Showing posts with label Brain Evolution. Show all posts
Showing posts with label Brain Evolution. Show all posts

Friday, May 22, 2009

An Evolutionary Perspective on the Human Brain

This term I wrote an essay on the topic of human uniqueness from an evolutionary perspective. As I drew on research that is also relevant to this blog, (and to some extent has already been covered here). I'll post some of it here.
In this post I'll have a short look at the human brain from a neuroscientific, a comparative, and an evolutionary perspective:

Human Evolution

We are evolved primates. (As are all other primates of course. So maybe it is better to say that we, like all other primates, are evolved beings with a unique set of specializations, adaptations and features. )

In our lineage, we share a common ancestor with orangutans (about 15 million years ago (mya)), gorillas (about 10mya), and most recently, chimpanzees and bonobos (5 to 7 mya). We not only share a significant amount of DNA with our primate cousins, but also major anatomical features (Gazzaniga 2008: 51f., Lewinn 2005: 61) These include, for example, our basic skeletal anatomy, our facial muscles, or our fingernails (Lewin 2005: 218ff.).


What most distinguishes us as humans on an anatomical level are our bizarre hair distribution, our upright posture and the skeletal modifications necessary for it, including a propensity for endurance running, our opposable thumbs, fat deposits that are unusually extensive (Preuss 2004: 5), and an intestinal tract only 60% the size expected of primates our size (Gibbons 2007: 1558).


Finally, there is also a distinguishing feature that is a much more remarkable violation of expectations – a brain three times the size expected of a primate our size. This is all the more interesting as primates are already twice as encephalized as other mammals (Lewin 2005: 217). A direct comparison shows this difference in numbers: Whereas human brains have an average volume of 1251.8 cubic centimetres and weigh about 1300 gram, the brains of the other great apes only have an average volume of 316.7 cubic centimetres and weigh between 350-500 gram (Rilling 2006: 66, Preuss 2004: 8). In a human brain, there are approximately a hundred billion neurons, each of which is connected to about one thousand other neurons, comprising about one hundred trillion synaptic connections (Gazzaniga 2008: 291). If you would count all the connections in the napkin-sized cortex alone, you would be finished after 32 million years (Edelman 1992: 17).


Expensive Tissue

The human brain is also extremely “expensive tissue” (Aiello & Wheeler 1995): Although it only accounts for 2% of an adult’s body weight, it accounts for 20-25% of an adult’s resting oxygen and energy intake (Attwell & Laughlin 2001: 1143). In early life, the brain even makes up for up 60-70% of the body’s total energy requirements. A chimpanzee’s brain, in comparison, only consumes about 8-9% of its resting metabolism (Aiello & Wells 2002: 330). The human brain’s energy demands are about 8 to 10 times higher than those of skeletal muscles (Dunbar & Shultz 2007: 1344), and, in terms of energy consumption, it is equal to the rate of energy consumed by leg muscles of a marathon runner when running (Attwell & Laughlin 2001: 1143). In all, its consumption rate is only topped by the energy intake of the heart. (Dunbar & Shultz 2007: 1344).

Consequently, if we want to understand the evolutionary trajectory that led to human cognition there is the problem that “because the cost of maintaining a large brain is so great, it is intrinsically unlikely that large brains will evolve merely because they can. Large brains will evolve only when the selection factor in their favour is sufficient to overcome the steep cost gradient“ (Dunbar 1998: 179). We have to come up with a strong enough selection pressure operative in the Pleistocene environment of evolutionary adaptedness that would have allowed such “expensive tissue” to evolve (Bickerton 2009: 165f.).


What About the Brain is Uniquely Human?


If we look to the brain for possible hints, we first find that presently, there is “no good evidence that humans do, in fact, possess uniquely human cortical areas” (although the jury is still out) (Preuss 2004: 9). In addition, we find that there are functions specific to humans which are represented in areas homologous to areas of other primates. Instead, it seems that in the course of human evolution some of the areas of the brain expanded disproportionally, “especially higher-order cortical areas, including the prefrontal cortex” (Preuss 2004: 9, Deacon 1998: 435-438). This means that humans do not only think in a better way, but that they think differently (Preuss 2004: 7). The expansion and apparent specializations of only certain kinds of neuronal areas could indicate a qualitative shift in neuronal activity brought about by re-organization of existing features, leading to a wholly different style of cognition (Deacon 1998: 435-438 Rilling 2006: 75).

This scenario squares well with what we know about the way evolution works, namely that it always has to work with the raw materials that are available, and constantly co-opts and tinkers with existing structures, at times producing haphazard, cobbled-together, but functional results (Gould & Lewontin 1979, Gould & Vrba 1982). Given the relatively short time span for the evolution of the “most complex structure in the universe, we have to acknowledge how preciously little time the evolutionary process had for ‘debugging.’ It could well be that make the human mind so unique is that it is a imperfect ‘Kluge:’ a clumsy or inelegant – yet surprisingly effective – solution to a problem,” like the Apollo 13 CO2 filter or an on-the-spot invention by MacGyver (Marcus 2008: 3f.). It may thus well turn out that what we think makes us so special is a mental “oddity of our species’ way of understanding” the world around us (Povinelli & Vonk 2003: 160). It is reasonable then to assume that human cognition did not just simply get better across the board, but that instead we owe our unique style of thinking to quite specific specializations of the human mind.

With this in mind, we can now ask the question how these neurological differences must translate into psychological differences. But this is where the problem starts: Which features really distinguish us as humans and which are more derivative than others? A true candidate for what got uniquely human cognition off the ground has to pass this test and solve the problem how such “expensive tissue” could evolve in the first place.


References:

Aiello L.C. and P. Wheeler 1995. ”The expensive tissue hypothesis: the brain and the digestive system in human and primate evolution.” Current Anthropology 36:199–221


Aiello, Leslie C. and Jonathan C. K. Wells (2002): “Energetics and the Evolution of the Genus Homo.” In: Annual Review of Anthropology 31:323–38.


