BECOMING HUMAN:

HOW EVOLUTION MADE US

Greg Downey
Published by Enculture Press
Smashwords edition

Copyright 2013 Greg Downey

Published by Enculture Press, Australia

PO Box 40, Berry, NSW 2535, Australia
ISBN: 978-1-925082-01-2 Smashwords edition
First edition 2013

Smashwords Edition, License Notes

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Table of Contents
Open2Study
Acknowledgments
Preface
Chapter 1: Traces

Chapter 2: Darwin
Chapter 3: Sex
Chapter 4: Brains
Photo credits
Glossary
About the Author
Enculture Press

~~~~~~~~

This work is licensed under the Creative Commons Attribution-NonCommercial 3.0

Unported License. To view a copy of this license, visit
http://creativecommons.org/licenses/by-nc/3.0/. For more information on Creative
Commons licenses, visit CreativeCommons.org.
Please contact Greg Downey for information on licensing: [email protected]
This ebook is distributed free through the support of Smashwords, Open Universities
Australia and Enculture Press. The text and resources accompany the Open2Study class,
Becoming Human: Anthropology.

Open2Study
Free online study for everyone!
The Becoming Human: How Evolution Made UseBook is an ideal companion to the free
online subject Becoming Human: Anthropology - presented by Greg Downey and offered
by Open2Study.
In Becoming Human: Anthropology, you can explore how evolution works and how
variation arises. Find out why, of all the orders of life, primates produced us. How did
apes start to look like us, walk on two feet and grow big brains that over the past 200,000
years have figured out where we came from? This subject will give you some thoughtprovoking answers.
Open2Study is brought to you by Open Universities Australia. Open2Study brings you
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Acknowledgments
Becoming Human started off as a kind of dare.
Well, more than one dare.
In a conference on the future of teaching, I dared my employer, Macquarie University, to
embrace open education more fully. I warned that open exchange of teaching materials
and online learning was inevitable.
My Associate Dean for Learning and Teaching, Prof. Sherman Young, and Macquarie
Universitys Provost, Prof. Judyth Sachs, dared me to back up my high-sounding call
through our partnership with Open Universities Australia. In April, 2013, we launched
Macquarie Universitys first MOOC, a massive online open classroom.
This book was prepared to give the students an opportunity to read more closely and to
find additional resources and links on the topics covered in that course, Becoming
Human: Anthropology.
I have to thank those who have helped at every step. Robert Parker, at Macquarie
University, for pitching in all over the place, from conceptualising the course to

struggling to reduce thirteen-weeks-worth of material down to six and then to four

to helping with the creation of the simulator and managing shooting in Melbourne
In Melbourne, Greg Bird did a masterful job, with tight time constraints, of tracking
down an immense variety of images to illustrate the lecture-videos and for this book. You
can see his work on many pages. He also gave some great feedback on educational design
issues.
Thanks to everyone at Macquarie University who made this possible, especially Sherman
Young and Judyth Sachs.
Open Universities Australia has been excellent support
Thanks to the crew at Smart Sparrow who produced the simulator that is used in
Becoming Human. That project had a very steep learning curve, and with David
Schnstein especially, that part of the project would never have gotten off the ground.
The video production team at Open University Australia in Melbourne was great. Thanks
to Asanka, Jessica, Evan, Roberto, Jason, Kate, and the entire crew that made the whole
production a really high-energy good time in spite of the fact that we were slowly baking
in a small studio together.
A special debt of gratitude to my wife, Tonia Gray, and my daughter, Mikhaela.

Preface
When colleagues find out I teach human evolution at Macquarie University and Open
Universities Australia, they often ask me, Why? Sure, its a fascinating topic, but its
one that I was not trained to teach.
Like many anthropology students in the United States, I took archaeology classes and
studied human evolution when I was doing my bachelors degree. Two of the many
standout classes that I took at the University of Virginia were Human Origins by Jeffrey
Hantman and an intense seminar on Mayan archaeology.
But I was trained as a cultural anthropologist in graduate school, prepared to make sense
of living peoples cultures, to pick up new languages, to live with a group who has a very
different way of understanding the world and over time make sense of their
perspective. In some places, committing to a career in cultural anthropology can mean
putting aside evolutionary research and theory because they can often seem only distant
causes of contemporary social life. Teaching human evolution, about bones and biology,
brains and genitalia, would seem to be a stretch, even for someone who didnt know the
field.
To someone who does know anthropology, to my fellow anthropologists, the jump from
cultural to biological anthropology can seem even greater. When I took my PhD back at

the very end of the 1990s, biological and cultural anthropologists were often convinced
that their greatest adversaries were in the other camp: cultural theorists had to argue that
culture overcame or transcended biology, biological researchers that culture was just a
superficial veneer on a universal biological reality. From this point of view, for a cultural
anthropologist to teach human evolution was a bit of betrayal. Do you talk about
genetics? some of my cultural colleagues would ask.
Fortunately, that tension has changed.
In the past decade or two, integrative approaches that bring together evolution and social
causes, nature and nurture, are in ascendance as the research problems we face illness,
cognitive abilities, differences between men and women make a mockery of trying to
divide up the facets of what make us human.
Im not alone in trying to bridge the gap these days between biological and cultural
approaches to the study of human nature. Anthropologists these days realize that we are
not really arguing most with each other: we are arguing primarily with people who pay
no attention to human diversity, with commentators who refuse to recognize the
importance of evolution, with overly simple explanations for human institutions that we
know to be complicated.
But most importantly, anthropologists are arguing with people who think they know how
evolution works or think they know about human nature, but dont really have a grasp
of the fascinating research results that our field has found over more than a century.
Often, I will read an article in the popular press where a science writer has been talking to
some other researcher who thinks he or she knows about human evolution or human
diversity (but really mostly does work in a laboratory or by surveying university
students). And the article will make proclamations about our species, throughout its
history, or Humans are all this way because evolution makes us this way, and I know
that the evidence the hard, cold bones in the ground, or the abundant ethnographic
record that comes from studying peoples all over the planet doesnt support what
theyre saying.
So thats why I teach human evolution, and why Ive written this book. To me, you
simply cannot understand how humans are today, right now, all around us, if you do not
understand evolution.
However, if you understand evolution badly (including listening to what some quite
famous commentators say about it), a little knowledge can be almost as misleading as
none at all. And what I read in the newspapers makes me feel like a lot of people out
there who think they understand evolution have just a little bit of knowledge and dont
even realize it. The closer you get to the scientists, the more interesting the research is,
and the less it boils down to a simple story.
The point is not that everything is complicated or that a particular scientist or
evolutionary theorist is dead wrong. Often, partial truths can become serious distortions
if over-applied; theories that are enormously helpful in clarifying how we see the world
can become constraints on our ability to see all the evidence if we cling to them too

tightly, as if they are sacred texts. Biology is endlessly surprising; thats one of the great
joys of studying evolution.
But human are even more surprising than most species. Human evolution is a very
special case, even if we share so much of our DNA with the other living things on the
planet.
~~~~~~~~
To make a long story short, I teach human evolution because, without it, we simply
cannot understand ourselves. But if we understand evolution badly, or we cling to a
particular theory even in the face of its exceptions, a little evolutionary thinking can also
be a problem. I needed an introduction to human evolution that prepared people for the
full breadth of anthropology, biological and cultural, archaeological and linguistic.
This electronic book is an attempt to provide a very basic knowledge of human evolution
that doesnt teach an overly simple version of evolutionary theory. Because Im a cultural
anthropologist, Im interested in how our evolutionary legacy continues to affect us to the
present day, including how it makes possible all of the technological achievements, social
complexity, and cultural variety around the planet.
If we understand evolution, we also understand two very important facts: first, that
evolution is certainly not over or finished with us. Second, we have the power to affect
our own, and other species, evolution. What we do can change the forces of natural
selection, although this may seem counter-intuitive.
In fact, were not the first species to have this power. Were only alive because other
generations of life, long before our ancestors arose, changed the very nature of the planet.
But were doing it now, more than the general public probably realizes, and we need to
be aware of this as we make important political, economic, and technological choices
going forward.
Im not just talking about the temperature of the planet or the range of species with which
our descendants will share our planet although both of these are important. One lesson
of evolution is that life changes. Recognising how we are participating in that change,
and may be affected by it in the future, is one of the great challenges of contemporary
anthropology, in cooperation with a lot of other researchers.

~~~~~~~~

Traces
One reason that we study human evolution is that the origin of our species is relevant to
the way we are today. We can see the effects of evolution in our bodies, brains,
behaviours, and so much else about us. But we are also left with some pretty challenging
questions, especially as biological research shows us the deep links connecting all life:
How did humans get to be so unusual?

Were 1-in-450-million
There are at least 9 million species alive on the planet Earth today. Some scientists say
this estimate is too low; they say the real number may be as high as 30 million species.
But this 9 or 30 million represents only a tiny fraction of all the species that have ever
lived. Some scientists argue that over 98% of all the species that have ever lived are now
extinct. Not just the dinosaurs, but countless, innumerable other species, some of them
killed off in five massive extinctions in the history of our planet. In the worst of them, the
Permian-Triassic extinction about 250 million years ago, over 90% of all marine species
died even insect species were decimated and it took the variety of life 10 million
years to bounce back.
So conservatively, and I mean very conservatively, theres been at least 450 million
different species, most of which are now extinct. All that variety. All these amazing traits,
from frill-necked lizards and towering sequoias and narwhals to penguins and armadillos
and giant colonies of fungi.
Animals that can live in the most hostile environments, such as bacteria that live by
oxidizing iron at the bottom of the sea; xerophyte plants that survive in deserts; microbes
that can live in deep salty basins in the Mediterranean, others that thrive in the hot springs
at Yellowstone National Park or deep under Antarctic ice. Life deep underground and in
the ocean, even inside other life. Creatures that blow our minds with their amazing ability
to survive and adapt, proving that evolution will find a way, even in conditions that we
think might make life impossible. Enough strange creatures and exotic species for a
lifetime of great nature documentaries.
And yet, in a very important way, were all alone. Humans are a one-in-a-half-billion
experiment. Conservatively.
You would get better odds playing Powerball or winning the lottery. No other species has
manufactured complex tools, built cities, gone to the moon, written novels, composed
symphonies, or produced Lady Gaga. Of all the 450 million or more species, potentially
many times more, we seem to be one of a kind. Utterly unique... well, on our planet at
least.
This is the central mystery of human evolution: how did evolutionary forces, forces that
equally affect the millions upon millions of other species, produce such a strange, balding

ape with such peculiar social habits and extraordinarily odd cognitive abilities?
How did evolution produce us from a set of biological building blocks that we
increasingly realize we share with other living things? We share a huge proportion of our
DNA with chimpanzees, our nearest cousins, for example some put the number at over
98%. We share a striking proportion of DNA, however, with every living thing. As
anthropologist Jonathan Marks put it, we arent just 98% chimpanzee, were 30%
daffodil or banana.
If we look at the cells of living things, the similarities are obvious, even between plants
and animals. The chemicals that make up DNA, the four base codes, are shared; the same
four bases combined over and over again in distinct ways to make all life that we know.
Evolution wrote the book of life with a remarkably small set of letters.
And yet, that difference, that 2% between us and chimpanzees, for example, is obviously
a giant gulf. What makes the gap so great?
Sure, its a narcissistic question. Yes, there are 9 or 30 million living species out there,
millions upon millions that have ever lived, but lets talk about us. Yeah, were
fascinating, especially to us. Well, and to our dogs, too, I suppose, ... although probably
not for the same reasons.
Its not just that evolution produced us. Evolution produced an animal capable of
understanding evolution. Thats extraordinary. A biological process, billions of years old,
somehow produced a creature with such extravagant cognitive and social abilities that,
working together over about the last 200,000 years or more, weve figured out where we
come from.
Thats what well be exploring over the course of this book: how does evolution work?
How does variation arise? Why, of all the orders of life, did primates produce us? When
did apes start to look like us, to walk on two feet, and why? How did our ancestors
reproduce, and why do we have such helpless babies, so fragile and needing attention?
And finally, whats the story of these big, versatile brains that we possess, that allow us
to do so much? And whats the price of these big brains, a price many of us may not even
realize were paying?
This book is about our origins. How did we get here?
Although this book will link to documentaries and websites, however, the materials are
part of a bigger project. This resource was created for an open classroom, which means
that we hope to provide you with a chance to talk with other people who are interested in
the same subject, to trade ideas, resources, advice, and to learn more together. This book
will present things quickly, as do the lecture videos that you can find at Open2Study, but
theres lots of space to go deeper into the issues or changes that you find most
fascinating. I hope to use this book to provide you with some great places to explore
further.

Introducing Anthropology
This course (and the accompanying book) is an experiment for us, both in the Department
of Anthropology at Macquarie University and at Open Universities Australia weve
never thrown open our doors quite like this, to anyone, anywhere. Were trying all new
sorts of technology and approaches to what were doing. We wanted to choose a topic
that is, at the same time, really exciting, with lots of new research and science, but also
philosophically deep and socially important, asking questions that go to the core of who
we are and what matters to us. We hope youll enjoy it and that youll really join in and
enrich the discussion. One thing I know from teaching human evolution to thousands of
students is that all kinds of people are fascinated by, well, humans. Our story. It may be
controversial, but Ill do my best to share how I think, and I really hope that youll
contribute as well.
Ill be your instructor, in part, although youll also learn a lot, maybe more, from each
other. Im Greg Downey, an anthropologist trained in the United States, but who moved
to Australia in 2005 because I married an Australian.
When I started out as an anthropologist, I didnt study or write about evolution. I went to
Brazil to study capoeira, an Afro-Brazilian martial art. Capoeira is a demanding,
acrobatic martial art that combines elements of dance and fight. Its really hard and takes
years to learn; all the training changes the people who do it. Capoeira practitioners build
muscles, become faster and more flexible. They even perceive, move, and react
differently like a lot of martial arts and sports and practices like yoga.
I worked for years on capoeira and wrote a book about it, moved on to study cage
fighting or mixed martial arts, and have recently been working on a range of subjects
including rugby.
The more I studied sports and how people learned skills, the more I realized that I was
studying perhaps the most important thing that makes humans different from other
species: we come into the world incomplete. Our bodies and brains can be trained to do
an incredibly wide range of things. And we change ourselves, forcing our bodies and
brains to adapt to everything from language and education to music, the internet, our
hobbies, along the way learning from each other and accumulating generations upon
generations of knowledge.
So a few years ago, I began to do neuroanthropology, to study the relationship between
brain and culture. Neuroanthropology includes how our brains produce culture, but also
how our cultures shape our nervous systems. For example, being born into a community
with language means that parts of our brains and ears get shaped by that language. By the
time youre a year old, your hearing can pick out speech sounds in your native language,
and starts to lose the ability to make fine distinctions important in other languages. To
learn to write, or balance on your head, or chase down your dinner on the savannah or use
a use a laptop computer means gearing up your body and brain for these purposes.
At the same time, all these activities leave their traces on your body and brain. The
illnesses we get, our aches and pains, our senses they all wind up being shaped by

patterns of activity.
The only way to understand this human process, the way we shape ourselves, in my
opinion, is to combine the study of culture and biology. We have to recognize that
evolution had made us with enormous potential but leaves a lot of the finishing off to our
complicated social and symbolic life. Evolution had made us biologically susceptible to
culture, just as it had made us capable of producing culture. We are the unfinished ape,
the self-made species.
Figuring out how that works, how evolution made us, in my opinion, requires an
anthropological perspective. Anthropologists are famous for three things in particular:
First, we study people in the field, where they live, or have lived, not in laboratories.
Anthropologists are good at messy, good at digging, good at hanging out and observing
what people do. We go to where people (or other primates) are, even if that requires a
plane, train and boat, even if that means learning difficult languages or exposes us to
nasty sicknesses. How are you going to study the ways humans evolved if you arent
ready to chase them to the four corners of the earth?
Second, anthropologists study human diversity. We dont sit at home or in a caf and
write deep thoughts about human nature well, sometimes we do that, I suppose and
we dont just study people like ourselves. We may start by studying a specific group, a
tribe, a community, or particular ways of life, sites where our ancestors have left remains.
We cast the net as wide as possible and explore the full range of the human condition.
And finally, anthropologists dont respect intellectual borders. Just as I had to start
studying biology to understand what I was learning from capoeira practitioners in Brazil,
teaching myself neurology and exercise physiology and a whole range of other things,
anthropologists believe that you have to look at all the data, even if it stretches your brain
to the breaking point. We call this a holistic approach. To study human evolution means
looking, not just at human skulls and bones and brains and DNA, but also at other
primates, at early tools, at languages, at child rearing, at human sexual variety.
So welcome aboard, and strap in. One thing youll learn, I hope, is that we dont yet have
the final word. New findings might still turn some of the things we thought we knew
upside down; I cant use all my slides from five years ago because we know more than
we did then.
But the evidence we have is making the story we can tell about our origin more and more
complete. I welcome you to bring to the discussions even perspectives that I dont
endorse 100% because its only by having a rollicking, open debate, by putting all the
evidence we have on the table, that were going to get a better and better understanding
of what it means to be human, and how we got here.
Evolution isnt over. Were still becoming human. And arguing about how we got here,
and what the implications are, is part of this process.

When you had a tail...

Once upon a time, you had a tail.
Im not singling you out. We all did.
Its nothing to be embarrassed about.
First off, we still have a tailbone, a set of fused vertebrae called the coccyx. If youve hit
it hard, such as slipping on ice or falling on your bum, you know what Im talking about.
At the very bottom of our spines, in the middle of our pelvis, there are these vertebrae
that are fused together into one triangular lump.

Figure 1: the coccyx

In x-rays, you can see where five vertebrae have fused together; the diagram shows the

suture points, where the bones knit together as you grow. They start out separate, like our
other vertebrae, but as we grow, they fuse into one chunk and dont grow as fast as our
other vertebrae. This makes the vertebrae that form the coccyx sort of useless, but they
remind us that our ancestors had tails, just like other mammals.
We call this a vestigial organ a body part that is a vestige of a previous structure.
Defining a vestigial organ is tricky, because organs like a tailbone are often not totally
useless they get reused by the body for new purposes. The key is that they do clearly
show their earlier function. In the case of the coccyx, it is that clear fusion of these
vertebrae, the coccygeal vertebrae, that lumping together as we grow of bones that would
have been the start of a decent tail for our ancestors. The lumping shows us the bone is a
vestige of our evolutionary past.
Truth is, weve got lots of these little traits in our body, weird design quirks that show us
that the current function of a body part is a new app, a new deployment of a trait. Weve
got furry bodies, remnant of a lot hairy ancestors, and we break out in goosebumps when
were cold or scared.
Goosebumps are that fur standing up. Fluffing their fur when cold would have kept our
mammal ancestors warmer. Back then, the hair was much thicker, and this fur would
have trapped a layer of air, like a puffy insulated coat.
Puffing up when youre scared would have made you look bigger, more intimidating.
Lots of animals try to look larger when theyre threatened, hopefully so that they look
less like a push over or an easy meal. Now, weve just got these mostly useless hair
follicles that some us insist on tweezing off or manscaping.
Some of these features are really vestigial in that they dont seem to have a lot of use
any more. Some people argue that our appendix is vestigial, left over from an earlier
model of a gastro-intestinal tract. You can take a persons appendix out, and they survive
pretty well. I should know, as I had mine removed twenty years ago. But some
researchers think that the appendix is a back up organ to aid in the recovery from severe
gastro-intestinal problems, a place where all the microbes in our guts that we need to
digest our food right can hang out in case of trouble.
But perhaps an even better example is the little bit of muscle at the top of the ear. If
youre sitting with someone, look at the top of the ear. In many, youll notice one part of
the top of the ear is slightly thicker than the rest.
Youve got a bit of muscle there, but, because its not connected to anything solid that it
can pull on, you cant use it to move your ears around. If you could, you could hear
better, shaping your ear to collect sound. Some of our evolutionary cousins can use that
muscle to change the shape of their ears, to hear better, to communicate with each other,
and to perform party tricks that we can only dream of.
Even structures like toes are vestigial to some degree, although theyre useful for some
kinds of things, like getting stubbed or wearing toe rings... or driving a car or braiding
your daughters hair.

Seriously.
Weve got these digits on our feet that most of us dont use much. Theyre vestigial from
our primate ancestors who could do all sorts of cool things with their prehensile toes, like
climb and hold their children.
But the truth is that we could do a LOT more with out toes if we didnt box them up in
shoes and let them get lazy. Some people, born without arms, or congenital amputees,
show us just what is possible, even with our short little toes you can find examples of
people playing guitar, driving cars, and caring for their children with their feet.
Stephen Jay Gould and Elisabeth Vrba (1982) wanted to call parts like our tailbones and
body hair and toes exaptations rather than vestigial organs. The word vestigial
sometimes gets taken to mean useless, but thats not true of either tailbones or toes.
Rather, tailbones and toes are exapted: they arise for one purpose, but they get
repurposed, recycled or reused for some new function. Exaptation points out that
evolution tends to be really good at putting old structures to new purposes rather than
starting from scratch. In evolution, you never get a complete redesign; you always get a
renovation.
How does the body do renovations? Well, lets go back to our tails. When I said you once
had a tail, I was just talking about it vaguely, like you, you descendant of monkeys. In
reality, however, Im talking about you, specifically.
Back when you were a little embryo about 30 days old, if we had good full-body pictures
of you, you would have looked something like this. Yeah, I know, not exactly the baby
picture you put up on Facebook is it?
You were only about 10 mm long back then, and more than 10% of your body length was
tail not five undersized fused vertebrae, mind you, but ten to twelve vertebrae in
embryo as impressive as the ones that went on to form your spine and neck. In this cross
section of the human embryos tail (Figure 2), you can see clearly that it has all the
precursor structures you would need to go on and form a fully functioning, waggable tail.

Figure 2: Cross-section of the human embryo's tail at Carnegie stage 14. Neural tube
(n), notochord (c), developing vertebrae (somites, s), gut (g), and mesenchyme (m).
So what happened? Well, your DNA and the interactions of your cells eventually told
your body to cut off those vertebrae, to starve them of resources, and the cells were
reconsumed by your body, especially your immune system. The reason is that the genetic
program to build you is quite similar to the ones that build other, similar animals, animals
that do go on to have fully formed tails. The ingredients can be quite similar even if the
instructions vary.
So, during human embryonic development, we have what could turn into a tail. In the
bodies of many species of monkeys, similar vertebrae go on to be a fully-fledged, treebranch-grabbing tail. But our DNA says, Nope. Scratch that. Thats not in our plan for
remodeling. So we dont get a tail, and the body closes up shop on the whole tailbuilding project.
However, every once in a great while, the renovations arent spotless, the instructions
dont quite get transferred flawlessly. A slight difference in the developmental recipe
leaves a bit of the tail behind, a bit of the tail that we all once had.
And then you get a fully fledged caudal appendage an actual tail. Theyre rare in
humans, really rare, but weve got a number of recorded cases. In this case, a posterior
aspect of a neonate thats science talk for a picture of a babys butt shows a clear
2.5 cm or about a one inch tail. The technical name is either a caudal appendage or a
saccrococcygeal mass because its actually easier to tell a mother, Your child has a

saccrococcygeal mass, than to say, Sorry lady, your babys got a tail.

Figure 3: X-ray image of an atavistic tail found in a six-year old girl. Image reproduced
from Bar-Maor et al. 1980, Figure 3.
And heres another great example (Figure 3): an x-ray of a 6-year-old girl with a full-on
soft tail in the middle of her back. Normally, S1-S5 would have all fused during
development in embryo into a little tailbone, about the size of S5 on this x-ray. Not only
had they not fused, and gotten really large, but this girl also kept C1 to C3, part of those
extra vertebrae that all of us have in utero but that disappear due to programmed cell
death in the vast majority of people.
We call this kind of trait an atavism.
An atavism is a sort of throwback; its the re-emergence of a trait that doesnt show up
in the immediate ancestors, but was in a more ancient ancestor. Its not a new thing or
some kind of innovative mutation. Its an old trait, a re-appearance, sometimes a very old
trait re-emerging.
These cases are really rare. Atavistic tails we probably have about 100 welldocumented cases. So why should we care about them? Arent they just freaks or
mutants?
I dont think so. Theyre unusual, but cases of atavistic tails, like our own tails in embryo
development and our vestigial tail bones traits all of us have show us that our

bodies are littered with traces of our evolution. They are an example Ill return to next
week when I talk about evolution and genetics because they show us two important
things about how evolution makes us:
First, atavisms and vestigial organs and exaptations show us that evolution works by
repurposing structures, using what it has lying around, not usually by making something
completely from scratch. One of the main ways development does this is by tinkering a
bit with the development program, cooking the basic ingredients like vertebrae in a
tail in slightly different fashions. In one case, you get an underdeveloped and pretty
useless vestigial tailbone. But in another related species, you get something else entirely
a magnificent tail thats useful for all sorts of things.
Second, traits like atavistic tails show us that we, like all life, have a lot of unexpressed
genetic potential lying around in our body. Unusual cases like an atavistic tail remind us
that some of these traits are there, dormant, maybe dismantled or checked by some
developmental process, what biologists call unexpressed, but that these developmental
programs can still produce a trait, given the right conditions. That said, youre not going
to get a tail just because you want one really bad, or because you buy lots of pants with
extra space in the seat youve got to get the right genetic material at the start.
This is one of the key points of studying human evolution. Human evolution is not just
the story of an ancient past; its also one of the only ways to understand the strange
bodies that we have, especially these design quirks.
The biologist Theodosius Dobzhansky (1973) famously titled a short essay on the
compatibility of science and theology, Nothing makes sense in biology except in the
light of evolution. When we look closely at the traces of evolution in our body, we
realize how right he was. Only evolution can explain why you once had a tail..

Getting hands from paws

Some of the best evidence we have of evolution, in our own skeletons and tissues, is
probably sitting right in front of you: your hands. On the one hand, human hands are so
unusual; we can type and play video games, fight and climb and carry our children and
even communicate.
On the other... hand, if we were to peel back the skin and look at the structure of our
hands, we find that they were similar, strikingly similar, not just to the hands of our
closest relatives the other primates like chimpanzees and monkeys but also to
animals that might surprise us, like dogs, cats, horses, whales, and even frogs.
Biologists call anatomical similarities like these, homologies. Homologies are shared
structures among species that are a result of evolution from a common ancestor. Even
though two species may look quite different, even have contrasting abilities, when we
look more closely at the internal configuration, the chemical and organic processes, the
anatomy of organs, we find amazing similarities.

British anatomist Richard Owen, the man who gave the dinosaurs their name, from the
Greek for terrible lizard, and who established the British Museum for Natural History,
originally defined the word homology in 1843. Owen coined a lot of terms, most of
which never caught on, but hes also given us a few rippers.
Owen wrote that a homology was the the same organ in different animals under every
variety of form and function. Anatomists of his day who systematically dissected
animals found surprising structural similarities underlying different animals organs.
When you peal the skin off, even of animals that look quite different, you often find that
the organs match up remarkably well. The skeleton of a mouse and an elephant, a bat and
a dolphin, often match up bone for bone, even though those bones have different
proportions and functions. If you line up the teeth of the elephant with other mammals,
for example, you find that the tusks actually match up to the incisors in other animals;
theyre massively extended front teeth!
To get a sense for how homologies work, lets take a look at my dog Louies front paw.
From the photo, you might not think it looks much like your hand. He cant pick anything
up with it, cant hold a pen, and certainly cant open a can of dog food for himself. He
mostly just runs around on it and uses it to scratch himself.

Figure 4. Louis, trusty dog assistant.

But if we were able to look at an X-ray of his paw, like this drawing, you would see some
striking parallels with your own arm and hand (see Figure 5). Like you, Louie has a

humerus connected to what we call our shoulder, and a pair of bones in the middle
segment that match up to your radius and ulna. Louis has what looks like an extra joint in
his leg that actually matches up very nicely to your wrist, with the carpals, or the wrist
bones, and the metacarpals, which go from your wrist to your first knuckle in your palm.
In Louie, however, the metacarpals, which seem like part of your hand, probably look
more like part of his leg than his foot. Louis even has a little fifth digit whats called a
dewclaw up a little ways on his leg. The dewclaw is a homologue to your thumb.

Figure 5. Homologies in the mammal forelimb.

If we look closely at a diagram, Louies front leg is almost like seeing your arm and hand
skeleton in a funhouse mirror, or using Photoshop to twist and distort the same basic
collection of bones into a shifted configuration.
Even in a mammal with a radically different looking limb, like a horse or a dolphin, we
find variants of this same basic skeletal anatomy (See Interactive 1.2). The structure is
adapted to serve slightly different purposes depending upon the needs of the animal.
Homologies are most impressive when we find similarities in what look like quite
different structures descended from a common ancestor.

Figure 6. Jeffrey, chillin...

This, for example, is Jeffrey (Figure 6). Again, Jeffreys front leg doesnt look much your
arm, but we find homologies with the human arm and the dogs leg. At the same time, we
also see how, because of the way the limb has adapted, the changes are greater. With
Jeffrey here, you have to realize that hes essentially standing on his middle finger, and
that his middle finger (that is, his hoof) doesnt have all the joints that you or I have; hes
just got one joint.
But hes got a humerus, a radius and ulna in the next section leading down to the knee or
what is called in veterinary anatomy, the horses carpus. The carpus contains a sevenbone configuration that is homologous to our wrist. The lowest part of the horses leg,
immediately above the hoof, is the canon. Its homologous to the middle bone in a
human palm, or the third metacarpal if youre counting in from the side. In the horse, this
third metacarpal is fused with the neighbouring two bones (the second and fourth
metacarpals in a human hand), like your five lowest vertebrae get fused into a coccyx.
The fetlock, this lowest joint on a horses leg, is like your first knuckle on your middle
finger.
To get the bones of this leg and hoof, however, you really have to put those homologous
structures we find in the human through some serious distortion. The phalange, for
example, above the horses hoof, is massive compared to the first digit in the human
hand. Similarly, theres a lot of extraneous little bones in the human hand, so essential to

doing what we do with our forelimbs, that never develop in the horses hoof. Just like
with the human tail, the developmental program of the horses leg stymies the full
development of these additional bits.
In contrast, in a dolphins fin or a whales front fin, you also find a pattern of
homologies, but you see contrasts: the humerus, radius and ulna, for example, are all
compressed to form a solid base for the limb when its a fin. Whereas in the horse, the
phalanges are simplified significantly, in whales and dolphins, the number of phalanges is
multiplied, like they have super-long, multi-jointed fingers. In the bats wing, again, we
find the five digits, as well as the radius, ulna and humerus all stretched to support the
wings. Here, the bats individual phalanges are incredibly elongated relative to other
mammals, because the surface area of the wing has to be so great, while still keeping the
skeleton light. The contrast couldnt be marked between the dolphins fin and the bats
wing, a result of one propelling itself through the air, and other needing to move through
the much thicker medium of water.

Direction in the selection

In all the cases of homologies we talked about in the last section, we see how different
species have taken a shared basic set of structures and adapted to a range of different
environments and functions. Homologies show us clearly how evolution doesnt tend to
make wholesale change, but rather, continually tinkers with structures and find ways to
adapt them for new functions. Adaptation isnt about a perfectly engineered, specialpurpose organ, but rather the gradual preservation of the best suited structures, and the
development of better and better suited variants.
We call this directional selection. When selection favours one extreme version of a trait,
like a strong structure in a fin, or a light, elongated one in a bats wing, that trait will get
pushed in the direction of greater and greater specialization.
In the case of the tetrapod forelimb, that is, the front leg of four-legged animals,
directional selection has driven different animals structures in diverse directions, leading
them to diverge in a wide range of ways. This kind of dispersal, we call adaptive
radiation: a small set of animals is replaced over time by a larger set as clusters of the
animal specialize and diverge in different directions.
We can also see how constrained adaptation can be by the process of directional selection
and homology. Bats do not develop the ability to fly by growing a new set of limbs; their
forelimbs are adapted for the purpose. So evolution can involve trade-offs. Is a change,
on the whole, more likely to allow a species to survive and reproduce successfully
because growing wings is going to cost the bat? If so, that variation will be preserved.
But that doesnt mean evolution is like Santa Claus: it doesnt give animals presents out
of the blue. It works one what they have, and is constrained by what is biologically
possible. Tetrapods dont grow extra pairs of limbs (although these kinds of extraordinary
changes can happen in some extreme cases).

Figure 7. Homologies in a variety of tetrapod forelimbs. 120, salamander; 121, sea

turtle; 122, crocodile; 123, bird; 124, bat; 125, whale; 126, mole; 127, human
Not every similarity between organisms is a homology; sometimes, the processes of
evolution, including natural selection, produce traits through convergent evolution.
Convergent evolution is a similarity that arises, not so much from shared ancestry as from

selective pressures, sometimes even quite similar selective pressures. For example, even
though wings in insects, bats, birds and pterodactyls developed quite differently, the
demands of flight led to some similarities, such as their lightweight structure, strength,
and flexibility.

Figure 8. Convergent evolution in wings. The wings of pterodactyls (1), bats (2) and
birds (3) are all light-weight with large surface areas, but their diverse structures show
that they evolved these traits through convergent evolution.
Unlike homologies, when you look closely at examples of convergent evolution, you
often find quite different underlying structures. For example, the structure of the wings of
insects, birds, bats and pterodactyl are quite different: the pterodactyl has a bit of a claw
on the middle joint because the wing is essentially a giant membrane stretched between
the animals body and a single, enlarged fifth digit. In contrast, a bats wing has four
digits with membrane stretched between them. The bird has yet another configuration
with a simplified skeleton and feathered skin to increase the surface area; feathers
actually decrease the need for the elongated digits in bats and pterodactyls. Flying insects
are wholly different because they are such a distant relative of the other three, with no
skeleton to reinforce their wings. Insects usually have two pairs of wings, composed of a
thin membrane supported by veins.
These structures are analogues because they have developed independently, although the
similarities between the tetrapods (the bird, bat and pterodactyl) and the radically
different structure of the insect wing, shows you that the first three are more closely
related.
Convergent evolution can produce fascinating examples of quite similar structures even
in distantly related animals. For example, in Australia, placental mammals only arrived
quite late in the continents biological development, first just bats and rats, and later,
tagging along with humans, the dingo and other invasives. Here in Australia, we have a
wide range of marsupials that resemble non-marsupial mammals occupying similar
ecological niches. For example, we have a marsupial possum, a marsupial mouse, a
marsupial mole, we even had a marsupial predator, like the Tasmanian tiger which was a

bit like a bobcat, and an extinct saber-tooth marsupial cat.

The similar pressures on these animals caused them to evolve in parallel ways to their
counterparts among their distant cousins in the placental mammals. Although its a little
more complicated than this, as well discuss next week, this kind of evolution, where
different paths lead to a similar destination, is an example of convergent evolution or an
evolutionary analogue.
In fact, the homologies across life can stretch much deeper. As our tools for genetic
research have gotten more sophisticated, we now find that some of the genes underlying
structures across virtually all of life are quite similar. In biological terms, they are highly
conserved, meaning that, even though they are in quite different animals, only distantly
related, the underlying structure in the genes can be really similar.
One of the most interesting examples of this sort of deep homology are the genes that
control overall structure of an animal, the Hox genes. The Hox genes specify which parts
of the body go in what order. Although most genes are scattered around, the Hox genes
are lined up pretty much in the order that they specify: the Hox gene that designates the
mouth first, followed by the head, thorax, and abdomen. If a Hox gene gets duplicated in
its genome, an animal often gets another copy of the body segment as the Hox gene
basically runs the developmental sequence for that body segment an extra time.
Well come back to it, but the Hox genes are basically the master control genes for body
configuration, switching on cascades of other genes at the right times, making sure that
an insect doesnt get legs on the top of its head, or a mouth in the middle of its abdomen.
Once geneticists figured out what Hox genes were in the 1980s, they went looking for
them in a range of animals. What was striking is how similar the Hox genes were across a
wide variety of species, even between simple worms and quite complex animals. For
example, the Hox gene Pax6/eyeless seems to control the formation of light sensitive
eyes in all species. There were differences, of course, such as the fact that mammals have
four copies of the whole Hox sequence, but the toolbox of these genes tends to get used
over and over again.
Although these discoveries have a number of implications, some of which well discuss,
perhaps the most obvious is that they are the trace of our shared origins: the highly
conserved, very ancient basic tools used to build all of the life we see around us. They are
evidence that evolution has made a multitude of forms from simple beginnings. Charles
Darwin suspected as much. In his book, On the Origin of Species, he wrote,...
Therefore I should infer from analogy that probably all the organic beings which have
ever lived on this earth have descended from some one primordial form, into which life
was first breathed. (Charles Darwin)
We are all related. Not just all humans, one to another, but all living things. Should make
for a hell of a family reunion.

Designed by evolution?
The last bit of evidence for human evolution that I want to bring up this week is the
blueprint or design of the human body. If you study the human body in functional
terms, looking at how it works, it turns out that there are lots of design glitches that point
to its evolution. The problems and flaws are telling, helping us to see the fingerprints of
evolution; in a sense, theyre just as interesting as the remarkable adaptations are.
In fact, I dont like using the word design at all to talk about the human body, or any
other evolved organism, because, to me at least, the word design suggests that theres
some kind of designer, some sort of plan.
Evolution has no plan. The body is not the product of a design, but rather of an organic
process, over millions upon millions of years.
But before we get into that, let me show you what I mean.
First up, your eye, if it was designed, could be better. Probably the most obvious flaw is
that the photoreceptors that detect light on the retina are all put in backwards. The layer
of nerves that connect the retina cells to your brain arent put where they obviously
should be, if the eye was actually well designed: the nerves are inside the eye, covering
the receptors, and not on the back of the eye toward the brain. So the light that gets to our
retina has to pass through the nerves that carry the signal on to the brain. It actually
makes our eyes less sensitive.
Moreover, it creates a real structural problem: all those nerve axons, as they come
together in a bundle, have to get out of the eye in order to get to your brain. So you have
a blind spot, a hole where the bundle of nerves goes out through the retina. Your brain
tricks you, covering over the blind spot by stitching together a virtual image, like
wallpapering over a flaw in your visual field with a bit of left-over information.
If you were a design student and you turned in the blueprint of the eye as your final
semester project, your instructor would probably say, What are you thinking? But the
glaring design error of putting the retina in backwards is only one of the problems we
have.
Our eyes could be so much better, more acute. For example, if we were sensitive to a
greater range of light, we might see in the ultraviolet or infrared portion of the spectrum.
Reindeer can see ultraviolet light, and important things like edible lichens, predator urine
and wolf fur stand out for them in the same conditions that might make us snowblind. A
bit more vision in the infrared part of the spectrum, and weve have night vision, the
ability to see the light put off by heated objects, including living things. But we make do
so well that we dont even notice the flaws.
Another flaw in the human body is the structure of the knee. No decent designer would
put in a flimsy hinge joint on this weight-bearing limb, especially given that humans are
bipedal, so weve got no spares. If there were a designer he (or she) would take one look
at the specs for the job, the stress that the knee would have to endure, and immediately
throw away the existing plan to bind together the joint with ligament. Back to the

drawing board.
Every time you see a sporting event where some poor athlete goes down in a crumbled
pile from a torn ligament at the knee, especially when they just tear it running or turning
or stopping quickly, you should realize that no decent designer would try to hold together
a knee with ligaments. The reason we have it though is because its descended from a
hinge joint in a fishs fin, where the flimsy construction would have been more than
adequate.
The point is that you dont get whats best from evolution; you get what development and
evolution can provide. Its good enough, or you wouldnt be here, but its not the best
possible structure.
Its often what engineers sometimes call a kluge, a kind of improvised, overlycomplicated, ad-libbed solution. The design is sub-optimal, or less-than-perfect, but
the key to understanding is to realize that its not designed at all. That less-than-perfectness shows us how evolution produces structures by re-purposing or exapting earlier
forms into new uses. Don Wise, a retired professor of geology who often argues with
creationists says that, when the creationists want to talk about intelligent design, he
brings up these sorts of examples of incompetent design.

Getting evolution wrong

Another example of suboptimal structure is the whole genital area, especially in men (Ill
come back to women in the sex chapter, so dont worry Im getting to you).
Theres an old joke that no right-minded designer would ever put your waste disposal
system right smack in the middle of one of your chief entertainment districts. The fact
that you have sex and reproduce with some of the very same equipment that you use to
get rid of waste is clearly not planned in advance. But thats not the only problem.
The way that testes get to where they need to be is also an exercise is jerry-rigging and
ill-thought-out, or simply not-thought-out design. The testes start out developing in utero
up in the abdominal cavity, but they cant stay there; theyre homologies to a womans
ovaries, so both organs start in the same place. If the male testes were to stay inside the
abdomen, however, theyd be too hot. The sperm would wind up defective from getting
lightly cooked by a mans body heat.
So the testes have to descend to their eventual location in the scrotum. But to do so, they
have to go through the lower abdominal wall. In the process, they leave two structural
flaws in their wake, little weak points that can lead to an inguinal hernia. This kind of
hernia is a painful and potentially life-threatening condition in which one of the flaws
tears, and you can push some of your intestines out through the hole. Trust me, if you
havent had one (or two), the flaw is no laughing matter.
In addition, however, if this werent enough of a problem the whole wastereproduction nexus, the descent of the testes in men, the urethra runs right through the

prostate. The urethra is the flexible tube that carries liquid waste out of us, and the
prostate is a donut-shaped gland that tends to swell up over time. Hows that for design?
Run a waste disposal pipe through an organ likely to swell over time and choke it off.
We could cite other examples: the fact that primates bodies cant make vitamin C even
though other mammals can, which means were susceptible to scurvy; too many teeth;
badly designed spine; an extra cavity between the fallopian tubes and uterus where an egg
can get stuck, leading to an ectopic pregnancy, we can go on...
But my last example is perhaps the most dangerous: human adults talk and breathe
through the pharynx, the same pipe through which we eat and drink. Other mammals
have this problem, too, but in humans its much worse. As the larynx or vocal cords drop
down the throat as you mature, the danger of getting food or liquid in your windpipe
grows greater; this is why infants can nurse and breathe through their noses at the same
time. The infant larynx is still high up in the throat.
But this design of the adult throat, with only the little epiglottis at the top of the larynx to
avert constant disaster, means that we can die from simply eating and drinking while we
talk. Evolution has come up with this kluge in part because we gain other advantages
the much greater range and control of sounds that we can produce as we grow into adults.
Its bad design, but were stuck with it because it produces other advantages, like a more
versatile throat for making sounds, even deeper voices.
The reason I bring these flaws up is because I disagree with the way some people talk
about the wonders of evolution. Sometimes, people even me will talk about how
amazing evolution is because it produces wonderful design. They will point to
extraordinary traits of humans or other animals birds ability to navigate on longdistance seasonal migrations, the extravagances of mating, zombie ants with mind control
fungus (no kidding), or the ingenuousness of adaptation and they will say something
like, Wow, evolution is a marvelous designer.
I think that this way of understanding or describing evolution is dangerously close to
believing that there really is a designer out there, some guiding plan or divine intelligence
that is shepherding along all these creatures, shaping their structures according to a
master plan.
In fact, living creatures are filled with bad design, in spite of their successful adaptation.
The sub-optimal design, the less-than-perfect structures, show that the process is not
being guided. No, evolution is tinkering with whats available. Here, evolution re-uses a
hinge joint where another design would work better; there, its stuck with a bit of muscle
thats no longer useful; in another place, it uses a development process that leaves behind
potentially fatal flaws.
Evolution doesnt give organisms the best design; it gives them tinkering and minor
adaptations, sometimes structures or traits that defy good design principles, precisely
because there is no designer.
This is important. Sometimes people think evolution has a plan, or that it makes
progress. Its what we call the teleological error, the idea that evolution has a telos

or goal. Sometimes people think this goal is increased perfection, or even that the telos is
us, humans. You hear it being implied when people talk about one species being more
evolved than another, which is simply impossible unless its sent back from the future
where its had more time to evolve. This kind of thinking can lead to people saying that
humans are the most evolved species, as if evolution where all some grand process
leading up to us.
Thats an anthropocentric view of the reality. Its totally natural, of course, just like
assuming were the centre of the universe, but its not scientific. We may be the most
intelligent or technologically advanced, but we are not the most evolved. That
expression doesnt even make sense. Every living thing is evolving all the time,
sometimes in ways that make it remain unchanged over time, and sometimes with rapid
change.
In fact, I would argue that thinking humans are more evolved is actually a hold over of
pre-evolutionary ways of thinking, especially of the idea of the Great Chain of Being.
The Great Chain of Being was a Christian concept other religions often have similar
ideas that humanity was the most perfect creation, just below spiritual or divine
creatures like angels. For Christians, humans were Gods most perfect material creation.
This way of thinking about our place in the universe has sometimes crept over into
popular understandings of evolution, but its not supported by the science. If we look at
evolution, certainly, we stand out as unusual forms of life, but so do lots of other species.
Which is best or most evolved depends entirely on the yardstick you use to measure.
What we do know is that every living thing today has beating out tons of competitors for
its niche; in that sense, its the most evolved for that niche. For now.
But theres no guarantee that, in the future, we or another species wouldnt follow the
countless other species that have gone extinct before us, the 98% or more of all species
that have gone extinct. We might get replaced by a new, tinkered-with version of
ourselves or simply turn out to be the last in a 7 or 8 million year branch on the family
tree of primates.
Studying human evolution requires a bit of humility: humility to recognize that were far
from perfect, humility to acknowledge that we are not rulers of all life simply because
weve got some unusual quirks, and the humility to realize that our continued place in the
web of life is far from permanent. There was a planet full of life before us; theres no
guarantee that they need us to stick around.

References and Resources

Key concepts from the chapter:
The great mystery of human evolution is how the same forces that produced millions of
other species, at the same, produced human distinctiveness.
Evolution has left its traces on modern human bodies and all living things both in

the way that they are well adapted for their current niches, but also in the vestigial traces
of past adaptations and in the counter-intuitive and overly complex adaptive traits.
Homologies demonstrate the way that underlying structures can be developed in
distinctive ways over evolutionary time.
Often, evolution is misunderstood to involve progress or striving for a goal; believing
that evolution has a goal is a teleological error.

Resources:
African Fossils, a project of Louise Leakey and the National Museum of Kenya. Virtual
lab with many virtual specimens. (link)
Human Evolution by the Smithsonian National Museum for Natural History. (link)
Evolution, Nova, PBS (USA). (link)
Jane Goodall, What Separates Us from Chimpanzees?, TED: Ideas Worth Spreading.
(link)

References:
Bergman, R. A., Afifi, A. K., and Miyauchi, R. Persistence of the Tail. In the Illustrated
Encyclopedia of Human Anatomic Variation: Opus V: Skeletal Systems. Anatomy
Atlases.org.
Bar-Maor, J. A., Kesner, K. M., and Kaftori, J. K. 1980. Human tails. Journal of Bone
and Joint Surgery Series B. 62-B: 508-510.
Gould, S. J. 1980. The Pandas Thumb: More Reflections in Natural History. New York:
Norton
Gould, S. J., and Vrba, E. S.. 1982. Exaptationa missing term in the science of form.
Paleobiology 8(1): 4-15.
Hall, B. K. 1984. Developmental mechanisms underlying the formation of atavisms.
Biological Reviews 59: 89-124. doi:10.1111/j.1469-185X.1984.tb00402.x
_____. 2003. Descent with modification: the unity underlying homology and homoplasy
as seen through an analysis of development and evolution. Biological Reviews 78: 409433. doi:10.1017/S1464793102006097 (download PDF by clicking here)
Marks, J. 2003. What It Means to Be 98% Chimpanzee: Apes, People, and Their Genes,
With a New Preface. Berkeley: University of California Press.
Pennisi, E. 2012. The Burdens of Being a Biped. Science 336 (6084): 974.

doi:10.1126/science.336.6084.974
Shubin, N. 2008. Your Inner Fish: A Journey into the 3.5-Billion-Year History of the
Human Body. Pantheon.
_____. 2008. Fish out of water: human ailments as varied as hernias, hiccups, and
choking are a legacy of our fishy ancestry. Natural History 117(1) (Feb 2008): 26(6).
(Available here)
Stringer, C. 2012. The Origin of Our Species. Penguin.
Stringer, C., and Andrews, P. 2012. The Complete World of Human Evolution (Second
Edition). Thames & Hudson.

Links
Evolution, a series by PBS (USA) (link)
The Multi-Dimensional Human Embryo. (link)
Douglas Theobald. 29+ Evidences for Macroevolution. Part 2: Past History. (link)
Greg Downey. Human (amphibious model): living in and on the water.
Neuroanthropology. PLOS Blogs. (link)
_____. Human, quadruped: Uner Tan Syndrome, part 1. Neuroanthropology. PLOS
Blogs. (link)
_____. 2 legs good, 4 legs better: Uner Tan Syndrome, part 2. Neuroanthropology. PLOS
Blogs. (link)
_____. Sympathy for Creationists. Neuroanthropology.net. (link)
Holly Dunsworth, Evolution is the only natural explanation. And it's all we need. The
Mermaids Tail. (link)
Marcelo Gleiser, Why Do So Many Have Trouble Believing In Evolution?, NPR News
(USA). (link)
P. Z. Myers, (2008) Hox genes in development: The Hox code. Nature Education 1(1),
http://scienceblogs.com/pharyngula/2007/09/the_hox_code.php (link, more links here)
Sarda Sahney, Why does my baby have a tail?, Fish Feet. (link)
Neil Shubin, lecture on the discovery of Tiktaalik, a species linking aquatic "lobe-finned"
fishes with early terrestrial tetrapods (YouTube). (link)
Kate Wong, The Most Fascinating Human Evolution Discoveries of 2012, Observation,
Scientific American. (link)

Carl Zimmer, Turning Fins Into Hands, The Loom (weblog), Discover. (link)

Weblogs
There are too many good weblogs out there by anthropologists on human evolution, but
here are a few of the ones that I consistently find great reading.
John Hawks Weblog. (link)
Greg Ladens Blog, ScienceBlogs (topic: Human Evolution). (link)
Dienekes Pontikoss Anthropology Blog. (link)
Context and Variation, Scientific American, by Kate Clancy. (link)
Powered by Osteons, Kristina Killgrove. (link)
Patrick F. Clarkin, Ph.D. (link)
Ancient Bodies, Ancient Lives, Rosemary Joyce. (link)
The Primate Diaries, Scientific American, by Eric Michael Johnson. (link)
The Mermaids Tale, group blog. (link)
Pop Anth: Hot buttered humanity! A collection of easy-to-read, fast-moving articles by
anthropologists. (link)
Anthropology in Practice, Scientific American, by Krystal DCosta. (link)
Carnival of Evolution is an itinerant, moving feast of links to new blog posts discussing
new research in evolution. You can find links to the most recent edition, and prior ones,
here. (link)

