The Cursorial Ape: a theory of human anatomy

Part Of: Anthropogeny sequence
Followup To: The Walking Ape
Content Summary: 2100 words, 21 min read

A Brief Review of Human Evolution

The most recent common ancestor of humans and chimpanzees lived 7 mya (million years ago). The very first unique hominin feature to evolve was bipedality, which was an adaptation for squat-feeding. The australopiths were bipedal apes. They could walk comfortably, but retained their adaptations for tree living as well. Dental morphology and microwear together suggest that australopiths acquired food from a new source: tubers (the underground storage organs of plants).

Climate change is responsible for the demise of the australopiths. Africa began drying out about 3 million years ago, making the woodlands a harsher and less productive place to live. Desertification would have reduced the wetlands where australopiths found fruits, seeds, and underwater roots. The descendents of Australopithecus had to adapt their diet.

The paranthropes adapted by promoting tubers from backup to primary food. These impressive creatures comprise a blend of human and cow-like features. In contrast, the habilines (e.g., Homo Habilis) took a different strategy: meat eating. These creatures had the same small bodies, but larger brains. Their hands show adaptations for flexibility, and their shoulders and elbows for throwing missiles. They began making stone tools (Mode 1 tools, the Oldowan industry). They presumably used these anatomical and cultural gifts to compete with other scavengers on the savannah (projectiles to repulse competitors, stone flakes to speedily butcher a carcass).

The habilines in turn gave rise to

  • [1.9 mya] The erects (H erectus)  with near-modern anatomies.
  • [0.9 mya] The archaics (H heidelbergensis) appear, who eventually give rise to the Neanderthals, Denisovans, and us.
  • [0.3 mya] The moderns (H sapiens) emerge out of Africa, and completely conquer the globe.

A Closer Look

Yes, humans are apes. But why do we look so different from our closest living relative, the chimpanzee?

I have previously explained why we are bipedal (flexible waist, straight backs, walking on two feet).

But why do we have scent glands in our armpits? Fat in our asses? Such weird hair? Hairless skin with massive subcutaneous fat deposits?

Most of these changes were introduced with Homo Erectus:

Born To Run_ Hominin Anatomy (4)

Natural selection explains why bodies change. Anatomical innovations are selected when they enable more efficient exploitation of some particular niche.

So what ecological niche forged the modern human body?

Where Homo Erectus Evolved

The australopiths never made it beyond the southern margins of the Sahara. Because the adaptation of equatorial species inhibits their colonization of temperate regions, the successful emigration of the erects out of Africa strongly suggests that this was a northern, not a tropical species.

To evolve adaptations to dry, open country, the erects would have had to suffer a period of isolation from other hominins, in an appropriately discrete habitat. There were few, perhaps no, places in tropical or Southern Africa that could have provided such a combination. Comparing these constraints with the distribution of Homo Erectus fossils, comparative zoologist Jonathan Kingdon submits there the two most plausible contenders where the erects could have evolved are the Atlas Mountains, or Arabia.

Nasal evidence corroborates the hypothesis that they evolved in a desert environment. The entry to the primate nasal passage is flat, with straightforward air intake. Erect skulls show the first evidence of a protruding nose. A protruding nose forces the air at a “right angle” before entering the nasal cavity.

One of the responsibility of the nasal passage is to humidify the air before it is passed to the lungs. The increase in room and turbulence serves to amplify the humidification of inhaled air. Our noses are adaptations for desert living.

A New Thermoregulation System

There are two things unique to human skin:

  • Functional hairlessness. We modern humans have hair, but it is so thin compared to chimpanzees that we are effectively hairless.
  • Eccrine sweat glands. Our skin also contains a novel approach to sweat glands.

These two features are linked: we now know in exquisite molecular detail how incipient hair follicles are converted into eccrine glands (Lu et al 2016).

Other primates rely on oil-based apocrine sweat glands. The emergence of water-based eccrine glands in humans led to the “retirement” of apocrine glands in our lineage. The distribution of odor-producing apocrine glands was ultimately confined to our underarms and pubic regions.

Born To Run_ Sweat Glands (1)

Losing our hair had two important side-effects:

  • Skin pigmentation. Fur protects against ultraviolet radiation. Without it, melanin was used as an alternate form of natural sunscreen.
    • Why do otherwise-bald humans have hair at the tops of their heads? This is the location of maximal radiation.
    • Why didn’t all humans remain dark-skinned? Melanin also inhibits the skin’s production of Vitamin D, and different locales have different radiation levels, requiring new tradeoffs to be struck.
  • Subcutaneous fat. Ever seen a hairless chimpanzee? Human skin is much less wrinkled than other skin. Why? Even in non-obese people, humans store more of their body fat below the skin (versus in the abdomen, or between the muscles). This change has three complementary causes:
    1. carnivores tend to store fat in this way,
    2. mitigate the hernia risk associated with bipedality
    3. replace the insulation services of fur, without interfering with sweat system.

We have reviewed four changes in human skin. Rather than a discrete event, these changes presumably evolved gradually, and in tandem.

Born To Run_ Evolution of Skin (2)

Yes, but why are we hairless? There are many competing theories.

Jonathan Kingdon claims these skin adaptations arose late, as a parasite avoidance mechanism induced by increased population densities. Two rationales are provided: hair is a potent vector of infection, and the eccrine sweat system also has antibiotic properties.

This interpretation is challenged by genetic evidence that shows hominins were naked at least 1.2 mya, if not earlier (Rogers et al, 2004).

However, given the evidence suggesting Homo Erectus evolved in a desert climate, the most parsimonious theory seems to involve thermoregulation. We were exposed to less direct radiation given our upright posture; fur no longer served as critical of a role. But the overall climate was warm and dry,  

Humans as Cursorial Species

A cursorial animal is one that is adapted for long-distance running, rather than animals with high acceleration over short distances; thus, a leopard is considered cursorial, while a cheetah is not. Other examples include wolves, horses, and ostriches.

Fit human amateurs can regularly run 10 kilometers, and longer distances such as marathons (42 kilometers) are achieved by tens of thousands of people each year. Such distances are unknown if not impossible for any other primate, but are comparable to those observed in specialized mammalian cursors on open habitats. African hunting dogs, for example, travel an average 10km per day.

Racing horses can gallop 10 kilometers at 9 meters per second. However, the sustainable galloping speeds in horses decline considerably for runs longer than 10-15 minutes. Well-conditioned human runners exceed the predicted preferred galloping speed for a 65-kg quadruped, and can even occasionally outrun horses over extremely long distances.

Thus, despite our embarrassingly slow sprinting speed, human beings can outcompete even cursorial animals at endurance running over large distances. How come? The answer has to do with our unique cooling system.

When other mammals trot, they cool themselves by panting. However, above certain speeds a quadruped transitions to a full gallop, which precludes panting. A horse can trot all day, but it cannot gallop continuously without overheating.

Human adaptations for running, and our unique eccrine sweat-based cooling system, meant that humans have a larger trot/gallop (jog/sprint) transition threshold. Our superior cooling technology is accentuated in high heat. We are literally the only mammal that can run a marathon in high heat.

Born To Run_ Trot-Gallup Transition (1)

Why are we Born to Run?

Our bodies are designed for endurance running. We are cursorial animals. But why?