Attwell, David and Simon B. Laughlin. (2001.) “An Energy Budget for Signaling in the Grey Matter of the Brain.” Journal of Cerebral Blood Flow and Metabolism 21:1133–1145.


Bickerton, Derek (2009): Adams Tongue: How Humans Made Language. How Language Made Humans. New York: Hill and Wang.


Deacon, Terrence William (1997). The Symbolic Species. The Co-evolution of Language and the Brain. New York / London: W.W. Norton.


Dunbar, Robin I.M. (1998)“The Social Brain Hypothesis” Evolutionary Anthropology 6: 178-190.


Dunbar, R. I. M. and Susanne Shultz. (2007)“Evolution in the Social Brain” Science 317: 1344-1347


Edelman, Gerald Maurice (1992) Bright and Brilliant Fire: On the Matters of the Mind. New York: Basic Books


Gazzaniga, Michael S. (2008): Human: The Science of What Makes us Unique. New York: Harper-Collins.

Gibbons, Ann. (2007) “Food for Thought.” Science 316. 1558-1560.


Gould, Stephen Jay and Richard Lewontin (1979). "The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme". Proclamations of the Royal. Society of London B: Biological Sciences 205 (1161): 581–98.


Gould, Stephen Jay, and Elizabeth S. Vrba (1982), "Exaptation — a missing term in the science of form," Paleobiology 8 (1): 4–15.


Lewin, Roger (2005): Human Evolution: An Illustrated Introduction. Oxford: Blackwell.


Marcus, Gary (2008): Kluge: The Haphazard Evolution of the Human Mind. London: Faber and Faber.


Povinelli, Daniel .J. and Jennifer Vonk. (2003) Chimpanzee minds: Suspiciously human? Trends in Cognitive Sciences, 7.4, 157–160.


Preuss Todd M. (2004): What is it like to be a human? In: Gazzaniga MS, editor. The Cognitive Neurosciences III, Third Edition. Cambridge, MA: MIT Press: 5-22


Rilling, James K. (2006.) “Human and NonHuman Primate Brains: Are They Allometrically Scaled Versions of the Same Design?” In: Evolutionary Anthropology 15: 67-77

Sunday, June 22, 2008

Thriller-Writer Lee Child and Linguist Dieter Wunderlich on the Evolution of Language


Lee Child is one of my favorite Thriller-authors. Normally I don’t find the time to read non-studying related fiction, but I especially enjoy his audio books because of the cool American accent of the narrator. However, when I stumbled on an essay by Lee Child called “The Origin of the Thriller”, I was a bit confused. Here’s his take on the evolution of language: According to Lee Child, a couple of hundred thousand years ago, Neanderthals and humans, the only homo “contenders” left were competing for resources, and Neanderthalers had the upper hand:

“They were heavier and stronger and faster. They were superb tool makers. They were much better equipped to survive the brutal conditions of prehistory.

But they didn't survive. We did. Why?

Because Homo Sapiens developed language. Many primitive species could communicate by making sounds — and many still do: prairie dogs make distinctive noises if a predator is spotted — one noise for a ground predator, and another for an airborne predator. But Homo Sapiens went beyond two words. After a random mutation our brains grew large and the new capacity was colonized by language, with a theoretically infinite number of available words, and more importantly with syntax, such that as well as reporting we could plan and speculate. Not just: a predator is coming, but also: a predator will come, or might come. Not just reaction, but also prediction: if we do this, we'll be OK, or if we do that, we'll be in trouble. “

But here’s the gem According to Child, humans are weak and fragile and in disorganized groups we would quickly be killed by predators:

“But a coordinated crowd of two hundred humans is the most powerful animal on earth. The heaviest, the strongest, the hardest to stop, the hardest to kill. Thus, grunting Neanderthals slowly died out, despite their muscle and bone and strength and speed, and talking humans marched on toward the present, despite our slender limbs and fragile skulls”
The last Sentence is faboulus isn’t it? We better don’t tell him anything about Neanderthal-DNA, Neanderthal Vocal Anatomy, FOXP2, Introgression, etc.

But I want to present a more plausible and worked out view on the topic. Yesterday I was searching for material on the evolution of syntax, because a friend of mine asked me to comment on a section of his MA-Thesis in which he shortly discusses the topic. (For a discussion of the speculation, that certain features of language are ‘living fossils’, or vestiges of an older proto-language, see this post).

On my search I found the slides of a talk given by German linguist Dieter Wunderlich on the evolution of language, that he held in Leipzig, Germany, in December 2007. The talk and the slides are in German, but because he offers a very comprehensive and nice review on some aspects of the current state of the art in language evolution research/evolutionary linguistics, I decided to write an English summary of it.


Key Questions

When looking at the evolution of language, there are three question that guide our inquiry:

1. How exactly did language get started? What is hotly debated is whether it evolved in a gradual manner (continuist/adaptationist position: see e.g. Pinker & Bloom 1990) or in a ‘sudden leap’ (discontinuist/ exaptationist: see e.g. Hauser et al. 2002)

2. When did language evolve? Depending on our answer to question 1., the question would be when each step toward language took place, or when the big leap happened.

3. Did language evolve only once or several times?


What is language?

Basically, language can be described as a system for expression, that relates Utterances and meaning to one another, in relation to certain contexts.

Morphosyntax governs the construction of complex utterances out of smallest units of language, namely lexical entries (=”words” in your mental lexicon) as well as morphemes (-s, -ed, un-, -er, etc.). Its system is compositional, that is every construction leads to complex expressions of meaning which are determined by the single lexical entries as well the rules that govern their combination. The rules governing the compositional construction of utterances are recursive.

There are two interfaces to non-linguistic abilities:

1. an interface with the system responsible for the production and perception of sounds and gestures (In generativist terminology this would be the Sensori-Motor System (SM) or the Articulatory-Perceptual (AP) System)

2. an interface with our mental representations and our discourse intentions (in generativist terminology: the Conceptual-Intentional (CI) system. cognitive linguists would probably call it our conceptual system)


Important events which shaped the current state of language

There are 5 important events that have shaped how languages and our mental structures that enable us to use it look:

In chronological Order:

1. There was an expansion of the cortex (frontal lobe) in early hominids such as homo erectus, who lived approx. 2 million years ago. In just 1 million years, brain size rose from 600 to 1400ccm. This may be related to the begin of an ice-age about 2.5 million years ago.

