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Full download The Tangled Bank: An Introduction to Evolution Second Edition – Ebook PDF Version file pdf all chapter on 2024
Full download The Tangled Bank: An Introduction to Evolution Second Edition – Ebook PDF Version file pdf all chapter on 2024
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To Grace, for growing our garden
together
IT IS INTERESTING TO CONTEMPLATE AN ENTANGLED BANK,
clothed with many plants of many kinds, with birds singing on the bushes,
with various insects flitting about, and with worms crawling through the
damp earth, and to reflect that these elaborately constructed forms, so
different from each other, and dependent on each other in so complex a
manner, have all been provided by laws acting around us. These laws, taken
view of life, with its several powers, having been originally breathed into a
few forms or into one; and that, whilst this planet has gone cycling on
according to the fixed laws of gravity, from so simple a beginning endless
forms most beautiful and most wonderful have been, and are being, evolved.
15 Evolutionary Medicine
Acknowledgments
Glossary
Index
Preface
FEW SOUNDS ARE AS JOYFUL for authors as the thud made by a mail
carrier dropping the first printed copy of their book by the front door. Years
of effort suddenly take shape as paper, ink, and covers. But for textbook
authors, there is another sound that’s just about as sweet: receiving the first
printed copy of the second edition of a textbook.
By the time a trade book about science reaches the bookstore, it’s probably
already out of date. Science rolls forward, as researchers perform surprising
new experiments or stumble across fossils lurking in the ground. Coaxing a
trade publisher to update a book is next to impossible, for a whole host of
economic reasons. Fortunately, textbook publishers think otherwise. A new
edition of a textbook can be an update on a scientific field, rather than a
cultural artifact. It is thus with great pleasure that I present the second edition
of The Tangled Bank: An Introduction to Evolution.
The first edition, published in 2009, was an experiment of sorts. A growing
number of colleges were offering evolution-related courses to nonbiology
majors, but they had no textbook specifically tailored for their students.
Having spent two decades reporting on evolutionary biology for newspapers
and magazines, as well as writing trade books on the topic, I decided to create
just such a textbook.
The response has been immensely gratifying, and it’s encouraged me as
I’ve revised The Tangled Bank for a new edition. Much of the work I’ve put
into this edition has involved surveying recent scientific literature for new
studies that advance our understanding of evolution. Those advances have
included everything from research on our Neanderthal relatives to
experiments on cancer-fighting drugs based on evolutionary principles.
The second edition contains new figures that better illuminate the concepts
I discuss in the text. I’ve also updated the overall organization. In response to
a number of requests, for example, I’ve written an entirely new chapter near
the end of the book dedicated to human evolution. There I integrate the
concepts introduced earlier in The Tangled Bank, such as phylogeny, genetic
drift, and sexual selection, to help readers gain a deeper understanding of our
own species.
From the start of this project, I’ve tried to make The Tangled Bank as
readable as possible. The second edition does not have some of the standard
pedagogical conventions. New terms are not set off in boldface, for example,
and the book does not contain problem sets or summaries. Instead, I define
new terms in the text itself. Students who want to check their understanding
can now consult a new study guide, produced by biologist and journalist
Alison Perkins. For more information on purchasing the study guide, visit
http://www.roberts-publishers.com.
Although the second edition of The Tangled Bank is new in some
important ways, it does not waver from the goals of the first edition. I hope
that it continues to help a wide range of students come to appreciate the
majesty and explanatory power of evolution.
One of the best feelings paleontologists can ever have is to realize that
they’ve just found a fossil that will fill an empty space in our
understanding of the history of life. Hans Thewissen got to enjoy that
feeling one day in 1993, as he dug a 47-million-year-old fossil out of a
hillside in Pakistan. As he picked away the rocks surrounding the bones
of a strange mammal, he suddenly realized what he had found: a whale
with legs.