~~~~~~~~

Darwin
Charles Darwins theory of natural selection was one of the most revolutionary ideas ever
put forward, rocking our understanding of ourselves and our place in the world. Its been
150 years since the publication of On the Origin of Species. Although Darwin could not

have anticipated all the discoveries that followed his work research from fields as
diverse as genetics and anthropology natural selection is still the cornerstone of our
understanding of evolution.

Charles Darwin sets sail

Charles Darwin, arguably, wrote the second most important book in shaping the Western
world, second only to the Bible. Published in 1859, the full title of the book was, On the
Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races
in the Struggle for Life. Today, we often refer to it simply as, On the Origin of Species or
just Origin.
By the time he published it, Darwin was already a famous scientist and naturalist. He had
been stewing on the ideas in On the Origin of Species for over two decades, afraid to
bring them out for a whole host of reasons, including the potential repercussions of his
ideas, repeated illness and family tragedy, and an intense desire to get it right, even if that
required patience. His patience, what we might even see as timidness, seems really quaint
today, when researchers often have PR people writing press releases heralding almost
every lab result as a major discovery.
Darwin was not the first to describe evolutionary processes, to argue that species were
not fixed, but that they changed and adapted. But Darwins account, especially On the
Origin of Species, seized the popular and scientific imagination in a way no previous
work had. Origin set biology on a whole new footing, offering the first comprehensive
account of the process we now see at the heart of evolution: natural selection. Darwin,
however, was a reluctant revolutionary: he once told a close colleague that sharing his
theories of natural selection felt like confessing a murder.
Darwins start wasnt terribly auspicious either: he was the fifth of six children, son of a
wealthy society doctor in England and grandson of a famous naturalist, abolitionist and
physician. He was sent off to Edinburgh Medical School to go into the family business,
but he was bored by medicine, freaked out a bit by surgery, and wound up studying
marine invertebrates instead: he discovered that the black spots on oysters were the eggs
of a skate leach. He was much more interested in pheasant hunting season than his
studies.
His father wasnt happy that he was neglecting his schooling, and so shipped young
Charles off to Christs College, Cambridge, to become an Anglican pastor. Darwin didnt
do any better there, instead developing a passion for collecting beetles, a fad at the time.
It may sound like an odd combination, but theology and naturalism were closely related
in English thought at the time, as many naturalists looked to the natural world for signs of
Gods design.
When Darwin returned from summer break in 1831, about to grab his guns and head out
hunting, he found that one of his professors had proposed him as the ships naturalist for
the HMS Beagle. The Beagle was set to embark on a journey around the world to map

coastlines, especially South America, and Captain FitzRoy thought they needed a
geologist and naturalist on board. Darwin was to be as much a companion to the captain
as a scientist and collector, as the captain couldnt be expected to socialize with the crew
during the long voyage (and the previous captain of the Beagle had gotten spectacularly
depressed and shot himself). Although his father was grumpy about it, and thought the
voyage a waste of time, a favourite uncle talked Darwins father into funding his sons
journey because Darwin wasnt even going to get paid for his work.
The Beagle departed in December on a five-year voyage. The ship mapped and charted
coastlines. Darwin spent a lot of time on shore collecting fossils, biological samples, and
geological observations as they went. When he recovered from seasickness, Darwin
proved an excellent naturalist, observing coastlines in Africa, tropical forests in Brazil,
and cliffs in Argentina. He collected marine life, lived through an earthquake in Chile,
helped return kidnapped Patagonians from a previous voyage, saw the diverse wildlife of
Easter Island and Australia and rode on horseback high into the Andes, hunting and
exploring. Darwin didnt always realize the importance of what he was seeing he ate
the turtles that would later prove important to his theories about evolution but he drew
and wrote and collected as he went, sending back samples from his voyage.
Darwins discoveries caused excitement among naturalists back home, as his samples and
journals were shipped back ahead of the Beagle while it slowly circled the oceans for five
years. By the time Darwin returned to England in 1836, his mentor had circulated his
letters, and was eager to help Darwin organize to have all the samples analyzed. Some of
the leading anatomists examined his finds: there were fossils of extinct animals from
South America and a host of new species. Darwins geological observations led him to
argue that South America was slowly rising, and to put forward a controversial theory
about how coral atolls, these fantastic circular islands, formed. Darwins travel memoir of
his voyage proved very popular and made him famous. He was a bit of a sensation and
became a gentleman scientist, elected into the leading scientific societies of England, but
mostly as a geologist.
But to truly understand the importance of Darwin, we have to look beyond his collecting.
What made Darwins impact so great was his imagination and grasp of the big picture,
including the way that he read across geology, biology and even economics. He saw a
pattern beyond the individual discoveries and samples. Darwins greatest discoveries, in
this sense, werent on the Beagle; they were in the years afterwards as he wrote and read
and gradually built up a theory that would embrace all of life, not just individual species.
As he voyaged, for example, Darwin read the first volume of English geologist, Charles
Lyells Principles of Geology, especially Lyells theories of uniformitarianism. Lyell
argued that natural forces were uniform across time. That meant that geological
formations like valleys had to be explained by forces that could still be observed,
working slowly over immense time rather than by one-off divine catastrophies. If
geologists found fossilized seashells high in the Alps, as they had, either oceans were
once much deeper, or the Alps had been pushed up, a little at a time, over countless years.
Darwin was thrilled by Lyells work as he now saw that the landscape told a story, telling
how it had been formed.

When Darwin returned, he talked with the leading biologists of the day, too. They were
already discussing the growing evidence that species could change or become extinct,
once thought impossible if God had created each one individually. Darwin brought more
evidence of species change, adaptation, and extinction.

The origin of On the Origin

To really understand how Darwin came up with his theory of evolution, you have to
realize that he really wrote it after he came home. By 1838, he read the work of Thomas
Malthus, An Essay on the Principle of Population. Malthus pointed out a grim fact of
economic mathematics: populations could easily double each generation, whereas food
production could not keep pace.
Darwin realized that species dont simply struggle against each other; the constant
potential for excess reproduction, the enormous fertility of life, also drove species
relentlessly, refining their adaptation. Darwin saw a relationship between the way farmers
selected seed from the best produce to replant, the way that pigeon breeders chose which
of their birds to cross-breed, and the natural process through which over-production could
produce better adaptation. For this reason, Darwin eventually referred to the process as
natural selection because he thought nature, too, was selecting among that excessive
fertility, like the pigeon breeders and farmers.
By the end of the 1830s and first years of the 1840s, Darwin was already thinking about a
theory of natural selection, but he continued to focus on his geological projects: reports
about the Beagles collections, a book about coral reefs, work barnacles. He wrote a 230page essay about natural selection, but he kept it secret, to be published only if he died
before completing the work he wanted to write: he knew that reception was likely to be
hard. His wife, Emma, a devout Christian, worried about the fate of his soul. But there
were still problems that troubled him intellectually; he knew so much about diverse
species that he kept analyzing more and more cases, reading widely and talking with
other scientists, farmers, fishermen and animal breeders.
Darwin also struggled constantly with poor health, and was devastated by the death of his
daughter, Annie. For twenty years he stewed on his ideas about natural selection, sharing
them selectively with colleagues in his learned societies and scientific circles, building up
a more and more massive case as he worked out his theories in notebooks dedicated to
what he originally called Transmutation.
Finally, in 1858, he was rocked by a paper shown to him by a young colleague, field
researcher Alfred Russel Wallace. Wallace and Darwin had been corresponding, but
Darwin was contending with an outbreak of scarlet fever that would eventually kill his
infant son. Wallace sent Darwin a paper outlining his own theory of natural selection,
threatening to scoop Darwin entirely. Fortunately for Darwin, two senior colleagues
decided that the two should present together to the Linnaean Society. Even when the day
came, Darwin was still too distraught to attend when the day came.

The 1858 presentation laid out the principles of evolution clearly, including parts written
by both Wallace and Darwin; the audience included many of the leading biologists of the
day. But the paper caused scarcely a ripple. In his annual report, the President of the
Linnaean Society of London wrote:
The year which has passed has not, indeed, been marked by any of those striking
discoveries which at once revolutionise, so to speak, the department of science on which
they bear.
But then, in 1859, Darwin published On the Origin of Species by Means of Natural
Selection, or the Preservation of Favoured Races in the Struggle for Life. He originally
wanted to title it An Abstract... because it was a cut down version of the much bigger
work he wanted to write. Luckily, the editor prevailed on Darwin to give it the shorter
title. Origin was an unexpected best-seller, the first printing selling out immediately. The
book went through six revised editions in Darwins lifetime.
What made the difference between 1858 and 1859?
First, Darwin was a great writer, and his work still is rewarding to read. He saw in
barnacles and beetles, seashells and orchids, peacocks and earthworms, a kind of
grandeur. On the Origin of Species has a staggering breadth, and drew on examples that
the broader population would know well, from the English countryside, farming and
popular hobbies like pigeon breeding. Instead of a catalogue of life, description, now we
have an explanation. Biology becomes an explanatory science instead of just a
descriptive one, much like Lyell had showed Darwin in geology. But it took a scholar
like Darwin, with his encyclopedic knowledge of life, to write a book with the kind of
sweep that the Origin has.
The irony, however, is that in spite of his popularity, Darwin was still uncomfortable.
Even though he we consider him the father of evolutionary theory, he didnt even like the
word evolution: he preferred transmutation, and kept writing descent with
modification in On the Origin of Species. Darwin thought the word evolution had too
much of a sense of direction or progress.
Although his supporters fought fiercely on his behalf, the center-piece of Origin, the
theory of natural selection, was still not widely accepted in Darwins lifetime, and really
wouldnt be for decades to come.
But his work was amazingly far-sighted. He foresaw some of the things we are
discovering today, like the deep connections among all forms of life that we see clearly in
genetic research. He pointed out that one reason that it is hard to see evolution is that
descent with modification is not a tree of life, at all; its more like a coral. The living
species are like the surface of the coral, still alive, atop the skeletons of dead
predecessors. We cant see the connections among the living because, like the hard
structure under the coral, the connections are dead.

Figure 9. Charles Darwin's 1837 sketch, his first diagram of an evolutionary coral
from his notebook
But ultimately, one of the reasons for the success of On the Origin of Species is because
Darwin himself was so fascinated with the range of life, the astonishing way that species
can adapt and change. Like the natural theologians, who saw the fingerprints of God in
the majesty of nature, Darwin still had the capacity to feel awe and to share that with his
readers.
At the end of On the Origin of Species, he shares this vision:
It is interesting to contemplate a tangled 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 upon each other in so complex a manner,

have all been produced by laws acting around us. . .

There is grandeur in this view of life, with its several powers, having been originally
breathed by the Creator into a few forms or into one; and that, whilst this planet has gone
circling on according to the fixed law of gravity, from so simple a beginning endless
forms most beautiful and most wonderful have been, and are being evolved.

Natural selection
How does evolution work? Lets take an example that we know is going on because its
happening right in front of us (and causing doctors all sorts of headaches).
In our hospitals right now, we are breeding super-bugs, strains of bacteria that are so
resistant to antibiotics that they produce hard-to-cure, even potentially, life-threatening,
infections. For example, there are now forms of staphylococcus, a bacteria that lives
normally on your skin and in your sinuses, that are not susceptible to penicillin; theres
even some varieties of staph that are resistant to multiple different antibiotics.
These varieties of the antibiotic evolved because, when they were doused with penicillin
or other antibiotics, those bacteria without resistance died. Left behind were the lucky
those who just missed contact or those bacteria with a bit of resistance. Over time, the
use of antibiotics made the population of bacteria more and more resistant by slowly
wiping out any bacteria that had vulnerability to the drug.
Staph evolved antibiotic resistance, not because it wanted to or because we wanted it to,
but because penicillin and other antibiotics slowly changed the population of bacteria,
making it more and more likely that vulnerable bacteria wouldnt make it to reproduce
and pass on their traits. Without the drugs, this trait wouldnt have been that important.
With the invention of penicillin, suddenly, whether a bacteria could survive being doused
with it became crucial to its survival, and to the evolution of the species.
Natural selection is the key to understanding evolution. The principle is deceptively
simple. What makes natural selection most difficult to understand, I would argue, is that
we tend to assume that the theory is saying things that it is not. One way to understand it
better is to see how you can get it wrong.
One of the most widely-accepted explanations for species change before Darwin, held
even by many of Darwins followers, for example, was the theory of French naturalist
Jean-Baptiste Lamarck. Lamarck believed in the inheritance of acquired traits. That is, if
an animal or plant acquired a trait through use, or if a trait atrophied because of neglect,
the change could be passed on to offspring.
An example of Lamarcks theory goes like this: If your father did hard labor and got
strong from it, you would be born with bigger muscles; he passed on his acquired
strength. The blacksmiths son would allegedly be born different than the shopkeepers
because of the parents occupation. In the classic example used by Lamarck, giraffes had
long necks because the ancestors of modern giraffes had stretched and stretched up to eat

the high leaves on tree branches. A trait became exaggerated through use, even through
aspiration or desire like the giraffes reaching and reaching.
On the other hand, disuse led to atrophy and the potential loss of a trait. Lamarck and
other naturalists knew that people grew weaker from inactivity; they just thought that this
could be passed on to an animals offspring.
In contrast, natural selection is deceptively simple. The process is, in four words: Nature
proposes. Selection disposes.
Darwin realized, from reading the works of Thomas Malthus, that population growth was
potentially geometric. Many more individuals were born than could survive and
reproduce; two human parents could easily produce four, five, six, even more children. In
other species, the potential growth rate was even more staggering. This constant increase
in population inevitably butted up against limits in habitat, food, and other resources.
This meant that there was always competition within a species, just as there would
always be predators, diseases, climactic change, and other forces of nature. Resources
were limited; life was potentially abundant.
In addition, living things vary. You do not look exactly like your siblings; more
importantly, you do not have the same abilities, traits or potential. You might even have
different resistance to disease. Members of the same litter of kittens or the same clutch of
eggs are not identical clones of each other. If every member of a species cannot survive
to reproduce, or reproduce as successfully, and they varied constantly, some variations
might be more successful than others.
By whatever mechanism Darwin didnt know anything about genes or mutation
natural processes produce abundant varied life; the growing population together with a
whole range of challenges finding food, staying safe, surviving the elements, finding a
mate sorts that variation out. If the variation can be inherited, and thats key to why
Lamarck was wrong about a lot of things, and the selective pressure continues over time,
slowly, the variants with better chance of survival and reproduction will grow more and
more common. A species will change and adapt by shifting toward one of its extremes, if
that extreme is more likely to survive generation after generation, like the staphylococcus
becoming resistant through generations of exposure to deadly penicillin.
If theres no variation in a species if theyre all identical clones then theres no
natural selection. Natural selection requires variation. Also, if theres no selective
pressure, theres no evolution. Thats sort of harder to imagine, but it comes up in the
sense that some situations are more or less intensely selective: super-high fertility and
really high mortality rates can set a species up for rapid evolutionary change. In times of
great hardship and mortality, selective pressure increases.
The expression survival of the fittest is sometimes used to describe natural selection,
and thats a good enough way to think about the process as long as you understand what
is meant by fitness. Fitness doesnt necessarily mean the strongest or biggest or
meanest or even healthiest survive. The idea of fitness is quite subtle; individuals
often dont compete directly with each other (although they sometimes do).

Which individual is most fit depends entirely on the selective pressures. Whatever
forces are most likely to prevent an individual from successfully producing offspring,
these are the selective forces. They can be predators, difficult to digest foods, competition
from another species, changes to the environment, diseases, competition to mate,
surviving lean periods and lasting through to mating season... Whatever force threatens
you, your ability to reproduce, or your offspring, that determines which variant is most
fit and which trait is under pressure to produce adaptation. In staph, its been resistance
to antibiotics, which didnt matter before the discovery of these drugs.
Another example: if a species is subjected to periodic famine, if the factor most likely to
stop an animal from reproducing is dying of hunger during a dry season, this pressure
would favour those individuals who are adapted to surviving near starvation. They are the
fittest. The pressure could favour the ones with the slowest metabolism, those able to
pack on reserves of fat, or maybe even a variant than just goes to sleep for long periods in
a safe place to avoid starving.
Different species sometimes compete head-to-head for the same resources. We call this
inter-species competition, like when two species of predators hunt the same food source.
Inter-species competition tends to lead to extinction of one specie due to competitive
exclusion; one species is inevitably going to have an advantage, and unless the less
competitive species can find a niche where it turns the table and hang on, its liable to
disappear.

Natural selection (part 2)

Most evolutionary change, however, is driven more by intra-specific competition. The
dynamic is not so much bats vs. flies, for example, but more among bats with all their
variation as to which bat will be most successful in getting food, reproducing, and
supporting more offspring. If flies get faster or evolve to better avoid bats, they wont
replace the bats in the ecological system; faster flies will put more pressure on bats,
making it harder for them to get food, perhaps driving bats to evolutionary change.
If you have competing directional selective pressures, you can often get long periods of
what look like no evolution, no change. Species can be very stable if they undergo
stabilizing selection. For example, it might make sense for a primate to get larger to
compete to reproduce, but at a certain point, the increased size might lead them to
struggle to get enough food, find tree branches strong enough to support their weight
when hiding from tigers, or even keep cool, because big bodies have more of a problem
with over-heating. Body size might hit a kind of adaptive sweet spot, remaining
consistent over long periods. The trade-offs involved may make a compromise between
the two directional tendencies optimal, and pin down a trait for long periods of time.
Natural selection can also lead to species change through directional selection.
Directional change occurs when an animal can gain advantage by a trait becoming more
exaggerated. If short-necked giraffes cant compete as well for food, they tend to die out,
leaving only longer-necked giraffes behind. In some cases, if selection keep eliminating

the shortest, the remainder get taller and taller until they hit some biological constraint or
a stabilizing point.
If you have competing selective pressures on a species, you can also produce diversifying
selection. The population may split, with those at each extreme better able to survive and
reproduce than more average members of the species. For example, for mice or rabbits
living in the arctic, being very lightly coloured to blend in with snow and ice makes some
animals more fit. They are camouflaged and less likely to be spotted by a predator.
The same colour pattern, however, is a real disadvantage if the animals lives in a
temperate climate, such as grasslands or a forest, where white fur will make the animal
stand out. In these areas, dark brown mice may have an advantage. In both settings,
intermediate coloured mice might be less well camoflauged, so they disproportionately
wind up getting eaten by owls or hawks. In the end, you can wind up with two separate
populations, one very dark and one light, each adapted to its environment. Given enough
time, if you keep them apart, and youll find that they turn into two separate species, what
we call speciation.
Diversifying selection can actually happen in multiple directions, into multiple niches, in
a pattern called adaptive radiation. A basic rule of thumb is that, if you see a pattern of
multiple species arising from a single ancestor type diversification you know that
the new variants are occupying different niches, maybe even dividing up the resources,
finding new ways to live, and specializing.
For example, all mammals alive today have descended from a common shrew-like
ancestor that lived in the shadows of the dinosaurs for almost 150 million years. With the
extinction of the dinosaurs about 65 million years ago, a whole lot of niches opened up
for mammals to fill, and our ancestors diversified into thousands of forms. As the number
of mammals increased, there even developed a niche for mammals that ate other
mammals. Over the next tens of millions of years, directional selection for a whole range
of different environments mammals that dug under grounds, mammals that lived in the
ocean, mammals that hunted at night, mammals that ate grasses produced an explosion
of adaptive radiation. Thousands of species.
To sum this up, natural selection is kind of like an equation:
(Variation + Inheritance + Selection) over Time = Adaptation
We can abbreviate this as VISTA to help us to remember. If any of the first four is not
present, you cant have natural selection. For example, if theres variation, but its not
inheritable, then it cant really be the target of natural selection. It can change, but its not
from natural selection. Many people argue that this is why culture and technology dont
really undergo natural selection, although they change and develop over time. You dont
inherit technology in the same was as a genetic trait: you dont inherit your parents
education, for example, or their inability to understand how their new phone works (so
thats a good thing!).
If theres not variation sufficient to create some kind of selective advantage or difference
in fitness, theres no natural selection. If a trait in an animal is uniform, selective pressure

on it cant make some more likely to survive than others. As Charles Darwin wrote in
1868, in his book, The Variation of Animals and Plants Under Domestication, it doesnt
take much: slight individual differences, however, suffice for the work, and are probably
the chief or sole means in the production of new species.
No diversity; no evolution. This is one reason that the idea of a genetically uniform
master race doesnt make evolutionary sense. Our evolutionary insurance policy is our
diversity.
Without sufficient time, natural selection cant bring about change. Or if conditions
change too frequently, then natural selection cant apply directional pressure long enough
to produce adaptation.
How fast can evolution happen? Well, that depends, especially on the severity of the
selective pressure and the rarity of the trait that confers an advantage. Some theorists
thought evolution was probably pretty slow, but we now have examples of changes
happening relatively quickly. One of my favourite examples is the case of the size of
snakes in Queensland since 1935.
In 1935, the cane toad Bufo marinus was introduced from Hawaii to Queensland by
sugarcane farmers who hoped that the toad would eat a type of native beetle that fed on
their crops. The experiment was one of the great disasters of human biological
engineering. Not only did the cane toad have no effect on the beetles, but the toxic frogs
started to cause a decrease in the native fauna, like quolls and even goannas and other
reptiles. They were incredibly fertile; a female B. marinus can lay up to 25,000 eggs at a
time.
The cane toad is a huge threat to biodiversity as most native animals like reptiles think
that it looks like food, but its loaded with venom, bufotoxin. One of the casualties has
been the local snakes. Eat a cane toad, and a snake is liable to die. There are so many of
the toads that virtually every snake will come across one and try to eat it. The only snakes
that were safe were those too small to eat the hulking toads: a form of intense directional
selection. Being large was virtually a death sentence as the snake found a cane toad lunch
irresistible. In less than eight decades, the introduction of cane toads changed the size of
snakes by introducing a new selective pressure.
The new variant of the snake was not an unprecedented mutation; it was just the runts of
the litter, the extreme of the species normal range. As large snakes died before they
could reproduce, their trait started to disappear from the population. Directional selection
concentrated and exaggerated the small size.
Some evolutionary theorists call this kind of pattern punctuated equilibrium. A species
might exist at a kind of equilibrium under stabilizing selection, with natural selection
favouring consistency generation after generation. A new selective pressure caused a kind
of punctuation, a rapid change to a new equilibrium, like a crash in the average size of
snakes that stabilizes once they are too small to eat the poison cane toad. The pattern
explains what we often see in the fossil record: long periods of consistency in some
species, interrupted occasionally by fits and starts of change.