2. Anatomically modern man (homo sapiens) probably evolved about 170.000 years ago in east Africa.

3. Starting from about 50.000 years ago, there is archaeological evidence of improved tool-making capacities, burials, ornaments, and figurative art in several parts of the world (the so-called Upper Palaeolithic Revolution). In Africa these features can be found even earlier, as is especially salient from about 85.000 years ago, which even led some researchers to call this period an “African Upper Palaeolithic” (see e.g. Dubreuil 2008).

4. From 12.000 years ago onwards, there was a spread in agriculture which seemed to have happened independently in several parts of the world (The Neolithic Transition)

5. Starting from 6000 years ago, writing systems were developed independently in several parts of the world.

Of these 5 events, the first three are biological of nature, whereas the last two are cultural events. A diversification of biological events brought about by mutations and selection led to one single event: genetically modern man. In principle, every human being can reproduce with every other opposite-sex human, which fulfils the criterion of biological species. Furthermore, every child, regardless of its parents, can learn every language of the world when it is put into the critical environment at the right age. Thus language is a common and uniting feature of humans, and we all share the genetic structure that enables us to learn language (but for evidence that there is still genetic variation on a small scale, see these two posts)


What are the differences between and non-human communication systems?

Influential linguist Charles F. Hockett has posed 13 ‘design features of language’, 7 of which are especially interesting regarding the evolution of language. These are:


1. For the majority of human languages, the vocal-auditory channel the basic mode of communication.


2. Parity/Interchangeability: hearer and speaker constantly switch roles during conversation, which also means that a communicator is able to both produce and receive the same signal, which isn’t the case with say, gender-specific calls in sticklefish.


3. Semanticity/Arbitrarines: specific signals can be equipped with specific meaning in a manner where there is no necessary or logical connection between form meaning (“table” has nothing to do with any actual properties of a table)


4. Displacement: humans can talk about things that are not present in their immediate environment, or even things that do not exist at all (say, for example, my comprehension of mathematics). Human reference thus extends “beyond the horizon” and the perceptual space, but extends to mental spaces and shared systemic spaces.


5. Honey bee communication about the location of nectar of course is also displaced, but it only built to convey very specific information, and is not unbounded and productive like human language.


6. Traditional Transmission/Learnability human children are able to learn the language of their environment. As W.v.O. Quine said “language is a social art”, which essentially depends on the ability to be part of and interact with a complex culture that transmits complex behavior and knowledge. This complexity and language-specificness is probably the dividing feature, because song-birds also have to learn their songs from other birds, vervet monkeys need to be reinforced in their innate tendencies to hiss at snakes and have to zoom in on the exact referent for eagle alarm callas through positive reinforcements by other group members, and chimpanzees have been shown to adhere to some forms of social transmission and conformity as well as for rational imitation (e.g. Whiten et al. 2005)


7.Discreteness/Duality of Patterning the smallest units of language combine with each other in a systematic and combinatorial manner (that is Phonemes/Sound units combine to form Morphemes) , which then also combine in a systematic fashion (Morphemes built whole words and then sentences). This method allows for “infinite expression by finite means” (Humboldt) and is responsible for the diverse displaced productivity of language.

Wunderlich argues that homo erectus already possessed most of these features, but maybe not 6. and 7.


So what could homo erectus do?

Although he certainly didn’t have full-fledged human language, he was able to perform complex and social tasks. From 1.5 million years ago onwards, he was able to keep a fire burning, and from 0.8 million years ago he could even make fire.

He was able to make hand-axes (1.4 million years ago) and was also able to throw them, which, according to William Calvin, is suggested by neuropaleontological evidence. He way able to hunt together with other group members. He was very mobile: signs of him can be found in Georgia (1.8 million), later on there are also signs in China, Indonesia, and Europe.

Homo erectus was divided in several species that probably lived alongside each other. The last ones co-existing with us were the Neanderthals.


Tool use

Chimpanzees and orangutans use tools (see here, for a cool post on spear fishing in orangutans)such as sticks for termite fishing, stones to crack open nuts, and leaves to soak up water, but these tools are contextually available, that is they do not have to be crafted in the same complex way as hand-axes, for example. Interestingly, there is population level handedness of 2:1 in chimpanzees. Termite fishers are mostly left-handed, whereas nut-crackers and wadge-dippers are mostly right-handed, this means that handedness seems to be heritable. In humans, the left-right ratio is 9:1. The distribution of handedness means that handedness is task-specific, and “

that the motor and cognitive demands of different tasks can have a significant influence on handedness in human and nonhuman primates” (Lanson & Hopkins 2005).
This means
“that antecedents of lateralization of function associated with hand use were present at least 5 million years ago, before the Pan-Homo split,”
and that due to tool-use the human-brain specialized, probably making possible further changes in human neurology and cognitive behaviour (for evidence that there is a neurological link between tool-making and speech, see this post). It is interesting to consider what the cognitive preconditions were that enabled tool-making. The Oldowan technique, which
can be found from 2.6 million years ago. This technique is used to make cutting tools by breaking off sharp-edged flakes from a larger stone. This is achieved by striking a core stone with a hammerstone, thereby knocking flakes from the core (see Ambrose 2001).


This technique definitely requires intentional planning that isn’t bound to the here-and-now, but directed at the future. Also, and more important for language, it requires the ability for complex propositional thought. Thus the following conscious key cognitive mechanisms found in language are also necessary for tool-making and thus already existed 2.6 million years ago:


1. Predication: the same referent (core stone), can be subject of different predications, a structure that can be called “Topic-Comment”-structure (see also this post):

George (Topic) is eating brains (Comment) / George (Topic) is a Zombie (Comment) / George (Topic) is (Comment).