A hundred million years ago, not a single whale swam in all the
world’s oceans. Whales did not yet exist, but their ancestors did. At the
time, they were small, furry mammals that walked on land. Over
millions of years, some of the descendants of those ancestors underwent
a mind-boggling transformation. They lost their legs, traded their
nostrils for a blowhole, and became creatures of the sea. This profound
change was the result of evolution.
FIGURE 1.1
Paleontologist Hans Thewissen has discovered many of the bones of Ambulocetus in Pakistan.
This book is an introduction to that process. Over the course of some 3.5
billion years, life has evolved from humble beginnings into the tremendous
diversity found today on Earth—from whales to tulips to viruses. Indeed, to
understand life on Earth—its diversity of forms, of biochemistry, of
behaviors—it’s crucial to understand evolution. The great twentieth-century
biologist Theodosius Dobzhansky summed up the importance of this branch
of science in 1971 when he declared, “Nothing in life makes sense except in
the light of evolution” (Dobzhansky 1973).
The Tangled Bank is intended for anyone with a curiosity about how life
works. To explain mathematical concepts, for example, it uses graphs and
diagrams rather than detailed equations. This book presents the research of
scientists like Thewissen not in technical detail, but through their own
experiences of discovery. When Charles Darwin formulated the theory of
evolution in the mid-1800s, the most sophisticated tool he could use was a
crude light microscope. As we’ll see in this book, scientists today can study
evolution in many ways—by analyzing our DNA, for example, by probing
the atoms of ancient rocks to determine the age of fossils, or by programming
powerful computers to decipher how a thousand different species are related
to each other.
Understanding evolution is too important to be limited only to evolutionary
biologists. Evolution underlies many important issues society faces, and a
better understanding of evolution can help us to grapple with them. The
world’s biodiversity faces many threats, from deforestation to global
warming. By studying past mass extinctions, we can better understand our
current crisis. Evolution has produced bacteria that can resist even the most
powerful antibiotics. Scientists can observe this resistance as it evolves in
laboratory experiments. The evolution of pests and pathogens poses a serious
risk to our food supply. But evolution can also help us to address some of the
biggest questions we ask about the universe and ourselves: How did we get
here? What does it mean to be human?
What Is Evolution?
The short answer is descent with modification. The patterns of this
modification, and the mechanisms by which it unfolds, are what evolutionary
biologists study. They have much left to learn—but at this point in the history
of science, there is no doubt that life has evolved and is still evolving.
The fundamental principles that make evolution possible are pretty
straightforward. Organisms inherit traits from their ancestors because they
receive a molecule called DNA from them (see page 115 for a discussion of
heredity). Cells use DNA as a guide to building biological molecules, and,
when organisms reproduce, they make new copies of DNA for their
offspring. Living things do not replicate their DNA perfectly; sometimes
sequence errors occur. Such errors are referred to as mutations (Chapter 5). A
mutation may be lethal; it may be harmless; or it may be beneficial in some
way. A beneficial mutation may help an organism to fight off diseases, to
thrive in its environment, or to improve its ability to find mates.
Evolution takes place because mutated genes become more or less
common within populations over the course of generations. Many mutations
eventually disappear, while others spread widely. Some mutations spread
simply by chance. Others spread because they allow organisms to produce
more offspring. This nonrandom spread of beneficial genes is known as
natural selection. The effect of a mutation depends on more than just the
mutation itself. It is also influenced by all the other genes an organism
carries. The environment in which an organism lives can also have a huge
effect. As a result, the same mutation to the same gene may be harmful in one
individual and harmless in another. Natural selection may favor a mutation in
one set of circumstances and may drive it to oblivion in another.
These processes are taking place all around us every day, and they have
been accumulating genetic changes ever since life began at least 3.5 billion
years ago (Chapter 3). Darwin argued that over such vast stretches of time,
natural selection could have produced complex organs, from the wings of
birds to human eyes. Today, the weight of evidence overwhelmingly supports
that conclusion. Changes in organs are not the only adaptations that have
emerged through evolution; behavior and even language have evolved as well
(Chapters 13 and 14).