Natural selection was happening all along; its just easier to see its effects when the
situation changes, and the selective pressures shift. Upheaval is more likely to produce
change, whether that upheaval is imposed on a species by external conditions, or whether
it comes about because of some change in the species itself.

Genetics
Charles Darwins theories were only partially accepted in his lifetime, even by some of
his fiercest supporters. Darwin convinced fellow naturalists that species changed, he even
convinced a lot of the public. But Darwins contemporaries, even his supporters, didnt
necessarily accept his argument that natural selection was the primary force. They
couldnt see how slight variations in a population, causing slightly different chances of
survival could, over time, add up to major change. The most innovative, and arguably
most important part of his theory was generally disregarded.
One reason is that Darwin couldnt explain how inheritance and variation worked no
one could at the time. Darwin, like many of his contemporaries, thought that offspring
had a mixture of the traits of their parents.
The problem is, if inheritance is just the mixing of the traits of the parents, then over
time, we would expect overall variation to decrease constantly in sexually reproducing
species. Any unusual plant or animal couldnt pass on its unusual trait fully to its
offspring because that trait would be constantly diluted by mixing with another, more
normal parents traits. Worse still, if all offspring are a mixture, then all offspring would
be alike. So variation decreases by half in each generation because two different parents
produce a group of intermediate and identical offspring.
Imagine that all the variation is different colours of paint: if you start mixing the paints
and you keep mixing randomly, time after time, generation after generation, you
eventually wind up with a collection of muddy brown mixed paints. The idea of the
mixing of traits seemed to predict the same outcome less and less variation. The
problem did not escape Darwin or his critics; one early reviewer of On the Origin of
Species pointed out the problem: how did inheritance produce variation?
The problem was not with evolutionary theory or natural selection, but with the very idea
that inheritance was a blending of traits. We wouldnt understand how inheritance and
variation worked until we learned about genetics.
At the same time that Darwin was working on his many projects, a German researcher
and friar, Gregor Mendel, was studying the reproduction of peas, working on what turned
out to be the solution to the problem of decreasing variation.
Mendel planted thousands of pea plants in an experimental five-acre garden at his
monastery. He wanted to understand how inheritance worked, so he carefully controlled
how these plants fertilized each other, mixing pure varieties to see how the hybrids would
turn out. His project was unprecedented. In less than a decade, he planted, carefully

fertilized and meticulously examined almost thirty thousand pea plants. His results were
surprising, so much so that he wasnt recognized until long after he had passed away.
Mendel discovered that the plants reproduced in a fixed and predictable way: if he
crossed two pure varieties of the pea plant, each with a contrasting trait, all of them
would predictably take on one of the parents traits. Cross pure-bred plants with
contrasting smooth and wrinkled-skin peas, for example, and you always got a smooth
pea off-spring. We call this the dominant trait because, when the two are both present, the
dominant trait gets expressed. In this case, the smooth skinned pea variety is dominant.
However, and this is where it gets interesting, if you take these second generation plants,
all with smooth peas, and cross them, you get something unexpected: three-quarters of
the plants have smooth skin peas and one quarter have wrinkled. So the smooth-skinned
second-generation plants are carrying, unexpressed, the trait that produces wrinkly skin
or they couldnt pass it on to the third generation. Keep crossing these hybrid plants, and
the non-dominant variety keeps popping up. The trait can skip a generation. The ratios in
each generation were crucial, because Mendel could work out that two different variants
together were causing the distribution (see Figure 2.20 for an explanation using red to
represent the dominant trait).
Mendel realized that the plants were getting what we now call a gene from each parent,
but the presence of the dominant allele, or gene version, was masking or hiding the other
allele, which we call recessive. The traits were not really mixing. They were being
passed intact to the offspring, lying in wait if they were ever to mix in an offspring with
the same recessive trait from the other parent. Variation didnt necessarily disappear,
even if it went underground.
This pattern is particulate inheritance, as opposed to mixing: you either get the gene, the
particle that determines this trait, or you dont.
The simplest example of particulate inheritance is sexual reproduction itself; in the vast
majority of cases, a child does not wind up a blending of the father and mothers sex.
Bodies are typically either male or female (with some exceptions well talk about in week
five). You either get a Y chromosome or you get an X; you dont get a mixture of the two
(although, as youll often find in biology, it can get more complicated than this and there
are almost always exceptions).
Mendel read a paper about his research at the time and published it, but it only got cited a
few times over the next three decades. Mendel got promoted to head of his abbey and
gave up his pea plants. What Mendel had discovered, however, were two key findings in
addition to particulate inheritance which we now refer to as the laws of Mendellian
inheritance:
First, every individual possesses two copies of the gene for a trait, but only passes one of
these along during sexual reproduction; a child gets one from each parent. You only pass
on half of your genetic material to your children if you reproduce sexually. If a hybrid
parent passes on a recessive gene, and it gets together with another copy of the same gene
from the other parent, the offspring can have a recessive trait that was being carried,

unexpressed in both parents.

Second, Mendel recognized that traits independently assort, that is, you dont get all
your traits from one parent or the other, or a blend, but different discreet traits from each
parent. Not all smooth skinned pea plants are tall and have big leaves and purple flowers;
each trait is determined independently during sexual reproduction by which particle of
inheritance you get.
Only some traits behave precisely in the way that Mendel described because many traits
are, in fact, polygenic; many genes affect them. For example, your skin colour is affected
by multiple genes. Mendellian traits, that is, those determined by a single gene pair, tend
to be discrete traits: either you get them or you dont, with no real in-between. In
contrast, many traits are continuous and affected by multiple genes for example, skin
colour comes in many, many shades, light and dark and everything in between.

The modern synthesis

The German evolutionary biologist August Weismann rediscovered Mendels work when
he was proposing his germ cell theory of inheritance. Weismann argued that inheritance
only happened from parent to offspring through the germ cells: the fathers sperm and
mothers egg. Nothing that happens to the bodily cells during your lifetime could affect
the germ cells, which produced body cells in your offspring. Transmission was one way;
from parent to child through germ cells, and from germ cells to body cells.
What came to be called the Weismann barrier meant that acquired traits could not be
passed on. The children of a weightlifter were not born extra strong. Lamarck was dead
wrong, according to Weismann.
To prove it, Weismann removed the tails of mice, and then of their offspring, over and
over again for multiple generations and yet every successive mouse was still born with a
tail, demonstrating that acquired traits in this case, being tail-less were not
inheritable. We now know that this is more complicated, but Weismanns ideas were
crucial for shaping research in genetics and for helping to seal the acceptance of natural
selection as the mechanism driving evolution.
By the early 1940s, biologists across a wide range of fields, from genetics to evolutionary
theory, could begin to see their work flowing together in what Julian Huxley called the
modern synthesis: that synthesis was the hybrid of Darwinian natural selection with
Mendel-inspired genetics.
According to the modern synthesis, to understand evolution, you had to focus on the
population as a whole, over many generations, not any one individual, nor any single
lifetime. Evolution was working slowly on a background of genetic variation in a species,
on the whole population, not on any one individual. Individuals lived and died,
successfully reproducing in some cases, passing on the genetic material in their germ
cells. What you did in your life didnt really matter to evolution it wasnt affecting

your germ cells except if you reproduced a lot, a little, or not at all.
Natural selection, over time, was shifting the total genetic profile of the whole
population, the gene pool, making the most fit genetic types gradually more and more
pervasive. If having wrinkly skinned-peas made a pea plant more successful, the gene
pool would slowly get filled up with that variation, squeezing out other, less-successful
varieties.
The discovery of DNA helped to confirm that biologists and evolutionary theorists were
on the right track. DNA or deoxyribonucleic acid is a complex organic compound that
can store information inside of living cells; its found in the nuclei of all living cells, and
even some viruses.
DNA is a very long molecule that looks like a twisting ladder, what biologists call a
double helix. The sides of the ladder are made up of a sugar-phosphate chain, but the
steps are what is crucial: the steps are base pairs of four different nucleotides, compounds
which are used in the DNA of all forms of life. The nucleotides are like four letters that
provide a code or pattern for building amino acids, which can then be assembled into
proteins to do all the sorts of things that cells need to do to live. The DNA molecule itself
is huge, with over three billion of these steps, the base pairs, on the human chain.
Geneticists realized that variation could arise if that code was not copied successfully.
When the DNA molecule unzips so that it can make two copies of itself, a mutation or a
change can be introduced to the sequence that might cause a new variation, even
something entirely unprecedented to arise. A part of the sequence could get accidentally
inserted, left out, copied twice, turned around the wrong direction, or any other sort of
DNA clerical error. Genetic mutation provided an important way to understand how
variation could arise; most were harmful, but some mutations might actually be
beneficial.
For example, we know that humans developed dark skin as we lost our hair in Africa; the
dark pigmentation protected our ancestors from over-exposure to ultraviolet light,
protection that we lost as our hair gave way to make it easier for us to cool our bodies. At
least three times since then, a mutation or genetic change has caused a population of
humans to develop much lighter skin: at least once in Neandertals, once in Europeans and
once in north Asians. A slight change in the genes linked to the production of melanin, or
the dark pigment in our skin, has proved to be an important change for surviving in
northern latitudes, where some of our ancestors got much less light, and would have
struggled to produce enough vitamin D to stay healthy.
That genetic code is crucial; sequencing the code has been one of the greatest scientific
achievements of the last twenty years. But its also produced some real surprises.
First off, in 2003, when the first human genome was sequenced, and researchers were in
for a bit of a surprise. Back in the 1960s, some scientists thought that the human genome
might have as many as 200,000 protein-coding areas. The number turned out to be only
about 21,000 or so sections that we had patterns for proteins about the same number as
the mouse. A lot of the DNA about 98.5% didnt seem to be active at first glance,

and some scientists started referring to it as junk DNA. We now realize that, in fact, this
DNA is quite busy, regulating and affecting how and when the DNA sequences that code
for proteins go about their work.
DNA, by itself, doesnt do anything. In fact, it can sit quite happily for a very long period
of time; thats why we can pull it out of a crime scene or even out of a bone of a longdead ancestor, in some cases. The cell itself has mechanisms for expressing different
genes, or preventing a gene from being expressed. This clearly happens in multicellular
animals where every cell has the same nuclear DNA, but they do very different things:
bone cells and muscle cells and brain cells result from the same DNA being expressed
differently.
Some of the inactive sections of our DNA are pseudogenes: sections of DNA in which a
mutation has knocked out the function. For example, humans have a large number of
areas of genetic material that look like the areas, in some species, that cause the olfactory
system to form in sophisticated ways. These genes appear mutated in us, so that they
dont serve the same purpose that they do in other species.
Some of the variation in the human genome also arises from copy number variation:
multiple copies of the same section of a DNA strand. Even identical twins can differ
genetically because they get different copy numbers of some sections of the same DNA.
We dont really know how all the variation works, although geneticists are making
enormous advances every year in helping us to better understand how inheritance and
variation both work. These discoveries are not over-turning natural selection, by any
means, and the vast majority of evolutionary theorists will still argue that it is the central
mechanism that is, over time, shaping the traits of all living species, including humans.
It may be hard to believe, but some of the most important parts of being human have
nothing to do with our consciousness or intelligence or even our intentions or motives.
Being human means passing on your genetic material or not, being part of a pool of
genetic material, constantly shifting and being tested by natural selection, never identical
from one generation to the next, even if you have absolutely no idea that its going on.

Epigenetic inheritance
In 1944, near the end of World War II, the Nazi army cut off supplies to the occupied
Netherlands to punish the Dutch for hindering German war efforts after D-Day. The
harsh winter of 44 to 45 became known as Hunger Winter as food and fuel ran out,
leading to a brutal famine until spring brought a thaw and resupply by Allied forces.
The mothers who were pregnant during Hunger Winter gave birth to infants that were
smaller than usual, and these infants were also more prone to certain health problems
later in life, like diabetes, cardiovascular disease and obesity. That was interesting, but
not a huge surprise.
What was more surprising is that the children of these children, born decades later, were

also more prone to certain health problems. Animal studies have confirmed that a
grandparents response to starvation can, in some cases, influence how their
grandchildren will develop. Increasingly, biologists are convinced that some acquired
influence can affect an individuals offspring; the question is, what sort of influences,
how important are they, and through what type of biological mechanisms?
The modern synthesis focused almost entirely on genetic inheritance. Some proponents
were absolutely steadfast: only genes could be inherited, nothing else. More importantly,
because only genes were stable over the period of time needed for natural selection, other
forms of inheritance didnt matter even if they did happen, or at least they couldnt
evolve from natural selection.
Some advocates of the modern synthesis went so far as to take a kind of genes view
of evolution. They argued that, because inheritance was genetic, you could model
evolution, not as a struggle for survival among organisms or between species, but among
genes themselves.
Theres some truth in this gene-level perspective, and its a necessary antidote to some
popular misunderstandings of how evolution works. For that reason, I think its
important. In a way, though, its kind of like economics: its a powerful way of looking at
the world, a way of blocking out a lot of confusing distractions and homing in on the
primary dynamic of evolution, which is genetic change of a population due to natural
selection. But the problem is, biology is seldom so simple.
Increasingly, evolutionary theorists are realizing that the modern synthesis needs to be
expanded to include the recognition of other mechanisms of inheritance not just genes.
Our VISTA model of natural selection doesnt say anything about genes: it only says that
inheritance matters. Clearly, genes are the primary vehicle for passing on traits, but
theyre not the only way. And with humans, these other mechanisms are really important.
First, genes are not the only thing you get from your mother; theyre not even the only
thing in a mothers egg. If an egg only had DNA, it would be like... well, more like
sperm.
There are other parts to these generative cells like the mitochondria, or the energyproducing part of the cell, which you only get from your mum. Researchers have realized
that theres DNA in the mitochondria 37 genes worth DNA that can give us
important information about your mother, your mothers mother, and so on. But your
mitochondria reproduce asexually from that egg cell, so you inherit your mitochondria
only from her.
Research in the past few decades on the DNA in the human mitochondria or mtDNA has
shown that we could use it as a molecular clock. Because you only get mtDNA from
your mum, and she from her mum, and so on, we can estimate how old a variant of
mtDNA is by how many mutations have occurred. We can estimate, for example, how
long ago two individuals shared a common ancestral form of mtDNA by comparing their
mtDNA and estimating for how long it would take for all the intervening mutations to
have arise.

Moreover, weve realized that Weismanns germ theory of inheritance wasnt entirely
correct; some of your experience could leave a trace in your germ cells so that some
kinds of change could be inherited, like the Dutch famine cohort passing on a pre-set
stress response. The study of epigenetics, or the way that embryos form from fertilized
eggs, helped us to see that the process was crucially shaped by DNA, of course, but also
by other material in the mothers egg, and by processes that happened in utero or
immediately after birth.
For example, a lot of the calibration of your immune system happens through breastfeeding, as your mothers milk affect how basic immune responses are sensitized to
potential threats. And, as weve seen, starvation of your mother or other ancestor can
prime your DNA to respond in distinctive ways, almost as if epigenetic processes were
giving you a flying start to face the same types of challenges your ancestors faced.
In another example, if you stress out a mother, you can make her children shift their
immune systems. The body uses inflammation to fight off bacterial infections, other
immune system responses to fight off viruses. Were beginning to realize in research
done on other primates with data to suggest the same thing happens in human that
if you stress out a mother, her childs immune system will be tuned more to fighting
bacterial infection, possibly arising from injury. In contrast, a happy, socially-engaged
mum tends to produce a child whose immune system is better tuned to fighting viral
infection, which youre more likely to get if mum is hanging out in a group with viruses
circulating among them.

Its not all in your genes

Inheritance also takes more circuitous routes, however, not just through direct biological
transfer. When you were born, you werent simply tossed out a window with your genes
and let to figure out things for yourselves. Like many species, you were cared for,
defended, fed, and nurtured. You came into an environment that was already shaped by
generation upon generation of humans who had come before you; otherwise, youd have
to be a lot more worried about predators, and thered be a lot less built up around you.
Some evolutionary theorists have argued that one important way to think about nongenetic inheritance is to look at the way that many species, not just humans, alter their
environment, changing it for future generations. They call it niche creation or niche
construction, or the way that selective pressures faced by one generation are changed
because of the actions of the previous generation. Not every species shapes its niche, but
some change their environment a lot, for better or worse, so that you have to think about
how accumulated activity alters the pressures that shape the species itself.
If the change to the environment is stable and persists over enough generations, it
becomes both selective pressure as well as being a form of inheritance. The theory of
niche creation is controversial; it make modeling evolution more complicated. If you take
niche creation into account, you have to acknowledge that the environment isnt just
driving an organisms evolution, but also being affected by that evolution. The influence

can go both ways.

With some species, the impact is small. With others, however, the investment of in
changing the environment is so huge, and the difference this change makes for the
survival of the species so great, that its impossible to ignore. For example, social insects
like ants and termites produce large mounds, a combination of fortress and sophisticated
breeding complex. The mound radically changes the survival rates of offspring and the
traits of those offspring.
Beavers create artificial wetlands by building dams, providing for their own defense,
multiplying the fish available to eat, and basically making an environment a kind of
beaver water park. A young beaver inherits, not just genes, but a world altered to favour
beavers; and their survival depends upon the interaction between their instincts and this
beaver-created environment.
With humans, the niche creation dynamics are even more extensive and unpredictable.
For example, our ancestors were really good at eliminating any animal that preyed on us,
with just a few holdouts, like crocodiles and sharks, who are almost all in danger of
extinction. Our ancestors actions mean that you and I go about our daily lives without
too much worry about attacks by bears or tigers or wolves.
Almost all forms of niche creation are unintentional, and some arent favourable in the
long run for the continued survival of a species. A species can change an environment to
such a degree that the niche actually becomes inhospitable to itself or other forms of life.
For example, many of the diseases that menace us today arise from our own domestic
animals, or are exaggerated by the ways we alter the environment; malaria, for example,
probably one of the selective pressure most affecting some pockets of humanity, is made
worse because we cut down forests for fields, and then create breeding grounds for
mosquitos in the stagnant water of irritation trenches.
Both epigenetic inheritance and niche creation are ways that acquired traits can, in some
circumstances, be passed on. Maybe Lamarck wasnt totally wrong. Your experience can
affect your offspring. But the most important way that this might happen is through
learning and teaching.
Learning and teaching are the clearest and most crucial form of non-genetic inheritance.
Its not just humans that learn: one of the most important reasons to have a brain at all, no
matter how small, is to learn. And all kinds of species learn, even some of their most
basic skills.
Song birds, for example, will start chirping as they develop as chicks, but they wont
develop the adult songs that they need in order to reproduce unless theyre exposed to
adults singing. Their brains and senses are ready to learn, primed to pick up the right
song. Theyre even instinctually experimenting with making sound. But unless they get
the right environmental stimulus, they cant learn what they need to know.
The key difference with humans, then, is not so much learning, as it is teaching. All kinds
of animals learn, but most have to do it for themselves. Although predators will
sometimes give their offspring wounded prey to play with in order to learn, and others

seem to show their young how to find food, humans build an elaborate social structure for
teaching so that this channel can pass on an immense amount of the information that any
human needs to survive.
Learning doesnt obey the strict inheritance patterns we find with genetics. In some
species, yes, learning happens entirely from mother to offspring. For example,
chimpanzees seem to learn foraging techniques and other social behaviours almost
entirely from their mums. Because of the way that chimpanzees interact, its hard to learn
a skill from an unrelated adult or one of your peers. But it does happen occasionally.
In humans, however, lateral transmission from peer to peer and diagonal
transmission from an adult to another adults child happens all the time.
Sometimes, its hard to draw a clear dividing line between learning and niche creation.
For example, you learn language mostly without anyone trying to teach you. You dont
sit down with a 6-month-old and say, Right, lets learn the words yes and no. In fact,
children learn to recognize their mothers voices and their native speech sounds before
theyre even born, when their mother and other adults have no idea that the unborn child
is listening in.
And humans are really good at teaching, even when they dont realize it. For example,
most adults unconsciously shift the way that they talk to young children. They raise the
pitch of their voices, simplify their language, and repeat themselves. If you didnt know
better, youd swear that they had lost it. Most adults have no idea that this is excellent
teaching, structuring language so that it is easier to acquire.
We instinctually create a kind of cognitive niche, a cut-down-to-child-sized version of
language and other activities that make it easier for a kid to pick them up: repetition,
face-to-face, simplified, interactive. Most children are primed for this, geared up and
receptive, with great social skills compared to other primates. And most parents are
primed, too: sleep deprived, sure, but excited and ready to interact in ways that heighten
how effective this channel of inheritance can be.
If a child has a problem with reception or social interaction for example, gets too
stressed out from face-to-face interaction or cant hear whats going on or if the parent
doesnt help to create a supportive niche for example, if a mother has severe post-natal
depression or people dont interact with the child this form of inheritance can get
undermined.
The good news, however, is that many of these developmental processes, these forms of
inheritance, are really robust. Because they dont go strictly from parent to child, and
they depend upon this whole human world weve built up, theyve got all kinds of
redundant channels.
The bottom line, however, from all these examples is that, if you want to understand us,
humans, and why were distinct, you have to realize that its not all in our genes. Our
genetic peculiarities create the possibility of these other mechanisms, which we can find
in other species, but not necessarily to the same degree that we find in ourselves. The
complete instructions to build a human arent just written in our genes; weve also carved

them into the world all around us.

Symbiosis & cooperation

Yet life is not all just competition. Our cells dont fight each other; colonies of ants and
termites work together on a massive scale, even laying down their lives for the good of
the nest; and primates and other animals lead complex social lives. Darwin recognized
this and drew attention to the way that the animal kingdom was not simply vicious
competition, but also involved dependence, inter-relations, even symbiosis.
Symbiosis occurs when two species live in close relationship and both derive benefit
from that relationship. A parasite does not have a symbiotic relationship because the
parasite, like a tick or a tapeworm, does not give any benefit back to its host. And both of
these relationships are different to a predatory relationship because both require that the
host remain alive in order to benefit. Symbiotic relationships can be quite close, or the
links can be more indirect.
For example, we have a gut full of helpful bacteria that weigh more than our brain, more
cells of bacteria than we have bodily cells because theyre so small. Thats right, youre a
mobile home for 100 trillion microbes of hundreds of species your microbiome.
Without them, youd survive, but it would be hard to digest everything you eat, and your
immune system would suffer. Thats a pretty tight symbiotic relationship you and your
(mostly) happy gut full of bacteria.
Other symbiotic relationships exist in the animal kingdom: ants and fungus that they
farm, sea anemones and clownfish, cleaner fish and larger sea creatures, even bees and
flowering plants. In all these cases, species co-evolve; that is, they become so adapted to
the opportunities presented by the other, that ones evolution shapes the evolution of the
other.
A more distant relationship is the link between living things through oxygen. Up until
about 2.4 billion years ago, the atmosphere of our planet had much less oxygen. The rise
of cyanobacteria, bacteria that could produce energy through photosynthesis, began to
shift the composition of our atmosphere. Photosynthesis allows plants, algae and bacteria
to create energy from light by combining water and carbon dioxide to produce a simple
sugar, but it leaves loose oxygen as a waste product.
Over millions of years, the cyanobacteria created more and more oxygen; at first, this
oxygen was mopped up by other geological processes, especially by oxidation, which we
see as rust in iron compounds. But eventually, after 200 million years, the cyanobacteria
filled these oxygen sinks, and the gas started to build up. In a sense, you can say that their
pollution laid the foundation for the rise of complex life. In that sense, we are symbiotic
with these organisms that can create energy from light because we literally depend upon
their waste in order to breathe.
But this shift would have been a catastrophe at the time; all that oxygen wiped out the life

that depended on that previous environment, and possibly contributed to a massive 300
million-year ice age. An ecological change was produced by a new form of life, laying
the foundation for life as we know it in the mass destruction of what had existed before.
Cyanobacteria are also important to understanding the really important ways that novelty
or variation can be introduced into species, through genetic sharing. Cyanobacteria are a
key player in what the late biologist Lynn Margulis called endosymbiotic theory.
Margulis first began proposing the endosymbiotic theory in the 1960s against enormous
opposition. She argued that complex, multicellular life was all the result of a form of
symbiosis or inter-organism cooperation. According to Margulis, a single-celled
organism engulfed its symbiotic partner; thats why we call it ENDO-symbiosis,
because one symbiot is inside the other.
Margulis took the work of earlier researchers who realized that parts of the process
through which a complex multi-cellular organism reproduce how one cell becomes
two and so on looked a lot like how bacteria reproduce. For example, the
mitochondria in our cells, the parts that produce energy, and the chloroplasts in plant
cells, the parts responsible for photosynthesis, go through their own reproduction process
as the cell divides. Moreover, if you look closely at these little organelles, or parts of the
cell, they have membranes around them like little Russian dolls, like they are
compartments inside the cell. Margulis argued that complex eukaryotic cells, like the
kind you and I have, were once cooperative little communities of independent parts that
had grown together into a permanent endosymbiotic relationship.
Margulis went on to argue that the modern synthesis, that combination of genetics with
the theory of natural selection, had over-emphasized competition and mutation as the
forces that drove evolution. Margulis described, instead, how complex life came about
through cooperation, mutualism, and interdependence, especially through symbiogenesis,
or the creation of new forms from the combination of pre-existing life, like the
cyanobacteria being captured as on endosymbiot by another bacteria. Margulis argued
that eukaryotic organisms multicelled organisms were the result of symbiotic
microbes becoming permanently linked so that they required each other to live.
Margulis theory, which had been suggested by Russian botanist Boris Kozo-Polyansky
in the 1920s, took an enormous leap forward in the 1980s when the technology was
developed to read the genetic material in cells. As Margulis theory predicted, some parts
of the cell, especially the energy producing organelle (the mitochondria) and the part
responsible for photosynthesis in plant cells, turned out to have their own DNA, different
to the nuclear DNA of the cell. Her theory was controversial at the time, and parts of it
rightfully still are, but most biologists now accept that symbiogenesis is crucial to
understanding complex multicellular life.
Like any radical thinker, Margulis may have over-stated her point, but symbiogenesis,
even if it is rare, encourages us to look for other mechanisms, other than just mutation
and competition, that might propel changes in life. Certainly, Darwin himself would have
been interested in this kind of observation, for he had noted in On the Origin of Species
how life grew in complex networks of inter-dependence.

Viruses & cooperation

One of the more controversial parts of Margulis theory, the theory of endosymbiosis that
we talked about in the last segment, was that variation in species wasnt just a result of
incorrect DNA copies; rather, new genetic material might be introduced into the nucleus
of a cell by either viruses or bacteria. Again, new research techniques have born out this
controversial idea. We now realize that single-cell creatures can sometimes pass DNA
laterally, that is, share amongst themselves, or even steal genetic material, rather than just
get it from their parent and pass it to their offspring. This genetic sharing helps us to
understand a whole series of evolutionary events, including even how sexual
reproduction itself evolved, as well come back to next week.
But perhaps the most startling is the discovery of large amounts of viral DNA in the
genomes of complex organisms, including our own. To understand how it got there, I
have to explain a little bit about retroviruses.
A retrovirus is a little bit of RNA, the simpler single-sided cousin to DNA, that inserts
itself into a living cell. The RNA then sort of reverses the usual order of things in the cell
nucleus, by building a DNA version of itself and clipping that into the cells DNA, using
the cells own reproductive mechanism to get itself copied. Some retroviruses are very
dangerous to their host, for example, HIV and some that cause cancer.
But some less virulent of these retroviruses are so successful that they become
endogenous. That is, like the endosymbiot, these viruses become permanently part of the
organisms genome so that they get passed on to the organisms offspring. They manage
to hack their way, not just into our body cells, or somatic cells, but into our germline
cells, the ones we use to make offspring, the ones that early twentieth century
evolutionary geneticists thought could not be affected by anything that happened during
an organisms lifetime.
In big, multi-celled animals like us, theres a lot of ancient retroviruses lying around,
artifacts of previous attacks which our ancestors survived, or of insertions by retroviruses
that werent all that dangerous in the first place. In fact, researchers now estimate that
there are about 100,000 pieces of viruses in human DNA, making up about 8% of all our
genetic material, getting passed on from generation to generation. Some of them can still
flare up into diseases; for example, endogenous retroviruses are now considered a
potential source of auto-immune disorders, when the immune system starts to attack the
body itself.
But some of this retroviral material gets used by the host in constructive ways, including
to regulate gene activity. In fact, endogenous retroviruses are crucial for the formation of
the placenta in all mammals. The placenta passes nutrients from the mothers body to the
embryo, and it protects the embryo from the mothers immune system. Without the
placenta, wed have to still be laying eggs. The same mechanism that allows the
retrovirus to latch onto a cell helps to fuse cells together in the placenta, so that mammals
can essentially have an internal egg. If the ancestor of the mammals had not caught this

retrovirus, we would still be laying eggs.

The point is not that this undermines the theory of natural selection. Retroviral insertions
and endosymbiosis help us to understand that variation can arise in many ways and that
selection may mean fitness for cooperation, or the ability to find the right partner, not
just competition. A living thing may find a niche to adapt to within the gut of another
living thing, or even inside its cells. If this relationship helps the host, the addition may
actually make that host more fit, and more likely to reproduce.
In social life, we see this even more; selective pressures may favor living things that
work well as a team.
Perhaps the most striking example of this is in eusocial animals, that is, animals in
which a substantial part of the population is sterile, like ants, termites and bees. Worker
animals typically serve a queen; the majority of the species never reproduce. How could
evolution favor individuals who cannot pass on their traits, for whom there is simply no
inheritance?
We realize now that, for some species, inclusive fitness is the key to understanding their
behavior; inclusive fitness recognizes that an individuals inheritance doesnt just come
from its own offspring. Because you are closely related to your siblings, their children,
too, can be seen as passing on a proportion of your genes. Inclusive fitness is one way to
explain apparently altruistic behavior; saving your brothers or sisters kids means that
you are creating more individuals related to you. Worker bees are an extreme example of
that.
Another possibility is group selection. Building a stable, highly-integrated community
may mean that your group is more cooperative; if resources are scarce, and communities
have to fight for them, then tight internal cooperation might make your group prevail
over rival groups. The idea of group selection leads some theorists to talk about multiple
levels of selection; that selection acts on individual genes, on individual organisms, and
on groups.
Both of these theories are themselves controversial, but no one disputes that cooperative
behavior, even the rare but extreme form of eusociality, can be an evolutionarily
successful strategy. The biomass of ants alone makes up more than half of all the insects
on the planet.
Arguably, humans, too, have benefitted from cooperation. From a precarious, small
population about 100,000 years ago, we have exploded in number, not just through
competition, but also through an extreme form of cooperation. The total mass of humans
on the planet is eight times greater than all wild vertebrates on land combined, and we
have about the same mass as all the fish and whales in the sea.
Symbiosis with other species hasnt exactly worked out too badly for us either: our
domestic animals outweigh all wild land animals by twenty times. Together, humans and
their herds all the pigs and cows and chickens and dogs make up more than a
quarter of all life on the planet by weight. Were even shaping each others evolution,
that is, co-evolving together.

We have gone from being a fragile, unusual ape to a world-changing organism in about
6000 generations, but this isnt the first time that a life form has worked this kind of
massive change to the planet. You should know. Youre breathing the evidence.

References and Resources

Key concepts
The basic theory of natural selection is that variation leads to difference in survival; if
survival is based on traits that are well suited to selective challenges, and those traits are
inherited, adaptation will occur over time.
The modern synthesis combined Darwins theory of natural selection with an
understanding of genetics, especially a focus on population-level dynamics.
Our increasing understanding of genetics helps us to see the complexity of inheritance,
including the possibility that gene expression may allow some experiential effects to be
inherited.
Multi-cellular life and social organisms also involve some degree of cooperation, not just
competition.

Resources
Darwin Online -- complete collection of manuscripts. (link)
Darwin exhibit (online) at the American Museum of Natural History. (link)
Interactive Voyage of the Beagle (online) at the Natural History Museum (UK). (link)
Ghost in Your Genes, NOVA, PBS (USA). Website supporting a special on epigenetics.
(link)

References
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Microbiome Project strategy for comprehensive sampling of the human microbiome and
why it matters. The FASEB Journal. doi:10.1096/fj.12-220806
Avise, J. C. 2006. Evolutionary Pathways in Nature: A Phylogenetic Approach.
Cambridge: Cambridge University Press.
Boyd, R., and Richerson, P. J. 1985. Culture and the Evolutionary Process. Chicago:
University of Chicago Press.

Boyd, R., and Silk, J. B. 2006. Adaptation by Natural Selection. In How Humans
Evolved. Fourth Edition. Pp. 1-23. New York and London: W. W. Norton & Co.
Darwin, C. 1868. On the Origin of Species by Means of Natural Selection; or, The
Preservation of Favoured Races in the Struggle for Life. New York: Appleton. [Note:
There are multiple editions.]
_____. 1901. Descent of Man, and Selection in Relation to Sex. London: Murray.
Eldredge, N., and Tattersall, I. 1982. The Myths of Human Evolution. New York:
Columbia University Press.
Feil, R., and Fraga, M. F. 2012. Epigenetics and the environment: emerging patterns and
implications. Nature Reviews Genetics 13, 97-109. doi:10.1038/nrg3142 (abstract)
Gilbert, S., Sapp, J., and Tauber, A. 2012. A Symbiotic View of Life: We Have Never
Been Individuals. The Quarterly Review of Biology 87 (4): 325-341. doi: 10.1086/668166
Jablonka, E., and Lamb, M. J. 2005. Evolution in Four Dimensions: Genetic, Epigenetic,
Behavioral, and Symbolic Variation in the History of Life. Cambridge, MA.: MIT Press.
Kirschner, M. W., and Gerhart, J. C. 2005. The Plausibility of Life: Resolving Darwins
Dilemma. New Haven, CT: Yale University Press.
McKinnon, S., and Silverman, S, eds. 2005. Complexities: Beyond Nature and Nurture.
Chicago: University of Chicago Press.
Mills, C. 2004. The Theory of Evolution: What It Is, Where It Came From, and Why It
Works. Hoboken, NJ: J. Wiley.
Morris, J. J., Lenski, R. E., and Zinser, E. R. 2012. The Black Queen Hypothesis:
Evolution of Dependencies through Adaptive Gene Loss. mBio doi: 10.1128/
mBio.00036-12
Morris, R. 2001. The Evolutionists: The Struggle for Darwins Soul. New York: W. H.
Freeman.
Odling-Smee, F. J., Laland, K. N., and Feldman, M. W. 2003. Niche Construction: The
Neglected Process in Evolution. Princeton: Princeton University Press.
Ridley, M. 2003. Nature via Nurture: Genes, Experience, and What Makes Us Human.
New York: HarperCollins.
Ridley, M., ed. 2004. Evolution. Oxford: Oxford University Press.
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and the evolution of gene regulation. Nature Reviews Genetics 13, 505-516. doi:10.1038/
nrg3229
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NJ: Wiley-Liss, John Wiley and Sons.