To achieve this, hominids exploited a new form of predation called persistence hunting. The most successful persistence hunts will involve:

  • Time: middle of the day (during peak heat)
  • Target: big prey (overheats faster)

If you chase a big animal above its trot/gallop transition speed, the animal will easily distance itself and begin panting. But you can track the animal, and chase it again before it has the opportunity to fully recover. Repeat this process, and after 10-25 km you will successfully drive the prey into hyperthermia. This style of hunting has a remarkable 75% success rate. Modern hunters typically prefer to use the bow and arrow, but persistence hunting is still in their repertoire. Before the invention of projectile weapons some 71 kya, persistence hunting surely played a larger role.


We know that habilines ate meat (many bones show signs of their butchery). But they likely acquired meat by scavenging, as they were not particularly effective carnivores. Their adaptations for projectiles were presumably used to repulse competitors, and stone tools certainly helped speedily butcher a carcass.

Of the dozens of running adaptations in our Homo Erectus, a substantial fraction already exist in habilines. Presumably the re-invention of our skin had begun too. These processes presumably began for simple reasons (it pays to move quickly, and have less fur, in the savannahs that emerged 3 mya).

Persistence hunting completely changed the game. Adaptations for running brought steep rewards. In a typical persistence hunt, the hunter averages an energy expenditure of 850 Kcal; they energy gains from big game is multiple times larger. Compare the calorie budget for a modern-day hunter-gatherer with that of chimps: in our prime, we produce twice as many calories as we consume!

Born To Run_ Calorie Budget (1)

Life is fundamentally about getting energy to make more life.

Persistence hunting was the turning-point in human evolution. Our species began winning, in terms of our reliably acquiring surplus energy. This surplus was the reason why our lineage could “afford” bigger brains, taller bodies, more frequent births, and longer childhood. All of these characteristics have improved gradually & continuously since the erects emerged.

Our Cursorial Adaptations

We have looked at the reasons behind our running. What does anatomy tell us?

First, let’s compare the physics of walking vs running:

  • Walking is an inverted pendulum mechanism.  Our feet and our hips alternate as the center of rotation.
  • Running is a mass-spring mechanism. Ligaments transfer foot-strike kinetic energy into tendons, which is released as we bounce onward.

Walking doesn’t require springs – but running does. And the bodies of erects have two new ligaments that serve precisely this purpose:

  • The Achilles’ tendon stores and releases 35% of energy expended while running (but not walking). In chimps, this tendon is 1cm long. In erects, it is 10cm and much thicker.
  • The dome-shaped arch of the foot is another spring, which lowers the cost of running by 17%.

During bipedal running the risk of falling and sprained ankles is high, which in the ancestral environment had adaptive consequences. Thus, the human body also developed many stabilization techniques:

  • Gluteus maximus. Barely active during walking, this muscle contracts forcefully during running to prevent the trunk from toppling forward. 
  • Various head stabilization devices. Promotes vision continuity and protects the brain (watch a runner with a ponytail sometime).
  • Enlarged semicircular canals (balance organs) in inner ear, which can be seen by measuring certain dimensions of fossilized skulls.

I have listed five features of our anatomy that relate to endurance running. Lieberman et al (2006) list twenty:

Born To Run_ Anatomical Comparison (1).png

As you can see, not all of these running adaptations emerged with Homo Erectus. Homo Habilis already shows adaptations for running. It would not surprise me in the slightest if that species also saw the beginnings of our skin trajectory.

Adaptations for running came at a price. We have lost our ability to climb trees. We are the first primate to lose this ability.


Why do humans look so different from our closest living relative, the chimpanzee?

Why do we have scent glands in our armpits? Fat in our asses? Such weird hair? Hairless skin with massive subcutaneous fat deposits?

Animal body plans are designed to excel in a particular niche. Our bodies are designed for persistence hunting. Compared to other primates, our anatomies optimize for thermoregulation, efficient energy transfer, and stabilization during running.

Born To Run_ Overview (5)

Chimpanzees don’t need to exercise to stay fit. We do. Our health sees dramatic benefits from aerobic exercise, especially running.


  • Bramble & Lieberman (2004). Endurance running and the evolution of Homo
  • Lieberman et al (2006). The human gluteus maximus and its role in running
  • Lu et al (2016). Spatiotemporal antagonism in mesenchymal-epithelial signaling in sweat versus hair fate decision.
  • Rogers et al (2004). Genetic Variation at the MCiR Locus and the Time since Loss of Human Body Hair

[Excerpt] How Language Evolved

Part Of: Language sequence
See Also: When Language Evolved
Excerpt From: (Johansson 2011) Constraining the Time When Language Evolved
Content Summary: 1600 words, 16 min read

The evolution of language had to involve at least a new ability to map concepts to sounds and gestures and to use these communicatively. But language actually consists of a good deal more than this: First, there is phonological structure—the systematized organization of sounds (or, in sign languages, gestures). Second is morphology—the internal structure of words, such that the word procedural can be seen as built from proceed plus -ure to form procedure, plus -al to form procedural: [[[proceed] [-ure]] [-al]]. Third is syntax, the organization of words into phrases and sentences.

One way to form plausible hypotheses about evolution is through reverse engineering: asking what components could have been useful in the absence of others. A primitive system for communicating thoughts via sound or gestures is useful without phonology, morphology, or syntax. The latter components can improve an existing communication system, but they are useless on their own. So if the components of language evolved in some order, it makes sense that the connection between phonetics and meaning came first, followed by these further refinements.

A system with a linear grammar would have words— that is, stored pairings between a phonological form and a piece of conceptual structure. The linear order of words in an utterance would be specified by phonetics, not by syntax. The individual words would map to meanings, but beyond linear order, there would be no further structure—no syntactic phrases that combine words and no morphological structure inside words (such as in the word procedural).

Language Evolution_ Linear vs Recursive Grammar (1)

Indeed, we can find evidence for linear grammar in many different contexts.