To trace the history of these traits, evolutionary biologists reconstruct how
species diverged from a common ancestor (Chapter 4). Natural selection and
other processes can make populations genetically distinct from one another.
Over time, the populations become so different from one another that they
can be considered separate species (Chapter 9). One way to picture this
process is to think of the populations as branches on a tree. When two
populations diverge, a branch splits in two. As the branches split again and
again over billions of years (and sometimes came back together), the tree of
life emerged.
To reconstruct the tree, evolutionary biologists identify which species are
closely related to each other. They do so by comparing anatomical traits and
DNA among many different species. Close relatives share more traits
inherited from their common ancestor. We humans have a bony skull, for
example, as do all other mammals, as well as birds, reptiles, amphibians, and
fishes (Chapter 4). We are more closely related to these animals than we are
to animals without a skull, such as earthworms and ladybugs.
Evolutionary biologists investigate not only adaptations, but the
shortcomings of those adaptations. Life does not evolve by steadily
progressing toward some particular goal. Our apelike ancestors did not
evolve big brains because they “needed” them, for example; the conditions in
which they lived—searching for food on the African savanna in big social
groups—were such that some traits were favored over others (Chapter 14).
Our ancestors benefited from a bigger, more complex brain, but in some
ways it’s a spectacularly maladapted organ. The human brain is so big that it
makes childbirth much more dangerous for human mothers than for other
female primates, for example. It also requires a huge amount of energy to
supply—one out of every five calories you eat is used to fuel your brain.
Evolution is imperfect, because it does not invent things from scratch: it
only modifies what already exists. Any organism can acquire only a limited
number of beneficial mutations, and so evolution can produce new forms
only under tight constraints. Because mutations can have several different
effects at once, evolution also faces trade-offs. Mutations that increase the
number of offspring an organism can have may also cut down its life span.
These tradeoffs underlie some of the diseases that are common in the elderly,
such as cancer. As a result, studying evolution can help researchers better
understand—and perhaps even treat—these diseases (Chapter 15).
Evolution has produced many helpful partnerships in nature, such as the
one between flowering plants and the insects that pollinate them—a
partnership we depend on for many of our crops (Chapter 12). But evolution
does not produce a peaceful balance in the natural world. The same process
species use in adapting to one another can also give rise to what looks to us
like cruelty. Predators are exquisitely adapted to finding and killing prey.
Parasites can devour their hosts from the inside out. Their adaptations are
finely honed, allowing them to manipulate individual molecules within their
hosts. Yet parasites and predators are not evil. They are just part of a dynamic
balance that is constantly shifting, driving the diversification of life but also
leading to extinctions.
The diversity of nature, in other words, is not eternally stable. More than
99 percent of all species that ever existed have become extinct, and, at some
points in Earth’s history, millions of species have disappeared over a
relatively short span of time. We may be at the start of another period of mass
extinctions, this time caused by our own actions (Chapter 11).
A Case of Evolution: Why Do Whales
Have Blowholes?
Evolutionary biology is a science of integration. To investigate questions
about life, evolutionary biologists integrate evidence from many different
sources, gathered with many different methods. Throughout this book, I take
a close look at many examples of scientists weaving together different pieces
of research to understand evolution. These examples run the gamut, from
venomous snakes to drug-resistant bacteria. But there’s no better way to
launch an exploration of evolution than by looking at whales and dolphins
(FIGURE 1.2).
FIGURE 1.2
Living whales all share a number of traits, such as blowholes and horizontal tail flukes. Some species
have filter-like growths in their mouths that they use to sieve small animals from seawater (left). Other
species, such as killer whales (center) and dolphins (right), have peg-shaped teeth that they use to grab
larger prey such as fish and seals.