2. Qualification of predications that have already been made and higher-order temporal hierarchical ordering.

To wit, what this basically means is that both in linguistics and tool-making you first have an initial structure which you change by commenting/acting on it. This new structure can then be the topic of new comments or actions, and the new structure arising from these computations/ actions can again be the topic of new predications and ad infinitum. Consider for example the sentence: The living homeless, who wanted money, had infested South Park, but then they went to California

A hierarchical behavioral sequence like toolmaking seems to have similar properties:


I don’t know about you, but for me this looks like recursion, the only difference being that for toolmaking the recursive operation are applied to a physical object, and for language they are applied to a mental object. However, if toolmaking requires intentional planning, there had to be at least some recursive mental operations that in mental action planning. This would mean that, pace Hauser et al. (2002), recursion is neither special to language nor recently evolved.

As Wunderlich writes in his (2006) paper “What forced syntax to emerge?”

"Both fast processing and mapping from hierarchical structure to temporal ordering, two other fundamental features of human language, can be attributed to progresses in the timing of actions necessary for producing and using hand axes, [as well as throwing] that is, to sensomotoric skills that could have been adapted for language.”

All this is made all the more exciting by converging evidence that combinatorial and hierarchical structure of any kind (be it linguistic, cognitive, or sensori-motor are constructed by the same part of the brain and via the same mechanisms:

“Similarities exist between the architecture for sentence structure and blackboard architectures for combinatorial structures in visual cognition, derived from the structure of the visual cortex” (van der Velde & de Kamps 2006: 1)

“Evidence is mounting that much temporally sequenced hierarchical structure is constructed by the same part of the brain – roughly Broca’s area and subcortical areas connected to it – whether the material being assembled is language, dance (Lieberman 2005: 297), hand movements (Lieberman 2005: 294; Wilkins 2005: 279), or music (Patel 2003). Nor, within language, is Broca’s area confined to syntax, as often asserted: it also appears to play a role in phonological and semantic combinatoriality, possibly in distinct though overlapping subareas (Hagoort 2005; Poeppel and Embick 2005)" (Jackendoff 2007: 388)

“On the basis of all these results, it can be hypothesized that [the recursive morphosyntax of modern language] is the computational output of a cortical premotor network originally evolved to control/represent the hierarchical structure of goalrelated action When in evolution, selective pressure led to the emergence of language, the same neural circuits doing computations to control the hierarchy of goal-related actions were ‘exploited’ to serve the newly acquired function of language syntax. A similar functional overlap between action and language acquisition is indeed evident during children’s development, i.e. children parallel their capacity to master hierarchical complexity both in the domain of language and goal-related action.“ (Gallese 2007: 666)”

That’s it for today. In my next post I will continue my summary of Wunderlich’s presentation.


References:


Ambr Ambrose, Stanley H. 2001. Paleolithic Technology and Human Evolution. Science 291, no. 5509 (March 2): 1748-1753. doi:10.1126/science.1059487.


Dubreuil, Benoît (2008): “What do modern behaviours in Homo sapiens imply for the evolution of language?”, in A. D. M. Smith, K. Smith, and R. Ferrer i Cancho (eds.), The Evolution of Language. Proceedings of the 7th International Conference (Evolang 7), World Scientific, 99-106.


Gallese, Vittorio (2007). Before and below 'theory of mind': Embodied simulation and the neural correlates of social cognition. Philosophical Transactions of the Royal Society B-Biological Sciences 362 (1480):659-669



Hagoort, Peter (2005). On Broca, brain, and binding: a new framework. Trends in Cognitive Sciences 9: 416–423



Hauser, Marc D., Noam Chomsky and W. Tecumseh Fitch (2002). “The Faculty of Language: What Is It, Who Has It, and How Did It Evolve?” In: Science 298, 1569-1579.


Jackendoff, Ray (2007): Linguistics in Cognitive Science: The State of the Art, The Linguistic Review 24, 347-401.


Lieberman, Phillip (2005): The pied piper of Cambridge. The Linguistic Review 22: 289–302.


Lonsdorf, Elizabeth V., and William D. Hopkins. 2005. Wild chimpanzees show population-level handedness for

tool use. Proceedings of the National Academy of Sciences 102, no. 35 (August 30): 12634-12638. doi:10.1073/pnas.0505806102.


Patel, Aniruddh D. (2003). Language, music, syntax, and the brain. Nature Neuroscience 6: 674– 681.


Pinker, Steven & Paul Bloom (1990). “Natural Language and Natural Selection.” In: Behavioral and Brain Sciences 13.4: 707-726.


Poeppel, David, and David Embick (2005). Defining the relation between linguistics and neuroscience. In Twenty-first century psycholinguistics: Four cornerstones, A. Cutler (ed.), Hillsdale, NJ: Erlbaum.


Van der Velde, F. & de Kamps, M. (2006). Neural blackboard architectures of combinatorial structures in cognition. In: Behavioral and Brain Sciences, 29, 1-72.


Whiten, Andrew, Victoria Horner & Frans B. M. de Waal. 2005. “Conformity to Cultural Norms of Tool Use in Chimpanzees.” Nature 437: 737-740.


Wilkins, Wendy K. (2005). Anatomy matters. The Linguistic Review 22: 271–288.


Wunderlich, Dieter (2006): “What forced syntax to emerge?” In H.-M. Gärtner et al. (eds.) Between 40 and 60 puzzles for Krifka. ZAS Berlin

Thursday, December 6, 2007

A Zombie’s Inquiry Into the Evolution of His Most Favorite Meal IV: Genes that Code for Tasty brains

There are two main approaches to look at the differences between humans and other non-human primates such as chimpanzees: ethological studies of animal behavior and their cognitive abilities (it looks like there is a difference between Cognitive Ethology, Comparative Ethology & Comparative Psychology, but as it seems this is more a matter of whether you emphasize the biological, cognitive science, or psychological aspect of behavior) and genomic comparisons.

