compromise. If you work your way through it, you might be startled to find how rewarding your labour turned out to be.
Smith and Szathmary deal with five major transitions in evolution, stages at which, without hurry or or obvious warning, certain emergent trends changed the game of life, or of a large part of life on the planet. They structure the material carefully and lucidly. Some readers might like to skip the first few, which dealt with chemistry and molecular biology. However, the chapters on Eukaryotes, sex, symbiosis, organisms, morphogenesis, societies, and language should enrich even people who shudder at the sight of a molecular structure.
In sum, in my opinion this is a great book, not in the sense of "Isn't it just AWESOME?", but really great. A landmark in the accessible documentation of the advance of evolutionary theory and insight.

[Xander · Rating 2 out of 5]

"This book has been about the major changes that have taken place in the way in which information is encoded, and transmitted between generations." So say both authors on the last page (p. 309) in The Major Transitions in Evolution.

The above citation is a good summary of the entire content of the book, in which evolutionary biologists John Maynard Smith and Eörs Szathmáry try to describe all the historical transitions that had to take place to get from a dead rock in space to the planet Earth that we are familiar with. Even though this sounds interesting, like most of 'big history', this book is not for lay people. It is much too technical and too professionalistic to be readable by wide audiences.

And this last point brings me to the most important point about this book. Maynard Smith and Szathmáry are professional scientists (mathematician and chemist respectively) and they focus on explaining all the important steps in evolution by giving an overview of the current (then-current: 1998) scientific status on the relevant topics. They continuously use jargon and, even while using much illustrations to make their points, are simply no story-tellers. It is, hence, ironic that in chapter 16, on the topic of the origin of societies they write:

[i]"The trouble with theories of society is that their formulations are so lengthy and so complex that they are hard either to grasp or to test. Theories should be formulated briefly, even if the facts to which they are relevant must be described at length. The Origin of Species [Darwin's book] is a long book, but no biologist regards it as impossible to outline Darwin's theory in a few sentences." (p. 273)[/i]

Perhaps. But at least The Origin of Species is easy to grasp because Darwin took the time to level with his readers: first introduce the concept in familiar terms (evolution by artifical Selection of livestock), then explain the concept is new terms (evolution by natural Selection) and afterwards adduce the concept by all the relevant facts (the importance of geographical distribution in the origin of species, etc.). This is something Maynard Smith and Szathmáry don't do: they briefly state their theories and claims, without introducing the reader to terms or context; they drop their ideas for the reader to grasp.

So be warned: the book is a struggle to work through. And perhaps a somewhat unnecessary struggle at that - most modern day handbooks on evolution explain all the facts from The Major Transitions in accessible terms and with (comprehensible) pictures and graphs to illustrate the processes.

Nevertheless, the main point of the book is strong: evolutionary history is littered with small steps and changes, that accumulated, over eons of time, in the (apparently designed) complexity that we observe today. The authors see the following (rough) important historical transitions as key steps:

1. The origin of replicating molecules from static molecules (this has, among other things, to do with polarity differences on molecules).
2. The origin of independent (i.e. enclosed) replicators: lipid membranes to shield the internal from the external.
3. The origin of RNA as gene (chromosomes) and calatyzing enzymes.
4. The origin of DNA and the protein-building processes (using RNA in new functions).
5. The origin of prokaryotes, and consequently eukaryotes.
6. The origin of sexually reproducting populations.
7. The origin of cell differentation, leading to plants, fungi and animals.
8. The origin of societies (for example, the social insects).
9. The origin of mankind.
10. The origin of language, coevolved with anatomical changes like the larynx, our cognitive abilites and consequently tools and culture.

(Although these transitions are all very interesting, the book concisely deals with the elemental ideas without elaborating much. Expect to be fed dry scientific statements about the current state of affairs in research about the topic involved; don't expect to be instructed in the topic.)

Once again, I cannot refrain from saying that one could better use a contemporary handbook of biology to study the workings of things like cells, DNA or the differences between plants and animals. This book is just too narrow and specialistic in scope. I understood the most important thoughts that both authors wanted to bring across, but I didn't really delve into all the details - they simply don't interest me and I've read about these exact same processes and phenomena in other places. I simply don't see the need for this obscurity...

I'd like to end this review by giving my personal opinion. Not on this book in particular, but on this kind of books in general. In my view, a lot of writing in science and philosophy is unnecessarily obscure. I cannot help but suspect that this is (at least partly) some form of aristocratic tendency in the writers of such works. They create a work that only 'insiders' can grasp; if you cannot grasp their texts, you're simply not up to the task. This arrogance annoys me ever more, the more I read books on these topics.

