Evolution: a primer
We are composed of cells that stick together like building blocks to create a living, functioning organism. There are more than 200 types of cells in humans including skin cells, liver cells, nerve cells, muscle cells, intestinal lining cells, bone cells and fat cells. These cells can group together to create tissues, which are ensembles of similar cells that collectively perform some function. For example, muscle tissue can contract and, by working against our skeleton (itself made of bone tissue), allow us to move, while lung tissue enables us to exchange oxygen and carbon dioxide with the atmosphere. The grouping of different tissues together creates complex organs, like the heart, lungs, stomach and skin, that are capable of performing all the functions we need to maintain life.
The organisation of around 30 trillion cells (see Chapter 4) into a living, breathing, fighting, feeding, fleeing, breeding, singing and dancing body relies on a tightly controlled and spectacularly diverse choreography of molecular interactions within our cells. The chemical reactions that digest our food, transform molecules and allow us to perform the functions we need to stay alive are collectively termed our metabolism. These reactions must take place at the right place and time, and at rates that are neither too fast nor too slow. To achieve this organisation and control, our cells contain complex networks of protein microtubules and fatty membranes, a host of enzymes and an array of tiny structures called organelles.
DNA is at the centre of all this complexity. This famous double-helix molecule is found within the nucleus, a large organelle found in most of our cells. A DNA molecule looks like a twisted ladder with each ‘rung’ on the ladder being formed from a pair of so-called bases, one on each ‘upright’, that reach across and weakly bond with each other. Each base is effectively a letter in the genetic code that directs our cells to make proteins. There are four bases: adenine, thymine, guanine and cytosine, often known by their initials A, T, G and C. The sequence of bases along the DNA molecule is a code that is used by structures in our cells called ribosomes to make molecules called proteins. Proteins have a huge range of functions in our body. They can be structural, like the collagen in our skin or the keratin of our hair; they are the bulk of our muscles; and, as hormones and enzymes, they help to control the many chemical reactions throughout our body. Proteins are built up from long chains of small chemical building blocks called amino acids. These amino acids (there are more than 20 to choose from) join together in a very specific sequence, which determines the properties and the function of the resulting protein. This precise sequence of amino acids is dictated by the order of letters within the DNA molecule. Each amino acid in the protein is coded for by a specific sequence of three bases, called a codon. All of this complexity and organisation arose through the process of evolution.
When we think of human evolution we are usually preoccupied with the more recent stages of our development as a species: characteristics like walking on two legs (bipedalism), our large brain and the evolution of language. These features are important, indeed defining, but we must also understand that most other aspects of our biology have been subject to evolution, in some cases occurring very early in the beginning stages of life itself. The wonderfully complex processes underpinning our metabolism, our nervous system and the synthesis of the proteins that build and control our bodies all evolved long before our ability to walk upright and talk about the weather. For example, the sequence of 10 reactions that comprise a process in our cells called glycolysis occurs in the same way in us, in pandas, in yeast and in bacteria. Glycolysis is the first step in converting the sugar glucose (from our food) into adenosine triphosphate (ATP), a kind of wonder-molecule that delivers energy to cells. This series of reactions and the enzymes that control it evolved long before skeletons and muscles, eyes and ears, logic and language. Likewise, the way our muscles contract at a molecular level involves biochemical pathways and molecules that can also be found in sea anemones, starfish and earthworms. The mechanics of our skeleton, how we digest food, the ways we respond to stress and a whole host of other processes happening at the level of cells, tissues and organs are fundamentally identical to those found in other animals, and indeed in some cases plants, fungi, single-celled organisms and bacteria. That is not to say that the last couple of million years of our evolution haven’t resulted in more than a few tweaks here and there, but for the most part the basics were laid down a very long time before we came onto the scene.
Biochemical pathways, enzymes, different cell types, bird bills, fish scales, hedgehog prickles and behaviour are all examples of adaptations: characteristics (or traits, to use the evolutionary term) that benefit those individuals possessing them. The development of such traits is usually moderated by specific sequences of DNA that we call genes. We have two copies of each gene, inheriting one from our mother via her egg and the other from our father through the DNA in his sperm. A sequence of DNA ‘letters’ making up a gene (or genes) results in some property of the individual bearing those genes. This property might result in some individuals surviving and breeding better than others who lack the gene. The fact that genes can be passed on from one generation to the next, and will be present in more offspring of successful parents, is what leads to evolution.