FIGURE 1.3
In the late 1800s, paleontologists began to discover early whales, such as Dorudon, a 5-meter-long
whale that lived 40 million years ago in Egypt. Dorudon was a completely aquatic whale and had a
number of traits found today only in living whales, such as the tooth and skull features noted in this
figure. The grape-shaped bone called the bulla contains bones of the middle ear; some anatomical
features within the bulla of Dorudon are also shared by living whales. On the other hand, Dorudon also
had some traits not found in living species, including a blowhole midway up its snout and complete
hind legs (albeit very small ones). Living whales have cone-shaped teeth, while Dorudon had more
complex ones that resemble those of certain extinct land mammals. This combination of traits reflects
Dorudon’s transitional position in the evolution of whales. (Information from Uhen 2010.)
In other ways, however, Dorudon was also different from any living
cetacean. Consider its teeth. Dorudon had incisors in the front and molars in
the back. Each tooth had a complex set of cusps and ridges. No living whale
has anything like this dental arrangement. One group of cetacean species,
known as odontocetes, has simple cone-shaped teeth. The other cetaceans,
known as mysticetes, have no teeth at all: instead, they filter out small
animals through baleen. The teeth of Dorudon are actually much more
similar to those of land mammals than of living cetaceans. This trait could
well be the result of the gradual evolution of whales: long after the rest of
their body had evolved many adaptations to the water, early whales still had
teeth like terrestrial mammals.
Cetacean evolution started to come into much sharper focus in the late
1970s. At the time, Philip Gingerich—a young paleontologist from the
University of Michigan—was paying regular visits to Pakistan to investigate
a geological formation from the Eocene period, 56 million to 34 million years
ago. Gingerich wanted to document the mammals that existed in that region
during that time. He and his colleagues brought back a number of fossils to
Michigan and then slowly prepared them in his laboratory, extracting the
fossilized bone from the surrounding rock. One 50-million-year-old fossil—
the teeth and back portion of a wolf-sized skull—was particularly intriguing.
It bore some striking similarities to cetaceans such as Dorudon, especially in
its teeth. Gingerich concluded that he had found the earliest known cetacean,
which he and his colleagues dubbed Pakicetus (Gingerich, Rose, and Krause
1980).
Gingerich’s discovery was dramatic not just for the great age of the fossil
but also for where he had found it. Whales and dolphins today live only in the
water—most live in the ocean, and a few species of dolphins live in rivers.
The rocks in which Pakicetus fossils formed offer clues to the environment in
which it lived. Geologists have determined that those rocks formed as
sediment accumulated in shallow streams that flowed only seasonally through
a hot, dry landscape (Thewissen et al. 2009). This evidence strongly suggests
that Pakicetus lived on land (FIGURE 1.4).
FIGURE 1.4
A: The 50-million-year-old Pakicetus was the first terrestrial whale ever discovered. This
reconstruction is based on fossils from several different individuals. The fossil material is colored red.
B: A painter’s reconstruction of what Pakicetus looked like in life. C: Pakicetus bears certain hallmarks
found in only cetaceans and no other mammals. On the right is a bone (from a dolphin) called the
ectotympanic. It is located in the bulla, which surrounds the middle ear cavity. In living cetaceans, the
inner wall of the ectotympanic forms a thick, dense lip called the involucrum. On the left is the
ectotympanic from Pakicetus. It has a similar involucrum. Such uniquely shared traits reveal that
Pakicetus was an ancient relative of living whales and dolphins. (Information from Cooper et al. 2009;
Gingerich et al. 1983; Luo and Gingerich 1999; Nummela et al. 2006; Thewissen et al. 2009.)
FIGURE 1.5
The skeleton of Ambulocetus shares the hallmarks found today only in whales, such as an involucrum.
It was also the first fossil whale ever discovered with complete legs. A reconstruction of Ambulocetus
is shown on page 2.
The early cetaceans evolved into many lineages, most of them long extinct.
The odontocetes and mysticetes, the two lineages of cetaceans still in
existence today, split from a common ancestor about 40 million years ago.
After that split, the odontocetes evolved muscles and special organs that they
used to produce high-pitched sounds in their nasal passages leading to their
blowholes. Listening to the echoes that bounced off objects around them in
the water, they could perceive their surroundings. Today, dolphins and other
odontocetes use these echoes to hunt for their prey. The mysticetes followed
a different trajectory: They didn’t evolve echolocation. Instead, they lost their
teeth and evolved huge, stiff pleats in their mouths that allowed them to filter
prey from huge volumes of water.