  1. As the early stages of contact languages, pidgins are often described as having no subordination, little or no morphology, no grammatical words like the, and unstable word order governed primarily by semantic principles like agent before action. If the context permits, the characters in the action can be left unexpressed. For instance, if the context had already brought the boy to attention, the speaker might just say girl kiss, which in English would require a pronoun—The girl kissed him. From the perspective of linear grammar, we can ask: Is there any evidence that pidgins have parts of speech like nouns and verbs, independently from the semantic distinction between individuals and actions? Indeed, there is no evidence for syntactic phrases, beyond semantic cohesion. Pidgin grammars are a good candidate for real-world examples of our hypothesized linear grammar.
  2. For a second case, involving late second language acquisition, Wolfgang Klein and Clive Perdue did a multilanguage longitudinal study of immigrants learning various second languages all over Europe. They found that all speakers achieved a stage of semiproficiency that they called the Basic Variety. Many speakers went on to improve on the Basic Variety, but others did not. At this stage, there is no inflectional morphology or sentential subordination, and known characters are freely omitted. Instead, there are simple, semantically based principles of word order including, for instance, agent before action.
  3. A third case is home signs, the languages invented by deaf children who have no exposure to a signed language. Susan Goldin-Meadow has shown that they have at most rudimentary morphology; they also freely omit known characters. In our analysis, home signs only have a semantic distinction of object versus action, not a syntactic distinction of noun versus verb. Word order is probabilistic and is based, if anything, on semantic roles. Homesigners do produce some sentences with multiple verbs, which Goldin-Meadow describes as embedding. We think these are rudimentary serial verb or serial action-word constructions, without embedding, sort of like the compound verb in English expressions such as He came running. So this looks like a linear grammar with possibly a bit of morphology.
  4. Another case is village sign languages, which develop in isolated communities with a significant occurrence of hereditary deafness. A well-known example is Central Taurus Sign Language (CTSL), spoken in two remote villages in the mountains of Turkey. CTSL has some minimal morphology, mostly confined to younger speakers. But there is little or no evidence for syntactic structure. In sentences involving one character, the word order is normally agent + action, and two-character sentences are normally (optional) agent + patient + action: girl ball roll. But if a sentence involves two animate characters, so that semantics alone cannot resolve the potential ambiguity, word order is not very reliable. For instance, girl boy hit is a bit vague about whether the girl hit the boy or vice versa, requiring a huge reliance on pragmatics, common knowledge, and context. In fact, there is a strong tendency to mention only one animate character per predicate, so speakers sometimes clarify by saying things like Girl hit, boy get-hit. So CTSL looks like a linear grammar, augmented by a small amount of morphology. Similar results have been obtained in Al-Sayyid Bedouin Sign Language (ABSL) and the earlier stages of Nicaraguan Sign Language.
  5. These less complex systems are not confined to emerging languages; they also play a role in language processing. Townsend and Bever (2001) discuss what they call semantically based interpretive strategies that influence language comprehension. In particular, hearers tend to rely in part on semantically based principles of word order such as agent precedes action, which is why (in our account) speakers have more difficulty with constructions such as reversible passives and object relatives, in which the agent does not precede the action. Similarly, Ferreira and Patson (2007) discuss good enough parsing, in which listeners apparently rely on linear order and semantic plausibility rather than syntactic structure. It is well known that we see similar though amplified symptoms in language comprehension by agrammatic aphasics. Finally, Van der Lely and Pinker (2014) argue that a particular population of children with specific language impairment behave as though they are processing language through something like a linear grammar. The literature frequently describes these so-called heuristics as something separate from language. But they are still mappings between phonetics and meaning—just simpler ones.
  6. We have also encountered a full-blown language whose grammar appears to be close to a linear grammar: Riau Indonesian, a vernacular with several million speakers, described by Gil (2005, 2009). Gil argues that this language has no syntactic parts of speech and no inflectional morphology such as tense, plural, or agreement. Known characters in the discourse are freely omitted. Messages that English expresses with syntactic subordination are expressed in Riau paratactically, with utterances like girl love, kiss boy. The word order is quite free, but agents tend to precede actions, and actions tend to precede patients. This collection of symptoms again looks very much like a linear grammar. Hence, this is a language virtually all of whose grammar is syntactically simple in our sense. Similar results obtain for the Piraha language, whose non-recursivity is well explained by the linear grammar theory as well.
  7. Another kind of linear grammar—that is, a system that relies on the linear order of the semantic roles being expressed to form conceptual relations—surfaces when people are asked to express actions or situations in a nonlinguistic task, such as in gesture or act-out tasks. Overall, there is a vast preference to gesture, or act out, the agent first (e.g., girl), and then the patient (e.g., boy). The action is usually expressed last (kiss), but when there is a potential ambiguity, people like to avoid it by expressing the action in the middle, between the agent and patient. Crucially, the ordering preferences in these tasks are remarkably stable, independently of the ordering preferences in test subjects’ native language. That seems to indicate that the capacity to map certain semantic notions to certain linear orders is at least partly independent from language itself.
  8. As a final case, traces of something like linear grammar lurk within the grammar of English! Perhaps the most prominent case is compounding, in which two words are stuck together to form a composite word. The constituents may be any part of speech: not just pairs of nouns, as in kitchen table, but also longbow, undercurrent, castoff, overkill, speakeasy, and hearsay. The meaning of the composite usually includes the meanings of the constituents, but the relation between them is determined pragmatically. Consider examples like these:
      • collar size = size of collar
      • dog catcher = person who catches dogs
      • nail file = something with which one files nails
      • beef stew = stew made out of beef
      • bike helmet = helmet that one wears while riding a bike
      • bird brain = person whose brain is similar to that of a bird

    The second noun usually determines what kind of object the compound denotes; for instance, beef stew is a kind of stew, whereas stew beef is a kind of beef. But this can be determined solely from the linear order of the nouns and needs no further syntax.

To sum up, remarkably similar grammatical symptoms turn up in a wide range of different scenarios. This suggests to us that linear grammar is a robust phenomenon, entrenched in modern human brains. It provides a scaffolding on top of which fully syntactic languages can develop, either in an individual, as in the case of the Basic Variety, or in a community, as in the case of pidgins and emerging sign languages. Furthermore, it provides a sort of safety net when syntactic grammar is damaged, as we have seen with aphasia and specific language impairment. We have also seen that it is possible to express a great deal even without syntax, for example in Riau Indonesian—though having syntax gives speakers more sophisticated tools for expressing themselves.

Language Evolution_ Linear Grammar

[Excerpt] When Language Evolved

Part Of: Language sequence
See Also: How Language Evolved
Excerpt From: (Johansson 2011) Constraining the Time When Language Evolved
Content Summary: 900 words, 9 min read

Speech is not impossible with an ape vocal tract, but merely less expressive, with fewer vowels available. Furthermore, the vocal tract in living mammals is quite flexible, and a resting position different from the human configuration does not preclude a dynamically lowered larynx, giving near-human vocal capabilities, during vocalizations.

Adaptations for speech can be found in our speech organs, hearing organs, the neural connections between these organs, as well as the genes controlling their development.