Evolution implies change and biological evolution is, at its most fundamental, a change in gene frequency over time. In other words, evolution occurs when genes become more or less common. Different forms of genes arise through mutations, changes in the sequences of DNA bases in the eggs or sperm of parents that are subsequently passed on to offspring. Such changes occur when mistakes happen in the process of copying DNA to make eggs and sperm, and mutations happen naturally at a low but appreciable rate. Many mutations have negative effects, but every so often a mutation may arise that provides some new way of doing things, some advantage to the offspring that inherit it. For example, perhaps a mutation slightly alters the enzyme for which it codes and the new version of this enzyme is just a little more effective in digesting food or, as we will see in Chapter 8, much more effective in breaking down alcohol.
One of the main ways in which gene frequencies can change, and therefore that evolution can occur, is natural selection. To understand how natural selection results in evolution let’s imagine a gene exists for some adaptation that causes individuals that have it to do ‘better’ than those that do not. We can define ‘better’ in many ways: perhaps individuals with our imaginary gene can gain an advantage in finding food, evading a predator or securing a mate. In evolutionary biology though, these abilities are all just proxies for the only currency that really counts: the ability to leave more offspring in the next generation than others manage. The relative ability to leave offspring in the next generation is what evolutionary biologists call fitness, although being ‘fitter’ is more complex than simply having more offspring than your neighbour. For example, a red deer female in good condition and capable of producing plenty of high-quality milk might achieve greater fitness by having a son rather than a daughter. Her excellent condition allows her son to be well fed, and by growing large and strong he stands a good chance of becoming a dominant male controlling the reproduction of a large harem of females. By having a son, the mother gains more grand-offspring and accrues greater fitness than if she had a daughter. Likewise, a nesting bird that lays a large number of eggs may end up with far more offspring leaving the nest than her neighbour with a more cautious egg-laying strategy. However, the fact that the ambitious mother has been run ragged feeding all of her chicks means that she may not have enough reserves to live through a cold winter to breed again next spring. Her death reduces her future fitness to zero. What is more, her many offspring may also not be in good enough condition to survive the winter. By balancing current reproductive output with the chance to breed again next year, and by having a smaller number of offspring that can each receive higher quality care, the mother with fewer offspring this year might actually achieve greater fitness over her lifetime.
Let’s assume that, all other things being equal, a gene leads to higher fitness for those that are lucky enough to have a copy of it. These individuals will have more offspring as a consequence of having that gene. Because these offspring have a good chance of inheriting the gene from their successful parents the gene will likely increase in frequency in the next generation. If the environment in which these individuals live, which includes physical factors such as temperature as well as biological interactions with predators, prey and parasites, doesn’t change, then it is quite likely that this new generation will also benefit from the gene they inherited. These offspring will go on to have their own offspring that will also have a chance of inheriting further copies of the gene. The inescapable, beautiful, logical result of these bouts of reproduction and natural selection across generations is that genes that confer a benefit to the individuals bearing them tend to increase in frequency. If on the other hand the selection environment changes, then genes (and for many traits we are usually talking about multiple genes) that code for particular adaptations may no longer be as beneficial to their bearers. In these circumstances we will see a reduction in the frequency of less beneficial genes, and a consequent reduction in the number of individuals with the adaptation that they code for. In other words, through natural selection, gene frequencies change and evolution occurs. This process happens whether you are a human drinking a martini on a luxury yacht or a bacterium digesting dog mess.
The evolution of modern humans
Natural selection can lead to the evolution of adaptations by selecting those individuals with genes coding for traits that increase survival and reproduction, as we have already seen. Evolution can also lead to the formation of different species through processes collectively known as speciation.
Only rarely will all the individuals that make up a species be able to interbreed freely with each other. An example would be a species found only on a small island. Most species, including humans, exist in different populations of individuals that tend to breed among themselves. The birds in your garden might be the same species as the birds in the garden of someone 1,000 miles away but in practice these two groups of birds will probably not breed with each other, even though, in principle, they could. Sometimes migration of individuals to and from different populations allows for interbreeding and gene flow between populations, but overall most individuals are likely to breed with members of their species that live close by.