Tooby J. and DeVore, I. 1987. The reconstruction of hominid behavioral evolution

through strategic modeling. In Primate Models of Hominid Behavior, W. Kinzey (Ed.).
New York: SUNY Press.
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Links
Rob Dunn, The Garden of Our Neglect: How Humans Shape the Evolution of Other
Species, Scientific American. (link)
Barbara J. King, Convergent Evolution: Hyenas Offer Clues To The Human Past, 13.7
Cosmos & Culture, NPR (USA). (link)
Carol Kaesuk Yoon. From a Few Genes, Lifes Myriad Shapes, The New York Times, 26
June, 2007. (link)
Dick Teresi, Discover Interview: Lynn Margulis Says She's Not Controversial, She's
Right, Discover. (link)
Carl Zimmer, How Many Species? A Study Says 8.7 Million, but Its Tricky, The New
York Times, 23 August, 2011. (link)
_____. How Microbes Defend and Define Us, The New York Times, 12 July 2010. (link)
_____. Mammals Made By Viruses, The Loom (weblog), Discover Magazine. (link)
_____. We Are Viral from the Beginning, The Loom (weblog), Discover Magazine.
(link)

~~~~~~~~

Sex
Natural selection is not simply about survival an animal must also pass on its DNA.
This creates the possibility of individuals competing with other members of the same

species through reproduction. Our bodies offer us clues about how our ancestors
reproduced, competed with each other, but also how they must have cooperated in order
to reproduce successfully.

Sex: Why bother?

Why bother to have sex at all?
I know what some of you are thinking: Youre obviously doing it wrong. But Im
serious.
From an evolutionary perspective, lots of living things reproduce without sex. All single
celled organisms can copy themselves. Even some very complex animals can reproduce
asexually or by parthenogenesis. Some species of lizards, amphibians and fish, for
instance, have no males; a female-only species simply lays eggs that are clones of
themselves. Other species can reproduce either sexually or flying solo such as the
komodo dragon and hammerhead sharks depending on the situation.
Although it might not sound like a lot of fun, from an evolutionary perspective, it seems
like a win. After all, if you reproduce asexually, you pass on all of your genes to the next
generation. If you reproduce sexually, you only pass on half of your genes; your DNA
gets mixed with the DNA of the other parent, so a child is only half of you. If your genes
were just being selfish, they wouldnt want in on that bargain.
In addition, sex limits your reproductive rate, slowing down the process, and it might
even make it impossible to reproduce, if for example, you get eaten by a predator before
you manage to find a mate. If you wash up on a desert island alone, youre not going to
leave any offspring if you reproduce sexually. In contrast, an asexually reproducing
animal might quickly fill the island.
Put simply: the cost of sex is males. If you reproduce asexually, you dont need them.
But sex has been around for over a billion years, so it must be good for something. We
dont know for sure why sex first arose, but over time, sexual reproduction has
accumulated a number of advantages, other than just having a good time.
The key is to understand that, from an evolutionary perspective, sexual reproduction is a
trade-off: on the one hand, you limit your ability to reproduce. On the other hand, there
are both genetic and ecological reasons why mixing together DNA from two parents, or
recombination, might be an adaptive advantage.
First, and probably foremost, recombination produces a lot of variation, which as we
know, is the raw material for evolution. Because of recombination, children are not
clones of either parent, but are each new combinations of genes; each siblings is also
different, even in cases of identical twins (although the differences are pretty slight).
Recombination allows traits that are beneficial to spread quickly through a population.

Even multiple traits can spread through a population as lines of inheritance cross and recross in a sexually-reproducing species, whereas reproduction by cloning asexually does
not produce this crossing of genetic material. This pattern is so beneficial that some
asexually reproducing single-celled organisms have come up with their own ways to do
this, swapping genetic material from one to the next.
Recombination also can clean up genetic errors. Getting one copy of a gene from each
parent may allow an offspring to overcome some copying errors. In an asexually
reproducing species, you only get genetic material from a single parent; if its defective,
youre stuck with it, whereas with recombination, you are likely to have two copies of
almost all your genes. For this reason, sex may decrease the possibility of extinction.
Where we dont have recombination like with the Y chromosome, because you only
can get it from your father we dont have this repair mechanism, and detrimental
mutations can start to pile up. In a few million years, this is going to be a problem for us
males of the species.
The advantage of sex, however, isnt simply that variation might produce a beneficial
new trait or repair genetic damage. Sexual reproduction also makes a species a kind of
moving target. All the time, other species are trying to figure us out. Viruses, bacteria,
parasites dont look behind you theyre all trying to crack your immune system, to
find a way to break down your defenses. Sexual reproduction means that every individual
is a new problem for the parasites and viruses to solve.
We call this the Red Queen theory after the Red Queen of Wonderland in the book,
Through the Looking Glass. Alice and the Red Queen were in a race, running as hard as
they could, but apparently going nowhere. Alice, panting, says,
Well, in our country,... you'd generally get to somewhere else if you run very fast for
a long time...
The Queen responded: A slow sort of country!... Now, here, you see, it takes all the
running you can do, to keep in the same place.
Evolution is a bit like this race; it never stops, and you have to run as hard as you can to
stay in place. Species that do not seem to change are undergoing constant selection, even
if it is stabilizing selection.
A species like the horseshoe crab, apparently unchanged for millions of years and quite
uniform one to the next, turns out to have a tremendous amount of genetic variation
underlying its anatomical uniformity. Those species that reproduce by cloning may have
their days numbers, as faster reproducing viruses are constantly generating new variants,
one of which might be especially virulent, and parasites may load up on a stationary
genetic target.
Sexual reproduction also produces its own evolutionary pressure: sexual selection. Sexual
selection is an evolutionary pressure that is produced by intraspecific competition in
order to mate: that is, animals must compete against members of the same species to
mate, rather than against predators or the environment or some other external factor.

Consider the peacock, an animal that gave Darwin fits of anxiety until he figured out
what was happening. A male peacock has an absurd tail. At the height of mating season,
Mr. Peacock drags around an enormous, brightly coloured fan of these incredible
feathers. They are so big that they make it harder to fly, so incandescent that they
virtually announce the birds presence to predators.
And if youve never been around them, male peacocks are even more absurd than they
look: they shriek at the top of their lungs and do this wild shimmying dance whenever
they see an eligible female peahen, or anything that vaguely reminds them of a peahen,
including horses and dogs and sometimes their own reflections. Seriously. As the male
peacocks dance back and forth and shimmy, they shake their enormous feather tails. Its
all a bit much.
We now realize that they are engaged in a type of competitive sexual display; they are, in
a sense, auditioning to mate with peahens. If a male doesnt have a terribly impressive
tail, or doesnt put on the right show, hes not going to be fathering any baby peacocks.
The female peahens exercise mate choice; if a trait is particularly attractive to the
females, that trait will be selected in the males and tends to get steadily more and more
exaggerated.
We know this because researchers have taken very successful males peacock studs
and lopped off the bright eyes of some of their tail feathers. That mating season, with
their reduced tails, the studs turned to duds, unable to impress with their paltry plumage.
Other male birds use their songs, build elaborate nests, or engage in ritualized dances to
try to become the preferred choice of a female. This type of sexual competition is one
way to make sure that a female gets the best possible mate; the healthiest, strongest
males, theoretically, should put on the best displays.

Sexual competition
Peacock males and most birds dont compete face-to-face, but some species do engage in
direct competition as a form of sexual selection. In bighorn sheep, for example, at the
start of the mating season, or rut, rams have to sort out a dominance structure to
determine who will mate. The male rams square off and literally go head-to-head,
slamming into each other to demonstrate that they are the strongest. The dominant males
tend to reproduce much more successfully, which can drive the evolution of the species.
In many primates, though not all, direct competition is also the rule. One of the most
dramatic examples is gorillas. If a male gorilla is going to reproduce, he has to take
control of a polygynous group of females. Polygyny is a reproductive system in which a
single male mates with multiple females, but not the other way around. A male gorilla
does this by deposing and taking the place of the male silverback, the only male who
can reproduce.
Among gorillas, only the strongest males pass on their genes to the next generation. The

rest are effectively sterile from the point of view of evolution. This brutal sexual
competition has produced extraordinary sexual dimorphism, or a difference between male
and female forms of the same species. A female gorilla is big, but a male is enormous
a silverback is up to two and a half times the size of his mates. In addition, he has an
outrageous skull, with terrifying canines and huge crests to anchor massive jaw muscles.
The irony is that the owner of this skull is a vegetarian.
Direct sexual competition has driven gorilla evolution to produce an extraordinary set of
weapons, all for the purpose of intraspecific competition, or fighting with other male
gorillas. Over and over again we see this: sexual selection, intraspecific competition, can
drive the production of more and more elaborate plumage, more extreme weapons, larger
size in males, and a host of other phenomena that we call secondary sexual
characteristics. We call them secondary because, unlike primary sexual characteristics,
they are not practically necessary in order to reproduce like testes and ovaries, although
they are used in the behaviours that lead up to reproduction like courtship dancing or
rutting.
Sexual selection tends to drive dimorphism, or create a marked difference in male and
female form or behavior. In the elephant seal, for examples, the males bulls are six
times larger than females.
In fact, the degree of difference between males and females, the degree of dimorphism, is
sometimes seen as a good proxy measure of the intensity of sexual selection. The more
intense the selection, the more sexually dimorphic a species will become, the differences
becoming more and more pronounced. Males will develop bright colours, elaborate songs
or dances, specialized weaponry, or greater size, which their female counterparts dont
have.
But wait a minute, some of you may be thinking: why is it that males have to fight over
females? After all, in Australia, we tend to read about supposed man droughts, and we
all know that women have to work hard to get a man, right? Beauty contests arent held
among men, are they?
Well, thats not what evolution and biology teach us.
To understand which sex is going to have to compete, we have to think about which sex
does the most investing in offspring. The evolutionary theorist Robert Trivers, one of the
most important evolutionary thinkers since Darwin, pointed out that in any species,
sexual reproduction is a kind of joint investment of the parents. Trivers realized that, in
most species, the parental investment of each sex is not equal.
For example, even in producing gametes, the gametes are not the same size: eggs and
sperm are not identical. Over the last billion and a half years, one sex has tended to invest
more heavily in each gamete, building a full mechanism to reproduce, a fully-fledged
cell, around it; we call this sex the female and this gamete the egg. The other sex has
tended to specialize in larger-scale mass production of little gametes, with little more than
the DNA being transmitted, and spread them around. We call this sex, male, and the job
of sperm is to get its half of the full complement of DNA to the egg, where most of

the equipment for producing life is waiting. The difference in gamete size is called
anisogamy, and wherever you have it, you start to have sexual difference.
In some species, the females invest more in each offspring; in some species, its the
males. Whichever sex invests the most becomes a limiting factor on the reproductive rate
of the species. Whoever has to sit on the eggs or look after the infants or carry them to
term if pregnancy was long or feed them, is going to be a kind of brake on the
reproductive rate of the species as a whole. Depending on how big the gap was, the other
sex is likely going to have to compete more or less severely in order to reproduce.
In all mammals, females tend to be the limiting factor, unless the number of males has
dropped off. That is, the rate of reproduction of a species is limited by the number of
females in the population; because female mammals carry their infants to term, rather
than simply laying a clutch of eggs and taking off, and because mammals feed maternally
its how they got the name mammals after all, because of the mammillary gland. So
its generally the females that engage in mate choice.
Trivers hypothesis, that differential investment determines which sex will have to
compete, predicts that human males will have to compete to mate, not females. You
would never know this if you listened to Western popular culture, of course; you would
assume that women have to try their hardest to attract a male, but thats not at all what
evolution suggests.
Like virtually everything with humans, we make things more complicated than they are.
Sex is certainly no exception.

Males and females

Our popular culture wants us to believe that men and women are fundamentally different,
even opposed: men are from Mars; women are from Venus. Boys are made of snips and
snails and puppy dog tails; girls, from sugar and spice and everything nice. Pink and blue.
Opposites attract. All that sort of thing.
In fact, we are not so different. Women and men are both from earth, not Venus or Mars.
And if we have a lot in common with other species, we have even more in common with
each other. We may have ideas about gender, what is masculine and what is feminine,
built up in our culture, but our biology tells a slightly different story.
First, male and female bodies are fashioned in the same workshop, tweeks on the same
basic design with a raft of surprising homologies at the same time that there are also some
really important differences. Weve got to remember this, even when we read research
about the evolution of human sex and reproduction because too often, this research starts
from our cultural attitudes or folk assumptions about the supposed gulf between men and
women.
Students are sometimes startled when we point this out anatomically. For example, the
testes and the ovaries begin life as the same organ, the scrotum starts out as the same

tissue as a labia, and we all have the same basic hormones, just in different proportion. If
you listened to popular culture, youd think testosterone was a kind of boy juice and
estrogen makes you female. In fact, women have testosterone, just as men naturally have
estrogen. Sure, levels are different in both, but take away either of these hormones from
men or women, and youll get health problems.
The point is that, although we tend to focus on and even exaggerate the differences
and Im going to discuss them, too male and female are homologous. In fact, up until
about six or seven weeks in utero, you cant tell whether a child is male or female from
its gross anatomy; you have to look at its chromosomes because the differentiation
processes havent kicked in yet. An embryo starts off as anatomically indeterminate.
All that changes in week six, when, in the case of boys, masculinization starts. On the Y
chromosome, present only in males, a gene imaginatively called the Sex-determining
region of the Y chromosome, or SRY, up-regulates certain developmental processes in a
kind of chain reaction that leads to testes formation. Cells responsible for a spike in
testosterone appear in the testes, which are still up inside the abdomen at this point, and
they start to pump out extra testosterone in week eight. Production doesnt peak until
around week twelve of uterine development. The penis and male gonads will mature until
around week fourteen, you get the first sperm starting to form. It becomes possible to tell
externally that an embryo is male as early as the third month of development.
Womens bodies start off a bit slower, but catch up quickly. About week eight and nine,
the vagina and uterus form if the embryo does not have that gene, SRY. The first ovarian
follicles, which will become eggs, appear around four months of development. If, for
some reason, the embryo is insensitive to hormonal effects, for example, does not have
receptors for the hormones that bring about masculinization, the default outcome looks
female, but will not be able to reproduce if it doesnt go through the feminization process.
From this perspective, female is sort of the default state for the embryo.
Every once in a while, a child is born with indeterminate sex; some of the more
conservative estimates suggest that the number is between 1-in-500 and 1-in-2000.
Although quite rare, what these individuals help us to recognize is that sex is not a switch
thats flipped either to male or female. Sexualization is a biological process and, in some
cases, doesnt go on to produce either male or female.
As part of this process, many of the secondary sexual characteristics dont show up until
puberty, starting around age 10 to 12; at puberty, another round of hormonal changes
propels sexual maturation. A spike in estradiol in girls cause the breasts to enlarge, body
shape and genitalia to change, and menstruation to start, the onset of adult fertility. High
androgen production in boys at puberty brings about a host of changes, including to the
sex organs, voice, body hair, overall build, and behavior.
Even after we mature, although we may believe that male and female are quite different,
our sexual organs behave in quite similar fashion, revealing the underlying homology.
When were sexually aroused, for example, blood rushes to the homologous organs
the penis and the clitoris causing swelling and flush, and the metabolic processes are
virtually identical in terms of heart rate and patterns of excitation.

There are important differences, of course. Men are, on average, larger than women, have
greater upper body strength, and seem to have a higher metabolic rate; women appear to
be better able to fight infection, mature younger, seem to be better at laying on body fat,
and, of course, lactate and menstruate.
A lot of evolutionary theorists say that, if you compare us to other primates, were a bit
more dimorphic than the purely monogamous species, but much less dimorphic than the
polygynous primates, like gorillas or orangutans. From the slight difference you might
predict a slight tendency toward polygyny, or a tendency for males to have more than one
mate.
In fact, for the past three or so million years, our ancestors have probably been staying
the same or, possibly, getting less dimorphic, closer to equal. If our ancestors were more
dimorphic and thats a topic of some debate then female hominins have been
getting larger faster, so that theyre about 80 to 85% caught up. That may not mean that
weve been evolving gender equality, but it does suggest that, if males are competing,
theyre not doing so directly. If males were engaged in sexual competition, wed expect
that some clearer sign of that competition from the past few million years.
In contrast, if we look at our most closely related cousins, studying the trajectory some of
the other great apes have gone through over the last ten or twelve million years, we see
that our species does not have the hallmarks of strong male competition.

Bodies dont lie

Just like our bodies offer details of past selective pressure, they also have clues of the
ways in which our ancestors reproduced, especially if we compare ourselves to other
species. Lets start with what mens bodies tell us to try to get some clues about the
evolutionary pressures on our ancestors from sex selection.
First, male gorillas have really small testicles and low sperm counts. No one really brings
it up around them because theyre these mountains of hairy muscle, but theyve got pretty
unimpressive balls.
In contrast, male chimpanzees have comparatively large testicles and high sperm counts.
Even though the male chimp is about a quarter of the weight of a gorilla, his testicles are
four times larger than his over-sized distant cousin. These disproportionately large
testicles allow a chimp to have sex multiple times a day without sperm count dropping
off too badly, possibly four or five times. (Of course, the chimp isnt all that impressive
when you realize that a lion can mate 75 times a day during the mating season.)
Why the difference between chimps and gorillas testicles? Because male chimps engage
in a form of sexual selection called sperm competition. When a female chimp is fertile,
she mates with a number of males. The dominant male in the troupe may try to keep other
males away, but its mostly a lost cause. Primatologist Jane Goodall observed one female
chimp mate 84 times with seven different males in her troop over eight days when she

was fertile. In a sense, each males chance at fathering a child is sort of like an
evolutionary raffle, with more sperm acting like more tickets in the raffle. Well come
back to why when we discuss womens bodies and children in the next video clip.
Chimpanzee males have faced so much to sperm competition that a large portion of their
sperm appear to be suited, not so much to impregnate a female, but more to block or
chemically kill sperm from other males. Some biologists suggest that as much as 99% of
chimp sperm is not there to impregnate a female, but is better designed for sperm
warfare.
Another way to look at it is that, gorilla males settle who is fittest directly, by fighting
violently to monopolize sex. In contrast, chimpanzee females mate with a bunch of males
and let their sperm sort it out. Since the last time that chimpanzees and gorillas shared a
common ancestor, the two have been under these radically different pressures of sexual
selection.
Human males dont look like either chimps or chimpanzees, and some of the data is
controversial, to say the least. On the one hand, weve got nowhere near the testicular
endowment of a chimpanzee.
On the other, mens bodies seem to anticipate the possibility that a partner has mated
with someone else, unlike the gorilla. In fact, the amount of sperm that a man ejaculates
is correlated, not with how long it has been since he last had sex, but rather with how
long hes been apart from the women with whom hes mating. If theyve been apart for a
while, he can up to triple the sperm he would normally pass on. The male body seems to
anticipate the possibility that a partner has had sex with another male and responds by
upping the ante, a clear sign of adaptation to sperm competition.
On the other hand (yes, thats at least three hands...), human males arent that well
endowed with testicles. After the first ejaculation of the day, sperm count drops off
precipitously; you can have another go if you like, but youre no chimp. More than 3.5
times a week, and your sperm count will suffer. In other words, our cultural ideas about
males may suggest that they want to sow their wild oats: evidence from our bodies is that
men may not have a whole lot of extra oats to sow.
But an even more interesting little comparative detail that may tell us about sex selection
is penis size. Male gorillas, on average, have erect penises that are about 3 centimeters in
length, or about an inch and a half. In contrast, the much smaller male chimpanzee has an
average erect penis length of around 8 cm. Humans usually come in around 13 cm
(according to scientists, not to the men themselves), disproportionately long and thick for
a primate.
Moreover, the human penis is unusual, with no penile bone to keep it erect other
primates have an actual bone men instead use a complicated hydraulic system.
Although it might be for sperm competition, some sex researchers point to the unusually
large, thick and hydraulic powered penis as a potential target for female preference, an
example of how womens mate choice may have affected our hominin ancestors.
If we turn to the evidence from mens bodies, then, we find clues about our ancestors

mating behavior, but not necessarily what some people might think. Back in 1967,
zoologist Desmond Morris, the author of The Naked Ape, could confidently write that the
evolutionary purpose of human sexuality was to strengthen the pair-bond and maintain
the family unit. Man, has that confidence taken a beating!
The truth is, genetics has delivered another blow to some of our more romantic ideas
about monogamy or mating for life, even in other species that we thought were models
of fidelity. Genetic testing in the field with wild animals has revealed that even some
species that we thought were thoroughly committed often have high cuckoldry rates. That
is, the chicks or pups or whatever turn out not to belong to the fellow who is hanging
around most diligently. Gibbons, for example, once held up as the model of a
monogamous great ape, turn out to be more wide ranging, the females especially
engaging in extra-pair couplings when they are most fertile.
Some researchers have controversially argued that evidence suggests human females may
have adapted to a pattern of seeking long-term support from a male partner but
occasionally getting genetic material from a different male. Or, as the scientists
euphemistically put it, they have a partner but, especially when theyre fertile, may
engage in extra-pair coupling. (If youre still not with me, that means theyre sleeping
around.)
Supporters of this theory have pointed to genetic evidence in Western countries that
many children have the wrong man listed on their birth certificate as their father. This is
hard research to do, but its well known in medical circles that a range of tests for things
like the inheritance of disease typically uncovers that somewhere between 5 and 10% of
the population in countries like the US and the UK and Australia could not possibly be
related to the father listed on their birth certificates.
Even female orgasm may be a mechanism for a woman to, inadvertently, give the sperm
of a preferred partner a greater chance of impregnating her. If so, this mechanism
assumes that a woman is mating with multiple partners.
The point is that, when we look more clearly at the evidence our bodies present, and
compare ourselves to those species to whom were most closely related, we may find that
evolutionary dynamics do not necessarily confirm our assumptions about the nature of
men and women. The pressures of sexual selection have left traces upon our bodies, but
they do not determine how we must act.

How our babies made us

When a horse is born, its almost ready to stand up. Within an hour, the foal is on its feet.
By the end of the first week, the babys running. Awkward, sure, but you try to catch one.
We say that a horse is a precocial infant. Precocious: quick to develop. Its mum looks
after it and will nurse it, but the newly born horse is quickly able to do some of its most
important tasks for itself.

In contrast, check out a human infant at birth. Yeah, cute, magical and all that. But totally
helpless, and it stays that way for a long, long time. Cant feed itself. Cant walk. Cant
go out and hunt, clean itself, dress.... Nothing. Dependent upon its parents until its what?
Five? Ten? Twenty-two?
No, seriously. Studies of hunters and gatherers reveal that a child will finally produce as
many calories as it consumes when it reaches no less than about 18 years of age in many
foraging groups. Up until then, it must depend on others for at least part of its nutritional
intake. One anthropologist has estimated that it takes 13 million calories to get a child to
a point where it can support itself.
This is an extraordinary ask. In most species, as soon as theyre weaned, animals can
provision themselves. In other great apes, for example, mothers nurse young for a long
time, sometimes longer than humans, but weaning means independence. Youre out on
your own, and the next sibling is there to take your place.
Not in humans. Getting food is so information dependent in foraging groups, and children
so immature relative to adult tasks, that it takes a lengthy apprenticeship to be able to take
care of yourself. So the children start to overlap, one being born before her or his
predecessor is independent. Orangutans can wait for a child to become nutritionally selfsufficient to have another baby it takes them about eight years between births but in
humans, we cant wait that long. Wed only be able to have a child every decade-and-ahalf or so. Our fertility is already low; that would probably drive us to extinction.
This patterns marks humans as extremely altricial. They require extensive care and
support for a long time.
We dont really know for sure why we give birth to such immature infants; we used to be
quite confident that it was because of the clash between the growing size of the infants
head and the mothers narrow pelvis, adapted to bipedalism. Anthropologists called that
the obstetrical dilemma, a conflict between the biomechanics of walking and the ability
to deliver a large-brained infant if you carried it for too long.
Now, its becoming clearer that the obstetrical dilemma isnt as bad as we thought; it
would be possible to grow a wider pelvis if that was really the issue. Instead, the issue
appears to be that this big-brained infant is an energy sponge in utero, creating a
metabolic crisis as it grows and grows. In a sense, we have to have our infants when we
do, even though theyre so helpless, because theyve hit a metabolic ceiling; any larger,
especially with their growing brains, and there just wouldnt be enough nutrients in the
system. What weve got to realize, and well come back to it next week, is that brains are
expensive in energetic terms; they gobble up calories at a high rate. The mother cant
keep up and must eject the child, where it will be possible to keep up with its voracious
appetite.
You may not have thought of it this way if you havent had a child of your own, but
theres actually a deep conflict between mother and child. The demands of our children
have shaped, not just mothers bodies, but the whole of our species, during evolution.
Although evolutionary stories about the origin of humanity used to focus almost

exclusively on mens business hunting, fighting, finding a woman to mate with

feminist criticism and better science have really turned the spotlight on women and
infants in the past few decades.
If you had asked an evolutionary theorist forty or fifty years ago, for example, about the
forces shaping womens bodies, he most of them were men would probably have
said something about attracting men or hanging onto a mate. Recent research on sexual
selection, however, turned that on its head. If we can see traces of our female ancestors
preferences in mens bodies, we can see the imprint of our infants demands on womens
bodies.
For example, one of the most obvious distinguishing secondary sex traits of women is a
higher body fat percentage than men and a distinctive body shape, with that fat deposited
on the breasts and hips. Human women, for example, have fatty breasts. No, Im not
talking about just some women: all human women. You dont need that fat to produce
milk; other primates produce milk without fatty tissue on the chest, no problem. Same
thing with the hips and buttocks. Again, unusual, compared to other species, and
apparently unnecessary.
Trivers explanation of sexual selection has taught us that this extra fat on breast and hips
likely isnt there to attract males. The cross-cultural evidence supports this hunch: men in
some societies may find one of these part parts erotic, but others dont. We realize that
males would be more likely affected by mate choice, not doing the choosing.
Increasingly, we think that the fatty deposits have more to do with infants hunger than
with potential mates desire. The extra tissue may be a kind of insurance policy against
low-level starvation that could interfere with a mother lactating. Our ancestors lived by
foraging, and food may not have always been easy to find. They probably had to survive
some pretty lean periods. An infants brain is growing so fast after it is born, at rates only
seen in utero in other species, that the infant needs constant top ups of energy. Extra fat
storage could be one way to make sure that you dont get a temporary milk shortage,
which could have a devastating impact on brain development.
Similarly, human infants are born disproportionately fat, as well. Other mammal infants
arent born such little chunks. Sarah Hrdy argues that the baby fat is not just there to
consume, but advertises to the mother, Hey, Im a very healthy, strong, cute baby. You
should invest your energy in me! Hrdy points out that, because the interests of infant and
mother do not perfectly coincide, and because human mothers have to make tough
choices about how to allocate scarce resources, a baby has to persuade its mother to fulfill
this challenging role as provider.
This conflict between an energy-hungry child and its mother, then, is part of a greater
pattern in human reproduction. The mother-infant relationship is the social centre of all
primates universes, but this centre is not harmonious. A mothers and an infants
interests, from an evolutionary perspective, do not perfectly coincide.
First, its in the infants interest to survive at all costs, regardless of the consequences to
the mother, to get as much nutrition as it can in embryo, for example, and be born as

large and healthy as possible. Babies want it all.

For a mother, passing on her genes may involve trade-offs, cutting losses with an embryo
that isnt optimal, and limiting her contribution to one offspring so as to support a number
of them. A mother wants to ration herself in the face of an infants demands because her
evolutionary legacy is not a single child, but all her children, including ones potentially
not yet born.
Part of what makes this difficult to see in Western societies and other wealthy groups is
that infant morality has been significantly reduced there, and fertility often quite low per
woman, even though our calorie-rich diets make modern women capable of having
babies very quickly.
For much of our existence as a species high infant mortality would have been an
important evolutionary pressure. Our fertility isnt very high compared to other species,
and it would have been a lot higher before the advent of birth control, but prior to
agriculture, mothers probably wouldnt have been able to have a baby but every five or so
years. In some foraging groups still alive today, mortality of infants is so high, and
fertility sufficiently low, that some women leave no descendants, even though they may
give birth to a number of children.

Struggles with children

From an evolutionary perspective, having a baby is not the goal getting that baby to
survive until you have grandchildren, preferably a lot of grandchildren, is. The problem is
how altricial our infants are. Recruiting help to keep a baby alive, and sometimes making
hard choices about how to allocate your resources, have probably been important to our
ancestors for a very long time.
One example of how a mothers body, without her knowledge, makes tough choices,
resists a babys demands, and weighs the viability a fetus is menstruation. Menstruation
happens because a woman, when shes fertile, builds a uterine lining during her cycle and
then expels part of it after her peak of fertility passes. Many mammals do not menstruate;
instead of building a uterine lining and then expelling part, they wait and only build one
if theres a fertilized egg. Or, if they build the lining, they reabsorb it if its not used. You
dont have to menstruate to reproduce.
We now realize that there are at least two good reasons to menstruate. First, menstruation
may be a kind of stress test which can expel embryos that are not sufficiently embedded
to be viable over a whole pregnancy. Menstruation checks a pregnancy very early to see
if its worth investing in, or if the process should just be started over. We know that a lot
of conceptions get naturally aborted early on; around 30% of fertilized eggs are expelled
in the first few weeks, with as many as 50% of all conceptions terminated early naturally.
The second reason women may menstruate is that they have to build a uterine lining in
advance to be ready when that energy hungry fertilized egg implants, so that the mothers

body can resist it getting too deeply embedded, too hot-wired into her system.
Evolutionary biologist David Haig, originally from Australia, but now teaching at
Harvard, has pointed out that many of the complications of pregnancy may arise from a
silent struggle between the mother and the fetus. Haig argues that the placenta, once it
forms at the end of the first trimester, is under the control of the fetus. The placenta
actually invades the mothers tissue and reroutes as much blood as possible by
restructuring the mothers arteries so that more blood flows to it. At this point, the mother
cant control the flow of nutrients to the fetus without cutting the nutrients in her own
blood, and the placenta can release hormones directly into the mothers blood to try to
wring out more nutrition.
The fetus and the mother get in a kind of food fight, battling after each meal for more of
the glucose in the mothers blood. The mothers body tries to absorb it quickly; the
placenta releases hormones to try to stymy the mothers ability to rob it of glucose so that
more winds up with the baby.
In some cases, this struggle can produce gestational diabetes in the mother. Although we
now think of this as a health problem, the condition leads to a particularly fat, healthy
baby, especially in an environment where obesity is not a danger. Gestational diabetes is
a sign that the fetus and placenta are getting the upper hand, undermining the mothers
ability to produce insulin, which would keep that blood sugar down.
Once the baby is born, its still terribly demanding, so much so that it can undermine a
mothers ability to reproduce as much as possible. Unless she can get a break, shes not
going to be able to have another child for a while.
One of the ways that human mothers respond to this demand is, unlike other primates,
they recruit a lot of help to raise children. Anthropologist Sarah Hrdy has pointed out that
most primate mothers jealously guard their offspring, and their offspring cling to them
constantly until theyre weaned, when a younger sibling takes their place. Humans are
part of a rare subset that shares childcare, handing off infants to others, even nursing the
infants of other mothers.
The pattern is alloparenting, or cooperative parenting by someone other than the two
biological parents; in humans across cultures, this includes especially, grandparents, the
mothers sisters, and even the childs older siblings. Hrdy says that to raise a human with
a human brain takes more than a mother; in a sense, we evolved to need more than the
usual primate single parent.
Hrdys research on primate childcare has helped us to see just how strategic reproduction
can be. She was the first to recognize in primates, that if a new male became dominant in
a group, he would kill all of the children of his deposed rival. The reason is that
infanticides brought all the mothers into fertility again. Hrdy also realized that females
had strategies of their own, such as mating with multiple males to confuse them about
which was the father; that way, a change in the dominant male was less likely to
infanticide.
Being a mother was just as strategic as fathering children, Hrdy explained. For example,

she argued that motherhood was not dominated by a maternal instinct but could be
much more strategic, involving a critical tradeoff between quantity of offspring and
quality of investment in each one.
Hrdy wasnt the first to argue that our infants needs were so great that it required
cooperative parenting, but she made it clear that this didnt necessarily mean the father
had to stick around, even though humans had evolved as cooperative breeders. Human
parenting could recruit sisters, grandmothers and even the childs siblings. This research
undermined the idea that the nuclear family mother and father with children only
was the natural evolutionary unit. The evidence was clear, not just from other primates,
but from a lot of non-Western societies, that childcare was often much more widely
distributed.
Hrdys theory of alloparenting also explained the strange, extended survival of
grandmothers, long after menopause, when you wouldnt think that their survival would
matter evolutionarily. In humans, staying alive to be a grandmother meant you could
pitch in to help make sure your genes survived in your grandchildren.
The bottom line is that we now realize that babies are not just packets of DNA. They are
an investment of parents, have their own evolutionary strategies for trying to stay alive,
and shape adults to a degree we may not have realized. Although many people believe
that unconditional maternal love is a kind of natural instinct, biological observations,
including comparisons with other species, remind us that reproduction is not so simple.

Long, slow sexual revolution

In 1672, after his brother fell off a horse and died, Moulay Ismail ibn Sharif became the
Sultan of Morocco. He was an extraordinarily ambitious man: under his rule, Morocco
conquered its neighbors, grew to an empire, fought off the Ottomans, and went through a
building spree to try to make the capital, Meknes, a rival to Versailles. Ibn Sharif also had
a prodigious capacity for cruelty: he legendarily ordered that the walls of Meknes be
decorated with the heads of 10,000 enemy soldiers.
But history will likely most remember Ibn Sharif because he fathered more than a
thousand children. Theres no definitive count because a European visitor recorded 867
in 1703, but Ibn Sharif was barely two-thirds through his reproductive campaign. He kept
at it for another 18 years. According to one estimate, he had sex twice a day, every day,
for his sixty-year reign, collecting approximately 4000 wives and concubines.
In contrast, history does not even record the name of the woman who gave birth to the
record for the most children; we know her only as the wife of Russian, Feodor Vassilyev.
During her lifetime, Mrs. Vassilyev gave birth to 67 children, including 16 sets of twins,
7 sets of triplets, and 4 sets of quadruplets.
From Ibn Sharifs selfish perspective, his approach to reproducing might make sense; you
might even think that evolution programmed Ibn Sharif to want as many children as

possible. But if you look at it from the point of view of the species as a whole, Mrs.
Vassilyev is more interesting.
Because, if Ibn Sharif had 4000 wives and concubines, and anything less than 2000
children, the fertility rate of women in Ibn Sharifs harem assuming that they didnt
have other children was more than 130 times lower than Mrs. Vassilyev. Tremendous
reproductive capacity got wasted in the palaces of Ibn Sharif. A population of Mrs.
Vassilyevs would have quickly swamped a population of Ibn Sharifs, outproducing them
spectacularly, if the behavior pattern was inherited (and Im not saying that it was, or is).
The point is this: from an evolutionary perspective, what might make sense for an
individual may not make sense for the good of a species.
But the other thing to notice is that, in both cases Mrs. Vassilyev and Ibn Sharif
their incredible fertility was not merely the result of desire or genetic selfishness, but of
the human capacity to cooperate and organise. Ibn Sharif attained his reproductive
success, not because of unbridled natural instinct and selfishness alone, but because of
quite sophisticated social and logistical tools: religion, government, military organization,
trade, large-scale networks. Likewise, Mrs. Vassilyev could only have been so fertile
given agriculture and a sedentary lifestyle. I daresay she probably didnt raise all 67
children by herself.
Both of them owe their fertility, not just to good genes, or to well-functioning
reproductive systems, but also to the extraordinary social skills of humans. They both
underline that reproduction isnt a solo operation, or even just a partnership; its a
collective project for humans that has shaped us.
Sometime in the past five million years, humans went through a profound sexual
revolution, a shift in reproductive strategies that didnt just change how we had children,
but made it possible to build this unprecedented human way of life.
Our current situation can make it hard to perceive the reproductive pressures our
ancestors faced. Like Ibn Sharif and Mrs. Vassilyev, we live with the social, cognitive,
and technological benefits that were won, in part, by this ancient reproductive revolution,
just like we live on a planet crowded by our reproductive success.
Today, if we think about human population at all, many of us worry about high rates of
reproduction, about the enormous number of us. But we, humans, were once small in
number, and may have nearly died out at one point. Our fertility was likely low
compared to today infant mortality high, and life hard; we had to cooperate to survive.
The success of our species has been won by such unusual social abilities that one of us
can even engage in the incredibly anti-social behavior of someone like Ibn Sharif.
Anthropologist Sarah Hrdy argues that cooperative breeding and alloparenting were key
to our ancestors success. Cooperative breeding means that a mother alone is not
responsible for raising an infant; other people, including perhaps her mate, a childs
grandmother, aunties, other adults, even older siblings what Hrdy calls alloparents or
other parents helped to raise the child.
According to Hrdy, cooperative breeding and alloparenting didnt just feed babies and

keep them safe. Cooperative breeding produced a human who was emotionally modern
long before we had anything like modern technology, probably even before our ancestors
developed sophisticated forms of cognition. We werent simply social like other apes; we
were hyper-social, overly empathetic, seeking approval from each other, cooperative
even when we must sacrifice for it... Basically, compared to other apes, were huge suckups. How could evolution produce such a bizarre, please-like-me ape?
Hrdy thinks that our pro-social nature is shaped by mating, in more ways than one. Were
made more social, not just by the way we reproduce, but by the difficulties of raising a
child, and being a child, trying to make sure the adults keep us alive.
Take, for example, our intelligence. In cognitive tests, chimpanzees, orangutans, and
human infants score quite close to each other in remembering where things are and the
basic way that objects work in the world; apes, including us, are good at problem solving
such as figuring out cause and effect.
However, human infants are much better at social cognition. Human infants read others
intentions better, figure out what people are thinking, and notice social cues like pointing
much better than other apes. Cognitive theorists call this theory of mind but we might
just call it social savvy. We seem to be primed to interact, even to manipulate and
deceive. Moreover, we create environments for our children where these skills are
nurtured; we interact with babies, play with them, talk to them and share with them, just
as we share them with other adults.
Getting passed around among different caregivers likely affected our intelligence. Most
great apes cling to their mothers until theyre weaned. Our ancestors likely got passed
from one adult or older sibling to the next, developing a rich social world and sensory
experiences. Instead of learning only from their mothers, out ancestors might interact
with and observe other adults. Cooperative parenting produced new lines of inheritance
just as it produced a hungry sponge of an infant.
Given this environment, infants social skills also improved: being a cute baby had a
survival payoff. Being able to read your caregivers emotionally and respond to them to
get what you wanted would have been a huge advantage.
Even things that may annoy us might have their roots in this social environment of shared
care. For example, some of our nearest ancestors, bonobos or what are sometimes called
pygmy chimpanzees, dont really cry; they just sort of whimper. It makes sense.
Producing a lot of noise if you live in a forest with predators can be a terrible survival
strategy. Besides, if youre a bonobo infant, youre basically hanging on your mother,
anyway, so you dont have to cry very loudly.
In contrast, our children can go off like a car alarm, sometimes crying for minimal
reason; thats a great strategy if youre fighting for attention in a group, or maybe need to
remind your older sibling to pay more attention to you. According to Hrdy, human infants
got better and better at manipulating their carers, provoking our sympathy they were
now in competition with each other for attention and support.

Confusion, with benefits

Getting help with childcare, passing your baby around, would not have been possible
unless a mother had a lot of confidence in the other people around her. If rival females
might kill an infant, or disinterested males neglect it, this kind of childcare wouldnt be
possible.
Our nearest relatives are not as socially cooperative as we are; chimpanzees have trouble
sharing food, even with their own offspring. Other monkeys who cooperatively parent
like humans, however, are similarly generous. Certainly, humans living together as
foragers share food, even with children who are unrelated to them, and even in times of
hardship.
Cooperative parenting allowed our children to grow up more slowly, taking the pressure
off us to mature as quickly as possible or risk dying before we had a chance to pass on
our genes. The cooperation also meant our mothers didnt have to delay going back into
reproduction. The window of development, when the brain was immature, started to grow
wider still, with consequences well talk about next week.
Without a change in the way that they reproduced, our ancestors wouldnt have
developed more advanced technology, larger and larger social groups, and our distinctly
human way of life. Our ancestors didnt know what they were doing this sexual
revolution was long and slow, so long and slow that our ancestors couldnt have observed
it but we inherit the legacy of this sexual revolution, including a set of important
confusions.
One of the most productive confusions is that we dont know when women are fertile.
I live with horses, and nothing could be clearer to them as when a mare is fertile. During
their reproductive season, when theyre in heat, all hell breaks loose. The stallions get
excited. Even the geldings get fired up, and theyve got nothing to work with.
And, if one of them, stallion or gelding, manages to get anywhere close to a mare whos
peaking, she will spray a pheromone-saturated fountain of urine in his direction, driving
him into a froth. The mares will sometimes even do it to each other, probably because we
keep them away from the stallions. If you actually let a stallion into a paddock with a
mare in heat, complete chaos: nothing else will matter to either of them except sorting out
whether or not they will mate.
Many animals are like this. Mating is a compulsory affair, even a kind of collective
temporary insanity.
In contrast, human females are pretty damn subtle and under-stated. We dont know,
without testing, when a woman is ovulating and most likely to conceive. We may notice
some subtle changes in womens behavior, and the behavior of those around them, when
women are at their peek of fertility we tend to get more flirty and think each other is
more attractive but this isnt conscious. Perhaps the most interesting thing about
humans is the degree to which were kind of oblivious to this really important basic fact
of reproduction.

We go through our daily activities unaware of whether the woman at the table next to us
in the cafe is reproductively primed or in an off-peak period, so to speak. I can teach a
classroom full of university students without anyone signaling that they have an egg
ready to fertilize.
As a species, human women have concealed estrus; they hide almost entirely their
fertility. Were not the only species that does this, but we conceal it to such a high degree,
especially given our intelligence, that we have to wonder how this is adaptive or at least
not maladaptive.
First, its obvious that human women dont need to signal estrus to get men interested in
them. Whereas males of other species can be pretty ambivalent about females, or
anyones company in general, without the right pheromones, human males dont have too
much trouble getting in the mood even without chemical encouragement. Same thing for
human women, too, of course; they dont have to be drunk on their own hormonal
processes to mate, and will even mate when they are not fertile.
This kind of readiness for sex, even when unlikely to result in pregnancy, could have
provided a social glue for increased pair bonding. If men didnt know when their mates
were fertile, then mating repeatedly might have been a more successful reproductive
strategy. Males also might have stuck around closely with females in order to try to make
sure that they didnt mate with a competitor when fertile. If that male was willing to help
with cooperative parenting, all the better, but it wasnt necessary as a mother had
alloparents.
Second, estrus has a cost: signaling fertility can be metabolically demanding. The most
obvious examples are female primates who develop extremely swollen, red buttocks
during estrus. This signal catches the males attention, but it sure takes metabolic energy,
makes mobility more difficult, and may even expose the animal to danger of infection or
injury.
Third, concealing estrus might have made it possible to conceal too the paternity of a
child. As Sarah Hrdy found with other primates, males are reluctant to mistreat children
that might be their own. If females were promiscuous, and males didnt know when they
were fertile, it might prevent infanticide; a guy wouldnt kill an infant if he thought he
might be the daddy.
Concealing estrus, however, may have more wide-ranging implications for humans
socially. In some species, estrus can be a rough time: sexual competition intensifies,
males fight to mate, and they may bully or isolate females in order to try to prevent
mating with other males. Its not just that the lady next to you has swollen buttocks or is
pumping out pheromones; its what those signals do to everyone else in your group.
In contrast, humans can gather in large numbers without anyone trying to mate with the
ovulating females well, you know what I mean. You have no idea in a group whos
fertile, so, if youre male, you wouldnt know who to fight for, even if you were so
inclined.
We can tell from the changes in our ancestors that the pressures of sexual selection did

not favor males built for fighting as in gorillas or males built for promiscuity as
in chimpanzees. In fact, the steady or decreasing dimorphism in hominins over the last
two or three million years suggests that direct competition hasnt been a strong force on
our development. Instead, theres strong evidence that our fragile, needy babies helped us
to learn to cooperate. And the way we took care of them drove their social intelligence
and skills to unprecedented levels.
Our babies helped to make us human.

Sexuality: it isnt natural

When Im teaching about human evolution, some student will almost always ask me what
I think is natural. Its a trick question, even if they dont mean it that way.
By natural, I think that they mean right or normal. Sometimes theyre worried about
something that theyre feeling, or just genuinely curious. They may have heard on the
news about how homosexuality or monogamy is supposedly unnatural because of
evolution. Or theyve read where an evolutionary psychologist said that rape is natural
or that, actually, humans are naturally mother-centred and poly-amorous, having many
sex partners, because bonobos or gorillas or chimpanzees are that way.
The problem is, if the antonym of natural is human-made, how can anything
associated with humans be natural? Im not just talking about sexuality, but about
emotions, maternal instinct, supposedly innate preferences for certain foods,
aggression, inequality...
I try to tell my students that I dont think weve been natural since we became human.
If we are the self-made species, why would our sexuality or our diet, or our social life,
or anything else be any different?
And if life is by nature diverse, if diversity and variation are, in fact, the precondition for
evolution, inherent in how every living thing is born, then why would any one way of
being male or female, married or sexual or anything else exist? Isnt our variation the
only insurance we have against changes in selective pressures?
And why should peeling away what makes humans human, complicated, get to what is
naturally human? Would peeling away language get to humans natural ways of
thinking? No, it would actually deprive us of one of the things that most makes us
distinctly human.
Students are never happy with my answers to this question. Theyre convinced that
theres a thing called human nature. So let me try a different tack to answer the
question.
Robert Sapolsky is a brilliant neuroscientist with a background in biological
anthropology. Bob studies, among other things, the way that brains and hormone systems
work in baboons. If youre interested, you should definitely check him out on YouTube

because hes a fantastic lecturer.

Sapolsky tells this amazing story about how he had been studying a troop of baboons for
years. They were sort of normal for baboons, with the dominant males behaving horribly,
battling each other for supremacy, beating up subordinates, attacking females, feuding
with other troops, and generally making life hell for the other baboons. Then, the
dominant male baboons took over a trash dump near a resort; they wouldnt let any of the
lower-ranking baboons near their tasty trash. But they caught bovine tuberculosis, so that
the most aggressive males were wiped out in a single epidemic.
Sapolsky was devastated; he thought his fieldsite might be ruined. He had been tracking
the hormone levels for markers of stress for years. The troop now had its gender ratio
totally screwed up, with twice as many females as males, and only the submissive males
from before the epidemic survived.
But heres where it gets interesting. The male behavior in the troop changed, and that
change persisted even when all the males in the group had been replaced more than seven
years later. Baboon males switch troops, so the troop is always recruiting. New male
recruits to the group didnt follow the nasty dominating male type, but all acted like
submissive males. New interaction patterns had been set up. Males were less aggressive;
they groomed the females kind of like the baboon equivalent of gossiping and
backrubs rolled into one.
And the baboons stress levels, measured by their hormones like cortisol, were lower. A
change in the groups interaction pattern, which remained stable even as the members
changed, caused a biological change in its members, including how they acted male and
female, how they raised children, how they mated. The aggressive pattern of male
dominance typical of baboons was missing.
The point is that, even with an animal much less complicated than us, expectations and
interactions have profound effects, not just on behavior, but even on biology. Our
relations are not just determined by the need to reproduce or hardwired into our instincts;
rather, we reproduce in a social environment, which shapes how we do it. Living things
adapt in their behavior, especially brainy living things, they dont just act out fixed
scripts.
Anthropologists like to draw a distinction between biological sex and cultural gender: sex
is the equipment youre born with; gender is the set of norms and expectations that you
think go with that equipment, but is really taught to you by your culture. The distinction
is messy, and some traits seem to blur the boundary, but its useful.
Being male or female in different places doesnt necessarily entail the same ideas about
what is masculine or feminine, and these ideas have consequences. Or as anthropologist
Agustn Fuentes puts it, with humans, who we think we are matters. It shapes us, our
behavior, our children, how we interact...
It turns out that in many primates sex is not only complicated, it is also frequently not
associated with reproduction. Many primate species (especially our closer relatives, the
apes) use sexual activity as part of their social repertoire. Social sex can make friends,

break friends, end fights, start fights, all in addition to the fact that sometimes it also ends
up in reproduction. Social sex is an important part of being a primate. It is no wonder that
primates have more sexually transmitted infections than most other mammalsthey have
more sex. (Agustn Fuentes, Why Is Sex So Complicated?)
Even in our societies, gender expectations have changed over recent generations. And
many of the traits that we think of as inherent the core of being either male or female
often turn out to be more flexible on closer inspection, even though they seem natural
to us.
Gender roles dont feel variable to us; they feel essential, unchangeable. The irony is that
our very adaptability makes a social pattern that is malleable feel like its inevitable and
unchangeable precisely because weve adapted so well to it. But that tells us more about
how we think and adapt, less about how we innately are or must be.

Sexuality 2: getting busy

You cant control what youre attracted to, no matter how you try to talk yourself out of
it. Man, would life be easier if you could!
Its one of the great tragedies, but also the really amazing things about humans we
seem drawn together, not just to reproduce, but even to try to build lives together, no
matter how much pain or discomfort it can cause.
Some evolutionary theorists argue that we are programmed with particular sexual
preferences or mating patterns by evolution in order to assure reproduction. They think
that men and women come into the world with a strong set of preferences for the best
possible partner who will help them pass on their genes.
I think that the preliminary data on this isnt that strong. We know that, around the world,
and even within our society, everyones preferences are not the same; some people are
attracted to members of the same sex, or even to no one at all.
Thank God! I say. If we all had the same preference, wed all be chasing after the same
person, and it would be even worse than it already is.
In fact, if we look closely at the evidence about human fertility, the most important thing
to point out is that sex and desire are simply not the limiting factors. The human desire
for sex is, to put it bluntly, excessive from an evolutionary perspective weve got
more than enough desire to get reproduction done.
Humans are interested in sex when theyre not fertile and have no chance of producing
viable offspring. Other animals generally mate when their bodies tell them that they can
conceive; humans do it all the time. And were interested in sex acts that have absolutely
zero chance of producing an offspring. Whenever someone says that sex should only be
for reproduction, I find it hard to take seriously. They do realize what theyre going to
have to give up? How little sex we need to get that job done?

Lets try to put some numbers to it: among our ancestors, women were likely to be
functionally non-fertile for about four years after giving birth, maybe even more. Our
female ancestors probably, like most hunters and gatherers, nursed frequently, day and
night, which suppressed fertility. Throw in nine months of pregnancy, an active lifestyle
and low body-fat percentage, and they probably were able to have a child every five
years.
We call that birth spacing, and its not out of line with other great apes: orangutans, for
example, have even longer birth spacing and even lower fertility.
Researchers estimate that having heterosexual, genital-genital intercourse randomly
during a womans cycle is a little more than 3.5% likely to result in a pregnancy. So, if
you assume a possibility of miscarriage, figure our female great-great grand ancestors
would have had about a 30-year reproductive life or so at most, youre looking at 5 to 7
pregnancies.
High end of the estimate: 7 pregnancies, 3% chance of getting pregnant... you need 250
or so heterosexual, reproduction-style sex acts over your entire lifetime to get close
to your maximum baby-making potential, if youre a woman living in those conditions.
You do the math, but its not a lot of sex. Less than 10 times a year, on average, with a lot
of variation and some really, really long dry patches.
I dont think humans have ever had that little sex. In fact, large-scale surveys in the US
suggest that even long-term married couples who are not trying to conceive have more
sex than this. The point is simply that human desire is not in short supply; its the babies
that are limiting our reproduction.
In other words, and to make this a lot simpler, some of us are having a lot of extra sex, a
lot of sex that we know damn well wont lead to reproduction, doing acts that cant lead
to reproduction, in combinations that will never produce children. And our ancestors
probably did exactly the same thing.
Consider one of our closest cousins, the bonobo or pygmy chimpanzee, an animal
beloved by anthropologists. Bonobos engage in all kinds of non-reproductive sexual
activity, in every conceivable combination female-male, female-female, male-male,
female-child, female-female-male-child... you get the idea. Whereas the common species
of chimpanzees builds social relations through grooming and aggression, bonobos seem
to use sexual interaction to settle fights, introduce themselves, calm their nerves, say hi
to a friend...
Were not bonobos, but bonobos show us that sex can be exapted or repurposed for all
sorts of other jobs: we use it to bond with our partner or close allies, to feel good, express
ourselves, ...
I think it makes more sense to realize that sexual behavior in humans, like in other
animals, is flexible and adapts to a situation.
We know that the pattern of mating in humans varies. For example, we find that a
number of societies practice polyandry one woman taking multiple husbands many

more than what Western anthropologists once thought. It seems to be a pattern that arises
when women are in short supply. And men often change their behavior if they are in
male-only situations, like at sea or in prison. Some societies even assume that a persons
sexuality will change over time, transitioning from one preference to another, or fear the
opposite sex. Fortunately for our continued survival, we have more than enough sex to
get the job done, no matter what crazy ideas we get about sex.
The thing is, this is not how we think about our sexuality or how it feels. Our preferences
feel to us to be innate, ingrained, and utterly unchangeable. Our preferences for one
type of relationship, for one way of being a man or a woman, for a particular sort of
physical interaction, for a partner who looks a particular way may feel like an essential
part of us because they are; sex is a crucial way that we define who we are, an activity
almost as important for humans as eating and drinking.
But we also confuse ourselves, mistaking the way we think things should be for the way
that they actually are. We think in norms, or ideals.
The norm of an institution like marriage is a complicated brew including religion,
tradition, economic relations, property and divorce law in a society. Its a bit of a fantasy,
but its a fantasy that we try to impose on a very messy reality.
For example, a classic survey of anthropological research found that 84% of all cultures
allow or encourage polygynous marriage: a man taking more than one wife. But even in
those societies that most strongly encourage it, the majority of men dont have
polygynous unions; for most, the cost is simply too high. Besides, you do the math; its
simply impossible unless a society figures out how to change the ratio of boy to girl
babies.
In summary, sexuality is not the same as sexual behavior. Once you have complicated
ideas about sex, ideas that shape your choices, youve got a really human way of being
sexual. And like the social apes that we are, we find it almost impossible not to worry
about what sex other people are having, how theyre doing it. You cant look to whats
natural to clear away the mess, because the mess is whats natural.