I can accept that some topics are simply abstract and hard-to-explain. I also agree that some ideas are just harder to grasp. I do also understand that the mathematics involved in some of the topics in this kind of books are necessary and make these topics harder to explain to lay people - a legitimate reason for making books more abstract. But I cannot (and will not) accept that things like cellular processes and the underpinnings of evolution are that hard, either to grasp, or to explain. Richard Dawkins is able to explain the key ideas of evolution in 250 pages; whereas someone like Maynard Smith needs 350 pages (his book The Theory of Evolution) and demands much more cognitive power of the reader; to explain the exact same thing! I find it funny to read reviews (afterwards) in which people claim "if you don't understand this book, don't get involved in arguments about evolution." The sheer arrogance.

(Scientists will object that science works only by meticulously defining your concepts and hence the need for obscurity. But please do this in your journals. As soon as you step into the world, make yourself understandable to your audience - this is the only purpose of language.)

[William Bies · Rating 4 out of 5]

The Darwinian mechanism of natural selection, positing that mutations occur at random and that, among them, those leading to adaptive characters become established in a population due to differential reproductive success, is on the face of it fairly plausible as a theoretical possibility, once the momentous idea had occurred to Darwin and Wallace in the mid-nineteenth century (nearly simultaneously). Whether it is the dominant factor driving evolutionary change and phylogenesis, in particular, is far more open to debate. Anyone can grant that something like this must be behind microevolution, as a contributing factor at least, and indeed laboratory experiments on rapidly reproducing bacterial cultures tend to confirm that it does take place, but it is another matter altogether to propose it as sufficient by itself to explain speciation or the radiation of entirely new higher taxa. That is why, contrary to the image presented by popularizers of evolutionary biology, Darwin’s theory did not by any means take the professional world by storm immediately upon its publication in 1859. To be sure, it generated great interest as a novel hypothesis, but comparatively few evolutionary biologists at the time were prepared to accept it wholesale (as notably did Huxley). For the better part of two generations, many competing proposals vied for attention, among them neo-Lamarckianism, orthogenesis and de Vries’ mutationism. Only beginning in the 1930’s, under the influence of the erstwhile rediscovery of Mendelian genetics and the rise of population biology, did the so-called neo-Darwinian consensus emerge, in which natural selection won general acceptance as probably the major driving force behind evolutionary change (though even then, few would go out on a limb and make it the sole such force).

The purpose of this preamble is to indicate why the major transitions in evolutionary history can exert such a fascination, for they serve as the best tests of our understanding of how evolution did in fact take place. Hence, the present work by the distinguished evolutionary biologists John Maynard Smith and Eörs Szathmáry will be most welcome to the interested and educated reader, who may be looking for a suitable entry into the literature on the subject. What are the major transitions? The origin of protocells and the genetic code, the origin of eukaryotes, the origin of sex, the rise of multicellular organisms, symbiosis, gene regulation, cooperation, societies and lastly, for Maynard Smith and Szathmáry, language in man.

The reader will find summary treatments of these and a few other topics in the present work, such as intragenomic conflict, comprehending meiotic drive, B chromosomes, control of transposable elements, repetitive DNA, chromosomal form, competition among organelles and between organelles and the nucleus (which affects the sex ratio). Each chapter tends to follow a similar format: a brief introduction to the problem followed by capsule summaries of leading ideas of researchers in the field. One should take note of the degree of compression involved; what in the journal literature would be entire papers are covered usually in a matter of a few pages. As a result, the authors have to skimp on derivations of results. Instead, one gets telegraphic accounts of the main concepts and ideas. As a rule, one can tell what the theoretical proposals are about and how they account for the observed data, without necessarily appreciating in full why, or the subtleties. Hence, one should look on this book as something in the nature of an extended review of the literature. As such, approached in the right spirit, it can be very helpful for orienting oneself and for gathering collected references to the original papers, should one wish to follow up on them. Certainly, one will become acquainted with a wealth of ideas and pertinent considerations.

Let us single out a few for comment. In discussing the preconditions on the early earth for life to appear, the authors bring up the example of a naturally occurring atomic fission reactor at Gabon in West Africa and known as the Oklo reactor. A uranium-rich seam embedded in sandstone at a 40-degree tilt, originally formed in a river bed under the conditions of increased oxygenation of atmosphere some 2 billion years ago, was exposed to percolating water in just such a way as to raise the concentration of uranium ore above the threshold for criticality, leading to a self-regulating reaction: if it went too fast, the water (which acted as the moderator) evaporated as steam; if too slow, water condensed. This system presents an interesting parallel to the presumed origin of life, with stringent geological requirements that could support chemical cycles involving catalysts. Then they introduce a simple model of a protocell, proposed by Gánti, known as the chemoton. It incorporates three necessary elements: an autocatalytic cycle to enable metabolism, a bilayer membrane and an information-bearing molecule to carry genetic instructions. The chemoton serves as a thought experiment in which one can speculate about the role of growth, replication by cell division and inherited genetic variation.