Different environmental pressures acting on populations living in different locations, or being active at a different time of the day or year, can cause selection for different adaptations in those populations. Perhaps in one valley where a species of mouse is found there is a different predator eating mice than is found in the rest of the mouse species’ range. The presence of this unusual predator has selected for more nocturnal behaviour than is usually found in the mice because mice that prefer to come out to forage when it is dark survive far better than those favouring daylight. The valley population was already perhaps somewhat geographically isolated from others of the same species. Now, the population becomes further isolated biologically because even nocturnal individuals that disperse outside the valley are unlikely to be able to meet and mate with the non-nocturnal individuals present in the rest of the species’ range. Over generations, further genetic differences can accrue as the nocturnal population evolves under the influence of the selection pressures of its new nocturnal niche. We might see the evolution of better low-light vision for example, or slightly different dentition to favour food more readily found at night. These genetic differences further increase the relative isolation of this population with respect to the rest of the species. Eventually the valley population becomes sufficiently distinct physically, ecologically and behaviourally for us to identify it as a new species.
To understand how our species, Homo sapiens, is affected by the evolution of adaptations unique to our species and by those adaptations that are common to a much wider group of organisms requires us to examine our evolutionary history. The problem is that our evolutionary history extends all the way back to the very start of life. Clearly, we are going to need to draw the line somewhere. To keep things manageable and relevant, I’m going to constrain this history to the emergence of what paleoanthropologists call anatomically modern humans. When necessary, for example when discussing the evolution of the mechanisms that lead to stress (Chapter 5), violence (Chapter 7) or addiction (Chapter 8), it might be necessary to delve a little deeper.
To define a species, and to classify it clearly and unambiguously as different from any other species, we need to define the characteristics that are shared by members of that species but not present in any other group, at least not in the same complete combination. You might think that defining a ‘human’ is neither especially difficult nor important. We never find ourselves in the position of wondering whether any given collection of flesh and bones in front of us is a human being or not and we don’t need to tally up anatomical features to be sure. In terms of an effect on our everyday lives, having a firmly nailed-down definition of what makes us part of the species Homo sapiens is indeed pretty inconsequential. However, by understanding the very specific features that make us ‘human’ we are much better armed to consider the role that those evolved features have in our everyday lives.
One problem with defining any species is that individuals vary. Humans are no different and any definition of our own species must account for this variation. Some of the variation in humans arises more or less solely from genetic differences between populations. Such differences might be highly apparent: the differences in skin colour between a white European and an African-American for example, or between the stocky stature and physique of an Inuit indigenous to Nunavut in northern Canada and the slimmer form of the Sān peoples of Namibia. You don’t need to travel to see other clear genetic differences between humans. The most profound variation is between males and females and these differences are also genetic, caused by the presence of a Y chromosome (male) or an X chromosome (female) in the sperm that fertilised the egg from which we developed. Other genetic differences might be less apparent and related to disease susceptibility or metabolism. A good example of a metabolic difference between populations that is genetic is the way our bodies deal with lactose in milk and gluten in wheat (Chapter 3).
While some variation is certainly genetic, other sources of variation are solely environmental. A good example of a highly variable characteristic that is determined purely by our environment is the primary way that we communicate. People growing up in different places and cultures speak different languages. While our linguistic ability is genetic and related to physical structures like the larynx (the ‘voice box’) and tongue, and to specific regions of our brain, the precise form our language takes is related to what language we hear as we develop from birth.
Yet more variation can arise from environment and genes interacting. Adult height, for example, has a genetic component, but without an adequate diet during childhood an individual bearing genes promoting tallness is unlikely to achieve their full potential height. The balance of genetics and environment, of ‘nature’ and ‘nurture’, is of great interest both scientifically and philosophically. It is also a topic of great relevance to this book, because evolution is fundamentally a genetic process. If we are to discuss the role that evolution has had in shaping us, and the mismatch between our evolved selves and the modern world, we are inevitably making assumptions that whatever features we are discussing have a genetic basis. As we will see throughout this book, despite being extremely seductive, such assumptions are not always easy to substantiate. So ‘nature’ is going to be critically important to our discussions, but so too is ‘nurture’, since it is the interaction between our present-day environment and our genes that leads to many of the mismatches that we see.
