The title of this book – Evolution evolving – can be read in two ways. The first captures the idea that the evolutionary process itself evolves over time, and to this day is still evolving. That implies that the way in which each organism evolves depends critically on how that organism works, and on the evolutionary mechanisms those characteristics afford.

“The evolutionary process itself evolves over time, and to this day is still evolving”

Without undermining the central importance of natural selection and other Darwinian foundations, a new understanding emerging within the contemporary evolutionary sciences implies that, say, yeast, oak trees, and human beings may each evolve in different ways; indeed, that all organisms possess a characteristic set of evolutionary mechanisms, contingent on how they develop.

The second reading follows from the first. Evolutionary theory is evolving, not just through the steady accrual of new data and technologies, but perhaps in a more fundamental way, with the emergence of a new way of explaining evolutionary change. That is a second key idea that we explore in this book. New data call for new ways of thinking: ways in which developmental processes are situated more centrally within evolutionary explanation than they conventionally have been.

“Evolutionary theory is evolving, not just through the steady accrual of new data and technologies, but perhaps in a more fundamental way, with the emergence of a new way of explaining evolutionary change”

What the two readings of our title have in common – and the principal thesis that we defend in this book – is that developmental processes do more than impose constraints on selection: they also help explain adaptive evolution.



“Evolution Evolving is a fascinating and original expansion of evolutionary theory, giving prominence to processes of embryonic development and the origins of novelty. The book goes well beyond the conventional narrative, which is still strongly rooted in population genetics and selection. The authors feature more recently developed concepts, based on our increasingly sophisticated knowledge of phenotypic plasticity, niche creation, extragenic inheritance and evolvability. Evolution Evolving is enlivened with many endearing examples and compelling stories but the authors leave no doubt that their serious purpose is to demonstrate that cellular and developmental mechanisms deserve a central place in the thrilling story of the origin of species.”

“A tour de force!”

Marc Kirschner, Harvard University

Assembling a rich toolkit of insights from dung beetles and orcas to foxes and humans, Lala and colleagues have begun a major renovation of the comfy theoretical edifice constructed by the architects of 20th-century evolutionary biology. Considering the impact of multiple inheritance systems (e.g., epigenetic, cultural, and microbial), plasticity, developmental biases, and niche construction on the core of evolutionary theory, they effectively raise the roof, knock down walls, and excavate the foundations, aiming to construct a more suitable theoretical structure for understanding the ‘endless forms most beautiful and wonderful’ of the 21st century. An engaging read, Evolution Evolving is an illuminating exploration for those curious about how all the new findings can fit into Darwin’s old house.”

Joseph Henrich, Harvard University

This book opens with the quip that ‘nothing in biology makes sense...anymore’. This riff on Dobzhansky’s dictum (‘....except in the light of evolution’) seems puzzling for a book on evolution. Why start here? The authors guide the reader through many of the seminal biological discoveries of the past century, with fascinating examples on each page. At first many discoveries don't make sense, especially when looking at them through a single lens. It is only by expanding our perspective, bringing together molecular, developmental, and evolutionary insights, can we come to fully understand and appreciate how novelty arises and how feedback and interactions — along with selection — shape the biological world. Beautifully illustrated, this book will make you come to love biology and its evolutionary underpinnings even more."

Sally Otto, University of British Columbia

An eloquent, example-laden, accessible narrative setting the stage and offering the story of a truly contemporary evolutionary theory. The book represents a myriad of theoretical frames and approaches united by the focus on developmental biology, with robust examples, well-argued theory, and a sincere, even compassionate, invitation to think together.”

Agustín Fuentes, Princeton University

This book is a signpost in the ongoing journey of evolutionary thinking, even though not everyone will agree with the direction it is pointing. Only a frank and open discussion will eventually lead to a renewed “synthesis” of evolutionary knowledge. For that this book will be an important point of reference.”

Gunter Wagner, University of Vienna

A thoughtful, interesting, and informative overview of major themes and research areas in contemporary evolutionary biology. Although some of the perspectives outlined in this book are controversial (and I often found myself disagreeing with the authors!), I would recommend this book to anyone seeking a better understanding of the ideas at the core of the ‘Extended Evolutionary Synthesis’.”

Russell Bonduriansky, University of New South Wales

Table of contents

Part 1: Introduction – why consider development?

1. Nothing in biology makes sense… anymore

Presents some striking new data that are challenging traditional understanding of heredity and evolution, including epigenetic and cultural inheritance

2. Rodents’ teeth and raptors’ toes

Uses a case study to show how a developmental perspective can enhance evolutionary explanation

3. How the turtle got its shell

Explains why genes and natural selection became central to evolutionary biology and why developmental processes remain less central

4. Understanding the debates

Not only do the traits of a given organism differ in their propensities to evolve, but organisms may themselves differ greatly in how effectively they are able to generate and find adaptive solutions.

Part 2: How development works

5. Opening the black box

Summarizes what evolutionists ‘need to know’ about development.

6. Five general principles of development

Presents five general principles of development relevant to evolution: development is modular, epigenetic, constructive, interchangeable with respect to genetic and environmental inputs, and prone to generate biased phenotypic variation

Part 3. The developmental bases of evolutionary processes

Chapter 7. Developmental bias

Explains how developmental mechanisms bias the phenotypic variation available to natural selection, creating admissible pathways along which selection can act.

Chapter 8. Plasticity-led evolution

Describes how developmental plasticity can lead genetic evolution, and when this will happen.

Chapter 9. The causes of selection

Explains how developmental processes, including plasticity and niche construction, create associations between traits and fitness, and thereby influence the dynamics of adaptive evolution.

Chapter 10. Inheritance beyond the gene

Summarizes and explains the significance of recent data demonstrating extra-genetic inheritance, including parental effects, epigenetic inheritance, the inherited microbiome and animal culture.

Part 4. Implications of the developmental perspective

Chapter 11. Novelty and innovation

Describes how developmental processes help account for evolutionary novelty and innovation.

Chapter 12. The developmental origins of evolvability

Discusses how the evolutionary process itself evolves and may have become more efficient with time, and how variation in the ability to respond to selection is partly explained by differences in the developmental mechanisms of organisms.

Chapter 13. Human evolvability

Explains how developmental bias, plasticity, niche construction and extra-genetic (particularly cultural) inheritance have played critical roles in human evolution and adaptation.

Chapter 14. The structure of evolutionary theory

Summarizes the arguments in the book, and points to exciting new opportunities that arise if researchers embrace a developmental perspective.