  • Speech organs. The shape of the human vocal tract, notably the permanently lowered larynx is very likely a speech adaptation, even though some other mammals, such as big cats, also possess a lowered larynx. The vocal tract itself is all soft tissue and does not fossilize, but its shape is connected with the shape of the surrounding bones, the skull base and the hyoid. Already Homo erectus had a near-modern skull base, but the significance of this is unclear, and other factors than vocal tract configuration, notably brain size and face size, strongly affect skull base shape. Hyoid bones are very rare as fossils, as they are not attached to the rest of the skeleton, but one Neanderthal hyoid has been found, as well as two hyoids from Homo heidelbergensis, all very similar to the hyoid of modern Homo sapiens, leading to the conclusion that Neanderthals had a vocal tract adequate for speech. The hyoid of Australopithecus afarensis, on the other hand, is more chimpanzee-like in its morphology, and the vocal tract that reconstruct for Australopithecus is basically apelike.
  • Hearing organs. Some fine-tuning appears to have taken place during human evolution to optimize speech perception, notably our improved perception of sounds in the 2-4 kHz range. The sensitivity of ape ears has a minimum in this range, but human ears do not, mainly due to minor changes in the ear ossicles, the tiny bones that conduct sound from the eardrum to the inner ear. This difference is very likely an adaptation to speech perception, as key features of some speech sounds are in this region. The adaptation interpretation is strengthened by the discovery that a middle-ear structural gene has been the subject of strong natural selection in the human lineage These changes in the ossicles were present already in the 400,000-year-old fossils from Spain, well before the advent of modern Homo sapiens. These fossils are most likely Homo heidelbergensis. In the Middle East, ear ossicles have been found both from Neanderthals and from early Homo Sapiens, likewise with no meaningful differences from modern humans.
  • Lateralization. There is no clearcut increase in general lateralization of the brain in human evolution — ape brains are not symmetric — and fossils are rarely undamaged and undistorted enough to be informative in this respect. But when tools become common, handedness can be inferred from asymmetries in the knapping process, the usewear damage on tools, and also in tooth wear patterns, which may provide circumstantial evidence of lateralization, and possibly language. Among apes there may be marginally significant handedness, but nothing like the strong population-level dominance of right-handers that we find in all human populations. Evidence for a human handedness pattern is clear among Neanderthals and their predecessors in Europe, as far back as 500 kya, and some indications go back as far as 1 mya. To what extent conclusions can be drawn from handedness to lateralization for linguistic purposes is, however, unclear.
  • Neural connections. Where nerves pass through bone, a hole is left that can be seen in well-preserved fossils. Such nerve canals provide a rough estimate of the size of the nerve that passed through them. A thicker nerve means more neurons, and presumably improved sensitivity and control. The hypoglossal canal, leading to the tongue, has been invoked in this context, but broader comparative samples have shown that it is not useful as an indicator of speech. A better case can be made for the nerves to the thorax, presumably for breathing control. Both modern humans and Neanderthals have wide canals here, whereas Homo erectus has the narrow canals typical of other apes, indicating that the canals expanded somewhere between 0.5 and 1.5 million years ago.
  • FOXP2. When mutations in the gene FOXP2 were associated with specific language impairment, and it was shown that the gene had changed along the human lineage, it was heralded as a “language gene”. But intensive research has revealed a more complex story, with FOXP2 controlling synaptic plasticity in the basal ganglia rather than language per se, and playing a role in vocalizations and vocal learning in a wide variety of species, from bats to songbirds. Nevertheless, the changes in FOXP2 in the human lineage quite likely are connected with some aspect of language, even if the connection is not as direct as early reports claimed. Relevant for the timing of the emergence of human language is that the derived human form of FOXP2 was shared with Neanderthals, and that the selective sweep driving that form to fixation may have taken place more than a million years ago, well before the split between Homo Sapiens and Neanderthals.

No single one of these indications is compelling on its own, but their consilience strengthens the case for some form of speech adaptations in Homo Heidelbergensis.

As the speech optimization, with its accompanying costs, would not occur without strong selective pressure for complex vocalizations, presumably verbal communication, this implies that Homo erectus already possessed non-trivial language abilities. While Homo erectus did not possess our species’ ability for ratcheting (cumulative) culture, it did exhibit art and sufficient skills to construct watercraft.

The Walking Ape: why our ancestors first stood up

Part Of: Anthropogeny sequence.
Content Summary: 1600 words, 16 min read

For all his noble qualities, godlike intellect, and exalted powers, man still bears in his bodily frame the indelible stamp of his lowly origin.

– Charles Darwin, Descent of Man

Setting The Stage

Common descent denotes the discovery that all species are related: that living organisms reside in a single tree of life. Homo Sapiens is no exception. We diverged from other hominoids (great apes) some 7 mya. During that time period, fossils more than 6,000 individuals from dozens of bipedal ape species.

Bipdality_ Hominin Phylogeny

Today, we explore why apes became bipedal. But first, the evolution of apes.  

Primate Evolution

Primates are mammals with flat nails instead of claws, grasping hands and feet, a highly developed visual system. They are highly iteroparous (long juvenile period) and have large brains to support the complex needs of group living. Primates are known for their symbolic dominance hierarchy, friendship mediated by grooming and mindreading (making inferences about the mental state of their peers).

Apes are primates that hang from branches (no tail), and even larger brains that promote behavioral flexibilities. Apes are known for coalitional warfare, group-specific cultural behaviors, flexible group signaling (e.g., mobbing), and tool-making.

The primate lineage emerged in the Paleocene (60 mya); apes in the Miocene (20 mya).

Bipedality_ Geological Periods (1).png

Without a tail (and in a “dead-end” body plan that precludes growing it back), apes increasingly relied upon behavioral flexibility to mitigate their comparative immobility. A monkey is an ecological specialist; the ape lineage was populated by generalists.

Apes flourished in the early and middle Miocene (20-10 mya). But they began to die out, starting in the late Miocene (10 mya). Today, there are hundreds of extant species of monkeys, and only five apes (gibbons, gorillas, orangutans, chimps and bonobos).

Evolution and progress are not synonymous. The ape branch of the tree of life is sparse because we are a failed lineage.

The failure of our ancestors seems to have been driven by a radiation from earlier primates (monkeys) in what can be called revenge of the specialized. It became increasingly difficult for generalized omnivorous species to find niches that were not more effectively exploited by a whole host of small-sized specialist monkeys.

Amidst this harsh inter-primate competition, it is interesting to note that modern apes are substantially larger than their Miocene ancestors. An increase in the body size of living apes and humans may well represent an evolutionary response to competition from monkeys.

We turn now to the question of bipedality. Before we can address why apes stood on two legs, we must first understand the anatomy of bipedality.

The Anatomy of Walking

The main anatomical structure that changed was the pelvis. The pelvis is not a single bone, but rather three bones glued together by cartilage. As we will see shortly, bipedality requires shortening of the ilium. 

Bipedality_ Pelvis Bones (1)

Walking is a pendulum-like motion. Most of the time one foot is off of the ground. This provides a stabilization problem. To solve this, bipedal animals have abductor muscles. You can actually feel these yourself: next time you walk around, feel the muscle on your hips flex (but only the muscle on the side of the weighted foot).

Bipedality_ Abductor Muscles (4)

Abductor muscles aren’t enough, however. In order to further stabilize a two-legged gait, the legs must be brought closer together. Adjusting the femur angle brings the center of gravity closer together:

Bipedality_ Knee

Finally, to improve the energy efficiency of walking, the human foot transitioned from a grasping surface to an energy-transfer platform.

Bipedality_ Foot Differences (3)

We have so far discussed four features of bipedal living. Here is a more complete list:

  1. pelvis shape (smaller ilium)
  2. pelvis musculature (abductor muscles)
  3. femur angle (more “knock-kneed”)
  4. feet (platform instead of grasping tool)
  5. foramen magnum angle (how the skull attaches to the spine),
  6. shape of the spine (bipedal spines are S-shaped), and
  7. reduced arm length (no longer needed to contact the ground)

The definition of hominin is bipedal ape. Little surprise then, that even the earliest hominin (Sahelanthropus Tchadensis) has at  least one feature associated with bipedalism. As we move to more recent species, we can see increasingly “classical” body plans:

Bipedality_ Anatomical Features (1)

Bipedality also explains why human beings suffer from:

  • Lower back pain. For hundreds millions of years, the spine was housed on a horizontal chassis. Switching to a vertical chassis places a lot of pressure on the lower spine. Zebras don’t suffer from lower back pain as much as human beings.
  • Hernias. The strain is not limited to the skeleton. Pressure also dramatically increases in the lower abdomen, causing an unusually high rates of hernias for human beings. In fact, one of the distinguishing characteristics of human beings is our smooth, fatty skin. We preferentially store fat subcutaneously to combat the pressure in our abdomen.