~~~~~~~~

James Trussell of Princeton University has pointed out that the probability of becoming
pregnant is a bit more complicated than my back-of-an-envelope estimate. He and his
colleagues, Germn Rodrguez and Charlotte Ellertson, in research on emergency
contraception, estimate that, if a woman had a 28-day cycle and randomly had intercourse
once in that cycle, her probability of pregnancy would be, on average, 3.82%. In order to
be 95% certain of achieving conception in a year (assuming this 3.82% rate), a woman
would have to have sex approximately 77 times. Of course, in many cases, a woman
would get pregnant on fewer attempts, but Trussell and colleagues suggest that my
estimate, although close, might be a bit on the low side.

References & resources

Key concepts
Sexual reproduction provides species with a number of benefits, including variation
through genetic recombination.
Sexual selection acts on the sex that is not the limiting factor on reproduction, and can
take the form of either direct competition or mate choice.
Human reproduction is shaped especially by the extremely altricial nature of human
infants; many social innovations seem to help our ancestors achieve shorter birth spacing
and higher-than-expected reproduction rates.
Sexual behaviour seems to be exapted into new forms and new functions, beyond simple
reproduction, in most species.
Human sexuality has been variable for a long time.

Resources
Wild Sex, series of web videos hosted by Dr. Carin Bondar on sex and reproduction
across species. (Beware: some content is not family friendly, but its all educational.)
(link)
Marriage and Other Arrangements. Open Anthropology, 1(1) April 2013. Collection of
classic anthropology papers on marriage, monogamy and related issues from the archives
of the American Anthropological Association. (link)

References
Bagemihl, B. 1999. Biological Exuberance: Animal Homosexuality and Natural
Diversity, New York: St. Martins Press.
Bailey, N.W., and Zuk, M. 2009. Same-sex sexual behavior and evolution. Trends in
Ecology and Evolution 24 (8): 439-446. (pdf available here)
Bogin, B., and Smith, B. H. 1996. Evolution of the Human Life Cycle. American Journal
of Human Biology 8(6): 703-716. (pdf available here)
Carothers B.J., and Reis H.T. 2013. Men and women are from Earth: examining the
latent structure of gender. Journal of Personal and Social Psychology 104(2): 385-407.
(download pdf from Harry Reis website)
Dunsworth, H.M., Warrener, A., Deacon, T., Ellison, P., and Pontzer H. 2012. Metabolic

hypothesis for human altriciality. Proceedings of the National Academy of Sciences.

doi:10.1073/pnas.1205282109
Fisher, H. E. 2004. Why We Love: The Nature and Chemistry of Romantic Love. New
York: Holt.
Fuentes, A. 2012. Race, Monogamy, and Other Lies They Told You: Busting Myths about
Human Nature. Berkeley: University of California Press.
Henrich, J., Boyd, R., and Richerson, P.J. 2012. The puzzle of monogamous marriage.
Philosophical Transactions of the Royal Society of London. Series B, Biological sciences
367(1589): 657-69. doi:10.1098/rstb.2011.0290
Hochberg, Z. 2012. Evo-Devo of Child Growth: Treatise on Child Growth and Human
Evolution. Wiley-Blackwell.
Hrdy, Sarah. 2009. Mothers and Others: The Evolutionary Origins of Mutual
Understanding. Cambridge: Harvard University Press.
Jolly, A. 1999. Lucys Legacy: Sex and Intelligence in Human Evolution. Cambridge,
MA: Harvard University Press.
Leavitt, G. C. 2005. Incest and Inbreeding Avoidance: A Critique of Darwinian Social
Science. Lewiston, NY: Edwin Mellen.
LeCroy, D., and Moller, P., eds. 2000. Evolutionary Perspectives on Human
Reproductive Behavior. New York: New York Academy of Sciences.
Martin, R. D. 2007. The Evolution of Human Reproduction: A Primatological
Perspective. Yearbook of Physical Anthropology 50:59-84.
Miller, G. 2001. The Mating Mind: How Sexual Choice Shaped the Evolution of Human
Nature. London: Vintage.
Ridley, M. 2003. The Red Queen: Sex and the Evolution of Human Nature. Harper
Perennial.
Ryan, C., and Jeth, C. 2011. Sex at Dawn: How We Mate, Why We Stray, and What It
Means for Modern Relationships. Harper Perennial; Reprint edition. (Sex at Dawn
website)
Smith, B. H., and Tompkins, R. L. 1995. Toward a Life History of the Hominidae.
Annual Review of Anthropology 24: 257-79.
Sun, J. X., Helgason, A., Masson, G., et al. 2012. A direct characterization of human
mutation based on microsatellites. Nature 44: 11611165. DOI: 10.1038/ng.2398
Sykes, B. 2003. Adams Curse: A Future Without Men. London: Bantam.
Trussell, J., Germn Rodrguez, G.,and Ellertson, C. New Estimates of the Effectiveness
of the Yuzpe Regimen of Emergency Contraception. Contraception 57: 363-369.

Links
Evolution, episode 5 (Why Sex?), part of a series by PBS (USA). (YouTube) (link)
Sex: An Unnatural History, episode 1, The Revolution (YouTube), SBS series, first
episode. (link)
Jason Antrosio, Mothers and Others, Testosterone Anthropology, Biocultural Synthesis,
Living Anthropologically. (link)
_____. Anthropology, Sex, Gender, Sexuality: Gender is a Social Construction, Living
Anthropologically. (link)
Rob Brooks, The Placenta and the Pace of Life, The Conversation. (link)
Kate Clancy, 5 Ways to Make Progress in Evolutionary Psychology: Smash, Not Match,
Stereotypes, Context and Variation, Scientific American. (link)
Patrick Clarkin, Part 11. Humans Are Blank-ogamous. Sexaptation: The Many Functions
of Sex, Patrick F. Clarkin, Ph.D. (link)
_____. Part 10: Humans are (Blank)-ogamous: Wondrously ComplexPaleo-Sex,
Patrick F. Clarkin, Ph.D. (Links to the previous NINE installments all available from the
most recent ones.) (link)
Greg Downey, The long, slow sexual revolution (part 1) with nsfw video,
Neuroanthropology (weblog), PLOS Blogs. (link)
_____. Sex on the brain & neuroanthropology on sex. Neuroanthropology.net (link)
Holly Dunsworth, How to Apply an Evolutionary Hypothesis about Gestation to Your
Pregnancy. Scientific American. (link)
Daniel Engber. Who's Your Daddy? The perils of personal genomics. Slate. (link)
Agustn Fuentes. Size Matters, or Does It? Busting Myths About Human Nature weblog
at Psychology Today. (link)
_____. Why Is Sex So Complicated? Busting Myths About Human Nature weblog at
Psychology Today. (link)
_____. Men and Women Are the Same Species! Busting Myths About Human Nature
weblog at Psychology Today. (link)
_____. On Marriage and Pair Bonds. Busting Myths About Human Nature weblog at
Psychology Today. (link)
David George Haskell, Natures Case for Same-Sex Marriage, The New York Times.
(link)