On the side of genetic comparisons, we already have the sequenced genome of humans, chimpanzees, and macaques, which, somewhere in the relatively near future, the future, are to be joined by the genomes of Neanderthals, bonobos, (both sequenced by our friends at the Max Planck Institute for Evolutionary Anthropology - gosh! it would really have been a tremendous loss for science if its members had been eaten by zombies… So thanks for that George) gorillas, and gibbons. As Kambiz Kamrani pointed out over at primatology.net, the more primate genomes we get together, the better we are able to make out human specialness (as well as Chimpanzees-Specialness, Bonobo-Specialness, Gorilla-Specialness etc.), as well as the things we share with other primates, in terms of specific genes.
This may indeed help us ““to find out what being human is.” James Watson originally hoped this would be the result of the sequencing of the human genome. (Pennisi 2007: 218) but now, with an ever-growing genetic database, we somewhere in the future we may indeed be able “to trace back the evolutionary changes that occurred at various time points, leading from the common ancestors of the primate clade to Homo sapiens,” as Bruce Lahn puts it. (Pennisi 2007: 218). Researchers all over the world further plan on sequencing the genomes of the orangutan, the marmoset, the tarsier, the mouse lemur, the galago, the tree shrew as well as the lemur in order get an ever broadening picture of our evolutionary history, ultimately tracing back 83 million years of evolutionary time.
Within this comparative context, we of course may really see the “dawn of cognitive genetics” (Pinker 2001: 465). In the field of language evolution, for example, we may finally establish the genetic foundations and extensions that made human language possible. But at the moment, it seems as if there are still so much things that are maddeningly unclear, (and I as layman, naturally don’ understand anything about “regulatory sequences”, “junk DNA” and messy genetic differences at the molecular level…) so it’s definitely still a very long way until we can go beyond FOXP2, MPH1, and ASPM (not to speak of understanding even the exact roles of these genes.)

On the side of ethological studies, the methods employed and results obtained (which I describe in my last post) by Herman et al. (2007) clearly show interesting avenues of future research, and hint at a possible meeting point between the two approaches:
“A major avenue of future research is thus to use [the research methodologies employed by Herman et al.] to characterize the behavioral-cognitive phenotype of a wide variety of primate species. This could be done through systematic testing of carefully chosen representatives of the more than 50 genera of primates, which should then enable us to map out cladistically the evolution of primates’ most important cognitive skills at the level of both the phenotype and, ultimately, the genotype.” (Herman et al. 2007: 1365)
For our Zombie-Scientist George the main question remains: “Why are human brains so tasty ?” (let’s presume that in our Parallel zombieverse, the zombie-gourmet is only fond of human brains and not that of other non-human brains.) The Theory by which George now arrives looks like this: “The unique tastiness of human brains basically must boil down to some uniquely human genes (or genetic combinations or gene expressions)” That’s why I will lok a bit at the differences between human and chimp-genes in my next post.

References:

Hermann, Esther Josep Call, María Victoria Hernández-Lloreda, Brian Hare, and Michael Tomasello. 2007. “Humans Have Evolved Specialized Skills of Social Cognition: The Cultural Intelligence Hypothesis” Science 317: 1360-1366.

Pennisi, Elisabeth. 2007 “Genomicists Tackle The Primate Tree.” Science 316: 218-221.

Pinker, Steven. 2001. “Talk of genetics and vice versa“ Nature 413: 465-466

Monday, December 3, 2007

A Zombie’s Inquiry Into the Evolution of his Most Favorite Meal III: What are Humans Good at?

Except running away from poor, starving zombies, that is (– the Selfish Bastards!)

A while back Juan Uarigerika wrote an article in Seed magazine about language evolution. In it he proposed that maybe language is responsible for most of our especially human intelligence, as well as precursor for our more advanced sensorimotor capacities, and that in the end it could turn out that research into our cognitive architecture would come up with the formula ‘Finch + Chimp = Human.’ Uarigerika’s article is written, for a magazine, so it’s clear that he doesn’t really do much in order of presenting evidence and arguments in a really ‘scientific’ way but rather presents his ideas in a in a popular style, but still I think his proposal is quite problematic (Mark Liberman has written a nice rebuttal of Uarigerika’s reductionsit view over at Language Log)

funny pictures
moar funny pictures

A better way to study the differences between human and animal cognition effectively is to compare differences and similarities of certain cognitive traits and analyze how these may come about and how the cognitive function in question is enabled in the given organism.
In a massive comparative study, Hermann et al. (2007) had Chimpanzees, Orangutans and 2.5 year-old children perform various task and then evaluated and compared the species’ qualitatively differing performances. The tasks were divided into two “domains”, physical and social, each consisting of three “scales” (physical: space, quantitiy, causality; social: social learning, communication, theory of mind). Among the 20 tasks there were such things as “using a stick in order to retrieve a reward which is out of reach.” (causality), “Locating a reward.“ (space), “Solving a simple but not obvious problem by observing a demonstrated solution” (social learning), “Following an actor’s gaze direction to a target” or “Understanding what an actor intended to do (unsuccessfully” (both Theory of Mind).

On average, the results of humans and chimpanzees were very similar in the physical domain, and scored much higher than the orangutans. In the social domain, however, humans outperformed chimpanzees and orangutans by far. The non-human apes were right only half as often as the human children. This means that chimpanzees outcompete orangutans when it comes to things as causal reasoning and quantities, but are equally bad at imitating others or assessing their intentions. Whereas in the physical tasks chimps sometimes performed better than humans (e.g. Tracking of a reward after location changes or Using a stick in order to retrieve a reward which is out of reach, something where human children performed much worse than both chimps and orangutans), interestingly
“Children were better than both ape species at the three causality tasks in which a judgment must be made before manipulation or choice, whereas chimpanzees were better than children and orangutans at the one causality task involving active tool use.” (Hermann et al. 2007:1362)
as well as in regard to inhibitory control, which could partly be due to the prominence and dominance of prefrontal circuitry in the human brain and its importance in cognitive control— “the ability of the brain to coordinate processing mong its millions of neurons in order to direct them toward future goals.” (Miller et al. 2002: 1131) — which I alluded to in my earlier posts.