Currently, there is only one living species in our genus (Homo), but in the geologically very recent past there were other Homo species wandering around that are now extinct, most notably Homo neanderthalensis. We know from genetic studies of modern humans and of well-preserved material from recent fossil records that Neanderthals and early humans interbred. We have also found other fossil material that indicates there were more Homo species in the mix, including Homo naledi in South Africa and Homo floresiensis, the famous ‘hobbit’ found in Flores, Indonesia. These other hominins, as members of our genus are known, were probably also interbreeding with each other in the run-up to the emergence of anatomically modern Homo sapiens. The piecing together of the emergence of anatomically modern humans and the reconstruction of our recent evolutionary past is a difficult, controversial and often fast-moving field; the discovery of a bone fragment here or a tooth there usually results in a big media splash and yet another rethink of sequences and dates. In short, it’s a bit of a mess and it’s likely to remain so for some time. Despite this complexity and confusion over some of the details of our recent evolution, we can reasonably safely say that archaic Homo sapiens were first striding around the Earth about 300,000 years ago and that fully anatomically modern humans arose around 160,000 years ago. The balance of evidence currently points to an African origin with a subsequent spread of our species into Europe, Asia and beyond. At the same time that our earliest modern human forebears were emerging in Africa, other Homo species were evident in more far-flung locations. Footprints of Homo heidelbergensis have been found in southern Italy for example, and Neanderthals were widespread across Europe and southern Asia as long ago as 400,000 years.
What makes us different
The anatomically modern humans that emerged around 160,000 years ago have numerous shared characteristics that differ from other hominins. First and foremost, the skull is a very distinctive shape. Unlike all other hominins (and indeed primates generally) we have a very weakly developed supraorbital arch, or brow ridge. A brow ridge is, however, present in remains of earlier Homo sapiens (those dating from between 160,000 and 300,000 years ago) and can also be seen in some humans alive today. Aboriginal peoples of Australia, for example, often exhibit a brow ridge, but it is very different from the ridge found in Neanderthals and other archaic members of the genus Homo. Where present, modern brow ridges are not generally complete across the eyes, and it is usually only the central section that is visible. The function of the brow ridge is to bolster the skull, providing structural reinforcement against the powerful forces generated by working the bottom jaw when chewing. It was a feature that was lost late in human evolution and its loss is related to another prominent feature of our skulls, a steep forehead.
Our almost vertical foreheads are in stark contrast to the sloping foreheads of other hominins. A surprisingly complex part of our anatomy, the forehead has a series of three muscles that allow us to display a range of expressions including quizzical and surprised. That the forehead is a useful billboard for our emotions, at least for those shunning the Botox needle, is a great example of evolution co-opting something that arose for an entirely different purpose, in this case related to housing our large and complex brain. To accommodate such an organ required a different type of space from the one that the sloping forehead of other hominins provided. Before we start getting too carried away about our big brains, though, it is worth noting that Neanderthals actually had a slightly larger brain. The general consensus from studies of their skulls suggests that Neanderthals were able to accommodate such a large brain despite possessing a sloping forehead by having skulls that extended lengthways as opposed to upwards, creating the more spherical skull shape that we have. It is this ‘rugby ball’ versus ‘football’ design that reveals the key advance in the modern human brain. Our brain has enlarged frontal lobes and it is this part of the brain that is heavily involved in all the cognitive processes that we tend to associate with modern humans: decision-making, planning, creativity, social behaviour and abstract thought. A big frontal lobe extends the brain forwards and upwards and to accommodate that you need a steep vertical forehead. That big frontal lobe was a driving force in our success and the sometimes subtle effects that it has on our fit to the modern world will be seen throughout this book.
Exemplified by the loss of the brow ridge, the overall structure of our skull is more delicate and fragile than the thicker and more robust skulls of other hominins. We have more delicate lower jaws with smaller teeth, especially our canines and incisors. We also have a pronounced chin that combined with our relatively short jaws and steep foreheads makes our proportionally smaller faces almost vertical. This gracility extends to the rest of our skeleton, with relatively longer limbs and thinner bones than any other hominin yet discovered. The extra length of our arms relative to our bodies does not come from equal (or isometric) growth of the limb bones but via disproportionate (or allometric) growth of those long bones nearest the hands (ulna and radius) and the feet (tibia and fibula). Overall, our build differs from the robust build of other hominins and it is thought that these more slender proportions are an adaptation to life in warm tropical regions. As a body becomes less stocky and more slender it reduces its surface area relative to its volume and loses more heat, a useful adaptation for an active animal in very hot regions.