Sarah Blaffer Hrdy, How Humans Became Such Other-Regarding Apes, On the Human
(National Humanities Center). (link)
Eric Michael Johnson, Raising Darwins Consciousness: Sarah Blaffer Hrdy on the
Evolutionary Lessons of Motherhood, The Primate Diaries, Scientific American. (link)
_____. The Case of the Missing Polygamists, The Primate Diaries, Scientific American.
(link)
Rosemary Joyce, Coupling and Culture, Psychology Today. (link)
Brandon Keim, The Surprising Ways Humans Continue to Evolve, Wired. (link)
Barbara J. King, Flexible Gender Identities Confound Expectations Of A Male And
Female World, 13.7 Cosmos & Culture, NPR (USA). (link)
_____. Promiscuous Males And Choosy Females? Challenging A Classic Experiment,
13.7 Cosmos & Culture, NPR (USA). (link)
Graham Townsley, The Evolution of Motherhood (an interview with Sarah Hrdy), Nova
(PBS). (link)
Jeremy Yoder, The intelligent homosexual's guide to natural selection and evolution, with
a key to many complicating factors, Denim and Tweed. (link)
_____. Is epigenetics totally gay?, Nothing in Biology Makes Sense. (link)

~~~~~~~~

Brains
Among all our traits, the human brain stands out as peculiar. Although our intelligence
may appear to us to be an unqualified good an all-purpose adaptation the rarity of
extravagantly oversized brains makes us wonder how evolutionary pressures could
produce such an outlier, especially given its cost to us. Moreover, once our ancestors had
out-sized brains, how did this affect evolution?

What are brains for?

What are brains for? Why dont we ask the sea squirt.

The sea squirt is a simple animal. It starts life as a larva in the sea looking like a tadpole.
This phase of its life is brief, however; the larva searches for an ideal place to anchor
itself on the ocean floor and then fastens itself where it will spend the rest of its life,
never moving again.
The sea squirt eats by filtering water, catching suspended particles. Theyre
hermaphrodites theyve got both male and female reproductive systems. They
reproduce by squirting their sperm out and letting the currents carry it to the ... other...
lucky... sea squirt.
But whats really interesting about sea squirts is that they are chordates. That means they
have a basic nervous system, even a kind of eye in some species. While they scout for a
place to cement themselves to the ocean floor, they have a basal ganglion, a cluster of
nerves like a simple brain. Once theyre planted, however, they start their adult lives, and
digest their own brains and nervous systems. They no longer need a brain, so its the first
thing that they eat as adults.
The point is simple. No, its not that adults have digested their own brains...
We may be impressed by our cognitive abilities, by our logic, creativity and advanced
intellectual skills. We may think that brains are for creating language and symbols, for
imagining, building technology, or plotting deviously against each other. But thats not
why we have brains, from an evolutionary perspective. We know this because lots of
living things have brains that cant do these sorts of complicated cognitive tasks.
From an evolutionary perspective, we have a brain for two reasons, both linked to
survival. First, if you dont move, you dont need a brain. The sea squirt shows us that.
Brains are for interacting with the world. If you dont perceive the world, or need to react
to it in order to eat or mate or defend yourself no brain. Trees and plants dont need
brains because, even if they could perceive their surroundings, they couldnt do much
about it. And they get along perfectly well.
Second, brains regulate bodies. They help an organism to achieve homeostasis, or healthy
balance. Lots of living things do this without a brain, but a brain can really help,
especially if your needs are complicated and your system needs some executive control.
A brain is your bodys way of telling yourself, Im hungry; Im going to go eat, or Its
too hot here; lets go in the shade, and then doing something about it.
We are separated from other animals by what may appear to be a mental chasm, a huge
gap. We have a host of cognitive abilities language, abstract thought, powerful
abilities to learn, self awareness that other species seem to only possess minimally, if
at all. We may be more than 98% similar genetically to chimpanzees, but whatever is in
that 2% must be pretty important.
When we look more closely at our brains, what we find can be startling. Like our
forelimbs, our brains are also full of homologies in fact, we cannot find a part of the
human brain that is not present in other species. The size and shape and functioning have
changed, sure, as well talk about in the next lecture, but theres an enormous amount of
consistency across species.

Once upon a time, for example, brain researchers went looking for the part of the brain
that allowed us to speak. They assumed that, because only humans could do it, we must
have some special language part of the brain. But the two regions that are most associated
with language, Brocas and Wernickes areas, both have homologues in chimpanzee
brains. They just get used for other purposes.
The structure of nerve cells is so consistent across living things that we can take brain
tissue from one species and implant it in a totally different species. If we do it early
enough in development, the implants will get incorporated into the animals brain, what
we call a chimera.
As with so many other parts of our body, evolution has not invented something wholly
new but rather tinkered with existing structures to produce innovative capacities. If there
is a gulf separating our cognitive abilities, there is also a shared foundation of brain
building blocks.
For this reason, we can really learn a lot from studying the brains of other animals
because of what we share with them. We can also see the ways we differ, how parts get
and adapted, to help us to understand what allowed our ancestors to survive and become
the way we are today.
So, to start off with, we need to remember the lesson of the sea squirt: when we look
across all species, brains are for moving and interacting with the world, not for silently
reflecting or deep thinking. Because of this, humans are good at some tasks, lousy at
others, even though we might not realize it.
The philosopher Andy Clark has said that our brains are, good at Frisbee, bad at logic.
Its a great point. You may not realize it, especially because were told our brains are like
computers memory or logic machines but youre actually not really all that good at
remembering or at logic. One of the reasons that computers or smart phones are so useful
is that our brains dont work like they do your phone is probably much better at
remembering phone numbers than you are! Your computer is much better at logic and
calculation than you are.
In contrast, youre likely really good at perceptual and motor tasks, pattern matching and
face recognition, for example, and possibly even frisbee. These tasks turn out to be really,
really hard. Its very hard to get a computer to do them. Weve learned this, in part, from
robotics. It was relatively easy to build a computer that can beat a human at doing math
operations, but have you seen the robots that they have tried to build to play soccer? They
couldnt play for the local under-10s soccer team!

Brain functions, old & new

We have to realize that brains were built by evolution, not to think, but to survive in the
world. We tend to be good at tasks that mimic the sorts of challenges our ancestors faced,
or that we can transform into these kinds of jobs, finding a way to hack around our

cognitive limits.
My friend at Macquarie University, biologist Ken Cheng, teaches about simple animal
brains to make this point. Among other things, Ken studies ants. He likes to point out that
animals can sometimes do amazing things with very small brains. Some can even do
things that we cant do with much bigger brains, because of the way that evolution has
shaped their mental equipment.
Ken discusses Cataglyphus fortis. Cataglyphus is a type of ant that survives in the desert,
in temperatures up to 50 degrees (thats 122 Fahrenheit). They eat other insects that keel
over in the heat.
Whats amazing about Cataglyphus is that, when it goes wandering around on the hot
sand or in a saltpan, it must look to the ant like a totally featureless landscape; the ant
might as well be walking on the surface of the moon. The ant will wander this way and
that, weaving around, searching, with no landmarks on the ground to guide them back to
the nest. When it comes upon some food, like a fly thats died from sunstroke, it grabs the
food and rushes straight back to its nest. Even if its a football-field length away from
home, and been wandering for a while, Cataglyphus will barely take a stray step.
What Cataglyphus has done is an example of highly modular intelligence. Its brain is
incredibly good at remembering how to get home, remembering each step, adjusting for
the direction, and then adding up all the little steps to figure out the path straight back to
the nest. Its eyes perceive patterns of light in the sky that are invisible to us so that it
always knows which direction north, south, east and west its stepping. The
evolutionary pressures of living in the desert have shaped a brain that is really
specialized, but Cataglyphus isnt good at learning new tricks.
Some brain researchers argue that the human brain, too, is built up of modules, specialpurpose cognitive tools that do a very specific task. In fact some of human intelligence
may work this way; for example, we may have special purpose modules for essential
tasks, especially social jobs, like understanding what other people are doing, imitating
them, or detecting each others emotions.
One of the interesting things about humans, however, is that, even with our most
specialized cognitive tools, we can often do quite strange things and build up new skills
from old modules. We repurpose or exapt some of the most basic brain tools into new
uses; some cognitive theorists call this neural reuse or neural recycling.
For example, when we learn to read, we train parts of our brain that could not possibly
have evolved for that task. Reading is just too new for a specialized part of the brain to
have evolved. We know, instead, that parts of the brain used by animals to recognize
basic shapes get repurposed in people who learn to read to pick out letters quickly. This
kind of neural reuse is widespread in our brains, and it involves both some of the newest
zones of the brain and some of the most ancient.
One thing we have to realize is that evolution has achieved some cognitive advances by
adding functions onto some brain areas, and layering new levels of control on old
systems.

For example, some of the oldest, most important areas of the brain, shared by virtually all
animals with a nervous system, are right at the top of the spine, at the brain stem. The
brain stem controls essential functions breathing, swallowing, sleeping, pain
perception and it connects your brain to the peripheral system so that you can feel and
control your muscles. Its ancient, but its also absolutely critical. Damage a part of the
neocortex of your brain, the outer layer, and youll get some weird memory or function
losses; damage the brain stem and youll be lucky to be conscious or breathing.
These parts of the brain are highly conserved across different species; evolution doesnt
mess too much with the control system for breathing or making the heart tick.
However, in humans, we actually find that we can override some of these automatic
systems, or drive them with new inputs. For example, we can drive ourselves to panic so
badly that we have trouble breathing and our heart races as if we were under severe
threat, even when were just watching a horror movie, or remembering a stressful event,
or seeing something that tips off a phobia. Theres even amazing examples, like being
able to hold our breath and talk, where we override one of our most basic automatic
functions breathing because of the power of other parts of the brain. Well discuss
this stuff in the next lecture.
Its been really important in our evolutionary history that the perfectly sensible automatic
responses of the midbrain and brainstem can get over-ruled, and that these responses,
such as strong emotions, can get recruited for new uses. Our early ancestors who made
the move from vegetarians and scavengers to hunters eventually had to overrule the most
reasonable demand of the automatic system to run like mad when a large animal started
to run toward them. It took an enormous act of imagination and self control to see a
charging animal as a potential meal and keep steady.
We recruited our basic responses into a whole new cognitive world: we learned to fear or
love imaginary creatures, suffer pain from disappointment, and long for justice or other
abstractions, not just a meal, a mate, or a place to sleep. We havent evolved beyond
primitive impulses; if anything, weve given them countless new targets, everything
from lusting after new technology to feeling real emotions when daydreaming.
One of the most widespread myths about the brain is that we only use a small fraction of
it. Nothing could be further from the truth.
We are constantly using virtually all of our brain, even if were not consciously aware of
it. The vast majority of our brain functions are going on without us knowing, but theyre
keeping us alive. Your brain is running your body, adjusting your organs and hormone
levels, tinkering with your muscles and posture constantly, and, occasionally, pointing
stuff out to your attention, alerting your consciousness to check something out.
Your brain is doing what your ancestors brains always did; keeping you alive in more
ways than you even realize.

Bigger isnt always better

Size matters. When it comes to brains, theres no doubt about it.
And humans have very large brains. Although theres some variation, we average around
1200 or 1300 cubic centimeters or ccs; thats about 1.5 kilos, or three pounds.
Compared to our earliest known hominin ancestors and to the other great apes, our brains
are triple in size.
However, if brain size alone determined intelligence, wed expect elephants, dolphins,
and whales to be far smarter than we are. Elephant brains are about three times heavier
than ours; a sperm whales, about six times.
Researchers who study brains across different species talk about encephalisation.
Encephalisation is the mass of the brain compared to the body. The clearest pattern
emerges if you log the two weights, that is, dont treat them as a straight 1:1 relationship.
As an animal gets bigger, its brain gets bigger too, but more slowly. It makes sense if you
recognise that a lot of the brain is responsible for keeping the body running and
controlling organs. Just because an animal gets larger doesnt mean it gets another heart
or a couple more pair of lungs for the brain to look after.
Some of the smallest animals have brains that are very large as a percentage of their
body; for example, the brain of the pocket mouse is 10% of its total bodyweight. In
contrast, the sperm whale has a giant brain, tipping the scales at almost 8 kilos (about 17
lbs.), but since the whale can weigh over 40,000 kilos, or 45 tons, that 8 kilos is a bit less
impressive; its about .02% of its bodyweight.
If you plot these comparisons, you find that they tend to be consistent for different
branches of the evolutionary tree. Most vertebrates, like lizards and fish, are at one level
of encephalisation; mammals are on a slightly higher curve. With a mammal and a lizard
or a fish of the same weight, the mammal will have a heavier brain. Mammals seem to be
more encephalised, or brainier, kilo for kilo.
We know that predators tend to be brainier than their prey; prey that actively evades
predators brainier than prey that just hangs out and hides. And diet and parenting style
also correlate with difference in encephalisation: fruit eaters and careful-parenting species
often have larger-than-predicted brains. Some of the most overly-well-endowed animals
are those with complex social lives.
Primates have an even higher encephelisation ratio than other mammals. But humans
stand out as an outlier, even among the primates. How much of an outlier depends upon
which species you compare us to. Even next to other really brainy great apes, weve got
two or three times more brain than youd expect given our weight.
Go back three million years ago, to the remains of Australopithecus afarensis, one of our
hominin ancestor, and you find a skull that still looks about the right size for a great ape:
smart, certainly, but not much more than a chimpanzee, probably. Were pretty confident
Australopithecenes were clever enough to use tools, but so are chimps, and even gorillas.

If you look at Lucy, the most complete Australopithicus skeleton we have, the most
dramatic differences between her and a chimpanzee are from the neck down, not up in
her skull. She was probably walking like a modern human, but she was still like our ape
cousins above her neck. Our hominin ancestors learned to walk upright before they really
became strange neurologically.
If we follow our ancestors from Lucy forward in time, towards modern humans, we see a
really startling changes. The brain size of our ancestors didnt grow steadily; it spiked,
apparently as a result of pretty intense selective pressure, or a relaxation of a constraint,
or some combination. Brains got big fast, at least from an evolutionary perspective.
The real increase in size, the break that created a brain different to other apes, starts a bit
before 2 million years ago. The remains we find from this time, usually referred to as
Homo habilis, have brain cases that are about 30-40% larger than earlier hominds, up to
about 600 ccs, at first. But over the next 300,000 or so years, things start to change
quickly.
By about 1.7 million years ago, we find evidence of a species we call either Homo
erectus or Homo ergaster, with brains as large as 900 cubic centimeters. Theres a lot of
variation around the average, a lot more than you find today, but the trend upward is clear
and quick. Youre probably thinking that 2 million years isnt so fast, but this is a huge
change for this kind of time frame.
Before I go on, anthropologists argue about naming and whether or not some of these are
separate species, and what to call them, but I wont get into that. I dont want us to get
bogged down in a lot of names, dates, and debates. If youre interested in this, and want
more, Ill post some great links for you to explore. The important thing for our discussion
of the brain is the trend over time and to understand why it might have happened and
how.
Some of this growth in brain size appears to be a result of an overall increase in body
size; Homo habilis was a bit bigger than Australopithecenes, and Homo erectus was even
bigger: about as tall as a modern person. But not all of the increase in brain size can be
explained as just the scaling of the brain along with the body.
Back 1.5 million years ago, Homo erectus still had some ape-like traits in its skull,
although its lower body looked modern, only tougher and more robust. They had smaller
teeth closer to ours flatter faces, less sexual dimorphism, longer legs and bodies
better built for walking, even running. But they also still had heavy ridges over their eyes,
no chin or forehead, and their brains were still shaped more like an apes than ours.
Homo erectus was incredibly successful. The species lasted about a million and a half
years, spreading out of Africa into Asia, and maybe even into Europe. They may have
been the first hunter-gatherers in our lineage; in fact, following game may have been how
our ancestors first go out of Africa.
To spread like this, they had to adapt to unfamiliar environments, to new food sources,
predators they had never seen before, a whole raft of new challenges. They had become
an invasive ape, not one tied to a particular ecological niche, and their brains obviously

had a lot to do with it. In some pockets in Asia, Homo erectus lasted until at least 300,000
years ago, with their brains getting even a bit larger towards the end.

The benefits of immaturity

In our ancestors tools, we can really see how the changing brain mattered. Up until about
1.8 million years ago, all the tools are pretty simple. We call them Olduwan or pebble
tools. They basically look like a river cobble with one side fashioned into a sharp edge by
some rough hammering. Theyre still better than anything any other living animal could
do, but if youre not an expert, you might walk by them without even realizing they were
tools.
About 1.8 million years ago, someone either Homo habilis or Homo erectus probably
started to fashion what we call Acheulean tools. We call them hand axes although
they were likely used for butchering and a whole host of other tasks, not so much cutting
down trees.
Theres two really interesting things about Acheulean tools: their regularity, and the fact
that they didnt change a lot over a very long time. Its hard to know for sure about the
cognitive abilities of the humans that made these tools its hard enough even to a
living things cognitive ability when theyre alive but they do offer us some clues.
Their regularity is remarkable because it shows that whoever made them was imposing a
mental model, an image of what a tool should look like, on the raw material, generation
after generation, for about a million years. Whoever made them didnt just whack
together something that would work, or do the minimum necessary. One of our ancestors
went out and found a proper stone for the job and imposed a design, one that was more
symmetrical and standardized than it needed to be, one that had been passed down to
them.
I think this shows the way increasing cognitive powers were giving our ancestors a more
sophisticated way of conceptualizing tools, learning and teaching, and preserving
knowledge.
But, and this is key, too, Acheulean tools show us the limits of this stage in cognitive
development. Although you can say lots of nice things about Homo erectus, they were
not innovative. They got something that worked and stuck with it. You could say that
they were good at passing on information and sticking to a model, but got mostly failing
marks at improving designs or coming up with new innovations.
Creativity and invention may have taken a bigger brain. But remember: bigger alone isnt
necessarily better in hominids either. Were pretty sure that Neandertals, who lived from
about a half-million years ago until about 30,000 years ago, and our own ancestors had
larger brains than we have, as much as 20% larger. The average brain size of a human has
actually been decreasing over the last 80,000 years or so, even more rapidly in the past
20,000 years.

Organization of the brain matters, just as size does. When brains get larger over
evolutionary time, every part does not get larger at the same rate; its kind of like the
relationship between brain and body weights. Some parts grow disproportionately.
In the case of primates and humans, especially, the brain structures that have grown
disproportionately are the outer layers, or the cortex and neocortex. The parts that control
the automatic functions didnt increase as much. I talked about this in the last lecture,
how we gain the ability to override perfectly sensible instincts, not because we get a
whole new brain area, but because the balance of power shifts when some parts grow
disproportionately strong.
Over-riding instincts may not sound like a big deal, but not always acting on your first
impulse creates a lot of space for flexibility in behavior. If every time you got frightened,
you could only get enraged or panic fight or flight youd never learn to ride a bike,
or talk in front of a crowd, and youd be pretty awful to live with.
These outer areas in the neocortex also do a lot of work making long-range connections
in the brain, helping us to build new cognitive associations. The point is not that the
neocortex makes us human; all mammals have one. Instead, the disproportionate growth
of the neocortex has made it possible to recruit other parts of the brain more easily, do
new things with our modules, control these responses better, and even use them to do
new tasks.
For example, we have parts of the brain that interpret what we see. They turn a whole
diffuse set of visual impressions into a kind of coherent image, and then recognize what
an object is. In humans, these visual parts of the brain can be driven, not just by the eyes
and sensory experience, but by will and imagination. You can run them backwards, if you
will, or top-down. You can think about what an imaginary creature might look like. You
can run different scenarios imaginatively in your brain, without having to act them out,
prior to deciding what you will do. The increased cortex makes our brains become less
automatic, more autonomous and capable of doing what we want them to, even if thats
really foolish and counter-productive.
How did our brains get this way? The key is by changing the developmental programs
and the way that the brain runs. That is, as weve been discussing, you dont need wholly
new genes or brain parts; you need to change the pattern of growth, the connections that
emerge, and how the brain runs. Thats affected by gene regulation as the individual
develops, first as an embryo, and later as a child and adult. And the developmental
trajectory of human brains is really unusual.
In most mammals, brain growth is over fairly early. In humans, the period of early, faster
brain growth appears to be prolonged. We stay immature longer, which gives our brain,
especially the parts that are last to develop, like the neocortex, more time to grow.
In fact, if you look at them closely, human skulls more closely resemble immature
chimpanzees than adult chimps. If you were a chimp looking at one of us, chimp-you
might say, Whats wrong with these humans? No fangs, no snout, flat face, big eyes,
sparse coat... They look like babies! When are those humans going to grow up? Humans

even mature more slowly sexually, and our teeth erupt later.
We call this pattern neotony, the retention of juvenile traits for an extended period of
time, even into adulthood. We can see something similar in domesticated dogs, who in
many ways are neotonous wolves who hang onto their juvenile traits.
Neotony hasnt just affected our faces; its also really affected our brains. When humans
are born, their brains are only about 25% of their adult weight. Chimps are born with
brains about 40% of their adult size. And macaques, who are still pretty brainy compared
to other mammals, theyve got 65% of their adult cranial capacity at birth. When were
born, chimpanzees and humans have very similar encephalization rates. Humans brains,
however, continue growing at accelerated pace, where chimpanzees brains slow down.
This long, slow maturation process means that our brains do much of their developing
outside the womb, bathed in the stimulation of adult interaction, full sensory experience,
and opportunities to learn. We retain youthful brain plasticity, the ability to adapt to new
challenges and learn skills more rapidly, for longer than other primates, and for a lot
longer than other mammals.
So the next time someone calls you immature, you can just smile and say. Yep, thats
the way evolution made me. Its one of the things that makes us human.

Expensive tissue
So, if having a big brain is such a good thing, if staying immature longer is so great for
learning, why arent all animals like us? Why doesnt every animal have a big brain and
take a slow train to adulthood?
We, humans, tend to be very impressed by our brains, thinking that theyre great for
every occasion. Whenever I ask students about what sort of change in an organism would
be a good adaptation for some selective pressure, virtually every time, a student will say,
Oh, a bigger brain will solve that problem. It doesnt matter what the problem is.
If large brains were so useful, such a no-brainer for evolution, wed expect heaps more
animals to have them. But they dont. Brains the size of ours are rare, even a freakish
adaptation, a strange outlier, and one that certainly does not work out for every animal.
In fact, the general rule of brain growth in evolution appears to be that an animal gets just
as much as it needs and no more.
The first reason is that brains are expensive tissue. You pay a high price for a big brain,
a price in calories. Your brain makes up only about 2% of your bodyweight, but when
youre sitting still, its sucking up at least 20% of your bodys energy. It takes a lot of
juice to power neurons.
As infants, its even worse. Towards the end of pregnancy, an embryos brain is eating up
60% of the calories going into the infant, not just to power the neurons, but also building

more and more of them. A shortage of calories during this rapid development can lead to
permanent neurological problems.
So we might say that the metabolic price of brains is high; it acts as a constraint on
growth in evolution. If a species cant afford a big brain, it wont get it, no matter how
nice it might be to have.
For example, gorillas appear to have gotten larger since we shared an ancestor with them
without their brains keeping pace. Their bodies have grown, possibly due to sexual
competition, and their brains have lagged behind. The problem is, a gorilla already has to
eat about 80% of the daylight hours to get enough calories to survive, and they dont even
move a lot maybe only a kilometer or less each day.
A gorilla would have to get another 700 calories or more a day to get enough energy to
support a human-sized brain. There simply arent enough hours in the day to chew up that
many leaves.
One additional problem for our ancestors in the past few million years, though, is that
usually when an animal needs more calories, it also develops a better set of equipment to
get that energy. If a species grows a larger body through selection to defend itself or to
fight for mates, it usually gets a larger jaw or bigger gut to pull in enough calories to feed
that bigger body.
Our ancestors, however, went through the opposite process: in Homo erectus and even
more so in Homo sapiens, our ancestors teeth got smaller and smaller, their jaws shrank.
We can see this clearly in their skulls: the way that the cheekbones grew smaller and the
sagittal crest, the ridge on the top of the skull, gradually disappeared as the jaw muscles
shrank. As the brain case was getting higher, rounder, larger marks of a bigger brain
the indicators of big jaws and powerful teeth were disappearing some theorists
even say that the brain case could only grow because the jaw shrank.
The growing brain, the shrinking jaw and a small gut made the food problem big, really
big. Primatologist Richard Wrangham has argued that you simply cant grow a humansized brain on the diet of a gorilla or chimpanzee; our ancestors had to change what they
ate. Gorillas, for example, are mostly raw food vegans; they eat leaves, fruit, and no
doubt the occasional insect. Chimpanzees prefer fruit, but arent as picky, but they still
only get occasional snacks of meat, perhaps less than 5%, maybe as little as 2%, of their
calories.
Wrangham, whos studied the evolution of diet, argues that, if our ancestors had stuck
with a chimpanzee-style diet, they would have had to eat 5 kilos of raw plant food each
day; thats about 11 pounds of fruit and veg. Theyd have to chew 6 hours a day,
minimum, even before they got to modern brain sizes.
In contrast to the other apes, humans are truly omnivorous. We eat a really wide variety
of food. Animal protein meat makes up anywhere from a quarter to nearly 99% of
the calorie intake of the foraging peoples studied by anthropologists around the world.
Why does this matter? Meat has more energy per kilo. To get the same calories you

would get from eating a kilo of lean game meat like kangaroo, you would have to eat
about twice as much fruit like grapes. And youd have to eat about 7 kilos of a leafy
green like kale to get the same energy.
We suspect that two million years ago, Homo erectus, already bigger brained but not yet
human, began to have an increasingly modern, varied diet. Big brains required rich food
if you didnt want to sit there eating all day. Homo erectus likely started to eat a lot more
meat, maybe just scavenged at first, but later hunted. Nothing about our bodies looks like
a straight-up carnivore; our guts are way too long and our stomach not nearly acidic
enough. Our ancestors had gone from being herbivores gradually to becoming high-yield
omnivores; they were foraging widely, and may have even cooked their food.
Cooking makes food easier to eat and digest. Cooking concentrates the calories and cuts
down the effort to chew, outsourcing the food processing duties from our jaws to our
technology, hands, or fire.
But it wasnt just meat or cooking alone that made the difference; our ancestors also got
better and better at spotting high-energy foods of all sorts, and then pre-processing them
before eating. They were substituting wide-ranging, high quality foraging walking
around finding the most high-calorie meals for short-range, low-quality grazing
like the gorilla, just sitting their eating whats around you. This involved learning how to
get energy-dense treats like honey, shellfish, more nuts, eggs and even stealing caches of
food left by other animals.
Our ancestors eventually, later, learned when seasonal sources of food were available,
like stands of fruit, migrating salmon, or calving animals. They had to devise new ways
to exploit these, especially as they spread into unfamiliar regions, and teach their young
to do the same. That is, their brains were hungry, demanding, but their increased
intelligence opened up ways to get foods densely-packed with the calories they craved.
By the time the first modern Homo sapiens show up a bit less than 200,000 years ago, we
see the beginning of a broad spectrum approach to diet. No longer strict vegetarians, no
longer big game hunting specialists, they were building versatile tools that suggest they
were learning to fish and trap, developing projectiles like spear-throwers and eventually
bows and arrows, sewing, trading over longer and longer distances...
I always warn my students who are interested in a Paleolithic diet that our ancestors
menus probably would have turned us off that is, unless you really like insects,
worms, small raw vertebrates like lizards, and basically eating anything and everything
edible that you can lay your hands on. Our ancestors probably were not terribly picky.
We know that most foragers studied by anthropologists actually eat a more varied diet
than most of us do, eating scores of animal protein types, a lot of them small animals, for
example. Bon apptit!

Even brains have limits

Bipedalism helped make this diverse, high-yield diet possible. Walking on two legs is
more efficient over long distances than getting about on all four: 35% more efficient
according to some research. We spend less energy moving around, going further on the
same tank of fuel. If youre living where food is scarce and widely spread, that extra
range might be the difference between getting something to eat and going hungry.
In addition, walking on two legs freed up our ancestors arms to carry food. They could
have ranged far and carried what they found back to a home base to share. Chimpanzees
and gorillas dont have to travel far each day to find enough to eat, and theyre not good
at sharing; our ancestors likely worked together better.
It wasnt all easy going, however. Scarcity would have stalked our ancestors constantly.
Especially in the period from around 800,000 to 200,000 years ago, a time when some
paleoanthropologists think that the human brain might have gotten larger more quickly,
they would have faced some serious climate fluctuations. They would have to endure
waves of cold and dry ice ages and wetter, warmer periods.
For the most adventurous bands of hominins, those families who moved to the fringe of
their habitable range, shifts in the climate ramped up the selective pressures. When we
look at our bodies, we see not just the effects of plenty, but also hunger shaping our
ancestors, such as our uncanny and frustrating ability to lay in a store of fat whenever we
get a chance. Dietary versatility also may have been one way that they survived repeated
waves of starvation.
Brains pose other challenges, however, not just diet. A large brain cranks out heat. Any
time living tissue consumes calories, it also warms up, so we know from the brains
appetite for energy that its also going to have had a cooling problem. In fact, one of the
greatest dangers of heat exhaustion and sunstroke is that your brain can roast, potentially
killing you. Your brain is the weak link in your bodys thermal system.
Our ancestors bodies adapted to deal with heat in a number of ways: they lost a lot of
their body hair, multiplied the number of sweat glands, and even rerouted blood into the
skull. The blood bathes our brains and draws off heat, pumping it throughout the rest of
the body, like a kind of radiator system.
Being bipedal also helped our ancestors to keep cooler. Standing up puts more skin into
contact with a breeze, and cuts down on how much sunlight hits a body when the sun is
high. You may have experienced this if youve ever been lying on the beach, getting
really, really hot, but stood up, and found that you were suddenly much more
comfortable. And there was a breeze you didnt even know about! The air moves more a
meter or two off the ground, and youre much less of a solar collector when youre
standing up the same factors may have helped our ancestors to overcome the
constraint posed by brain heating.
One of the more controversial theories, but one that I quite like, is that this advanced
cooling system didnt just keep us alive, it helped us to go from long-distance walking to

running. According to anthropologist Daniel Lieberman, we didnt just evolve to walk;

our bodies are well built from running long distances without having to stop. Other
animals run much faster, but they have to stop periodically or they risk over-heating.
In contrast, humans can run and run and run without stopping, even in the heat. Our
ancestors might have developed the capacity to be persistence hunters, literally running
prey to death. Some hunting people can still do it. The trick isnt to go fast, its just to
prevent your prey from resting and cooling down. Eventually, if the animal cant stop
long enough to bring its temperature back down, it will simply pass out from heat stroke.
Our brains, however, arent just a challenge because of the heat and the need for energy.
The slow development of our brains, although it makes more learning possible, also
imposes its own costs. A human baby is a bit like an extra-uterine fetus; even basic motor
control and the senses are undeveloped. Because our brains have only achieved about a
quarter of their final size when were born, our nervous systems are especially
underprepared for life outside the womb. And that slow maturation process, although it
creates opportunities to learn, places an extra, long-term burden on the adults who must
care for the baby.
The pattern of growth means postponing sexual maturation, increasing the risk that the
child dies before passing on its genes. If our ancestors mortality was higher, the risk
would have been too great, and the slow growth trajectory would not have been possible.
Some researchers even think that this slow development makes humans more susceptible
to brain disorders, especially those that dont seem to show up in other animals.
Finally, big brains also have certain inherent engineering problems that come from the
way that theyve evolved. The signaling mechanism we use all animals with brains
use is about 600 million years old, and may have evolved first in a kind of jellyfish
that didnt even have a brain. The bigger the brain gets, the harder and slower it is to
move messages around. The more neurons you get what we call gray matter the
more you need connecting material, or white matter. The connections actually increase
more quickly than the number of neurons because theres a lot more connections needed
for each extra bit of neurons.
In humans, the number of connections hasnt kept pace with the number of neurons, so
our brains are less densely-connected, in some ways, than simpler brains. This creates
some quirks for the human brain; lower communication density means that local areas of
the cortex can specialize more, which may create more cortical zones, so that we can
learn more skills, more complicated behaviours. Weve also adapted by splitting up our
brain functions even more between the two sides of the brain, the right and the left.
The bottom line is that, like every other part of the body, the brain has limits and imposes
costs. We may tend to look at em and think, Man, our brains are great, but theyve also
been a real burden. Without our technology, our intelligence, our sophisticated ways of
getting food, and our ability to cooperate, we wouldnt be able to afford to pay the
evolutionary bill on these brains. The price would simply be too high. Thats the lesson
other animals, with their smaller, less costly brains teach us. Whatevers most unusual
about us, in some sense, needs the most explanation.

Brains change everything

Brains change everything. They affected the selective pressures our ancestors faced, our
patterns of inheritance, even our likelihood of surviving into the future.
But one of the most curious things about our big brains is that, although our ancestors
looked very modern in the brain case as much as 170,000 years ago, it really still took
them a while to start acting fully modern. Finally, about 50 to maybe as long ago as
70,000 years, they came out of Africa, and by 30,000 years ago, they were already
surviving on the cold coasts of the Arctic Ocean. How did they get so smart, but also,
why did it take so long, if the brains were just as big 100,000 years earlier?
To really understand how our brains affect us, we also have to think about why we got
them. Their origin helps us to understand how bigger brains eventually, eventually, lead
to sophisticated, modern lifestyles.
There are two basic types of theories that explain how we got such large brains:
environmental and social. Theres variations in each, even combinations, but the question
is really what role do we think external factors or selective pressures had, and to what
degree internal factors, competition among humans, drove changes in the brain.
Environmental theories often follow along the lines that weve already been discussing
about the brain and diet. According to these theories, a change in brain size allowed
human ancestors to change their foraging behavior, or shifted the relationship between
our ancestors and the environment, like the way that humans dealt with predators or
climate change or some other external factor.
The improved diet, for example, had a kind of cyclical effect; it demanded greater
intelligence, memory, and problem solving, but it also provided the calories that we
needed to feed a bigger brain. Throw in scavenging meat, and eventually hunting, and
invading new ecological niches, tool use, cooperation, and a whole range of cognitive
skills, and youve got a big payoff, a complete shift in how humans relate to external
selective pressures. These are all environmental explanations.
I dont think theres any way to dispute the importance of external relationships in brain
evolution diet, self-defense, expanded range, resilience against climate change.
However, the ecological model alone doesnt seem to explain the large brains: if
ecological forces, alone, drove brain evolution, youd expect that other animals that went
through the same sorts of processes might also wind up with big brains.
There were lots of other animals around in Africa going through the same changes, even
other apes, but they didnt get this spike in brain growth that weve seen in humans in the
last 2 million or so years. Also, our ancestors had the big brains before they really had the
complex foraging behaviors that eventually allowed them to spread throughout six of the
seven continents.
Increasingly, many evolutionary theorists think that we cant really understand that spurt

in brain growth or the slower effect on our ancestors behaviour unless we focus on social
forces. Social forces are forms of intraspecific competition, that is, human against human,
not just human against environment. Our brains got bigger, not because of the cognitive
demands of our diet or disease or climate or predators, according to social explanations,
but because we used our brains to compete with each other.
One of the earliest versions of this theory was known as the Machiavellian intelligence
hypothesis, after the political philosopher Nicola Machiavelli. Machiavelli wrote a book
called The Prince in the sixteenth century that offered cold, ruthless, calculating advice to
a political leader, how to use cunning to stay in control and expand his power, with every
strategy at his disposal.
Primatologist Frans de Waal, who studied chimpanzees, first pointed out that their social
lives demanded strategic skills, alliance building, conflict negotiation, and deception. De
Waal wrote that chimpanzees needed Machiavellian cunning, just like humans.
Its not just that primates lived in groups; ants lived in large groups too, and they didnt
get particularly intelligent. De Waal and other theorists recognized that primates scheme
and cooperate. The idea is that the complex and shifting nature of primate daily life, the
fluid structure of rivalries and changing alliances, taxed our ancestors intelligence. As
hominins or other apes grew smarter, they became more devious in everyday politics.
We now generally refer to this as social intelligence theory, especially as cooperation is
just as important as deception.
Social intelligence theory suggests that human brain evolution has been driven by a
cognitive arms race among our ancestors, especially most recently, as the most socially
adept out-performed those who couldnt keep up. Brain growth was driven by social
cognition.. If you want to look at it this way, our obsessive interest in each others
business, gossip and in-group maneuvering has a long evolutionary history; it may have
forced us to evolve. Next time you see a gossip mag, realize that this may be one of the
reasons that weve gotten as smart as we have weird thought.
The advantage of the social intelligence theory of brain growth is that, because its an
intraspecific form of competition, or one amongst the members of a species, any
individuals gain becomes a challenge to others. This kind of internal evolutionary arms
race can drive rapid change because it keeps cranking up the pressure with each
improvement.
If, at the same time, evolution loosened a constraint on a trait thats under this kind of
pressure like fixing limits on a big brain from diet and heat you can get a pretty
intense skewing of the trait through directional selection. This is the kind of pressure and
release from constraint that together might produce a big spike in a metabolically
expensive trait.
Relaxed limit, plus internal competition, plus big external pay-off can make for a real
evolutionary upheaval.
Another possibility that some theorists have raised as an explanation for our brain is
sexual selection: could women have preferred more intelligent mates, and their

preference driven our ancestors cognitive explosion? Well, its possible, and the
argument is not really opposed to the social intelligence thesis. The two could have
worked together.
But what would we expect if sexual competition among men was the dynamic driving
our increasing intelligence? Well, we would probably expect sexual dimorphism; mens
brains could get much better developed than womens, like a peacocks tail or a
silverback gorillas enormous muscles.
Thats not really what we find, though. Men and women are quite close in brain size,
when you take into account the difference in their body size. There are subtle differences
between mens and womens brains, but nowhere near what youd expect in a trait driven
by mate choice, especially a trait thats undergoing such intense selective pressure that
its changed radically in a few million years.
But the reason that I think its still worth considering sexual selection is this: ask
yourself, how would one of our female ancestors know which male was more intelligent?
She would have to be intelligent too, especially if some of the less-bright hominid males
were trying to fake it. If some of our male ancestors were trying to fake being brighter
than they really were, youd have created a selective advantage for smarter women.
If thats the case, you might wind up with symmetrical sexual selection, mate choice
driving both the chosen and the chooser to evolve more and more intelligence, putting
pressure on both men and women.
I suspect that these forces were working in conjunction: our ancestors increasingly
sophisticated strategies for getting and preparing food, changing ecology and technology,
social competition, alliances and deception, and even sexual selection for intelligence,
among men and women.

Ratcheting up intelligence
Why did out ancestors get bigger so much faster though, if all these selective forces
were there before?
One possibility is that about two million years ago, there was a real shift in the way that
these pressures interacted which caused the spike in growth. The idea is that, at a certain
point, our ancestors became so capable of getting food, protecting themselves from
predators, and shielding themselves from the environment that they achieved a kind of
ecological dominance. Once external selective forces were relaxed, sexual selection and
social competition could exercise greater influence on brain development.
The problem for our ancestors is that, with ecological dominance, the population of
humans started to increase. At first the excess population spread to new territory,
especially because meat-eaters need a bigger range than vegetarians. But, eventually, our
ancestors own success started to bite; the crowding ramped up pressure on each other.

First, it drove them out of Africa, as they followed the animals that they hunted. Later on,
much later on about 15,000 years ago the increased population would start to really
prove a problem, when our ancestors suffered through another brief ice age that dried out
the planet. This made it harder and harder to find the foods that they had learned to eat.
Our ancestors success, their growing numbers, would eventually lead to the slow dawn
of agriculture, then to the domestication of animals.
Most anthropologists dont think that our ancestors figured out agriculture because they
just thought it was a great idea. We think they were forced into it. Their own success at
getting food and growing their numbers, even setting up permanent places to live, would
become a burden when the climate changed. During one of these bottlenecks, some of
them would start to figure out how to help the environment to provide them with more
food.
In the Middle East, and elsewhere, our human ancestors started to learn how to
manipulate that environment, so that it provided more food. They started to change the
way water flowed, used fire to shift the types of plants in the forest, hunted more
carefully and entered a new kind of relationship with nature, where nature itself was
changed. We dont know exactly when it started, and it probably wasnt totally conscious
when it did start, but when humans started to modify the environment they accelerated
the processes of niche creation.
But that social crowding may have also produced a new kind of relation, an
intensification of human communication. We dont know when language started, but
once humans had it, they began to pass on to their children how they saw the world in a
new and more powerful way.
Language demonstrates clearly that the accomplishments of human intelligence are not
simply the product of the human brain alone, in isolation. No one invents his or her own
language. We are born immersed in an environment in which language is a constant
presence, and we gradually take it on board. Adults help us along the way, simplifying
how they talk to us, pointing things out, correcting us, providing role models of how to
use language.
Our brains, including those hyper-social instincts, like paying super close attention to
other people, make us really good at imitating compared to other primates, but our
environments help us along. If you show a child and a chimpanzee how to use a tool to
do something like get a treat, they can both learn how to use the tool.
But if you show them both how to use it with a strange and inefficient technique, the
chimp will probably just play with the tool until it finds a way most likely a better way
to get what it wants. The human child, on the other hand, will copy your technique,
usually with excellent fidelity, even though your technique is lousy. Human social skills
are stronger than our own ability to problem solve as children.
In the process of social learning, then, you inherent with language a way of seeing the
world, with all sorts of cognitive skills and information embedded in it, but also biases
and blindspots. We benefit from the accumulated knowledge and skill of our ancestors,

just as we benefit from the way that they have changed the environment: eliminating
predators, building up permanent shelter, exploring the places we live and finding the
resources. Niche creation is environmental, but its also cognitive; we create a niche to
grow a human brain around the infant.
Add together increased population density, excellent transmission of information, and
language you have a really strong channel for inheriting knowledge. Less gets lost each
generation.
Most chimpanzee culture and technology can basically be re-invented in a generation. Its
not cumulative. You dont need to figure out how to make one tool, to make another tool,
to make another tool. Human culture, in contrast, is the result of many generations of
transmission and gradual improvement; you couldnt re-invent it by yourself, alone, no
matter how smart you were. The point is, to truly understand how humans have evolved
to become so peculiar, you have to recognize that our social skills are as important as our
brains because our social skills allow our brains to work together over generation upon
generation.
Michael Tomasello, a psychologist and evolutionary theorist, has called this the ratchet
effect.
A ratchet wrench is a wrench that can only turn one way; as you tighten a bolt, you move
it freely one way and the other, but it only twists the bolt in the direction you want it to
go. It doesnt twist backward; you cant lose progress.
Culture and technology develop a ratchet when innovations dont get lost. If an
individual has an insight or makes an invention, social ties preserve that information so
that the innovation will not die with the inventor. At that stage, technology only goes in
one direction.
The ratchet takes a while in human evolution to get going. At first, we have the
information getting transferred successfully, but the tools remained the same for a very,
very long time a million years, really. But eventually, when we start to increase the
intelligence, add in a bit more creativity, improve the problem solving, and multiply the
number of potential improvers who are communicating, the cultural ratchet starts to turn
at first slowly click, click, click.
For example, when I show students how our tools changed, I think that theyre
underwhelmed; some of the most sophisticated stone tools, the last to develop, are the
microliths. These are tiny stone tools, often glued with tree resin into wood handles or
used for very specialized purposes. But these tools were the sign of a truly diverse,
innovative tool kit.
Special tools for hunting big game, special tools for fishing, special tools for catching
birds, digging up roots, making clothing, preparing skin, cutting down grass harpoons
and fish hooks and needles and saws with teeth tons of techniques and bits of advice
that have to get passed down from parent to child. Theyre a sign of that ratchet click,
click new inventions, modifications and improvements on old tools. Now, weve got
something that looks like human culture.

Probably with the development of Homo erectus, we began to have a kind of bio-cultural
evolution, but it would really take off with modern humans in the last 200,000 years.
Learning would be absolutely essential to survival, and the demands of culture making
tools, finding food, fashioning shelter, maybe even making fire and cooking would
have become a selective pressure on our brains. These skills demanded intelligence and
ability, but they also demanded intense social skills learning, imitating, teaching,
cooperating, even specializing within a group.
Youve got a fully human mind at this point, but you also see that our intelligence is not
just determined by whats encased in our skulls. We inherit so much from our ancestors
because we have the cognitive, social and organizational skills to take it on board. We
have to be immature when were born because we have so much to absorb after we get
here.
The funny thing is, the social nature of our intelligence is our great advantage, but its
also our Achilles heel. We dont just get correct ideas or accurate insights. Sometimes
we absorb mistakes, biased perceptions, or obsolete patterns of thinking. Yes, we stand
on the shoulders of our ancestors, so we can see further. But we also have to realize that
we sometimes see through their eyes, even when they didnt perceive things clearly.

The end of evolution?

So, thats how evolution made us. Weve blown past some details, rushed past some great
debates, and left out a lot of unanswered questions, but the last four weeks have been the
basics of how humans have evolved.
You might asking, What next? Is evolution over? Surely, natural selection cant be able
to get to us, protected as we are with our advanced medicine, our hygiene, and our
sophisticated technology. I mean, its not like theres bears prowling the streets, picking
off the weak and slow, right?
For a long time, anthropologists thought human evolution was over, or at least that we
had fundamentally changed the dynamics. We were so convinced that technology and
culture buffered us against selection that many theorists came to the conclusion that the
biological chapter of our history was already written. If we were to evolve any further,
they thought, our technology, culture, or overall enlightenment would be the avenue.
Well, the genetic evidence has come back, and its pretty clear that selection is not done
with us. In fact, by most genetic measures, the past 20,000 years or so may have been
some of the most rapid evolution in our species history, with all sorts of selective
pressures on our genome. Preliminary results suggest that pressure especially seems to be
on genes linked to our immune system, brain function, metabolism and digestion.
Turns out that the only time a species stops evolving is when its extinct, and were not
dead yet.
If you think about it more deeply, it makes sense. If evolution is variation, inheritance,

and selection over time leading to adaptation VISTA the basic factors are all still in
place. Theres no reason to think theyve stopped, is there?
First, weve got more variation than ever before. Theres over seven billion of us on
Earth today, having a lot of babies; thats up from two billion less than a century ago, and
only 5 million back in 9000 BC. Population increased 1000 times!
The expansion of our species around the globe and the growing size of the population
have already produced some interesting quirks. Anthropologist John Hawks points out
that ten thousand years ago, no one on the planet had blue eyes; that gene has developed
recently, meaning that only 400 generations has produced a noticeable change.
In fact, the rate of mutation may actually be going up because older fathers tend to have
more mutations in their sperm. In some countries, the average age of fathers has
increased dramatically, suggesting that we might be producing more genetic innovation
than before.
Inheritance hasnt changed. We may know whats going on, and in some of the wealthier
countries, were doing genetic counseling for couples at risk of having babies with certain
birth defects. But we havent replaced sexual reproduction and recombination with
cloning or cryogenic freezing, and its not clear that we will anytime in the near future.
But, aha, you say, surely theres no selection going on? After all, everyone survives to
reproduce, if they want to, dont they...?
Actually, selection is going on. You just have to know where to look for it.
First, any time theres a radical change in the environment, youd expect that selective
pressures would also shift quickly, tossing any existing stabilizing selection and
equilibrium out the window. No one can doubt that our environment right now is in a
state of flux. The only question is how much change and what kind.
Right now, for example, the forces of erosion are only moving around one-tenth the
amount of soil that humans are moving around. Researchers in the US estimate that 30
tons of earth get moved for every individual. Were changing the amount of heavy metals
in the atmosphere, dumping waste products at an unprecedented pace over one billion
tons last year and some of this waste has no foreseeable way to decompose before its
going to wind up in the food chain.
Most biologists are of the opinion that we are producing a sixth great extinction event on
the planet. The first five wiped out at least half of all the species alive at the time, and it
probably takes at least 10 million years for diversity to recover, as new species emerge
out of the survivors.
Species go extinct all the time, its true. Thats the way evolution works. Its normal. But
right now, the extinction rate appears to be way above normal. Scientists estimate that
40% of the species may be in danger of dying out in the near future. The rate is at least
100 times higher than normal; the famous biologist E. O. Wilson has argued that the
actual rate is probably 1000, even 10,000 times above what is normal. Humans are

having an impact similar to the meteor strike that wiped out the dinosaurs 65 million
years ago.
Youd have to be pretty confident about our situation as a species if you looked around
and said, yeah, all those other animals are evolving or going extinct, but nothings
happening to us.
Youd also have to be pretty oblivious to the way that most of us in the world today live.
Last year, 7 million children under the age of five died; thats around 5% of all the
children in that age bracket. That number is down a lot from what it was even fifteen
years ago, which is great, but there are parts of the world right now, countries like
Afghanistan, Mali, and Somalia, where infant mortality gets as high as 10% before a
childs first birthday. Im not even got to talk about this from a human rights
perspective; from an evolutionary perspective, it should certainly put to rest the idea that
our species has universally escaped from selective pressures. Weve made huge strides in
global health, but pneumonia, malaria and diarrhea are still working their way through
some parts of our gene pool, viciously weeding out those who are particularly
susceptible.
Some evolutionary theorists are discussing the possibility that humans will have to
evolve resistance to HIV, probably in those places where infection rates are the highest.
We know that some people are particularly resistant to developing full-blown AIDS from
the virus, and theres lots of evidence that chimpanzees had to do something similar a few
million years ago; their genome is littered with retroviral DNA from an attack by a
similar disease.
Weve done a great job of getting rid of some selective pressures. And its not just bears.
But it turns out that living close together was a really rough transition for humans that
subjected us to a whole new set of pressures. For most of its history, London has been a
population sink; more people have died there than been born, the countryside supplying a
steady stream of immigrants to reinforce the population. When Darwin was writing On
the Origin of Species, infant mortality may have been as high as 50% in England.
And those cities bred some pretty amazing diseases; just ask the Native Americans.
When Columbus and the rest of the crew from Spain, Portugal, England and the other
maritime powers showed up in the Americas, they brought with them diseases that had
evolved in the streets and slaughterhouses and sewers of European cities. The Europeans
had developed resistance over centuries of suffering from plagues and epidemics; the
Americans had no resistance. In some cases, the mortality rate following colonization
was over 90%, people dying, not just from disease, but from over-work, war, and the
complete collapse of all their social structures.
Talk about a bottleneck even for genetic diversity in the Americas! It may seem like
ancient history to us, with a world that seems to change every decade, but in an
evolutionary framework, a few dozen generations isnt much time at all. Were in a
super-heated phase of evolution.

Where to from here?

When we look around, if we live in wealthy countries, sure, we can be forgiven for
thinking that the laws of natural selection have been repealed. In some rich countries,
weve done a great job on infant morality. In parts of the industrialized world, it has
dropped below 1%. Thats a huge accomplishment. Absolutely, in these pockets of
human population, the selective pressures of pneumonia, malaria, tuberculosis, and other
killers of children have been held at bay.
But even the way that weve abolished certain diseases and malnutrition may be
rebounding back around to create new selective pressures on those people and dont
forget, relaxing the pressure on one part of the gene pool doesnt mean its been
abolished for the whole of the species, by any stretch.
For example, as many of you probably know, our widespread use of antibiotics to keep us
safe, even to keep our domestic animals safe, is starting to really scare some
epidemiologists. Weve created a tough world to be an infectious bacteria. But that means
that some of the strongest are surviving, adapting, and getting harder to kill. Our growing
population, more and more densely packed cities, and all the crowded agriculture
facilities where we raise our food, may also be breeding the next major selective event in
our own stockyards or hospitals.
Among the wealthiest people, weve abolished malnutrition as a selective pressure, for
certain, but weve also placed ourselves under pressure to adapt to a rapidly changing
diet. People may not starve today in some places because of agriculture, but agriculture
makes demands on our bodies, including adapting to a shift in the type of food and the
profile of nutrients we get. The evidence from genetic research is that our ability to
process food is under intense selective pressure; remember, 10,000 years ago, virtually no
adult humans could digest milk.
Were also staring down the barrel of an obesity epidemic in some parts of the world, a
reminder of how our bodies have not all successfully adapted to our changing way of life.
We know that obesity affects fertility in both men and women. Fat produces estrogen, so
obese men have lower levels of androgens and can even wind up with erectile
dysfunction. Obesity in women can mess with their reproductive cycle, even leading to
amenorrhea, or the interruption of the menstrual cycle.
We suspect that male sperm count has dropped in recent years. Boys alive today tend to
have lower sperm counts than their grandfathers generation, although this varies a lot
from one to the next, and it may be affected by changes in the way we measure the count.
However, if a French research team that studied data from more than 100 fertility clinics
is correct, the average sperm count may be dropping by 2% a year there. We dont know
for sure why some reproductive scientists worry its synthetic chemicals that mimic
hormones in our bodies but the European Science Foundation report that one in five
young men had a sub-fertile sperm count.
If its happening and this is controversial some individuals may be more resilient,
meaning that they have a trait that will likely be selected for by reproductive success.

Remember, its the population as a whole and not any single individual were more
concerned about, so there will be lots of exceptions even if the trend is very clear across
the whole group.
My goal is not to scare people about their weight or their sperm counts or anything else.
The point is that, given our changed, unprecedented nutritional, chemical, and
technological environments, theres whole new forces shaping our reproductive success.
Depending upon how your body responds to the new environment, you may be more or
less successful at surviving and reproducing. We may not be able to see it happening
because, even though the pace of evolutionary change has accelerated, no one is going to
change in front of your eyes. The shift will be slow, from one generation to the next,
almost, almost imperceptible.
However, recently in the UK, some researchers may have caught it. Using widespread
surveys of health and basic measurements, the researchers found that UK women were
becoming shorter, stouter, with lower cholesterol levels, lower blood pressure, and
extended windows of fertility. We cant tell on such a short time scale, but these women
may be, as a group, adapting to our radically changing environment.
All of these changes punch huge holes in any simple distinction between natural selection
and artificial or human-driven processes. We are having such a massive impact on our
environment, doing so much niche creation, changing our way of life so much, that its
really hard to argue that anything around us is natural anymore. We may think of
nature as flowers and birds and forests, but right now, for the purposes of natural
selection, its everything around you that affects your survival or your ability to
reproduce.
This doesnt mean were doing anything intentional to make ourselves evolve. Im kind
of suspicious of anyone who says that they think were going to be able to plan our future
evolution. In fact, most of us have no idea how we are changing the selective forces that
are shaping us. In addition, were probably not even used to thinking of evolution this
way many of us are more used to getting excited about technology when we think
about our future, not the way that this technology might affect our reproductive success.
Theres really no way to know what is going to happen. Or, as the great philosopher Yogi
Berra, warned: It's tough to make predictions, especially about the future.
In spite of the difficulty of predicting the future, however, I want you to think about it.
And I want to leave you with the words of Jonas Salk, the discoverer of the polio vaccine,
and a man who refused to patent the discovery because he thought it was better for the
good of humankind. Salk was of the opinion that:
The most meaningful activity in which a human can be engaged is one that is directly
related to human evolution. This is true because human beings now play an active and
critical role not only in the process of their own evolution but in the survival and
evolution of all living beings. Awareness of this places upon human beings a
responsibility for their participation in and contribution to the process of evolution.
Evolution shows how we were all stitched together at the start: all humans, all living

things, arisen from shared ancestors. But it also shows how we are tied together in the
future. What we do now affects how other life will evolve, whether some forms of it will
disappear. But it also affects our descendants, and whether they would recognize us, or
prefer not to know about us.

References & Resources

Key concepts
Human brain evolution is unusual, especially the degree of encephalisation, requiring us
to think about how such a metabolically expensive organ could be afforded by our
ancestors.
Brain function tends to involve repurposing of old brain areas and layering new types of
control, even over automatic functions.
Human intelligence is not just the result of a large brain, but also a modified social life
and developmental trajectory that encourages learning.
Genetic and other evidence suggests human evolution is not over; natural selection is
liable to remain an important process in human genetic change.

References
Bradshaw, J. L. 1997. Human Evolution: A Neuropsychological Perspective. Hove:
Psychology Press.
Calvin, W. H. 2004. A Brief History of the Mind: From Apes to Intellect and Beyond.
New York: Oxford University Press.
De Waal, F. 2007 (1982). Chimpanzee Politics: Power and Sex Among Apes. Baltimore,
MD: Johns Hopkins University Press.
Donald, M. 1991. Origins of the Modern Mind: Three Stages in the Evolution of Culture
and Cognition. Cambridge, MA.: Harvard University.
Dunbar, R. I. M., and Shultz, S. 2007. Evolution in the Social Brain. Science 317:13441347. doi: 10.1126/science.1145463 (pdf available here)
Flinn, M.V., Geary, D.C., and Ward, C.V. 2005. Ecological dominance, social
competition, and coalitionary arms races: Why humans evolved extraordinary
intelligence. Evolution and Human Behavior 26: 10-46. doi:
10.1016/j.evolhumbehav.2004.08.005 (pdf available here)
Geary, D. C. 2005. The Origin of Mind: Evolution of Brain, Cognition, and General
Intelligence. Washington, DC: American Psychological Association.

Hawks J, Wang ET, Cochran G, Harpending HC, and Moyzis RK. 2007. Recent
acceleration of human adaptive evolution. Proceedings of the National Academy of
Sciences, USA 104: 2075320758. Available from: http:/?
dx.doi.org/10.1073/pnas.0707650104
Herrmann, E., Call, J., Hernndez-Lloreda, M. V., et al. 2007. Humans Have Evolved
Specialized Skills of Social Cognition: The Cultural Intelligence Hypothesis. Science
317:1360-1366. doi: 10.1126/science.1146282 (pdf available here)
Rilling, J. K. 2006. Human and NonHuman Primate Brains: Are They Allometrically
Scaled Versions of the Same Design? Evolutionary Anthropology 15(2): 65-77. doi:
10.1002/evan.20095 (pdf available here)
Russon, A. E., and Begun, D. R., eds. 2004. The Evolution of Thought: Evolutionary
Origins of Great Ape Intelligence. Cambridge: Cambridge University.
Somel M, Creely H, Franz H, Mueller U, Lachmann M, et al. 2008. Human and
Chimpanzee Gene Expression Differences Replicated in Mice Fed Different Diets. PLoS
ONE 3(1): e1504. doi: 10.1371/journal.pone.0001504
Tomasello, M. 1999. The Cultural Origins of Human Cognition. Cambridge, MA:
Harvard University Press.
Wynn, T., and Coolidge, F. L. 2004. The expert Neandertal mind. Journal of Human
Evolution 46:467-487. doi: 10.1016/j.jhevol.2004.01.005 (pdf available here)

Links
The Childs Developing Brain (interactive infographic), The New York Times. (link)
Juan Enriquez: Will our kids be a different species?, TED: Ideas Worth Spreading. (link)
Mark Changizi, Bursting the Bubble of Human Intelligence, The Crux, Discover. (link)
Greg Downey, Food for thought: Cooking in human evolution, PLOS
Neuroanthropology. (link)
Andrew Glikson, Views on the future of Homo sapiens, The Conversation. (link)
_____. Is another mass extinction event on the way?, The Conversation. (link)
John Hawks, Ninety percent of your brain is (not) useless. John Hawks weblog. (link)
_____. Why human evolution accelerated. John Hawks weblog. (link)
Jennifer Jacquet, How Culture Drove Human Evolution: A Conversation with Joseph
Henrich. The Edge (video and podcast available). (link)
Brandon Keim, Humans Evolving More Rapidly Than Ever, Say Scientists. Wired. (link)

Greg Laden, Is Childhood The Most Important Human Adaptation?, Greg Ladens Blog,
Science Blogs. (link)
Gary Marcus, Where Uniqueness Lies. Nautilus. (link)
Brian Palmer, The Real Caveman Diet: Did people eat fruits and vegetables in prehistoric
times?, Slate. (link)
H. L. Shapiro, Man500,000 Years From Now, Natural History. (link)
Peter Tyson, Are We Still Evolving?, Nova (PBS). (link)
Christie Wilcox, Expensive Organs: Guppies Reveal The Cost Of Big Brains, Science
Sushi (weblog), Discover. (link)
Ed Yong, Q: Why dont apes have bigger brains? A: They cant eat enough to afford
them, Not Exactly Rocket Science (weblog), Discover. (link)
_____. A duplicated gene shaped human brain evolution... and why the genome project
missed it, Not Exactly Rocket Science (weblog), Discover. (link)
Carl Zimmer, First Comes Global Warming, Then an Evolutionary Explosion.
Environment 360 (Yale University). (link)