Theories of Bipedality

The fact that African apes became bipedal around 6 mya is not particularly interesting. A more interesting question is why African apes became bipedal. How did bipedality amplify the hominin niche?

There is no shortage of theories. Here are six:

  1. Brachiation (arm-based locomotion via branch-swinging) responsible for the postcranial features we share with apes.
  2. Arboreal apes modified their vertical climbing to walk bipedally along thick branches in the canopy.
  3. Bipedalism emerged from the need to carry babies, food, and other objects back to base.
  4. An aquatic phase of foraging and avoiding predators in water.
  5. Predator avoidance in the savannah with frequent peering over tall grass.
  6. A thermal theory whereby savanna dwellers stand up to keep cool.

These theories leave much to be desired, however.

First, some disregard ecological data entirely. The last two theories rely on the savannah hypothesis: that standing on two legs was made advantageous as forests increasingly disappeared. But the savannah hypothesis is wrong. Bipedalism emerged 6 mya, but the savannah grasslands only appeared 2-4 mya.

Second, they disregard the incrementality of natural selection. Two-legged standing preceded true bipedal walking and should not be lumped with it.  We must conceive of an ape that can stand but not walk (Orrorin tugensis?), and an ape that can walk but not run (Australopithicus afarensis).

More generally, whenever we see a complex adaptive package like walking, it is immediately useful to explore prerequisite abilities. One natural way to conceptualize the increments is as follows:

Bipedalism_ Incremental Improvements (2)


The above image identify anatomical increments with each new behavioral capability. 

We are not looking for a single ecological incentive for bipedalism; rather, we need individual motives for each increment in the journey to bipedality.

What kind of niche would reward flexible hips and a straight back?

The Primacy of Ecology

To answer this question, we need to get familiar with African geology and ecology.

As the most common promoter of diversity, allopatric speciation occurs when some population becomes isolated from the broader gene pool. Typically, these episodes are caused by climate change: the species gets “locked in” to a particular area by encroaching deserts, and then expands to surrounding habitats once the desert recedes.

The African continent contains wet-spots (equational rain) and hotspots (deserts). During cold glacial periods, these wet-spots expand along an east-west axis. For warm interglacial periods, the hot-spots expand along a north-south axis.

Bipedality_ African Hot-Dry Cycles.png

There are two primary forests in Africa:

Bipedality_ Two Forests

During the most arid climatic phases, the desert corridor separating these forests would close, leading to genetic isolation and speciation.

Squat Feeding in the Eastern Littorals

What kind of ape would emerge from the Main Forest Block? Such species would remain conservative (change slowly) because their much larger range embraces a much wider range of different types of wooded habitats. In fact, we know that modern-day gorillas derive from this ecosystem.

What kind of ape would be forged by the Eastern Forest Littoral? This smaller, fragmented ecosystem would cause both selection and genetic drift to accelerate. There are several peculiarities to this ecosystem worth pointing out:

Bipedality_ East African Littoral Ecology

In short, apes isolated in East African littoral forests seem likely to have found a niche on the forest floor. The natural distribution of resources favors this interpretation; and the growing competition from monkeys would have made the canopy increasingly infeasible.

These ground apes faced strong selective pressure to improve their foraging efficiency. The chimp pelvis has a very long ilium, which “locks into” the ribcage. There are clear foraging benefits for a reduction in the ilium (flexible waist), and straightening of the back (improved visibility). 

Bipedality_ Squatting Posture.png

In short, the squat-feeding hypothesis explains why flexible hips and straight spines were selected in ground apes of the early Pleistocene.

Other adaptive explanations only become relevant in further increments of the transition to bipedality. In particular, starting around 4 mya, the African continent began to dry. This made fruit increasingly less concentrated, and more seasonal. Locomotion thus became increasingly necessary to get enough calories. 

In modern humans, walking is four times more efficient than chimpanzee knuckle walking. Of course, very ancient hominins like Ardipithecus Ramidus could walk, but were less efficient than the Australopiths (and us, for that matter). But clumsy walking merely needs to improve upon the kinematic efficiency of knuckle walking, which as we have seen is not hard to do.

Bipedalism is not universally advantageous. Hominins like us are half as fast as other apes, and we have lost the ability to gallop. Greatly reduced ability to change direction while running.  The earliest bipeds probably avoided open habitats because of their increased vulnerability to predation, preferring forest and riverine habitats instead.

Bipedalism_ Ecological Pressures (2)

The facilitation of walking and running was not the ecological reason why our ancestors began the journey towards bipedality. But once they started on this particular anatomical pathway, these applications became possible. Thus, it is only with hindsight that we can say that the ultimate worth of standing up, the hidden evolutionary prize, was the ability to find the way out of a sort of ecological cul-de-sac.

Concluding Thoughts

The squat feeding theory of bipedality, as well as several of the images of this post, are credited to Jonathon Kingdon, African zoologist and author of Lowly Origin. I highly recommend this text, for those curious to learn more.

Until next time.

Cooking and the Hominin Revolution

Part Of: Anthropogeny sequence
See Also: Born to Run: a theory of human anatomy
Content Summary: 2100 words, 21 min read

The Universality of Cooking

Cooking is a human universal. It has been practiced in every known human society. Rumors to the contrary have never been substantiated. Not only is the existence of cooked foods universal, but most cuisines feature cooked foods as the dominant source of nutrition.

Cooking_ A Human Universal (1)

Raw foodists comprise a community dedicated to consuming uncooked food. Of course, compared to historical hunter-gatherers, modern raw foodists enjoy a wide variety of advantages. These include:

  1. Elaborate food preparation (pounding, purees, gently warming),
  2. Elimination of seasonal shortages (supermarkets)
  3. Genetically enhanced vegetables with more sugar content and fewer toxins.

Despite these advantages, raw foodists report significant weight loss (much more than vegetarians!). Further, raw foodists suffer from increasingly severe reproductive impairments, which have been linked to not getting enough energy.  

Cooking_ Consequences of Raw-Foodism (1)

Low BMI and impaired reproduction are perhaps manageable in modern times, but are simply unacceptable to hunter-gatherers living at subsistence levels.

The implication is clear: there is something odd about us. We are not like other animals. In most circumstances, we need cooked food.

The Energetics of Cooking

Life exists to find energy in order to make more copies of itself. Feeding and reproduction are the twin genetic imperatives.

Preferences are subject to natural selection. The fact that we enjoy cooked food suggests that cooking provides an energy boost to its recipients. The raw-foodist evidence hints towards this conclusion as well. But there is also direct evidence in rats that cooking increases energy gains.

In the following experiments, rat food was either processed/pounded, cooked, neither, or both. After giving this diet over the course of four days, rats in each condition were weighed.

Cooking_ Energy Benefits of Cooking (1).png

For starches (left) and meat (right), cooking is by far more effective at preventing weight loss and promoting weight gain. Tenderizing food can sometimes help, but that technique pales in comparison to cooking.  

The above results were taken from rats. But similar results have replicated in calves, lambs, piglets, cows, and even salmon. It seems to be universally true that cooking improves the energy derived from digestion, sometimes up to 30%.

How does cooking unlock more energy for digestion?

First, denaturation occurs when the internal bonds of a protein weaken, causing the molecule to open up. Heat predictably denatures (“unfolds”) proteins, and denatured proteins are more digestible because their open structure exposes them to the action of digestive enzymes.