Scientists are now beginning to find important new clues to the origins of
both groups. Fossils have revealed that the earliest mysticetes still had teeth,
for example, and probably grew only small patches of baleen at first (Uhen
2010). Only later did their teeth disappear. Mysticetes still carry genes for
building teeth, but most of them have acquired disabling mutations.
Biologists have long been impressed with the size and complexity of whale
brains (FIGURE 1.9). Aside from humans, dolphins have the biggest brains in
proportion to their bodies of any animal (Marino 2007). Dolphins can also
use their oversized brains to solve remarkably complicated puzzles that
scientists make for them. A number of studies suggest that big brains evolved
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s’ouvrait sur une grande arsa [25] aux vertes perspectives
mystérieuses. Mais je ne songeai plus à regarder nulle chose
lorsque parut Lella Kenza. Car elle est plus belle et charmante
qu’aucune des « vierges aux yeux noirs » dont les bons Musulmans
goûteront les délices dans les « jardins élevés, pleins de sources
vives, où les fruits seront à portée de la main [26] ».
[25] Verger.
[26] Koran.
Lella Kenza est presque une enfant, mais elle possède déjà les
grâces troublantes de la femme. Ses yeux profonds, ombragés par
de longs cils bruns, s’ouvrent, candidement étonnés, sous l’arc
parfait des sourcils. Le nez est petit et droit, la bouche vermeille
comme une fleur fraîche éclose, le teint doré, l’ovale exquis… Des
nattes sombres, piquées d’agates et d’émeraudes brutes, encadrent
son visage, et vont se perdre dans un volumineux turban d’étoffe
dorée. Elle est mince, souple, et chacun de ses mouvements révèle
l’harmonie du corps sous les brocarts aux plis lourds. On dirait une
vivante petite idole égyptienne. C’est la perle soigneusement
cachée [27] qui fut connue par un seul… : Mouley Abbas est son
époux.
[27] Koran.
Lella Kenza m’installa tout près d’elle, à côté de son lit. Elle me
comblait d’amabilités et se penchait constamment vers moi pour me
désigner ses parentes ou me faire remarquer un détail de la fête.
Pourtant je lui trouvai un air soucieux, malgré son apparente gaîté.
— Comment va ton fils ?
— Grâce à Dieu !… L’assemblée est belle, n’est-ce pas ? Tu
resteras toute la nuit.
— Non, non, c’est impossible.
Elle en fut désolée, et, à force d’instances, obtint de me garder
jusqu’au moghreb.
Les invitées ne se départissaient pas de leur attitude rigide,
tandis qu’à l’autre extrémité de la pièce, les cheikhat
accompagnaient rageusement, de leurs instruments, des chants
nasillards. On ne s’entendait plus… il me fallait parler très haut à
Lella Kenza et je perdais la moitié de ses phrases. Elle semblait, du
reste, de plus en plus lasse et préoccupée.
Quelques vieilles femmes, accroupies autour de l’accoucheuse,
tenaient de longs conciliabules. Elles firent apporter sur le lit un petit
canoun allumé, dans lequel on jeta divers ingrédients qui dégagèrent
une âcre fumée. L’enfant fut exposé au-dessus des charbons, puis
frotté avec un liquide mystérieux. Il poussait de faibles cris en
s’agitant.
Lella Kenza le regardait d’un air inquiet.
— Que lui fait-on ? — demandai-je.
— Rien… des choses à nous… — me répondit-elle évasivement,
et elle détourna mon attention sur le thé, le lait d’amandes, les
sucreries et les parfums que les négresses passaient à la ronde.
L’une d’elles offrait aussi de la gouza [35] en poudre, dont les invitées
avalaient une pincée, tandis que leurs regards devenaient plus
vagues et leur expression plus hébétée.
[35] Noix de muscade avec laquelle les Marocaines se
donnent une sorte d’ivresse.