Interestingly, stockier proportions are apparent in populations of humans who evolved to live in the cold northern Polar Regions today where, conversely to living in the tropics, heat retention is a good thing. This is sometimes said to be an example of Bergmann’s rule, one formulation of which states that individuals within a species (especially warm-blooded mammals and birds) tend to be larger at higher latitudes.* For humans though, it is not body size but body shape that is the key player here and Allen’s rule, that body shape tends to be more rounded and compact in colder climates, is a better fit.† That we can understand human evolution and variation using the same rules that we apply to other species further reinforces the point that we are just animals, subject to the same pressures of selection and processes of evolution as all other organisms.
An obvious feature of hominins compared to most animals is bipedalism, the ability to walk upright on two legs as the preferred gait. Bipedalism is not unique to hominins, as even the most casual observation of birds reveals, but it is unusual in mammals. Kangaroos and wallabies, for example, are bipedal and so too are the springhares, a curious group of African rodents superficially resembling rabbits. A number of mammals, including bears and many primates, can walk on two legs for a short time and some tree-dwelling primates, notably the gibbons, exclusively use a bipedal gait when walking on the ground. Outside of the mammals and birds, basilisk lizards can run on their back legs at such a rate that they can achieve the feat of walking on water (a feat that earns them the name Jesus Christ lizards), but this can only be maintained for short periods. The elegant, permanent bipedalism of humans is only possible because of the configuration of our backs, pelvis and supporting ligaments, tendons and muscles. To walk upright in any sort of convincing fashion it is necessary to take long strides without falling over. This is far from easy and the problem of bipedalism still vexes many of those working in the robotics industry although (pun intended) great steps forward have been taken and we can now see the development of some very effective bipedal robots. Key to solving the problem of bipedalism is finding a way to balance the upper body over the legs in a manner that allows the legs to move and develop a stride pattern without toppling. We achieve this mainly through a curved spine and changes to our pelvis that together make our skeletons very distinctive.
So far all the features we have explored that define humans have been skeletal. Clearly, our skeletons and related features are of great importance in defining our physical abilities and limitations, but modern humans cannot be defined by bones alone. Complex tools, the use of fire, the development of art, music, language and abstract thought are all human cultural features that have resulted from our large, complex and powerful brain. Given that we are often interested in comparing modern humans with other hominins, a focus on skeletons, hard parts that fossilise and persist until their discovery, is unsurprising. Increasingly though we are taking more interest in whatever products of hominin culture we are able to discern from the sparse archaeological and fossil record. The findings are often surprising. Burial rituals equating to what we would surely regard as a funeral involving fire and the seemingly symbolic use of animal horns can be inferred from Neanderthal sites, for example.2 The discovery of Homo naledi, a hominin with an intriguing combination of modern and primitive features (notably a rather small brain), in a cave in South Africa in 2013 revealed perhaps the most surprising insight into possible non-human hominin culture. The remains were pushed far into a chamber accessed through a series of climbs, crawls and squeezes; the evidence points to the body having been interred there deliberately. Such behaviour is important because it implies the capacity for abstract thought and symbolism, traits usually ascribed only to Homo sapiens.
The fact is we are still learning what truly makes us human and we are still unravelling the complexities of our evolutionary past. This book is about mismatches between that evolutionary past and the environment we have created, and exploring these mismatches is going to require some serious intellectual juggling. The story of why we are unfit for purpose in the modern world is a story of genetics, natural selection, evolution, biogeography, genome analysis, biochemistry, sexual selection, archaeology, psychology, sociology, politics and much more besides. So, pull up a chair, grab some calorie-dense snacks and let’s start with a very ‘big’ mismatch; why the modern world conspires with our evolutionary heritage to make us fat.
Notes
* Bergmann’s rule is named after German biologist Carl Bergmann (1814 – 1865). Bergmann noticed that within groups of related species there was a tendency for those species in colder regions to be larger. This is now often applied to patterns found within species, where individuals tend to be larger and more robust in colder climates. Like many rules in biology it doesn’t always hold true, but nonetheless it does describe the patterns found in a diverse range of mammals and birds
† Named after the American zoologist Joel Asaph Allen (1838 –1921), this rule states that animals adapted for colder climates tend to have shorter limbs and appendages (such as ears) than animals adapted for warmer regions. It is better supported for individuals within a species, including us, than for patterns between species where other factors (including Bergmann’s rule) tend to become more important.