Besides heat, three other techniques promote denaturation: acidity, sodium chloride, and drying. Cooking experts constantly harp on these exact techniques, because it aligns with eating preferences.

Second, tender foods is another boon to digestion, because they offer less resistance to the work of stomach acid.  If you take rat food, and inject air into the pellets, that does not augment denaturation. Nevertheless, softening food in this way improves the energy intake of the rat.

Cooking does have negative effects. It can cause a loss of vitamins, and give rise to long-term toxic molecules called Maillard compounds, which are linked to cancer. But from an evolutionary perspective, these downsides are overshadowed by the impact of more calories. In subsistence cultures, better fed mothers have more, happier, and healthier children. When our ancestors first obtained extra calories by cooking their food, they and their descendants past on more genes than others of their species who ate raw.

A Brief Review of Human Evolution

The most recent common ancestor of humans and chimpanzees lived 6 mya (million years ago). But the first three million years of our heritage are not particularly innovative, anatomically. The australopiths were essentially bipedal apes. They could walk comfortably, but retained their adaptations for tree living as well. There is some evidence that australopiths acquired food from a new source: tubers (the underground energy storage system of plants).

Climate change is responsible for the demise of the australopiths. Africa began getting dryer about 3 million years ago, making the woodlands a harsher and less productive place to live. Desertification would have reduced the wetlands where Australopiths found fruits, seeds, and underwater roots. The descendents of Australopithecus had to adapt their diet.

The paranthropes adapted by promoting tubers (underground storage organs of plants) from backup to primary food. In contrast, the habilines (e.g., Homo Habilis) took a different strategy: meat eating. These creatures inherited tool making from the late australopiths (Mode 1 tools, the Oldawan industry- was discovered in Ethiopia 2.6 mya), and used these tools to scrape meat off of bones). The habilines are more ecologically successful, and lead to:

  • 1.9 mya: The erects (e.g., Homo erectus/ergastor) with significantly larger brains and near-modern anatomies.
  • 0.7 mya: The archaics (e.g., Homo Heidelbergensis) appear, who eventually give rise to the Neanderthals, Denisovans, and us.
  • 0.3 mya The moderns (e.g., Homo Sapiens) emerge out of Africa, and completely conquer the globe.

Here is a sketch of how our body plans have changed across evolutionary time:

Cooking_ Hominin Anatomy Comparison

Explaining Hominization

The transition from habiline to erects deserves a closer look. We know erects evolved to be persistence hunters. But a number of paradoxes shroud their emergence:

  1. Digestive Apparati. The erect diet appears to be mainly meat and tubers. Both require substantial jaw strength and digestive apparati. Yet the Homo genus features a dramatically reduced digestive apparatus. How was smaller mouths, weaker jaws, smaller teeth, small stomachs, and shorter colons an adaptive response to eating meat and starches?
  2. Expensive Tissue. Australopiths brain size stayed relatively constant at 400 ccs (10% of resting metabolism). Erect brains began to grow. This transition ultimately yielded a 1400 cc brain (20% of resting metabolism) in archaic humans. How did the erects find the calories to finance this expansion?
  3. Time Budget. The above anatomical features of erects are geared towards endurance running, which suggests that their lifestyle involved persistence hunting. Chimps have about 20 minute intervals in between searching for & chewing food. Thus, chimps can only afford to spend 20 minutes hunting before giving up. How did erects perform the risky behavior of persistence hunting, which consumes 3-8 hours of time?
  4. Thermal Vulnerability. As part of their new hunting capabilities, erects became the naked ape (with a new eccrine sweat gland system to prevent overheating). But Homo Erectus also managed to migrate to non-African climates such as Europe. How did these creatures stay warm?
  5. Predator Safety. Erects lost their anatomical features for arboreal living, which suggests they slept on the ground. Terrestrial sleeping is quite dangerous on the African savannah. How did erects avoid predation & extinction?

All of these confusing phenomena can be explained if we posit H. erectus discovered the use of fire, and its application in cooking:

  1. Digestive Apparati. As we have seen, the primary role of cooking is to “externalize digestion”, and to increase the efficiency of our digestive tract. Cooked meat and starches are incredibly less demanding to process than their raw alternatives. This explains our reduced guts. By some estimates, the decrease in digestive tissue corresponds with a 10% energy savings by our erect ancestors.
  2. Expensive Tissue. Cooking increases the metabolic yield of most foodstuffs by ~30%. For reference, a 5% increase in ripe fruit for chimpanzees reduces interbreeding interval (time between children) by four months. 30% is an absurdly large energy gain, enough to “change the game” for the erects..
  3. Time Budget. Cooking freed up massive amounts of time otherwise spent chewing. Chimpanzees can take 4-7 hours per day chewing; humans only need one hour per day. This frees up massive amounts of time, which can be used for e.g., hunting.
  4. Thermal Vulnerability. It is very difficult to explain a hairless Homo Erectus thriving on the colder Asian continent without control of fire.
  5. Predator Safety. It is very difficult to explain how erects were not preyed upon to extinction without fire to identify & deter predators. Hadza hunter-gatherers comfortably sleep through the night, typically by taking turns “on watch” while the others rest.

Cooking_ Overall Argument (3)

The Archaeological Record

We are positing that erects learned to create and controlling fire 2 mya. Is that a feasible hypothesis?

Habilines had learned how to create stone tools 2.6 million years ago. By the time of the erects, techniques to create these tools had persisted for 600,000 years. So it is safe to say that our ancestors were able to retain useful cultural innovations.

Independent environmental reasons link fire-making with H Erectus. The Atlas mountain range is the most likely birthplace of this species, and this dry area fires triggered by lightning are an annual hazard. Hominins living in such environments would be more intimately familiar with fire than those with less combustible vegetation zones.

Erects would have seen sparks when they hit stones together to make tools. But the sparks produced by many kinds of rock are too cool to catch fire. However, when pyrites (a fairly common ore) are hit against flint, the results are used by hunter-gatherers to reliably produce fire. The Atlas mountain range is renowned for being exceptionally rich in minerals:

Why is Morocco one of the world’s great countries for minerals? No glaciers! Many of the world’s most colorful minerals are found in deposits at the surface, formed over time by the interaction of water, air and rock. Glaciers remove all of that good stuff (as happened in Canada recently, geologically speaking) –  and with no recent glaciation, Morocco hosts many fantastic occurrences of minerals unlike any in parts of the world stripped bare during the last Ice Age.

Since this mountain range contains pyrites, early erects could have found themselves inadvertently making fire rather often.

Once it is created, fire is relatively easy to keep going. And it does not take much creativity to stick food a fire. Moreover, modern-day chimps prefer cooked food over raw; it is hard to imagine H Erectus finding cooked food distasteful. All of these considerations suggest an early control of fire is at least plausible.

We can consult the archaeological record to see record of man-made fire (i.e., hearths). This is bad news for the cooking hypothesis! There is strong evidence for hearths dating back to 800 mya and the advent of archaic humans. Before then, there are six sites that seem to be hearths; but these are not universally acknowledged as such.

Cooking_ Archaeology Evidence (1)

But absence of evidence isn’t evidence of absence, right?

No! That idiom is wrong. Silence is evidence of absence. It’s just that the strength of the evidence depends on the nature of the hypothesized entity.