This case nicely illustrates Maynard Smith and Szathmáry’s modus operandi—for each major topic, kick off with some general points that place it in context, then review in a fair amount of detail several contributions to the research literature that have stood the test of time, whether the Urey-Miller experiment, Eigen and Schuster’s hypercycle and error-threshold for fidelity in replication, the stochastic corrector model, the possibility that RNA can act as an enzyme (so-called ribozymes) and speculations about the RNA world that preceded DNA, autocatalytic protein networks, Cairns-Smith’s ideas about clay performing a role as the original template that got replication of larger oligomers going, the optimal size of the genetic alphabet, the nature of the codon assignments in the genetic codon-to-amino-acid table and how it may have been modified, the need for active compartmentation and the origin of protocells, the origin of the nucleus in eukaryotes and Margulis’ theory of the endosymbiotic origin of mitchondria and chloroplasts, the cellular mechanisms behind the haploid-diploid cycle and what they can tell us about the origin of sex, mutualism and the division of labor in multicellular eukaryotes, gene regulation and the origin of cooperation—the reader can expect to have the issues laid out for him and examined in the light of some substantial work in the field, occasionally, though not always, concluding with a few summary observations on the authors’ part.

What one might miss in all the profuse riot of perspectives are any well-developed overarching themes, of the kind Stephen Jay Gould is so good at pursuing in his magnum opus, The structure of evolutionary theory (q.v., this recensionist’s review elsewhere on this site). Scientists seem to come in two sorts, those who always keep the big picture in mind and those who excel at going into minutia. Maynard Smith and Szathmáry definitely belong to the latter category, and could thus be labeled technicians rather than seers (to borrow terminology employed by Lee Smolin in the context of theoretical physics, but which is apposite here). In consequence, the book stops abruptly and there is no ringing conclusion that would bring everything together. Let the reader beware what he is getting into! If there are any lessons on the nature of evolution to be found in the extensive and circumstantial commentary in this work, he will have to draw them out for himself. Thus, with respect to the question with which this review started out, concerning what we can learn about the nature of evolution from a study of the major transitions, we are left in the lurch. But, let us speculate based on what we have learned from Maynard Smith and Szathmáry. The major evolutionary transitions strike this reviewer by their spontaneity. They could scarcely have been foreseen; one could well imagine other habitable planets on which, say, life remains arrested at the unicellular stage and never undertakes the fateful departure into multicellularity. The earth, therefore, appears very special in that it has witnessed a series of major transitions, any one of which might very well be extremely improbable (for this reason, many evolutionary biologists tend to be skeptical of extraterrestrial life in our galaxy, at least of the form that could eventuate in intelligence and advanced civilization). Now, what follows from this reflection? To this reviewer, it supports a case for what the complexity theorist Stuart Kauffman terms organizing principles of life. That is to say, strict neo-Darwinian doctrine is insufficient, for it fails to account for the fact that the supposed undirected random walk of orthodox natural selection takes place in a context of order that is inherent in the world process due to the very laws of physics. Hence, neo-Darwinism is incomplete as an explanatory schema for evolutionary change; without the mooted ordering principles, it, by itself, would be incapable of generating the genuine novelties associated with the major transitions in life’s history (or the timescale for doing so would be too indefinitely long). Would Maynard Smith and Szathmáry agree to this statement? A hint in their preface suggests perhaps so: they are inclined to regard the major evolutionary transitions as non-equilibrium phase transitions, in which an order parameter suddenly becomes non-zero and, thereafter, free to grow. For an adequate handling of this rather intricate issue, one could turn to Kauffman’s own monograph, Origins of order: Self-organization and selection in evolution, for which, accordingly, this recensionist will have to promise a review here in due time.

[Jafar Isbarov · Rating 4 out of 5]

Scientific literature is sharply divided into relevant and outdated. There is little room for classics, yet The Major Transitions in Evolution is a certain one. After death of both authors, it has been revised by a group of scientists to keep it up-to-date with rapid scientific progress. Value of the original work, however, goes beyond its relevance.

Book is essentially a composition of eighteen distinct essays, save for references to each other. One can single out any of them, but they come together to form a brilliant piece. Book is ordered chronologically, covering some ten evolutionary transitions in eighteen sections. It is by no means an effortless read, but every page is priceless.

My first attempt to read Major Transitions in Evolution was almost a year ago. I was occupied with exams then, so I decided it is not worth to push trough such a dense book. Two months ago I picked it up again. This time I was too busy waiting for the summer vacation, so I had to abandon it for another month before I finally decided to finish it. I think I will never finish it, unless I actually study some biology.