~~~~~~~~

Photo credits
Book cover
Top photo by Sergiu Bacioiu. Creative Commons licensed (BY NC). Original location:
http://flickr.com/photos/sergiu_bacioiu/4444496049/
Second photo by Greg Downey, Creative Commons licensed (BY NC)
Third photo by belgianchocolate:, Creative Commons licensed (BY). Original location:
http://flickr.com/photos/frank-wouters/36720564/
Bottom photo by Greg Downey, Creative Commons licensed (BY NC)
Figure 1
The coccyx, from Grays Anatomy. Image in the Public Domain.

Figure 2
Cross-section of embryonic tail. Source: (Modified from plate 22 of Fallon and Simandl
1978). http://www.talkorigins.org/faqs/comdesc/section2.html
Figure 3
X-ray image of an atavistic tail found in a six-year old girl. Source: reproduced from BarMaor et al. 1980, Figure 3. http://www.talkorigins.org/faqs/comdesc/section2.html
Figure 4
Louis, trusty dog, photo by Greg Downey, Creative Commons (BY NC).
Figure 5
Homologies, image by Vladislav Petrovich Volkov, 2011, released into the public
domain.
Figure 6
Jeffrey, absurdly charismatic Shetland Pony, photo by Greg Downey, Creative Commons
(BY NC).
Figure 7
Tetrapod forelimbs, image by Wilhelm Leche, 1909. Public domain.
Figure 8
Convergent structure in analogous wings, diagram by John Romanes (1892): Darwin and
after Darwin, Public domain. Source:
http://commons.wikimedia.org/wiki/File:Homology.jpg
Figure 9
Drawing from Charles Darwins First Notebook on Transmutation of Species (1837)
Public domain. Original location:
http://commons.wikimedia.org/wiki/File:Darwins_first_tree.jpg

Glossary
Acheulean
(Sometimes Acheulian) Derived from the name Saint-Acheul, a suburb of Amiens in
France. These tools are also sometimes referred to as Mode 2 technology.
First developed around 1.8 million years ago, Acheulean stone tools are associated first
with the rise of Homo erectus/Homo ergaster. These tools tend to be more thoroughly

worked, on all sides, unlike earlier Olduwan tools. The distinctive teardrop-shaped
Acheulean hand-axe, a symmetrical tool, is found over a wide geographic range.
Related Terms: Olduwan

Adaptive radiation
A pattern of diversifying or disruptive evolutionary change, leading from a single species
into multiple different species over time, with each adapted to a different niche. The rise
of multiple different specialised mammal species from one or a small number of
mammals into thousands is often cited as an example of adaptive radiation.
Related Terms: Diversifying selection, Homologies

Allele
One of the variants that can occur at a particular gene or genetic locus in a cells DNA. In
some cases, an allele will produce an actual observable difference in an organisms
phenotype or traits, but in others, an allele will not produce an expressed difference.
Related Terms: Continuous, Discreet, DNA, Dominant gene, Recessive

Alloparenting
A pattern of multi-individual parenting or assistance to a mother in raising offspring,
from either unrelated individuals or individuals related to the child (e.g., siblings, father,
parents siblings like aunts, grandparent... etc.). Alloparenting distributes the demands of
altricial human infants and can thus lead to shorter birth spacing and greater fertility for
women.
Related Terms: Birth spacing

Altricial
An individual who requires great assistance, especially an infant who is particularly
helpless and unable to care for her- or himself. Humans are very altricial relative to other
more precocial (or self-sufficient at an early age) species infants.
Related Terms: Metabolic ceiling, Obstetrical dilemma, Precocial, Sexual revolution

Anisogamy

Anisogamy is sexual reproduction between two sexes who do not contribute similar
gametes. The smaller gamete, referred to as the sperm, determines which sex is male; the
female contributes the larger gamete, or egg.
Related Terms: Sex

Anthropocentric
A tendency to see humanity as the most important topic of biological research and even
to over-ephasise the importance of humanity. Anthropocentric theories often fail to
recognise how unusual humans are or the complexity of non-human life.
Related Terms: Great Chain of Being, Teleological error

Atavism
A trait in an individual that does not appear in its parents or immediate ancestors, but was
present in a much older ancestor. A recurrence of an ancient trait that no longer manifests
in other contemporary members of a species. In laymans terms, an evolutionary
throwback of a trait to an earlier variant.
Related Terms: Vestigial organ

Birth spacing
The interval between births, often affected by breast-feeding (feeding on-demand can
suppress fertility), maternal body weight and exercise levels, and overall fertility.
Humans have shorter birth-spacing intervals than some of the other apes, like orangutans,
leading to higher fertility in our species. Some credit our relatively shortened birth
spacing to alloparenting.
Related Terms: Alloparenting

Bodily cells
The cells of the body, opposed to germ cells, which are the reproductive cells (and thus
the first cells of your future offsprings body). According to Weismann, body cells could
adapt or be affected by the environment, but any changes to bodily cells would not be
passed on to offspring, because the communication between your germ cells and bodily
cells was one-way only.
Related Terms: Germ cells, Weismann barrier

Bonobo
Pan bonobo. The less common species of chimpanzee. Only discovered by Western
scientists in the twentieth century. Although physically similar to common chimpanzees
(Pan paniscus), bonobos have distinctive patterns of social interaction.

Brain stem
In vertebrate neural anatomy the brain stem (or brainstem) is the rear part of the brain,
connected to and structurally continuous with the spinal cord. The brain stem
communicates be- tween the rest of the brain and the nervous system and provides
regulation of some automatic functions, like cardiac rhythm and breathing.

Brocas area
Brocas area is a region of the hominid brain cortex in the left hemisphere linked to
speech production. The area is named for French neurosurgeon Paul Broca, who
discovered its function while examining the brains of patients with brain injuries and
language difficulties. The area is on the surface of the brain, right above the left ear.
Related Terms: Wernickes area

Capoeira
An Afro-Brazilian martial art and dance, involving a distinctive musical style and
vigorous, acrobatic movement. Greg studied capoeira in Brazil while working on his
doctorate degree in order to understand how culture affected physical and psychological
development of the people who practiced it.

Chimera
A chimera is an organism composed of two or more genetically distinctive populations of
cells. Chimeras can be formed through the fusion of two initially separate fertilised eggs.
Some cases of chimera have been reported in humans. Unlike a hybrid, in a chimera, the
origin of each cell can be clearly determined as the two types are clearly distinct.
Chimeras can also be created artificially; in some cases, biologists have placed foreign
cells into an organism and these cells are assimilated into the animals functional
architecture.

Chloroplasts
Organelles found in plant cells and some other eukaryotic organisms. They are the parts
of the plant cell responsible for photosynthesis, converting energy from sunlight into
stored molecular energy.
Related Terms: Endosymbiotic theory, Photosynthesis, Symbiogenesis

Chordates
The phylum of animals which has a notochord, or a form of central nervous system.
Vertebrates are a subphylum of the chordates, which also includes some animals, like the
sea squirt, that have a notochord for part of their lives, but do not have a permanent
central nervous system or vertebrae.

Coccyx
The tail bone, or the final three to five spinal vertebrae, below the pelvis, that fuse into
a single bone in primates without tails.

Cognitive niche
A concept proposed by John Tooby and Irven DeVore to explain human distinctiveness.
According to these theorists, humans exploit an ecological and evolutionary niche made
available by their cognitive skills. The concept of the cognitive niche also can be linked
to the idea of niche construction -- see glossary -- to capture the way that individuals are
born into a personal environmental setting or niche where these distinctive cognitive
skills are encouraged and developed.
Related Terms: Niche creation (or niche construction), Ratchet effect

Competitive exclusion
Sometimes referred to as Gause's law of competitive exclusion. The ecological principle
that two species cannot compete stably for the same resources. One species will likely
have an advantage, however slight, leading this species to dominate in the long term. The
other competitor will either become extinct or be forced into a different ecological niche.
Related Terms: Interspecific competition

Concealed estrus

The lack of signaling in the females of a species when they are ovulating and fertile or in
heat. Unlike chimpanzees, who signal when they are fecund with genital swelling,
human women offer no signal detectable to men when they are fertile. Gorillas are semiconcealed, suggesting that both chimpanzees and humans have evolved derived traits, or
adapted with their own distinctive patterns since they shared a last common ancestor.
Related Terms: Estrus, Sexual revolution

Conserved
A conserved trait is stable over evolutionary time, or shared between descendants of a
common ancestor. Often, conserved traits are believed to be important for survival, so
they cannot vary without decreasing an organisms chance of survival. Some traits or
genes are highly conserved across multiple different species, owing to their essential role
in organism functioning.
Related Terms: Hox genes

Continuous
A trait that varies with a smooth distribution and many intervening variants. For example,
human skin colour and height are both continuous traits, with various individuals
distributed across the whole range of possible phenotypes. Continuous traits tend to be
determined by multiple genes (polygenic) rather than a single, discrete allele, which is
more likely to produce a discreet trait.
Related Terms: Allele, Discreet, Polygenic

Convergent evolution
The evolution of similar features in organisms from distinct lineages. The example used
in the text to explain convergent evolution is the wing: wings have developed in a range
of species from different lineages (insects, pterosaurs, birds, and bats). Convergent traits
often have quite distinct underlying structures, but the demands of the environment can
shape them to have similar features. For example, wings are anatomically varied, but they
all have certain shared traits because of the demands of flight: large surface area,
flexibility, strong mechanisms for moving them, light weight.
Related Terms: Homologies

Copy number variation

A structural variation in the genome caused by an unusual number of copies of a section

of the DNA sequence. A section may be copied an extra time, or missing. The human
genome contains a large number of these duplicate areas; sometimes identical twins vary
in the number of copies that they have of a section of DNA.
Related Terms: Junk DNA, Mutation

Cyanobacteria
Also called blue-green algae, cyanobacteria produce energy through photosynthesis.
They take their name from their colour (cyano is from the Greek word for blue).
Endosymbiotic theory holds that plants gained chloroplasts by capturing symbiotic forms
of cyanobacteria. Cyanobacteria are also widely credited with oxygenating Earths
atmosphere.
Related Terms: Chloroplast, Endosymbiotic theory, Photosynthesis

Diagonal transmission
A term used to describe transmission of traits from adults of a species to children who are
not their own. That is, diagonal transmission occurs when behaviour, traits or practices
can be communicated to genetically unrelated young of the species, either through
copying, teaching, or a mechanism like cognitive niche creation. Without diagonal
transmission, traits are only inherited from ones parent or parents.
Related Terms: Cognitive niche, Inheritance of acquired traits, Lateral transmission,
Ratchet effect

Dimorphism
Phenotypic difference between the males and females of a species; that is, a dimorphic
species has two forms, one for each of the sexes. (Some species are polymorphic.) The
greater the difference in the sexes, in some species, the clearer indication that individuals
are engaged in sexual competition.
Related Terms: Direct competition, Sexual revolution, Sexual selection

Direct competition
A form of sexual selection in which members of one sex compete directly through
confrontation or ritualised combat. In elephant seals, for examples, males fight for control
of breeding females, a process which has favoured the development of extremely large
males (and thus increased sexual dimorphism).

Related Terms: Dimorphism, Sexual revolution, Sexual selection

Directional selection
A pattern of natural selection in which one variant of a species is favoured, leading a
species to skew over time so that this variation becomes more and more predominant.
Over time, directional selection can lead a species to adapt and change significantly.
Related Terms: Stabilising selection

Discreet
A discreet trait typically has two alternative expressions, with no or little intervening
expression or what is sometimes called discontinuous expression. A trait like biological
sex (male and female) is discontinuous, with any mixed expression extremely unusual.
Discontinuous expression or discreet traits are typically understood to be the result of a
trait determined by a single allele. Gregor Mendel first proposed genetic inheritance
through his research on discreet traits in pea plants.
Related Terms: Allele, Continuous, Dominant gene, Recessive

Diversifying selection
A pattern of natural selection in which different parts of a population adapt in distinct
ways, often leading to a disruptive pattern for key adaptive traits. For example, if a
species exists across an ecological range with a variety of niches, members of the species
may be subject to different selective pressures, leading to a variety of traits being
selected. In extreme cases, diversifying selection can lead to speciation, especially if the
different parts of the population are isolated from reproducing with each other,
preventing gene flow.
Related Terms: Adaptive radiation, Speciation, Stabilising selection

DNA
Deoxyribonucleic acid (DNA) is a molecule that encodes the genetic instructions used in
the development and functioning of all known living organisms, founding in the nucleus
of cells. Genetic information is encoded as sequences of four nucleotides guanine,
adenine, thymine, and cytosine (frequently indicated by their initials: G, A, T, C). Most,
though not all, living cells have their DNA as a kind of spiraling, wound up ladder shape,
with two spine-like chains of sugar-phosphates molecules linked by rungs of the
nucleotides.

Related Terms: "Junk DNA, Allele, Double helix, Epigenetics, mtDNA, Mutation

Dominant gene
With a discreet trait, one allele is dominant when it masks the presence of the other,
recessive allele, in an organisms phenotypic expression. An organism can carry the allele
for the recessive trait without it being expressed if it occurs alongside the dominant allele.
Related Terms: Allele, Discreet, Particulate inheritance, Recessive

Double helix
The double helix is the structure of the DNA molecule in most, though not all, living
things. The term describes the ladder-like structure of the nucleotide bases linking two
sugar-phosphate chains in a spiraling structure.
Related Terms: DNA

Ecological dominance
Originally suggested by Richard Alexander, the theory of ecological dominance suggests
that our hominin ancestors, at some point in evolutionary history, became so capable of
resisting selective forces in nature hunger, predators, climate that intraspecific
competition became disproportionately important in shaping human evolution. Whether
or not our ancestors became dominant, the theory draws attention to the important roles
played by social competition, inter-group rivalry, and cooperation in human evolution.
Related Terms: Environmental theories, Intraspecific competition

Emotionally modern
Sarah Blaffer Hrdy has suggested that one of the key changes in human evolution is a
shift in the motivational and emotional structure of our ancestors, diminishing withingroup aggression, allowing suppression of instinctive responses, and allowing for greater
cooperation. Her theory suggests that these changes may have preceded other cognitive
changes such as language or complex learning laying the social and emotional
foundation for sub- sequent cognitive shifts.
Related Terms: Social cognition

Encephalisation

The process over evolutionary time of species developing larger-than-expected brain size
given their body weight. A precise measure of encephalisation is difficult, however, due
to the allometric relationship between body size and brain size, and disagreement about
how to measure relative encephalisation.
Related Terms: Expensive tissue

Endogenous
Refers to substances, processes or events that are initiated from within an organism or
entity. In this book, endogenous viruses are viral sequences of DNA that are carried into
the zygote with the germ cells, either the sperm or egg. In other words, endogenous
substances are passed on from parent to offspring as a fact of reproduction.
Related Terms: Retrovirus

Endosymbiotic theory
First proposed by Russian botanist Konstantin Mereschkowski, endosymbiotic theory
argues that some of the structures in complex living cells developed through evolution
from symbiotic lifeforms that were originally separate. Later advocated most notably by
Lynn Margulis, endosymbiotic theory proposes specifically that eukaryotic cells the
foundation of complex life began as symbiotic communities of less complex
structures. Most notably, the theory has been given support by the study of chloroplasts,
mitochondria, and other organelles that contain their own sequences of DNA, shorter than
nuclear DNA, and seem to go through their own forms of reproduction as cells reproduce.
Related Terms: Chloroplasts, Mitochondria, Symbiogenesis, Symbiosis

Environmental theories
Theories of evolutionary change that privilege external forces as shaping an organism. In
the case of humans, especially the development of human encephalisation and cognitive
elaboration, environmental theories suggest that the driving factors were the need to
adapt to a variety of ecological niches, survive climactic change, forage for varied food
sources, convert to a hunting-based diet, or contend with other external threats. Often
contrasted with social theories of intelligence or accounts that focus on intraspecific
competition.
Related Terms: Ecological dominance, Intraspecific competition, Machiavellian
intelligence

Epigenetics

The study in genetics of changes in gene expression in phenotype that arise from nongenetic sources. Epigenetics investigates how functional changes can occur in the
genome without changing the sequence of nucleotides, through such processes as DNA
methylation and histone modification, both of which can lead to alterations in gene
expression while leaving the underlying DNA sequence unchanged.
Related Terms: DNA, Hunger Winter, Inheritance of acquired traits, Weismann barrier

Estrogen
Also spelled, oestrogen (primarily in UK). A group of compounds that are essential in
moderating female reproduction and the menstrual cycle, found in vertebrates and
insects. The widespread presence and conserved nature of estrogens across species
suggests that these sex hormones have a deep evolutionary history and profound
importance in reproduction and survival. Although at much higher levels at women,
especially during child-bearing years, estrogens also appear in men, and are essential to
normal body function.
Related Terms: Sex, Sex-determining region of the Y chromosome (SRY), Testosterone

Estrus
The phase in a female animals menstrual cycle when she is fertile and sexually receptive
(sometimes referred to popularly with horses and other domesticated animals as in
heat). In some species, estrus is accompanied by visible signals of receptivity, such as
swollen or brightly coloured genitalia or by distinctive behaviour, such as the lordosis
reflex, when receptive females present themselves for mounting to males. In humans,
estrus is largely concealed, although some researchers argue that there are changes in
female behaviour.
Related Terms: Concealed estrus

Eukaryotic cells
A eukaryote is an organism whose cells contain complex structures enclosed within
membranes, especially a separated cell nucleus. In contrast, prokaryotes, including
Bacteria and Archaea, do not have internal, membrane-enclosed structures. All
multicellular organisms are eukarylotic, as well as many single-celled organisms.
Related Terms: Symbiogenesis

Eusocial

The most complex forms of social organisation in a species are eusocial, and they involve
high degrees of specialisation, often including some members of a group who are nonreproductive. Bees, termites, wasps and ants are the clearest example of eusociality, with
large numbers of sterile drones, workers or warriors supporting a much smaller group of
individuals who specialise in reproduction, including a queen. The evolutionary success
of eusociality is explained in large part by inclusive fitness; even though sterile workers
will not pass on their own genetic material, their cooperation is essential to the survival of
closely-related, reproductively-specialised siblings who will.
Related Terms: Group selection

Exaptation
Also sometimes called a pre-adaptation, an exaptation is a trait that shifts function over
time, leading to successive cycles of modification. For example, feathers first arose on
flight-less dinosaurs, likely to serve either in sexual selection or as insulation and thermal
regulation. Only later were those feathers exapted to serve for flight. The process of
exaptation can leave imperfectly adapted traits or structures that clearly evidence earlier
functions. Al- though the concept preceded their discussion, Stephen Jay Gould and
Elizabeth S. Vrba introduced the specific term, exaptation.
Related Terms: Neural reuse or neural recycling

Expensive tissue
The expensive tissue hypothesis was first proposed by proposed by Leslie Aiello and
Peter Wheeler in 1995. They proposed that our ancestors underwent an evolutionary
trade-off between the size of the brain and that of the digestive tract, which is smaller
than expected for a primate of our body size. Since both brains and digestive tract use
more energy than other bodily tissues for their weight, Aiello and Wheeler proposed that
our ancestors had to develop new techniques for getting calories because the ration
between brain and gut-size had departed so much from what we find in other mammals.
In this book, I use the term expensive tissue more loosely, in part because the argument
made by Aiello and Wheeler has come in for criticism. The fact that brain tissue is
metabolically expensive is indisputable, and in most animals, this high metabolic
cost (especially the need for energy) leads to a constraint on brain growth.
Related Terms: Encephalisation

Fitness
In evolutionary theory, the term fitness describes an individuals or species ability to
survive and successfully reproduce. Over time, an organisms fitness is its ability to pass

its genes on to the next generation. The popular understanding of fitness as overall
health or physical vitality is not the same, and can cause confusions in students of
evolutionary theory.
Related Terms: Gene pool, Gene-level perspective, Inclusive fitness, VISTA

Gender
A range of behavioural, physical, and psychological traits which distinguish from
masculinity from femininity. Social scientists and feminist scholars use the term gender
to distinguish social expectations and learned roles from biological differences linked to
reproduction. In the course of their research, anthropologists and historians have found
that gender norms what is considered normal or acceptable for men and women are
subject to quite wide variation. Although sex and gender are linked and clearly affect
each other, the process of socialization can often lead people to mistake gender traits
(learned social normals) for sexual traits (innate and unalterable products of physiological
difference between men and women).
Related Terms: Norms, Sex

Gene pool
The total sum of genetic information and variation in a species at any given time. The
gene pool is the underlying genetic material in any population, and evolution works over
generations to alter this gene pool through processes like natural selection and genetic
drift (relatively random changes in gene frequency due to differential reproduction,
without clear selective pressure). The term highlights both that species are inherently
varied, and that evolution operates, not on an individuals traits, but on the total gene pool
of a population over time.
Related Terms: Fitness, Gene-level perspective, Population

Gene-level perspective
Essential to the modern synthesis in evolutionary theory, a gene-level perspective focuses
on the evolutionary effects of natural selection, not on individual organisms, but on the
frequency and variation of genes in a population. Taken to an extreme, a gene-level
perspective can lead to the exclusion of other phenomena that shape evolution, and a
treatment of the gene as the only unit of selection. But the approach is a powerful tool for
modeling and making predictions about evolutionary processes, and it helps to explain
why some genes replicate successfully even if they are detrimental to the well-being of an
individual.
Related Terms: Fitness, Gene pool

Germ cells
Germ cells are those cells that give rise to gametes the sperm and egg in sexually
reproducing organisms. The germ line cells can give rise to new organisms, whereas
somatic or bodily cells cannot. Genetic changes to germ line cells will be passed on to
subsequent generations.
Related Terms: Bodily cells

Gestational diabetes
In some women who have not had diabetes previously, the blood glucose levels can
become very high during pregnancy, especially during their last trimester. Medical
specialists debate whether this should be considered a disorder, as the high glucose levels
seem to be caused by the placenta interfering with the insulin receptors in the mother.
The high blood glucose levels can lead to the infant be larger than normal for its
developmental age, and to other complications.