  • If you think an unidentified planet orbits the Sun, a lack of evidence would weigh heavily against the hypothesis.
  • If you think an unidentified pebble orbits the Sun, a lack of evidence doesn’t say much one way of the other.

Wrangham argues that evidence of hearths are more fragile than e.g. fossils, and points to facts like there are zero hearths recorded for modern humans during European “ice ages” – but we know these must have existed. It is possible that the contested hearth sites will ultimately be vindicated, and that we just can’t see much evidence.

Despite these claims about evidential likelihood, the silence of the archaeological record is undeniably a significant objection to the theory.

Weighing The Evidence

Is the cooking hypothesis true? Let us weigh the evidence, and contrast it with alternative hypotheses.

The most plausible alternative hypothesis is that archaic humans H. Heidelbergensis discovered cooking. But the emergence of that species involved an increase in brain size, and more sophisticated culture & hunting technology.  Neither adaptation seems strongly connected to cooking. In contrast, the H. Erectus adaptations would have all been strongly affected by cooking. 

Moreover, alternative hypotheses must still answer the five paradoxes of hominization:

  1. Digestive Apparati. Why did erects evolve smaller mouths, weaker jaws, smaller teeth, small stomachs, and shorter colons?
  2. Expensive Tissue. How did the erects find the calories to finance more brain tissue?
  3. Time Budget. How could erects afford spending 3-8 hours per day engaged in the risky strategy hunting?
  4. Thermal Vulnerability. Erects also managed to migrate to non-African climates such as Europe. How did these creatures stay warm?
  5. Predator Safety. Erects slept on the ground. How did they avoid predation?

The habilines ate meat. It is unclear how they did so (hunting or scavenging), but we have strong evidence that they did. Meat is a much higher quality food than tubers (cf. paranthropes) or fruit (cf. chimpanzees). The meat-eating hypothesis argues that meat eating was the primary driver of hominization.

Meat-eating resolves the Expensive Tissue paradox (meat allows for brain growth) and Digestive Apparati (carnivores are known to have smaller guts). But it doesn’t address why a meat-eater would develop smaller canines. And it struggles to explain how the reduction in gut size is compatible with the tuber component of the erect diet. And what about time budget, thermal vulnerability, and predator safety? The meat eating hypothesis fails to address these paradoxes entirely.

Which is more likely to occur in the next twenty years: undisputed evidence for early control of fire, or an alternate theory that resolves all five hominization paradoxes?

My money is on the former.


  • Wrangham (). Catching Fire: How Cooking Made Us Human
  • Aiello & Wheeler(1995). The expensive tissue hypothesis: the brain and the digestive system in primate and human evolution.

An Introduction to Domestication

Part Of: Anthropogeny sequence
Content Summary: 1300 words, 13 min read.

The Domestication Syndrome

Since our emigration out of Africa 70,000 years ago, Homo Sapiens have domesticated many other species, including

  • dogs (18 kya, first domesticated in Germany)
  • goats, sheep (11 kya)
  • cattle, pigs, cats (10 kya)
  • llamas, horses, donkeys, camels, chickens, turkeys (5 kya)
  • foxes (50 years ago)

Consider the domestication of wolves into dogs. An important part of the environment of a species is other species- not merely its predators or pathogens but its symbionts. In this case, canines began to get food from human campsites. Dogs that were less aggressive were (by unconscious preference and conscious intent) more successful at extracting resources. This process is known as artificial selection.

Most ancient dogs kept by hunter-gatherers share a common body shape. More recently however, humans have conducted pedigree breeding: influencing the morphologies of different dog breeds. We have used this power to sculpt breeds as diverse as the Chihuahua and the Great Dane.

The defining feature of domestication is docility: a reduction in reactive aggression. All domesticated species exhibit this feature, in comparison to their wild counterparts. Not all species are capable of this sort of control. For example, humanity has tried for centuries to domesticate big fauna such as zebras, lions, and hippos. However, some breeds have reproductive and aggressive styles that prohibit domestication.

But domestication doesn’t just bring about a change in behavior. It also brings with it a bewildering number of anatomical changes, to essentially all domesticated species. The domestication syndrome include:

  • Docility (agreeableness, reduction in irritability)
  • Depigmentation (especially white patches, brown regions)
  • Floppy ears
  • Shorter ears
  • Shorter jaws
  • Smaller teeth
  • Smaller brains (10-15% reduction in volume)
  • More neotenous behavior (juvenile behavior that extends into adulthood).
  • Curly tails

Most domesticated species express some aspect of the domestication syndrome, as we can see in the following table:

Self-Domestication_ The Domestication Syndrome (1)

Three Theories of Domestication

The sheer complexity of the domestication syndrome requires an explanation. What is the link between floppy ears and docility?

Three hypotheses suggest themselves:

  1. Multiselection. Are the symptoms of domestication all expressions of human preferences? Do we simply like curly tails and floppy ears?
  2. Environment. Is there something about proximity to humans that incentivizes these changes?
  3. Byproduct. When the genes for aggression are altered, does that somehow incidentally cause these other changes?

Animal husbandry practices are lost to the sands of time. Nevertheless, there is a way to test multiselection directly: by creating a domesticated species in the laboratory.

In 1959, Dmitri Belyaev began trying to domesticate silver foxes. He used exactly one criterion for selection: he only bred pups that exhibited the least aggression. Skeptics thought it would take centuries to complete the domestication process. But changes in temperament were seen after only four generations. At twelve generations, “elite” foxes began to emerge with dog-like characteristics: wagging their tails, allowing themselves to be petted etc. At twenty generations, the entire population was considered fully domesticated.

Despite only selecting for docility, Belyaev’s foxes exhibited the full domestication syndrome. The foxes inexplicably developed floppy ears, curly tails, white patches, etc etc. The multiselection hypothesis is false.

Is there something about proximity to humans that selects for the domestication syndrome? The environment hypothesis seems false for two reasons.

  1. First, when they return to the wild, domesticated species take a long time reverting their characteristics. In fact, often domestication gives them a selective advantage over their wild cousins.
  2. Second, as we will see in the next section, self-domesticated species such as bonobos exhibit the syndrome despite their evolution not being influence by hominids.

The byproduct hypothesis is our only remaining explanation for the domestication syndrome. But what specific system produces these changes? 

The Biological Basis of Domestication

In order to fully explain aggression reduction, we must understand it at a biological level.

The primary basis of aggression reduction is a shrinking amygdala and periaqueductal gray (PAG). These modules comprise the negative valence system which learn which stimuli are negatively-valenced, and forward them to the mobilization system (e.g., snake → bad → run away). Serotonin inhibits the negative valence system, and domesticated animals have much high concentrations of serotonin receptors in these regions. Finally, it appears that these changes mostly act across development. The negative valence system comes online only slowly: there exists a socialization window in the first month of a wolf’s life, where it can learn “humans are okay”. Domestication primarily acts by increasing the socialization window from one to twelve months. If a dog isn’t exposed to a human in its first year, it’s now-active fear system will kick in: it will be wild for the rest of its life.

So what biological system is able to a) expand the socialization window, and b) induce the rest of domestication syndrome? The leading hypothesis involves a feature of development called the neural crest.