Don't get me wrong, this is not an unreadable book -- neither in the sense of being badly-written nor in terms of toughness. I cannot call it thoroughly enjoyable, but it was not meant to be anyway. What we are dealing with is an original scientific work -- conclusion of years of theoretical research by two vigorous scientists. You don't read it on a beach with a coke on your side. You read it in an abandoned, dusty biology lab only to be distracted by drop of a limb of a skeleton.

[Carmen Álvarez · Rating 5 out of 5]

Libro clásico sobre las major transitions. En su lectura se nota por qué se ha convertido en un clásico del campo. Ahora bien, tiene algunas partes en las que más bien plantea las cuestiones. Esto no es algo malo ya que lo veo como un gran libro para introducirse en las major transitions y luego, según los intereses de cada lector profundizar más en unas u otras por la lectura de papers. Además, hay que tener en cuenta que el libro es de finales de los 90 y desde esos años hay nuevos descubrimientos.

Aún así, un gran libro y con una lectura cómoda.

[Pratik · Rating 4 out of 5]

I regard "The Major Transitions in Evolution" as a foundational text that profoundly reshapes our understanding of life's history. Smith and Szathmáry present a cohesive framework that connects key evolutionary events, emphasizing the role of cooperation in driving complexity—from the emergence of chromosomes to multicellularity. Their insights into molecular evolution and social structures, including the development of human language, extend the concept of major transitions beyond traditional biology. While some ideas have evolved since its publication, the book remains essential for anyone seeking to grasp how life's complexity has unfolded over billions of years and continues to inspire research across various subfields of evolutionary biology.

[A · Rating 4 out of 5]

This is a difficult book to read, since you need a strong background in chemistry and biology to understand it fully.

Although a bit outdated, in general it presents great questions about biology, such as how replicators appeared, compete and collaborate. The only weak part in my view is the part of language, that seems too anglocentric and derives universal claims that are most likely not true. Overall an interesting book.

[David Hunter · Rating 4 out of 5]

A really difficult read. The last chapter covers linguistics, which I happened to study in college. That was the only chapter that did not require heavy mental lifting to make sense of.

[Taveri · Rating 3 out of 5]

This is an earlier, longer, more elaborat version of the same authors' "Origins of Life". It has better drawings and better explanations on how natural selfotion might account for transitions such origin of eukaryotes, sex and symbiosis (the last is a chapter not in the shorter book). Lynn Margulis, who was mentioned in "The Prophet and the Wizard" shows up numerous times. The telling is single minded in presenting just the evolutionary belief system (I subscribe to neither that nor the Creationist viewpoint) and the gyrations and convulutions presented are reminiscent of the attempts of the earth is the centre of the solar system thinking to try and explain why the sun and planets revolve around terra firma. Here is a sampling from page 152:

"A possible way out of this difficulty is to suggest a eaploid-diploid cycle proceeded by the loss of a rigid cell wall and the evolution of phagotrophy. Although conceivable the the explanation is not attractive. An alternative, more general explanation is that the genetic load at equilibrium between mutation and selection is lower in haploids. (The mutation rate per genome is twice as great as in adiploid. If deleterious mutations are not completely recessive, they will be removed from a diploid population in heterozygotes, one mutation per selection death. Thus it take twice as many selective deaths in a diploid population to balance mutation.) Simulations show that if there is recurrent deleterious mutation, and periods of DNA damage, a population with a haploid-diploid cycle, syncrtonized so that the diploid phase coincides with periods of DNA damage, has a higher mean fitness than a population that is either permanently diploid or permanently haploid."

[Liam · Rating 3 out of 5]

Very difficult read and highly technical at the beginning. Towards the end, where the authors declare they are not experts on the subjects, it is much better. I wish they were less expert on biochemistry.

[Peter Riboprotein · Rating 5 out of 5]

One of my favorite books about abiogenesis

[Bria · Rating 4 out of 5]

The first half or so of this book brought the immense achievement of the existence of life and cells to the forefront of my attention, making me feel if not guilty, at least a little strange, every time I would blow my nose or anything. Billions of years to reach the state where such a complicated thing as a cell was possible - and I'm just expelling them by the million. If I had had a proper biochemical and molecular biological training then I'm sure I would have been even more struck, as the arguments about what was necessary and how it could have happened would have made more sense to me.

[Rodrigo · Rating 4 out of 5]

I found it difficult to follow and too geared towards specialists in evolutionary biology. I'm reading their "origins of life" next so that the ideas hopefully sink in a little more. It's interesting to see just how complex evolution gets when you get into the nuts and bolts of it. It's far more complicated in practice than just saying "natural selection".

[mensch · Rating 3 out of 5]

Despite the reviewers blurbs mentioning that this is a very readable book, this is not a title for the regular lay person. It's very readable though, just very hard to understand when not an expert on the matter.