Chimps and orangutans both performed a little better than human children when it came to “Producing communicative gestures in order to retrieve a hidden reward.”, which was the only social domain task in which the difference between the human and non-human primates wasn’t significant. The authors conclude that
“the current results provide strong support for the cultural intelligence hypothesis that human beings have evolved some specialized social-cognitive skills (beyond those of primates in general) for living and exchanging knowledge in cultural groups: communicating with others, learning from others, and “reading the mind” of others in especially complex ways“ (Hermann et al. 2007: 1365).
But they caution against the conclusion that social intelligence as a whole, or a “Theory of Mind-module” is the distinctive property separating humans, chimps and orangutans. Instead, taking into account that human children were better than chimps in causality tasks that didn’t include the active manipulation of tools, they speculate that
“what may be distinctive is the ability to understand unobserved causal forces in general, including (as a special case) the mental states of others as causes of behavior. Even in this case, however, it is a plausible hypothesis that understanding hidden causal forces evolved first to enable humans to understand the mental states of other persons, and this generalized only later to the physical domain.”
Which fits well with the evidence that humans are especially good at displaced mental and conceptual simulation (Miller et al. 2002, Barsalou 2005) and such things as mental time travel (Gilbert & Wilson 2007).
In my next post I will expand a bit on complementary approaches to comparing human and other non-human primate cognition and differences in general.

References:

Barsalou, Lawrence W. 2005. “Continuity of the conceptual system across species.” Trends. Cog. Sc. 9.7: 309-311.


Gilbert, Daniel T. and Timothy D. Wilson. 2007. “Prospection: Experiencing the Future.” Science 317: 1351-1354.

Hermann, Esther Josep Call, María Victoria Hernández-Lloreda, Brian Hare, and Michael Tomasello. 2007. “Humans Have Evolved Specialized Skills of Social Cognition: The Cultural Intelligence Hypothesis” Science 317: 1360-1366.


Miller, Earl K., David J. Freedman and Jonathan D. Wallis 2002. “The Prefrontal Cortex: Categories, Concepts and Cognition.” In: Phil. Trans. R. Soc. Lond. B 357: 1123–1136

Thursday, November 29, 2007

A Zombie’s Inquiry Into the Evolution of his Most Favorite Meal II

In my last post I listed some important factors in the evolution of the human brain, or better imagined what a zombie evolutionary biologist, named George, might dig up when investigating the evolutionary path of his Dinner Nr. 1. As Terrence Deacon (1997) puts it, there is no escaping the fact that human brains are unusually large. There are several factors why humans could develop large brains, but what is still at stake is the question why they actually did, and how they came into a position that allowed them to devote so much energy to a walnut-shaped pink lump of tissue with the consistency of a half-baked egg.
This question is critical, because organisms do not normally develop new traits just because they can. This of course also happens, in combination with random genetic drift and populations bottlenecks. But because evolution is a ‘miserable and greedy tinkerer’, or put more nicely, an economical process, it is highly unlikely that it produces needless and incredibly complex capacities that are extremely costly to maintain. As a consequence,
“some proportionately beneficial advantage must have driven brain evolution against the steep selection gradient created by the high costs of brain tissue.” (Dunbar & Shultz 2007)
Why, Dunbar and Shultz ask further, do primates have so much bigger brains than squirrel, with both facing about the same foraging decisions? (Dunbar & Shultz argue that ecological explanations fail to account for this differences, but their Chimpanzee-Squirrel dichotomy nevertheless is a bit hyperbolical, given that, among primates, those whose diet includes insects and fruits show higher encephalization rates than leaf-eaters, and strategic hunting and gathering of food and prey places additional demands on navigational, representational and other cognitive skills. However, their general argument is still valid. (Park et al. 2007))

To shed light on this issue, we can divide the big picture into several smaller ones. Useful questions include: What are we good at? Split into What are we (primates) good at? And What are we (humans) even better at than other primates? What could the ecological niche favoring big brains in humans have looked like? How exactly does our brain differ from that of other primates?
These questions essentially depend on comparative ethology (how do our minds work compared to how the minds of other animals work?), comparative (neuro)anatomy (On which evolutionary foundations are our modern cognitive abilities, and other phenotypic traits built upon?), and the kind of scenario we envision or infer from these observations togther with the fossil record and other lines of evidence. Of course it is also crucial what we think what the most salient and essentially aspects of our ancestors were. Do we see our ancestor as “Man the Tool Maker”, “Man the Hunter” or “Man the Social Animal”, or just as “Man with the extraordinarily big & expensive (and extremely delicious, George might add) brain”.