A blastocyst has no brain. To correct this unfortunate situation, every vertebrate genome contains instruction for constructing a neural tube. This structure emerges via folding.

The neural crest resides between the epidermis and the neural tube. These neural crest cells (NCCs) then proceed to migrate to a certain number of other anatomical structures to assist development. When the NCC migration malfunctions, the resultant disease is called a neurocristopathy. Many neurocristopathies result in outcomes similar to the domestication symdrome! For example, here is the effect of piebaldism:

Self-Domestication_ Piebalism

The mild neurocristopathy hypothesis (Wilkins et al, 2014) holds that domestication syndrome is a byproduct of changes to the NCC migration pattern.

Self-Domestication_ Mild Neurocristopathy Hypothesis

The hypothesis, however, is not very detailed (how exactly is NCC migration changed? What are the genomic and epigenomic contributions?). It is more of a promissory note than a mechanistic account. And there are other holistic hypotheses on offer, including genetic regulatory networks (Trut et al 2004) and action of the thyroid gland (Crockford 2000). It seems clear that, in the coming decades, a detailed mechanistic theory of domestication will emerge to vindicate the byproduct hypothesis.

Two Kinds of Domestication

Natural selection explains why the “design requires a designer” trope is obsolete. For the same reason, domestication can occur in the absence of a domesticator. More precisely, change in a species ecological niche can itself select against aggression.  Because aggression is very relevant to survival, we see plenty of species that have increased, and plenty that have decreased their rates of aggression. We call those less aggressive species self-domesticated: they became more peaceful in the absence of humans. What’s more, these species also exhibit the domestication syndrome.  

Another example is embedded in Foster’s Rule. Islands tend to be geologically more recent than continents, so their populations derive from the continent rather than vice versa. Islands tend to have fewer predators, but also fewer resources. Reduced predation increases the size of small animals (e.g., dodos evolved from pigeons), but limited resources decreases the size of big animals (e.g. the 3ft tall dwarf elephant).  

Self-Domestication_ Foster's Rule

Because islands have fewer predators, they also tend to have higher population densities; as such, reactive aggression is a less useful strategy. Selection favors the less aggressive. And we can see the domestication syndrome in island species. For example, the Zanzibar red colobus monkey has diverged from the continental red colobus along the same trajectory as dogs diverged from wolves.

Other examples of self-domestication can be found with group size reduction (ungulates, seals) and low-energy habitats (extremophile fish).

Finally, bonobos provide a particularly relevant example of self-domestication. Because food is more plentiful (don’t have to compete with gorillas for vegetation), females can spend time close to one another. Proximity produces bonding, and female coalitions exert pressure on bonobo behavior.

  • In chimps, bullying women increases reproductive success. Chimps will systematically beat up all females in their group as a coming-of-age ritual.
  • In bonobos, female coalitions retaliate against male aggression, making it unprofitable. Sexual selection then acts against reactive aggression.

So we can see that domestication (i.e., reduction in aggression) can come in two flavors: traditional vs self-domestication.

Self-Domestication_ Categories of Aggression Reduction (1)

As we will see next time, Homo Sapiens is yet another example of a self-domesticated species. See you then!

Related Resources

  • Wilkins et al (2014). The “domestication syndrome” in mammals: a unified explanation based on neural crest cell behavior and genetics

[Excerpt] Replicators and their Vehicles

Original Author: Richard Dawkins, The Selfish Gene
See Also: [Excerpt] The Robot’s Rebellion
Content Summary: 800 words, 4 min read

The First Replicator

Geochemical processes gave rise to the “primeval soup” which biologists and chemists believe constituted the seas some three to four thousand million years ago. The organic substances became locally concentrated, perhaps in drying scum round the shores, or in tiny suspended droplets. Under the further influence of energy such as ultraviolet light from the sun, they combined into larger molecules. Nowadays large organic molecules would not last long enough to be noticed: they would be quickly absorbed and broken down by bacteria or other living creatures. But bacteria and the rest of us are late-comers, and in those days large organic molecules could drift unmolested through the thickening broth.

At some point a particularly remarkable molecule was formed. We will call it the Replicator. It may not necessarily have been the biggest or the most complex molecule around, but it had the extraordinary property of being able to create copies of itself.

A molecule which makes copies of itself is not as difficult to imagine as it seems at first, and it only had to arise once. Think of the replicator as a mold or template. Imagine it as a large molecule consisting of a complex chain of various sorts of building block molecules. The small building blocks were abundantly available in the soup surrounding the replicator. Now suppose that each building block has an affinity for its own kind. Then whenever a building block from out in the soup lands up next to a part of the replicator for which it has an affinity, it will tend to stick there. The building blocks which attach themselves in this way will automatically be arranged in a sequence which mimics that of the replicator itself. It is easy then to think of them joining up to form a stable chain just as in the formation of the original replicator. Should the two chains split apart, we would then have two replicators, each of which can go on to make further copies.

Replicator Competition

The primeval soup was not capable of supporting an infinite number of replicator molecules. For one thing, the earth’s size is finite, but other limiting factors must also have been important.

But now we must mention an important property of the copying process: it is not perfect. mistakes will happen. I hope there will be no misprints in this book, but if you look carefully you may find one or two. We do not know how accurately the first replicator molecules made their copies. Their modern descendants, the DNA molecules, are astonishingly faithful compared with the most high-fidelity human copying process, but even they occasionally make mistakes, and it is ultimately these mistakes which make evolution possible. Mistakes were made, and these mistakes were cumulative.

Replicators with a comparatively worse design must actually have become less numerous because of competition, and ultimately many of their lines must have one extinct. There was a struggle for existence among replicator varieties. They did not know they were struggling, or worry about it; the struggle was conducted without any hard feelings, indeed without feeling of any kind. But they were struggling, in the sense that any mis-copying which resulted in a new improved level of stability, or a new way of reducing the stability of rivals, was automatically preserved and multiplied.

This process of replicator improvement was cumulative. Ways of increasing stability and of decreasing rivals’ stability became more elaborate and more efficient. Some of them may even have ‘discovered’ how to break up molecules of rival varieties chemically, and to use the building blocks so released for making their own copies. These proto-carnivores simultaneously obtained food and removed competing rivals. Other replicators perhaps discovered how to protect themselves, either chemically, or by building a physical wall of protein around themselves. This may have been how the first living cells appeared.

Replicator Self-Improvement

Replicators began not merely to exist, but to construct for themselves containers, vehicles for their continued existence. The replicators that survived were the ones that built survival machines for themselves to live in. The first survival machines probably consisted of nothing more than a protective coat. But making a living got steadily harder as new rivals arose with better and more effective survival machines. Survival machines got bigger and more elaborate, and the process was cumulative and progressive.

Was there to be any end to the gradual improvement in the replicators’]techniques? What weird engines of self-preservation would the millennia bring forth?  Four thousand million years on, what was to be the fate of the ancient replicators?

They did not die out, for they are past masters of the survival arts. But do not look for them floating loose in the sea; they gave up that cavalier freedom long ago. Now they swarm in huge colonies, safe inside gigantic lumbering robots, sealed off from the outside world, communicating with it by tortuous indirect routes, manipulating it by remote control..

They are in you and in me; they created us, body and mind; and their preservation is the ultimate rationale for our existence. They have come a long way, those replicators. Now they go by the name of genes, and we are their survival machines.