Great Chain of Being

Originally from Greek philosophical thought, the Great Chain of Being was an
understanding of all things being arranged in a strict hierarchical order, ordained by the
divine, from God down through the angelic beings, celestial bodies, kings and nobility,
normal humans, wild animals, domesticated animals, trees, all other plants, precious
stones, precious metals, and other minerals. For medieval thinkers, human occupied a
crucial place, poised between the divine and the earthly, with flesh that could die but, at
the same time, immortal souls. This model of the ladder of life was even adopted by the
early natural historians, including Linnaeus, the creator of our system of classifying
species. We can hear echoes of these ideas in some popular understandings of evolution.
Related Terms: Anthropocentric, Teleological error

Group selection
A set of controversial theories that natural selection can act, not just on an individual, a
gene or a population, but on competing groups, if they are organised. Group selection is
often part of multi-level selection theories, where selective processes are seen to be
operating simultaneously on multiple different levels of organisation (the gene, organism,
group, species). Group selection might help to explain why socially advantageous traits
could arise traits like altruism, empathy, willingness to sacrifice oneself that clearly
might undermine individual fitness. One alternative explanation is inclusive fitness.

Related Terms: Eusocial, Inclusive fitness

Hermaphrodites
A hermaphrodite possesses structural traits of the reproductive systems of both males and
females. Most researchers now use inter-sexed instead when talking about humans, as
many individuals who have traits of both males and females do not have reproductive
structures of both. That is, some inter-sexed individuals are not hermaphrodites, but
rather have more ambiguously sexualised organs. In some species, hermaphrodism is
normal, with either sequential change between sexes, or the possession of both sets of
reproductive organs.
Related Terms: Sex, Sex-determining region of the Y chromosome (SRY)

Holistic
In anthropology, the term holism is used to describe an approach that does not seek to
isolate a single facet of existence biological, cultural, economic, social, symbolic, etc.
but rather seeks to understand how the various dimensions of human life interact. In a
research environment where specialisation is essential for training and other
practicalities, holistic approaches can require a significant degree of cooperation among
specialists. In a discussion of human evolution, a holistic approach seeks to take all
available data to understand how humans evolved, assuming that no single cause or type
of process will likely be able to explain all the changes.

Homeostasis
A system that works to maintain consistency over time, especially in response to
changing conditions. Usually applied to living things, the concept of homeostasis points
out that organisms must actively work to achieve consistency as both their internal state
and external environment are subject to change. Various monitoring, control and
feedback mechanisms help organisms to maintain homeostasis.

Homologies
Structural similarities underlying dissimilar traits that demonstrate that two organisms
have descended from a common ancestor. The dissimilarity is often a sign of adaptive
radiation, as one structure, over times, is shaped by different selective pressures into
diverse homologous structures.
Related Terms: Adaptive radiation, Hox genes

Hox genes
Hox or homeobox genes are a group of related genes that control the body plan of the
embryo along its axis, from top to bottom. Hox genes produce transcription factors which
bind to other parts of an organisms DNA, encouraging or suppressing expression of
other genes. Often referred to as control genes, Hox genes designate which part of a
body a cell is building, so defective Hox genes can lead to conditions like antennapedia,
in which a defective signal leads to the growth of legs where antenna should be on an
insect. Although Ho genes are fascinating for many reasons, we discuss them here to
show two things: 1) the highly con- served deep structure of life, as Hox genes are quite
similar across all bilateral forms of life (that is, life with two sides that mirror image each
other); and 2) to show how regulation of genetic expression can lead to diverse structures
growing from quite similar protein-coding sequences of species DNA.
Related Terms: Conserved, Homologies

Hunger Winter
The Hunger Winter (Hongerwinter) was a famine during the winter of 1944 to 1945 in
the German-occupied parts of the Netherlands toward the end of World War II. Although
relatively few died, due to the work of collective soup kitchens, millions of Dutch
citizens suffered severe malnutrition, leading to long-term health consequences, even in
the children whose mothers were carrying them during the famine.
Related Terms: Epigenetics

Inclusive fitness
Inclusive fitness is the recognition that an organisms fitness is really all of its genes
that get passed on to the next generation, even if those genes are carried by its siblings or
siblings offspring. As evolutionary theorist W. D. Hamilton argued, an individual might
be more fit by this measure if it acted on behalf of its closely-related kin, not just in
pursuit of its own survival or that of its children. Hamilton suggested that inclusive
fitness, also sometimes referred to as kin selection when focusing only on close kin,
might explain altruism, cooperation, and self-sacrifice. An animal might give off a
warning signal, for example, when a predator approached, putting itself in greater danger,
but potentially protecting its group, including its close kin.
Related Terms: Fitness, Group selection

Inheritance of acquired traits

Suggested by French naturalist Jean-Baptiste Lamarck, and for this reason often called
Lamarckian evolution, the inheritance of acquired traits is the idea that organisms pass

on the adaptations that they have made during their lifetime to their offspring. Although
severely criticised by proponents of the modern synthesis, some contemporary
evolutionary theorists now argue that some forms of inheritance, including epigenetics
and cultural transmission, may demonstrate Lamarckian properties.
Related Terms: Epigenetics, Lateral transmission, Weismann barrier

Interspecific competition
Competition in evolution between different species, often trying to exploit the same resource. The success of one species can generate greater stress on its competitor. In an
extreme, inter-species competition leads to competitive exclusion, in which one species
will become extinct. Traits like high reproductive rates can aid a species in inter-specific
competition.
Related Terms: Competitive exclusion

Intraspecific competition
Competition within a species for resources, reproduction or survival. In this form of
competition, one members of a species success threatens other members of the same
species. For example, sexual competition is a form of intraspecific competition, as
members of one sex vie to reproduce at the expense of other members of the same group.
Related Terms: Ecological dominance, Environmental theories, Sexual selection, Trivers
hypothesis

Junk DNA
A popular name for non-coding DNA, or sequences of nucleotides on DNA that do not
encode for a particular protein. When the human genome was first decoded, 98% of the
DNA was found to be non-coding, and was initially referred to as junk DNA in some
reports, especially in the popular press. Subsequent research showed that this non-coding
DNA often had important functions, including regulating the expression of the coding
sections of DNA. Current estimates are that approximately 80% of the human genome
serves some biological function for the organism, although much of this research remains
to be done.
Related Terms: DNA

Kluge
Sometimes written, kludge. A clumsy or awkward solution to a problem, such as a

temporary, jury-rigged repair or hastily-made modification to a system. Contrasted with a

de- signed solution, in which forethought goes into the creation of a solution which is
appropriate and well-designed for the purpose it serves. In evolutionary theory, a kluge
refers to the odd, inelegant ways that organisms have often adapted, including in traits
like the skeletal adaptations humans have for walking bipedally.
Related Terms: Teleological error

Lateral transmission
Inheritance of traits within a generation, through mechanisms like imitation, learning,
or in some single-celled organisms exchange of genetic material. In sexuallyreproducing species, lateral transmission is a way in which non-parents influence the
development of off- spring, especially through culturally learning.
Related Terms: Diagonal transmission, Inheritance of acquired traits, Ratchet effect

Machiavellian intelligence
Proposed by Frans de Waal in his book, Chimpanzee Politics, Machiavellian
intelligence is a type of political savvy or social cognition that allows an individual to be
a successful actor in a social group, including building alliances, recognising deception,
and scheming against rivals. An early version of a social intelligence theory, de Waal
pointed out that the greater cognitive abilities of chimpanzees were used more for
intraspecific competition than for control of the environment.
Related Terms: Environmental theories, Social cognition

Marsupials
A group of mammals, related to placental mammals, that carry their offspring, who are
born quite early compared to other mammals, in a pouch. The absence of the placenta
makes the growing marsupial embryo more vulnerable to the mothers immune system,
forcing an earlier birth and a longer period of development in the mothers pouch.
Marsupials and placental mammals split at least 140 million years ago, and most
marsupials are found in the Southern Hemisphere, especially in South American and
Australia, the latter of which has been especially isolated from the other continents.
Related Terms: Placenta, Placental mammals

Mate choice
A form of sexual selection in which one sex (usually the female in mammals and birds)

chooses which male with whom to mate based upon some kind of display, such as an
elaborate mating dance, brightly-coloured feathers, sophisticated nest, or other indirect
means of competition. Mate choice can lead the preferred trait to become quite
pronounced, for example, in the peacocks tail.
Related Terms: Sexual selection
Index Find TermChapter 3 - Sex: why bother?

Melanin
A naturally-occurring pigment found widely in living things (although spiders, curiously,
dont seem to have it). Melanin is responsible for human skin colour, varying from deep
saturation in populations who have long resided near the equator to quite pale shades in
populations closer to the poles. Melanin protects the skin against ultraviolet light but can
make it hard for the body to metabolise vitamin D in low-light situations.
Related Terms: Pigmentation

Menstruation
The periodic discharge of blood and mucosal tissue in women from the uterus during
their reproductive life, stretching from menarche (the onset of menstruation) through
menopause. Not all mammals undergo menstruation; why human women do, and whether
the blood loss has an adaptive function, are the subject of debate in anthropology.

Metabolic ceiling
Dunsworth and colleagues (2012) have argued that human babies are born extremely
altricial, not because of the obstetrical dilemma or limits set by the size of the birth canal,
but because of energetic constraints. According to these researchers, humans can only
maintain a metabolic rate 2.1 times the basal metabolic rate, the ceiling for metabolic
exertion. In other words, the human fetus is so energy hungry that a mothers body
simply cannot feed it internally beyond our nine-month gestation period.
Related Terms: Altricial, Obstetrical dilemma

Microbiome
The total population of microbes that live in and on the human body, including all over
the skin, in the mouth and sinuses, under the eyelids, and in the gastro-intestinal tract.
Currently, research is ongoing to understand the functioning of this diverse microbial life
which outnumbers the cells that make up our body including its roles in helping to

maintain health and well-being.

Related Terms: Symbiosis
Index Find TermChapter 2 - Symbiosis & cooperation

Microliths
Extremely small stone tools that are an indication of some of the more sophisticated lithic
technology or stone age tools. Microliths very small points or flakes of stone
were often used in projectile points, including arrowheads, and in the manufacture of
composite tools, that is, tools made from multiple different substances, such as stone
glued with resin to wood handles.

Mitochondria
Small, membrane-enclosed organelles within most eukaryotic cells that they generate
most of the cell's supply of chemical energy. Mitochondria are in the cells cytoplasm,
outside the nucleus; their number varies depending upon the cell type. Mitochondria
undergo their own processes of reproduction and have their own, circular chromosome,
providing evidence for symbiogenesis and allowing mitochondrial DNA (mtDNA) to be
used as a molecular clock for estimating how long ago individuals or species have
shared a common ancestor.
Related Terms: Endosymbiotic theory, mtDNA, Symbiogenesis

Mixed martial arts

A fight sport allowing competitors to use a wide range of techniques, including kickboxing, wrestling, and other martial arts. Since the early 1990s, mixed martial arts have
become one of the fastest-growing sports, both at the professional and at the participatory
level, although the sport has sparked controversy because of its inherent violence and the
perception that contestants might be in danger.

Modern synthesis
Christened by evolutionary theorist, Julian Huxley, the modern synthesis was the
attempt in the twentieth century to combine Darwinian theory of natural selection with
emerging theory in population genetics. Prior to the modern synthesis in the 1930s and
1940s, some biologists thought that findings in genetics were inconsistent with Darwins
theories. Although there have been significant additions, the modern synthesis is still the
foundation for modern evolutionary theory.

Modular intelligence
Modular intelligence operates over a narrowly-defined domain, or a particular type of
function. Some animals are quite good at highly specialized cognitive tasks, but cannot
use their perceptions or intellectual abilities in novel fashion. One approach to
understanding human intelligence is to point out that humans, too, have modular forms of
intelligence, even a large collection of special-purpose, innate cognitive tools. In contrast,
other theories argue that human intelligence is more general purpose or that, although
specialized, our cognitive tools are not innate, but learned.
Related Terms: Neural reuse or neural recycling

mtDNA
mtDNA is mitochondrial DNA, the small circular chromosome found in the mitochondria
of our cells. mtDNA is distinctive from nuclear DNA in eukaryotic cells, providing some
sup- port to the theory of symbiogenesis. mtDNA is also inherited only from your
mother, so it can be used to estimate the relatedness of individuals (at least along this
maternal line), as the mtDNA only changes through mutation.
Related Terms: DNA, Mitochondria, Symbiogenesis

Mutation
A mutation is a spontaneous change to the sequence of nucleotides in DNA. A mutation
can occur because of damage to the DNA (such as from radiation), because of an error in
transcription or gene copying, or because of an accidental deletion, reversal, or insertion
of a part of a sequence of DNA. If a mutation occurs to a coding portion of the DNA, that
is, to a section that carries a gene for a protein, most often that mutation is detrimental. In
some cases, however, mutation may produce a genetic variation with a selective
advantage.
Related Terms: DNA, Recombination, VISTA

Neotony
The preservation of juvenile traits for an extended period, or even into adulthood, by an
organism. Neotony is one way that variation can be induced through changes in an
organisms developmental timing.

Neural reuse or neural recycling

The idea that novel human cognitive abilities make use of brain regions or circuits that
were used in our ancestors for other purposes, potentially biasing how they perform. For
example, neuroscientist Stanislas Dehaene has argued that learning to read requires
recycling regions of the brain used in other primates for small object recognition. The
neural reuse theory is one way to explain novel brain capacities in a modular brain. Other
theorists argue that neural reuse is just a subset of a larger pattern of exaptation found
throughout evolutionary theory.
Related Terms: Exaptation, Modular intelligence

Neuroanthropology
The anthropological study of human brain functioning, including especially a
consideration of the ways that human neurological traits underwrite human diversity, and
how diverse cultural and developmental contexts affect brain functioning. In some cases,
neuroanthropologists seek to use field research to test hypotheses generated by
laboratory-based neurosciences.

Niche creation (or niche construction)

A concept used by some evolutionary theorists to explain non-genetic ways that
organisms may pass on influences to their offspring, through modification of the
environment. In some cases, organisms build nests or structures that create conditions
that facilitate reproduction and infant development. In other cases, an organisms activity
modifies the environment so that the selective pressures on future generations will
change. In the case of humans, our ability to modify our niche has changed profoundly,
especially since agriculture, and moreso with industrialisation.
Related Terms: Cognitive niche

Norms
Social norms are collective principles, sometimes not made explicit, about how people
should behave or what they should believe. Although the processes of socialisation and
enculturation tend to make norms appear inevitable, or at the very least logical, they can
often differ significantly among groups. For example, in one group, the norms for
childcare might appear negligent or even cruel in another group.
Related Terms: Gender

Obstetrical dilemma
The conflict posed by the narrowing of the human pelvis, due to its adaptation for

bipedalism, and the growth of the human brain, leading to larger skulls in infants. These
two processes lead to a conflict around the size of the birth canal and potentially have
made child- birth more difficult for modern women than for our more ancient hominin
ancestors. For a long time, anthropologists thought that the obstetrical dilemma explained
the extremely altricial nature of human infants; they had to be born before their heads
grew too large to pass through the birth canal. Increasingly, we are also realising that the
metabolic demands of the growing fetus may pose a more serious limit on the length of
gestation prior to birth.
Related Terms: Altricial, Metabolic ceiling

Olduwan
Sometimes spelled, Oldowan. The oldest and simplest types of stone tools, sometimes
called Mode 1, are Olduwan tools, named for the Olduvai Gorge in Tanzanias Rift
Valley region where the first examples were found. Olduwan tools are characterised by
minimal working, usually along only one or two edges, and tend to vary with the type of
material being used. The oldest examples of Olduwan tools are 2.6 million years old, but
these were likely not the first tools; the evidence of tool use in other apes suggests that
the last common ancestor may have already used simple tools made from perishable
materials.
Related Terms: Acheulean

Parasite
A parasite is an organism that lives, in part, from a host, who does not gain a benefit from
the relationship. Some parasites require a host for a stage of their life. Unlike symbiots,
parasites tend to decrease the fitness of their host and, in extreme cases, may kill their
host. Many parasites are quite specialised, living only in a particular host species, and
have short life cycles so that the hosts defenses cannot evolve resistance to the parasite.
Related Terms: Symbiosis

Parental investment
Any energy, resources, time or other cost from a parent in the production and care for one
of its offspring. Theoretically, the minimal investment would be the time and energy to
copulate or produce sperm and egg, but in many species, the investment is much greater.
Parental investment helps us to understand sexual selection because the sex that must
invest more in reproduction acts as a limiting factor on the other sex in reproduction,
possibly leading to competition among members of the lessor-investing sex. Parental
investment also helps us to understand how evolution might produce high investment
reproductive strategies, if the result is a marked increase in survival or reproductive

success for the offspring.

Related Terms: Sexual selection, Trivers hypothesis

Parthenogenesis
Some species reproduce through parthenogenesis, a form of asexual reproduction in
which the embryo is grown from an unfertilised egg. In some species, parthenogenesis
produces a full clone of the mother, with eggs containing the full complement of the
mothers chromosomes.

Particulate inheritance
Discovered by Gregor Mendel, the theory of particulate inheritance explains that traits
are passed on to offspring through sexual reproduction, not by simply mixing the traits of
the parents, but by passing on discrete particles or factors of inheritance. We now refer to
these particles as genes. Mendel recognised that, even if a particle was not being
expressed (that is, was not shaping an organisms appearance), it might still be passed on,
only to reappear in the appearance of later offspring.
Related Terms: Dominant gene, Recessive

Permian-Triassic extinction
The largest extinction event yet on Earth, around 252 million years ago. Although
scientists disagree about the finer details, including the timing, rapidness, whether the
event was a single one or multiple pulses of extinction, and the causes, evidence does
suggest that severe climate change was involved. The extinction event may have been the
result of a combination of forces, including a major asteroid impact, a period of increased
volcanic activity, or the release of stored chemicals in the Earths crust. The most severe
extinction occurred in the oceans, as chemical changes in the atmosphere led to increased
levels of carbon dioxide and decreased oxygen.

Photosynthesis
The process used by plants and other photosynthetic life to create chemical energy from
light, building carbohydrates like sugars from combining carbon-dioxide and water. In
plants, photosynthesis usually happens through a green pigment called chlorophyl in the
chloroplasts in the plant cells.
Related Terms: Chloroplasts

Pigmentation
Human skin pigmentation or colouration is provided by different types of melanin, a
naturally-occurring dye. Human skin pigmentation mostly likely became darker as our
ancestors body hair decreased and became finer than other apes as the colour provided
protection from equatorial sun. In contrast, other apes have light-coloured skin under
their fur (al- though their faces and hands can also be quite dark).
Related Terms: Melanin

Placenta
A structure in the uterus that connects the growing embryo to the mothers body.
Nutrients and oxygen pass through the placenta to the embryo, and waste products are
removed from the babys body through the placenta. The human placenta is particularly
invasive, penetrating more deeply into the mothers endometrium, rerouting some of
her arterial blood flow, than happens in some other mammals.
Related Terms: Marsupials, Placental mammals

Placental mammals
One of the two major branches of the mammalian family; the other is composed mostly
of the marsupials. Some other animals, including some reptiles, have structures that
resemble placentas, but a true placenta is one of hallmarks of placental mammals.
Related Terms: Marsupials, Placenta

Polyandry
A relationship in which a single female mates with multiple males. In some human
societies, marital relationships can be polyandrous, with women taking more than one
husband, in some cases, marrying brothers. Polyandry is one form of the more general
phenomenon: polygamy, or systems in which individuals can take multiple spouses.
Related Terms: Polygyny

Polygenic
A polygenic trait is controlled by two or more genes. In some cases, a large number of
genes can affect a trait, and those genes can themselves be pleiotropic, affecting multiple
traits. The point is that the folk understanding of genes for example, that for every trait
there is one and only one gene can often be markedly different from the way the

genome actually functions. Polygenic traits are often continuous, rather than discreet.
Related Terms: Continuous

Polygyny
Polygyny is a system of mating or marriage in which a single male individual takes
multiple female mates or wives. Polygyny is a more specific term than polygamy,
which includes both polygyny and polyandry. Polygyny in animals can often lead to
direct competition among males as a dominant male may seek to monopolise a number of
females, preventing rivals from mating with them.
Related Terms: Polyandry

Population
A population is group or whole species, including all living members. We use the term in
anthropology to focus our attention on the group as a whole, including its internal
variation and even unexpressed genetic traits. As individuals are born, grow, reproduce
and eventually die, the population as a whole evolves, with traits that are selected for
becoming more pervasive and traits that are being selected against decreasing in
frequency or even disappearing.
Related Terms: Gene pool

Precocial
An adjective used to describe infants that are neurologically and behaviourally mature,
especially that are able to achieve some degree of self-sufficiency or ability to participate
in a group similar to an adult at an early stage. Precocial infants, like horses, cows and
sheep, al- though small at birth, are often able to keep up with adult members of their
species quite quickly.
Related Terms: Altricial

Pro-social
An adjective that describes the extreme social nature, including cooperation and social
intelligence, in some species that cannot simply be explained as the pursuit of ones own
self interest. Some anthropologists argue that pro-social traits are not strictly beneficial
that is, a pro-social animal may be more empathetic or altruistic than strictly
advantageous for its own survival and reproduction because the overall payoff in
terms of selective advantage outweighs the individual cost of altruistic acts.

Pseudogenes
Pseudogenes resemble genes that code for proteins, but they have accumulated enough
mutations that they no longer function to produce those proteins. Pseudogenes are
sometimes taken as evidence that the protein for which they coded is no longer necessary
for the species survival nor sufficiently advantageous to prevent the transmission and
spread of the now-dysfunctional version of the gene (because of gene duplication, some
pseudogenes are former extra copies of still-functioning genes). Although they do not
code for proteins, pseudogenes sometimes serve other functions or affect the functioning
of other genes.

Punctuated equilibrium
Punctuated equilibrium is an evolutionary model suggesting that, over time, species can
re- main consistent for long periods of time, punctuated occasionally by periods of rapid
change, especially when selective pressures change or a new variant arises within a
species with a significant advantage. First proposed by paleontologists Niles Eldredge
and Stephen Jay Gould, the punctuated equilibrium model argued against the assumption
that evolutionary change would always be gradual and consistent, also seeking to explain
why some species often appeared to be unchanged for very long periods in the fossil
record.

Ratchet effect
A ratchet is a type of wrench or lever that can only turn in one direction; a locking
mechanism prevents the bolt or pin being turned from reversing and undoing its progress.
Psychologist and evolutionary theorist Michael Tomasello used the term ratchet effect
to describe a cultural process through which technology, invention and knowledge no
longer get lost with the death of a generation of individuals because the mechanisms for
cultural transmission are sufficiently strong to prevent their loss. Advanced cultural
learning, including lateral cultural transmission, meant that human knowledge became
shared and cumulative, allowing our species to make great technological advances.
Related Terms: Cognitive niche, Lateral transmission

Recessive
A recessive gene is a gene that only affects phenotype, or the appearance of an organism,
in the absence of the dominant gene. Usually, this requires that the offspring receive a
recessive gene from both parents because the presence of a single dominant gene will
mask the presence of the recessive variant although, because of particulate inheritance,
the recessive trait might still be passed on to offspring.

Related Terms: Allele, Discreet, Dominant gene, Particulate inheritance

Recombination
The production of new, novel DNA sequences through reproduction, either sexually or
asexually. During meiosis, the cell division process that produces germ cells (sperm and
egg) with only half of a full complement of chromosomes, parts of the chromosomes can
cross over, exchanging genetic information. The process can lead to novel sequences
and combinations of the two parents traits.
Related Terms: Red Queen theory, Mutation, Sex

Red Queen theory

Taking its name from Alices race with the Red Queen, in Lewis Carrolls Through the
Looking Glass, the Red Queen theory points out that, in evolution, species must evolve
and change constantly because other species, including competitors, predators, and
parasites are all evolving at the same time. The Red Queen theory explains why species
that were once well adapted can still go extinct, and why sexual reproduction, including
the variation produced by genetic recombination, may provide a decisive advantage for a
sexually-reproducing species.
Related Terms: Recombination

Retrovirus
An RNA-based virus that builds a DNA version of itself (the reverse of the usual process,
thus the name retro-virus), so that it can insert itself into the nuclear DNA of a cell. A
special variant of a retrovirus is an endogenous retrovirus, a virus that successfully writes
itself into the germ-line DNA, thus being there from the conception of the host.
Related Terms: Endogenous, RNA

RNA
Ribonucleic acid (RNA) is a family of large biological molecules like DNA. Unlike
DNA, RNA tends to be a single-strand structure of nucleotides. RNA serves to send
messages and mediate the actions of DNA, controlling gene expression, signaling within
a cell, or helping to build proteins from the coding portions of DNA. Some viruses are
RNA strands, rather than DNA.
Related Terms: Retrovirus

Secondary sexual characteristics

Traits that differ between males and females of a species, but are not directly linked to
reproduction are called secondary sexual characteristics. For example, in humans, men
and women differ on a range of traits not directly linked to reproduction, such as the
distribution of fat on their bodies, the over-body musculature, and the pattern of hair
growth on their bodies. In some cases, secondary sexual characteristics are the result of
sexual selection.
Related Terms: Sexual selection

Sex
In species that reproduce by combining gametes to form offspring, some contribute
identical gametes (isogamy). When the two types do not contribute similar gametes
they are characterised by anisogamy, or difference between sperm and egg -- the sex
contributing the larger gamete or egg is female, and the sex contributing the smaller
gamete or sperm is male. In many animals, females possess XX chromosomes, and
males possess the XY chromosomes (birds, some fish and crustaceans have a different
chromosomal mechanism for determining sex).
Related Terms: Anisogamy, Estrogen, Gender, Hermaphrodites, Recombination, Sexdetermining region of the Y chromosome (SRY), Testosterone

Sex-determining region of the Y chromosome (SRY)

The sex-determining region of the Y chromosome, or SRY, codes for a protein that
initiates development of the male testes. SRY is (as its name suggests) located on the Y
chromosome, so is present in most males; if the gene has a mutation or is defective, even
though a persons body has the Y chromosome, male genitalia will not develop (one form
of this is Swyer syndrome, in which individuals look female but are sterile). A woman
with XX chromosomes (normal) can have the SRY gene, leading to de la Chapelle
syndrome, a rare chromsomal disorder in which individuals appear male but are infertile,
and some have other physiological feminsation. SRY is not the only gene that can
determine male sex, as other mechanisms do so in some species.
Related Terms: Estrogen, Hermaphrodites, Sex, Testosterone

Sexual revolution
Usually, social scientists use the term sexual revolution to refer to the period from the
1960s through 1980s, during which changes in reproductive technologies (such as birth

control), human rights, social mores, and laws in many Western countries led to a rapid
change in sexual behaviour, including increased divorce, sex prior to marriage, increased
numbers of sexual partners, greater liberty for women, and decreased repression of
homosexuality and other alternative lifestyles.
In this course, we argue that humanity has undergone a number of sexual revolutions,
including a radical but slower change in the way that our ancestors reproduced to support
our increasingly altricial infants, more complex and enduring social networks, and
changed pat- terns of childrearing. Biological factors such as decreased sexual
dimorphism and concealed estrus, as well as less evidence for direct physical competition
or sperm competition among hominin ancestors unlike gorillas and chimpanzees, for
example also point to this distinctively human type of reproductive strategy.
Related Terms: Altricial, Concealed estrus, Dimorphism, Direct competition, Sperm
competition

Sexual selection
An intraspecific form of evolutionary competition in which individuals compete to outreproduce other members of their sex. Sexual selection helps explain why members of a
species may develop traits that are not necessary or advantageous for survival; they may
be used, instead, to compete for mates or to reproduce more. Sexual selection takes two
forms: direct competition and mate choice. One hallmark of sexual selection is that the
process tends to produce dimorphism, as the selective pressures in reproduction on males
and females are not the same.
Related Terms: Dimorphism, Direct competition, Intraspecific competition, Mate choice,
Parental investment, Secondary sexual characteristics, Sperm competition, Trivers
hypothesis

Social cognition
Forms of intelligence, perception, memory and other cognitive abilities that are used
specifically in relations with conspecifics, or members of the same species. Humans
appear to have more sophisticated social cognition than other apes even at a very young
age, a gap even greater than that found in our ability to anticipate cause-effect, for
example, in the physical world. The greater social cognition may be a result of intense
intraspecific social competition, mate choice, or inter-group competition, with benefits
for those hominins who were better able to perceive each others thoughts and intentions,
strategise, and cooperate.
Related Terms: Emotionally modern, Machiavellian intelligence

Speciation

The evolutionary process that turns a population, originally of a single species, into two
or more distinct species, usually through a combination of reproductive isolation (which
halts gene flow) and diversifying selection or genetic polymorphisms. Often, determining
precisely when speciation has occurred is difficult because populations may be
geographically isolated, for example, if a population has occupied a new niche.
Related Terms: Diversifying selection

Sperm competition
A form of direct sexual competition in which, because females mate with multiple males
while fertile, male sperm competes directly with other males sperm. The result can be
extremely high sperm production or defensive mechanisms like semen that acts to plug
the vagina or attacks subsequent sperm. Many primatologists point to the large testes of
male chimpanzees, and the fact that females copulate with many males while in estrous,
as an example of sperm competition.
Related Terms: Sexual revolution, Sexual selection

Stabilising selection
Stabilising selection leads to a decrease in genetic variation over time as selective
pressures produce an ideal state for a trait, making variation a selective disadvantage.
Stabilising selection can produce long-term consistency in a species trait, unlike
directional or diversifying selection. Because most traits in organisms are consistent over
time, stabilising selection is thought to be the most common type of selective pressure.
Related Terms: Directional selection, Diversifying selection

Symbiogenesis
The merging of two separate, symbiotic species into a single organism, first proposed by
Konstantin Mereschkowsky and later advocated by Lynn Margulis. Advocates of
symbiogenesis argue that slow genetic change through mutation and selection is
insufficient to ex- plain evolution, especially the rise of multi-cellular organisms and
complex biological structures. Symbiogenesis gained additional support from the close
study of chloroplasts and mitochondria, two organelles found in complex cells, which
suggests a symbiogenetic origin for eukaryotic life.
Related Terms: Chloroplasts, Endosymbiotic theory, Eukaryotic cells, Mitochondria,
mtDNA, Symbiosis

Symbiosis

A close, long-term relationship between two species in which both gain benefits from the
presence of the other. In some cases, symbiosis is obligatory: one species cannot survive
without the other. Symbiosis can include a species living on or inside another species, or
less intimate relationship.
Related Terms: Endosymbiotic theory, Microbiome, Parasite, Symbiogenesis

Teleological error
Telos is the Ancient Greek word for goal or purpose. The teleological error in
evolutionary thinking is assuming that evolution itself has a goal, or that evolution has a
clear direction or inevitability, which is most likely in thinking of humanity is the end
or highest expression of evolution. As a natural process, evolution does not have a
design, that is, a fore- thought or intentional plan. Thinking teleologically about
evolution, including assuming that humanity is the highest expression of evolution, a
form of anthropocentric thinking, is a common error.
Related Terms: Anthropocentric, Great Chain of Being, Kluge

Testosterone
Testosterone is a steroid hormone of the androgen group, found in a wide range of
vertebrates. Testosterone is secreted primarily by the testes in men, and by the ovaries in
women although some is also produced by the adrenal gland. Testosterone is essential in
the masculinisation process through which male bodies form and mature, and testosterone
levels tend to be around seven or eight times higher in men than in women, although they
fluctuate in both.
Related Terms: Estrogen, Sex, Sex-determining region of the Y chromosome (SRY)

Trivers hypothesis
Robert L. Trivers recognised that, because females typically invest more in their
offspring (see parental investment), males will have to compete for a chance to
reproduce, unless the parental investment of males is as high or higher than female
parental investment. Trivers hypothesis predicts which sex will be subject to sexual
selection, helping evolutionary theorists to better understand intraspecific competition in
humans and others species.
Related Terms: Intraspecific competition, Parental investment, Sexual selection

Vestigial organ

Vestigial organs have lost a particular function or changed function significantly over the
course of an organisms evolutionary history. Although some people think of vestigial
organs as useless, even organs that have lost a major function often continue to serve a
different purpose subsequently. In humans, for example, the appendix has lost most of its
older function in the digestive system, but it continues to serve an immunological
function and also can provide a reservoir of gut flora in event of a serious gastrointestinal infection. Unlike atavisms, vestigial organs are common to all or nearly all
members of a species.
Related Terms: Atavism

VISTA
Variation + Inheritance + Selection over Time leads to Adaptation
One way to remember the factors that need to be present for natural selection to occur.
The VISTA acronym highlights the roles of variation and the necessity of inheritance; the
absence of either (such as non-heritable variation) prevent adaptation from occurring
through natural selection.
Related Terms: Fitness, Mutation

Weaning
The transition from nursing to subsisting on an adult diet and eating independently in
mammals. The appropriate age to wean humans is the subject of great controversy, and
norms for weaning as well as strategies to do it differ widely between and within
many cultures. In Australia, almost half of all children are weaned by six months of age,
the minimum recommended by the World Health Organisation, with some starting to
wean their children al- most immediately after birth. Anthropologist Kathy Dettwyler,
using comparative research with other species and growth trends in humans, suggests that
a minimum age for weaning in our hominin ancestors may have been around 2.5 years,
but with a maximum of 7 years of age.

Weismann barrier
The Weismann barrier is a principle argued by August Weismann that germ cells create
body cells, but that what happens to bodily cells (somatic cells) cannot influence the
constitution of germ cells. Put simply, nothing you do during your life except survive and
reproduce influences your offspring because the germ cells in your sperm or eggs is
unchanged byyour experience. The Weismann barrier was a definitive rejection of
Lamarcks principle of the inheritance of acquired traits. The Weismann barrier was once
thought to be unbreachable, but new findings, especially in epigenetics, suggest that some
acquired traits may affect germ cells or transmit mechanisms that affect gene expression.

Related Terms: Bodily cells, Epigenetics, Inheritance of acquired traits

Wernickes area
A region of the human brains cortex behind the ear on the dominant side (left
hemisphere in most people) that has an important role in the comprehension of spoken
language. The area is name for German psychiatrist and neurologist Carl Wernicke, who
first hypthesised about the regions role in speech comprehension after examining
patients with brain injuries leading to aphasia, a disruption in speech abilities that does
not affect other forms of intelligence. In particular, a lesion in Wernickes area can lead
to Wernickes aphasia, an inability to understand spoken language while still being able
to speak.
Related Terms: Brocas area

Xerophyte
An organism adapted to survival in extremely dry environments (deserts).

###

About the author

Greg Downey is Associate Professor in Anthropology at Macquarie University in
Sydney, Australia, where he teaches about human evolution, psychological and economic
anthropology, and human rights. Greg did his doctoral research at the University of
Chicago before teaching at the University of Notre Dame (USA). He relocated to
Australia in 2005.
Greg has done research on capoeira in Brazil, with mixed martial arts fighters in the US,
and with rugby players and coaches in Australia. He writes extensively on the
relationship between brain and culture, especially skill acquisition, sensory training, and
developmental dynamics. You can read about his work on the weblog Neuroanthropology
at the Public Library of Science (PLOS). This book contains some links to his relevant
online writing at the PLOS Neuroanthropology weblog.
Greg is co-editor, with Daniel Lende, of the recent book, The Encultured Brain: An
Introduction to Neuroanthropology (MIT Press). Gregs previous books include
Learning Capoeira: Lessons in Cunning from an Afro-Brazilian Art (Oxford University
Press) and, with Melissa Fisher, Frontiers of Capital: Ethnographic Reflections on the
New Economy (Duke University Press).

When hes not being an anthropologist, Greg, his wife, Tonia, and their daughter,
Mikhaela, breed horses on the south coast of New South Wales, helped with unceasing
enthusiasm by their dogs, Louis and Roxy. Although proudly from St. Louis in the U.S.,
Gregs now a dinky die Australian, even a mad keen cricket fan.
If you enjoyed this book, please consider learning more, either about the author, Greg
Downey, or about Enculture Press.
You can learn more online at:
Twitter: http://twitter.com/gregdowney1
Neuroanthropology: http://blogs.plos.org/neuroanthropology
Enculture Press: http://enculturepress.com

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