Well, of course Man should probably rather be seen as “Man the cooperative, competitive, tool-making, hunting, {…}, articulate social animal.” And all of these property probably contributed (co-evolutionary, we might say, without adding much in terms of explanatory adequacy) to our cognitive abilities and brain size, but in which order? And which driving forces were a little more pushy than others?
As Cheney & Seyfarth (2007) have show in baboons, interactions in primate groups are cognitively highly demanding and require sophisticated representational and predictive abilities, because of the intricate and complex networks and ‘friendships’ they inherit. Thus rising complexity in social life could be seen as a key selection pressure in the evolution of cognitive abilities in primates in general, and especially in humans. (Lewin 2005: 220f.)
Depending on which aspect one wants to stress, this correlations can be described in different terms. Scholars who wanted to stress the competitive aspect of social life dubbed it the “Machiavellian Intelligence Hypothesis.” (Byrne & Whiten 1988: who themselves, interestingly, didn’t want to stress the competitive aspect by giving the hypothesis the title). Now it is most widely called the “Social Brain Hypothesis” to emphasize the general complexity of primate groups including all arising affordances (Dunbar 1998, Dunbar & Shultz 2007).
Unfortunately, this is still rather vague. To get a clearer picture, it is important to make explicit the advantages and disadvantages of large social groups and the specific problems which need to be solved. However, group size indeed seem to contribute advantageously to genetic fitness by minimizing predation risk, but paired with greater ecological and reproductive competition and suppression, affording higher behavioral flexibility (Dunbar & Shultz 2007). Brain expansion theories stressing the importance of ’technological intelligence’ as a driving force. (without neglecting the importance of social factors, but seeing the latter as less crucial). According to these views, the ‘behavioral drive’ for cultural transmission and innovation is more frequent in species with large brains. As a consequence these species are led to exploit the environment in new way, opening up new possibilities regarding new selective pressures. ´(Reader and Laland 2002). Certainly, these tendencies were important, but where do they come from? Big-brains seem to be a prerequisite for ‘technological intelligence’, but how did these evolve in the first place? Rather it seems probable that
“Although innovation, tool use, and technological invention may have played a crucial role in the evolution of ape and human brains, these skills were probably built upon mental computations that had their origins and foundations in social interactions.” (Cheney & Seyfarth 2007: 283).
Supporting Reader and Laland’s emphasis on the importance of technological aspects on human cognitive evolution, Tomasello and his colleagues propose that human’s advanced Theory of Mind-skills were amplified not in the context of intention-reading present in great apes, but rather during learning and imitation of hierarchical planned and structured tool-making and tool-using. (Tomasello et al. 2005: 687). I’m not sure whether George would like this speculation. Probably, he would argue this to be a ‘just-so story’ and propose that all scientist coming up with these should be eaten. So thank God scientists are not really zombies, I wouldn’t miss the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany (and especially its co-director) for anything in the world (OK, except for the really important things such as love, life, family, donuts.

I haven’t addressed much of the questions stated in the beginning, especially What we as humans are especially better at than other primates. I will come to this issue in my next post (relying again on research done by scientists from the Max Planck Institute for Evolutionary Anthropology, so again, glad they haven’t been eaten.)


References:

Cheney, Dorothy L. and Robert M. Seyfarth. 2007. Baboon Metaphysics: The Evolution of a Social Mind. Chicago: University of Chicago Press.

Deacon, Terrence William 1998. The Symbolic Species. The Co-evolution ofLanguage and the Brain. New York / London: W.W. Norton

Dunbar, Robin I.M.1998.“The Social Brain Hypothesis” Evolutionary Anthropology 6: 178-190.

Dunbar, R. I. M. and Susanne Shultz. 2007.“Evolution in the Social Brain” Science 317: 1344-1347

Park, Min S., Andrew D. Nguyen, Henry E. Aryan, Hoi Sang U, Michael L. Levy, Katerina Semendeferi. 2007. “Evolution of the Human Brain: Changing Brain Size and the Fossil Record.” Neurosurgery 60:555–562.

Reader, S.M. and K.N. Laland. 2002. “Social Intelligence, innovation, and enhanced brain size in primates” PNAS 99: 4436-4441

Tomasello, Michael, Malinda Carpenter, Josep Call, Tanya Behne, and Henrike Moll. 2004. “Understanding and Sharing Intentions: The Origins of Cultural Cognition.” Behavioral and Brain Sciences 28

Monday, November 26, 2007

Zombies have Taste

In my last post I wrote about the fact that human brains are selfish energy-hungry little bastards, which makes the stuff they’re made of extremely ‘expensive tissue’ (Aiello & Wheeler 1995). This means that zombies have a quite extraordinary taste, equivalent to a caviar-gourmet (either that, or they are ‘informavores’ just like we are (Miller 1991)).
Now imagine (instead of the oft-cited martian scientist) a zombie-evolutionary biologist (Insert joke about the parasitic tendencies of the ‘mindless new atheism’ and/or Intelligent Design, the Idea of theistic evolution, greedy reductionism, Evolutionary Psychology or whatever floats you boat here) puzzling over the evolutionary emergence of his most favorite meal. Let’s take it for granted that our zombie-scientist is not easily satisfied by zombie-centric evolutionary concepts, just as Steven Pinker warns us that, if Elephants were the most culturally advanced species (well, and maybe they are, who knows), their evolutionary biologist (albeit only the bad ones) would probably search for the evolutionary path that inevitably climaxed in the highest form, the evolutionary optimum of trunkitude. Let’s also assume our zombie-scientist isn’t a friend of ‘just-so’ stories like ‘humans evolved bigger brains to run away from zombies more effectively’. Assuming, too, that human scientist like Aiello, Wheeler, Dunbar and others weren’t eaten before publishing their caveats about the expensiveness of brain evolution and maintenance, or that some other zombie-scientists could hold back their hunger long enough to test human subjects before eating them, coming to similar conclusions as Aiello and others did – or, rather would have come if they hadn’t been eaten beforehand. Assuming this, we could be sure that our zombie-scientist would not regard the evolution of a ‘general being-eaten-avoidance intelligence’ as unlikely.

What then, our zombie-scientist, call him George, would ask, was the reason humans developed such large, specialized brains. Looking for homologues or convergent evolution in other (hopefully not entirely eat… I mean extinct) species, and considering what makes the human mind special. George could come up with a lot of possible hypotheses as driving forces and triggers of brain evolution, and other facts he would have a hard time to make sense of such as:
  • positively selected genes involved in regulating (Microcephalin: Evans et al. 2005) and determining brain size (ASPM: Mekel-Brobov et al. 2005), development of the human neocortex (HAR1F: Pollard et al. 2006), and playing a part in progressive changes in cognitive abilities (Neuropsin: Li et al. 2004)

  • cooking, paired with gastrointestinal shrinkage (our intestinal tract is only 60% the size expected of a primate with similar size) may have saved energy from digestion which in turn could be used to help fuel the brain. Together with the possible role of meat and more efficient upright walking and running, this could have expanded the human energy budget significantly (Gibbons 2007)

  • supporting this hypothesis, AMY1, a gene improving the digestion of food containing starch, is found in much greater numbers in humans than in chimpanzees (Perry et al. 2007)

