The essential one-volume reference to evolution
The Princeton Guide to Evolution is a comprehensive, concise, and authoritative reference to the major subjects and key concepts in evolutionary biology, from genes to mass extinctions. Edited by a distinguished team of evolutionary biologists, with contributions from leading researchers, the guide contains some 100 clear, accurate, and up-to-date articles on the most important topics in seven major areas: phylogenetics and the history of life; selection and adaptation; evolutionary processes; genes, genomes, and phenotypes; speciation and macroevolution; evolution of behavior, society, and humans; and evolution and modern society. Complete with more than 100 illustrations (including eight pages in color), glossaries of key terms, suggestions for further reading on each topic, and an index, this is an essential volume for undergraduate and graduate students, scientists in related fields, and anyone else with a serious interest in evolution.
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Jonathan B. Losos is the Monique and Philip Lehner Professor for the Study of Latin America and professor of organismic and evolutionary biology at Harvard University.
"The marvelous new tools and theoretical approaches that have so greatly deepened our understanding of evolution over the past few decades are clearly and concisely presented in this splendid new guide, which should be of use to everyone involved in the field."--Peter H. Raven, president emeritus of the Missouri Botanical Garden, St. Louis
| Preface.................................................................... | vii |
| Contributors............................................................... | ix |
| Section I Introduction..................................................... | 1 |
| Section II Phylogenetics and the History of Life........................... | 47 |
| Section III Natural Selection and Adaptation............................... | 189 |
| Section IV Evolutionary Processes.......................................... | 305 |
| Section V Genes, Genomes, Phenotypes....................................... | 363 |
| Section VI Speciation and Macroevolution................................... | 483 |
| Section VII Evolution of Behavior, Society, and Humans..................... | 605 |
| Section VIII Evolution and Modern Society.................................. | 727 |
| Index...................................................................... | 837 |
Introduction
I.1
What Is Evolution?
Jonathan Losos
OUTLINE
1. What is evolution?
2. Evolution: Pattern versus process
3. Evolution: More than changes in thegene pool
4. In the light of evolution
5. Critiques and the evidence for evolution
6. The pace of evolution
7. Evolution, humans, and society
Evolution refers to change through time as species becomemodified and diverge to produce multiple descendantspecies. Evolution and natural selection are oftenconflated, but evolution is the historical occurrence ofchange, and natural selection is one mechanism—in mostcases the most important—that can cause it. Recentyears have seen a flowering in the field of evolutionarybiology, and much has been learned about the causes andconsequences of evolution. The two main pillars of ourknowledge of evolution come from knowledge of thehistorical record of evolutionary change, deduced directlyfrom the fossil record and inferred from examination ofphylogeny, and from study of the process of evolutionarychange, particularly the effect of natural selection. It isnow apparent that when selection is strong, evolution canproceed considerably more rapidly than was generallyenvisioned by Darwin. As a result, scientists are realizingthat it is possible to conduct evolutionary experiments inreal time. Recent developments in many areas, includingmolecular and developmental biology, have greatly expandedour knowledge and reaffirmed evolution's centralplace in the understanding of biological diversity.
GLOSSARY
Evolution. Descent with modification; transformationof species through time, including both changes thatoccur within species, as well as the origin of newspecies.
Natural Selection. The process in which individuals witha particular trait tend to leave more offspring in thenext generation than do individuals with a differenttrait.
Approximately 375 million years ago, a large and vaguelysalamander-like creature plodded from its aquatichome and began the vertebrate invasion of land, settingforth the chain of evolutionary events that led to the birdsthat fill our skies, the beasts that walk our soil, me writingthis chapter, and you reading it. This was, of course, justone episode in life's saga: millions of years earlier, plantshad come ashore, followed soon thereafter—or perhapssimultaneously—by arthropods. We could go back muchearlier, 4 billion years or so, to that fateful day when thefirst molecule replicated itself, an important milestone inthe origin of life and the beginning of the evolutionarypageant. Moving forward, the last few hundred millionyears have also had their highs and lows: the origins offrogs and trees, the end-Permian extinction when 90percent of all species perished, and the rise and fall of thedinosaurs.
These vignettes are a few of many waypoints in theevolutionary chronicle of life on earth. Evolutionarybiologists try to understand this history, explaining howand why life has taken its particular path. But the studyof evolution involves more than looking backward to tryto understand the past. Evolution is an ongoing process,one possibly operating at a faster rate now than in timespast in this human-dominated world. Consequently,evolutionary biology is also forward looking: it includesthe study of evolutionary processes in action today—howthey operate, what they produce—as well as investigationof how evolution is likely to proceed in thefuture. Moreover, evolutionary biology is not solely anacademic matter; evolution affects humans in manyways, from coping with the emergence of agriculturalpests and disease-causing organisms to understandingthe workings of our own genome. Indeed, evolutionaryscience has broad relevance, playing an important role inadvances in many areas, from computer programmingto medicine to engineering.
1. WHAT IS EVOLUTION?
Lookup the word "evolution" in the online version of theOxford English Dictionary, and you will find 11 definitionsand numerous subdefinitions, ranging from mathematical("the successive transformation of a curve bythe alteration of the conditions which define it") to chemical("the emission or release of gas, heat, light, etc.") tomilitary ("a manoeuvre executed by troops or ships toadopt a different tactical formation"). Even with referenceto biology, there are several definitions, including"emergence or release from an envelope or enclosingstructure; (also) protrusion, evagination," not to mention"rare" and "historical" usage related to the conceptof preformation of embryos. Even among evolutionarybiologists, evolution is defined in different ways. Forexample, one widely read textbook refers to evolution as"changes in the properties of groups of organisms overthe course of generations" (Futuyma 2005), whereasanother defines it as "changes in allele frequencies overtime" (Freeman and Herron 2007).
One might think that—as in so many other areas ofevolutionary biology—we could look to Darwin forclarity. But in the first edition of On the Origin of Species,the term "evolution" never appears (though the lastword of the book is "evolved"); not until the sixth editiondoes Darwin use "evolution." Rather, Darwin'sterm of choice is "descent with modification," a simplephrase that captures the essence of what evolutionarybiology is all about: the study of the transformation ofspecies through time, including both changes that occurwithin species, as well as the origin of new species.
2. EVOLUTION: PATTERN VERSUS PROCESS
Many people—sometimes even biologists—equate evolutionwith natural selection, but the two are not thesame. Natural selection is one process that can causeevolutionary change, but natural selection can occurwithout producing evolutionary change. Conversely,processes other than natural selection can lead toevolution.
Natural selection within populations refers to the situationin which individuals with one variant of a trait(say, blue eyes) tend to leave more offspring that arehealthy and fertile in the next generation than do individualswith an alternative variant of the trait. Suchselection can occur in many ways, for example,...
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