  • Correlations between group size and neocortex size (Dunbar 1993, Dunbar & Shultz 2007) on the one hand, and significant positive correlation between innovation, social learning, tool use and brain size on the other, (Reader and Laland 2002, Reader 2003), making it likely that social and technological innovative intelligence (mediated by social learning) both played a crucial and inseparable role in human brain evolution (Cheney & Seyfarth 2007)

  • The possibility that the ability to evolve fat babies was the precursor for the evolution of the big and metabolically expensive brain (The article proposing this hypothesis is called ‘survival of the fattest’, What a great pun! Er… or maybe not) . In a resting newborn baby, the brain consumes 74% of the baby’s energy intake. In a 4-6 months old baby the rate is 64%, further dropping during ontogenetic development until reaching a rate of about 23% in adults. And whereas in chimpanzee infants, there is virtually no body fat, in human infants body fat contributes about 11-14% of the baby’s weight (as does the baby’s brain) (Cunnane & Crawford 2003)
So one thing is clear: a stable high-energy food supply was essentially necessary for human brain development, as were the possibility for longer ontogenetic development (as often observed, human (and generally primate) newborns are pretty much helpless compared to newborns of other species, with some even able to walk following almost immediately after birth).
Another important aspect is the general tendency in mammals to develop bigger brains compared to other species (they are about 10 times ‘brainier’ than amphibians or reptiles). Then, humans are part of the order of primates, which (along with toothed whales) have bigger brains than other mammals. And among primates, monkeys and apes have the biggest brains. But, as I said, our brains are even three times bigger than that expected of an ape of similar size. Another factor is the fact that the pre-natal rapid brain growth observed in other species whose infants are relatively helpless continues post-natally in human babies for about twelve months instead of changing into a slower pace.
As a consequence, human infants are even more helpless than that of other primates. This requires a much greater devotion of time, energy and other resources from the parent’s side. (Lewin 2005: 217f., John L. Locke and Barry Bogin (2005) make a similar argument concerning the unique human life history and ontogenetic development, but extending it not only to brain growth in general, but also to the evolution of language).
Making such a list, George would probably have a lot of trouble to distinguish preconditions, epiphenomena, co-evolutionary processes and driving forces of brain expansion. In my next post I will try to shed some light on this issue (Of course I will fail even more grotesquely than someone who is not a complete layman, but I hope that I will at least clarify some points)

References:


References:

Aiello L.C. and P. Wheeler 1995. ”The expensive tissue hypothesis: the brain and the digestive system in human and primate evolution.” Current Anthropology 36:199–221

Cheney, Dorothy L. and Robert M. Seyfarth. 2007. Baboon Metaphysics: The Evolution of a Social Mind. Chicago: University of Chicago Press.

Cunnane Stephen C. and Michael A. Crawford. 2003. “Survival of the fattest: fat babies were the key to evolution.” Comparative Biochemistry and Physiology Part A: 136.1: 17-26

Dunbar, R.I.M. 1993. “Co-evolution of Neocortex size, group size and language in humans.” Behavioral and Brain Sciences 16.4: 681-735

Dunbar, R. I. M. and Susanne Shultz. 2007.“Evolution in the Social Brain” Science 317: 1344-1347

Evans, Patrick D., Sandra L. Gilbert, Nitzan Mekel-Bobrov, Eric J. Vallender, Jeffrey R. Anderson, Leila M. Vaez-Azizi, Sarah A. Tishkoff, Richard R. Hudson, Bruce T. Lahn “Microcephalin, a Gene Regulating Brain Size, Continues to Evolve Adaptively in Humans” Science 309: 1717-1720.

Gibbons, Ann. 2007. “Food for Thought.” Science 316. 1558-1560.

Lewin, Roger. 2005. Human Evolution: An Illustrated Introduction. Fifth Edition. Suffolk: Blackwell.

Locke, John L. and Barry Bogin. 2005. “Language and life history: A new perspective on the development and evolution of human language” Behavioral and Brain Sciences

Mekel-Bobrov, Nitzan, Sandra L. Gilbert, Patrick D. Evans, Eric J. Vallender, Jeffrey R. Anderson, Richard R. Hudson, Sarah A. Tishkoff, Bruce T. Lahn. “Ongoing Adaptive Evolution of ASPM, a Brain Size Determinant in Homo sapiens.” Science 309: 1720-1722

Li, Yi, Ya-ping Qian, Xiao-jing Yu,* Yin-qiu Wang, Ding-gui Dong’ Wei Sun, Run-mei Ma and Bing Su. 2004. “Recent Origin of a Hominoid-Specific Splice Form of Neuropsin, a Gene Involved in Learning and Memory.“ Molecular Biology and Evolution 21.11: 2111-2115.

Miller, G.A. 1991. The Science of Words. New York: W.H. Freeman

Reader, S.M. 2003. “Relative brain size and the distribution of innovation and social learning across the nonhuman primates.” The Biology of Traditions: Models and Evidence. Eds. D.M. Fragaszy and S. Perry, 56-93.

Reader, S.M. and K.N. Laland. 2002. “Social Intelligence, innovation, and enhanced brain size in primates” PNAS 99: 4436-4441.

Pollard, Katherine S., Sofie R. Salama, Nelle Lambert, Marie-Alexandra Lambot4, Sandra Coppens, Jakob S. Pedersen, Sol Katzman, Bryan King, Courtney Onodera, Adam Siepel, Andrew D. Kern, Colette Dehay, Haller Igel, Manuel Ares Jr, Pierre Vanderhaeghen & David Haussler. 2006 “An RNA gene expressed during cortical development evolved rapidly in humans.” Nature 443: 167-172.

Perry, George H, Nathaniel J Dominy, Katrina G Claw, Arthur S Lee, Heike Fiegler, Richard Redon, John Werner, Fernando A Villanea, Joanna L Mountain, Rajeev Misra, Nigel P Carter, Charles Lee, & Anne C Stone. 2007 “Diet and the evolution of human amylase gene copy number variation“ Nature Genetics Advanced Online Publication doi :10.1038/ng2123