Language Evolution and Levels of Explanation

A somewhat contentious debate among the behavioural sciences is currently underway concerning Mayr’s division of causal explanations in evolutionary theory. Here I’m going to give you a brief rundown of two papers in particular, before I chip in my two-cents about how other insights from the theoretical literature can inform this debate. It seems the discussion is just getting started with respect to cultural evolution, so it’d be interesting to hear other peoples’ comments from either camp.

Over the years, evolutionary theorists have tried to make logical divisions between the kinds of things we can ask about, with a view to making it clear what exactly scientific studies can tell us. A dominant paradigm dividing two levels of causation for biological features we see in the world is Mayr’s distinction between ultimate and proximate causes.  Ultimate causation explains the proliferation of a trait in a population in terms of the evolutionary forces acting on that trait. For example, peahens that prefer peacocks with larger tails (an honest signal of fitness following the handicap principle) will have stronger or more successful offspring, and so this preference proliferates along with larger peacock tails. Proximate causation uses immediate physiological and environmental factors to explain a particular peahen’s penchant for a large-tailed peacock in a mate choice trial, where the signal of the peacock’s large tail elevates the hormone levels in the peahen and copulatory behaviour ensues. Although the behaviour in both of these examples is the same, the levels of explanation are based on different sets of factors.

In Perspectives on Psychological Science last year, a paper by Scott-Phillips, Dickins and West voiced some concerns about these two levels of causation being conflated in the behavioural sciences. In particular, they addressed instances where proximate explanations of traits are being framed as ultimate ones. The paper points specifically to studies of the evolution of cooperation, transmitted culture and epigenetics to illustrate this. Regarding the evolution of cooperation, they point to an instance where ‘strong reciprocity’ (an individual’s propensity to reward cooperative norms and sanction violation of these norms) is purported to be an ultimate explanation of why humans cooperate, rather than a proximate mechanism that enables such cooperation.

Table of proximate and ultimate explanations. For the topic of 'linguistic structure', cultural transmission is listed as a proximate explanation, and the adaptive benefit of cooperative activity through communication is given as an ultimate explanation.,
Table 1 from Scott-Phillips et al. (2011), highlight added.

Among the examples was the feature of linguistic structure (see table 1 from paper above), where several studies pointed to the cultural transmission process as an ultimate explanation of linguistic structure. They suggest that cultural transmission constitutes a proximate process, because it gives the means by which linguistic structure is expressed – and this is how cultural transmission contributes to what the linguistic structure looks like. One analogy might be that the vibrating of my particular vocal cords is a proximate mechanism giving rise, in part, to how my voice sounds, rather than an ultimate explanation of why I vocalise. Since an ultimate account must suggest how a trait contributes to inclusive fitness in order to explain its prevalence in humans, they uncontroversially venture that the ultimate rationale for the ubiquity of linguistic structure is that it greater enables communication (and therefore increases inclusive fitness by enabling cooperative activity).

An opposing view was later published in Science by Laland, Sterelny, Odling-Smee et al., who suggest that the use of Mayr’s division of ultimate and proximate causation is not helpful to all evolutionary investigations, and even hampers progress. The grounds for rejecting Mayr’s paradigm seem to lie largely in what Laland et al. term “reciprocal causation”. That is, that “proximate mechanisms both shape and respond to selection, allowing developmental processes to feature in proximate and ultimate explanations”. After aligning proximate explanations with ontogeny and ultimate explanations with phylogeny, they suggest that what we may have called ultimate and proximate features are no longer sharply delineated, and that these reciprocal processes mean that the source of selection sometimes cannot be separated. They present an idea from the field of evolutionary-developmental biology that, if a developmental process makes some variant of a trait more likely to arise than others, then this proximate mechanism helps to construct an “evolutionary pathway”.

Flow charts presenting alternative processes for Biological evolution, Cultural evolution and Gene-culture coevolution
Figure 2 from Laland et al. (2011)

The paper also highlights developmental plasticity, and gene-environment interaction more broadly (see fig. 2 from paper, above), as a process where reciprocal causation offers an evolutionary explanation conceptually comparable to ultimate causation. Talking specifically on the topic of linguistic structure, they present the debate about whether specific design features of language are attributable to biological or cultural evolution. The paper points out that cultural evolution determines features of linguistic structure – for example, word order – and that the existing word order determines that of future speakers. Indeed, at the Edinburgh LEC we know that transmission by iterated inductive inference under general conditions can explain particular structures in languages. That cultural evolution determines the variation between languages, Laland et al. say, provides evidence that it is an evolutionary force comparable to natural selection (and, therefore, ultimate explanation).

What follows is a collection of my thoughts on the matter, which are (spoiler alert) largely in support of the Scott-Phillips et al. paper. I hope others more experienced in cultural evolution studies than I will contribute their perspective.

It seems to me that there are a few assumptions made in the Laland et al. paper that are not quite in line with how Mayr himself understood the paradigm, and perhaps much can be learned from this debate’s previous incarnation when Richard C. Francis made similar arguments against the ultimate/proximate distinction in 1990. In his critique, he equated ultimate causation with phylogeny and proximate causation with ontogeny – an approach that was rebuked by Mayr in 1993, who made the point that “all physiological activities are proximately caused, but is a reflex an ontogenetic phenomenon?” Mayr’s response is actually rather unhelpful in addressing the arguments fully, and this statement is particularly dense. But what he is getting at here is the idea that interaction with the environment that gives rise to adaptive behaviours (such as recoiling instantly from a hot stove) is itself subject to selection, and thus constitutes a proximate explanation of causation. Relatedly, he points out that most components of the phenotype are indeed the result of genetic contribution and interaction with the environment, which has been successfully explored in biology within the traditional theoretical paradigm.

A perhaps more nuanced account of how we can divide the possible explanations of biological phenomena is offered by Tinbergen in his “four questions”, where ultimate explanations are further subdivided into Function (concerning the adaptive solution to a survival problem favoured by natural selection) and Phylogeny, which is a historical account of when the trait arose in the species, and importantly includes processes other than natural selection that give rise to variation – such as mutation, drift and the constraints imposed by pre-existing traits (see blind spot example below). Proximate explanations are further split into Mechanism (immediate physiological/environmental factors causal in how the trait operates in the individual) and Ontogeny (the way in which this trait develops over the lifetime of the individual). As a simple example, here is the paradigm applied to a trait like mammalian vision that I lifted from Wikipedia:
Function: To find food and avoid danger.
Phylogeny: The vertebrate eye initially developed with a blind spot, but the lack of adaptive intermediate forms prevented the loss of the blind spot.
Causation: The lens of the eye focuses light on the retina
Ontogeny: Neurons need the stimulation of light to wire the eye to the brain within a critical period (as those awful studies of blindfolded kittens illustrated).
A schematic below, adapted from Tinbergen (1963) shows how these levels of causation may interact with one another, which appears to communicate something roughly comparable to the importance Laland et al. place on “reciprocal causation” in the formation of adaptive variants:

Flow chart showing the interaction of functional, phylogenetic, mechanistic and ontogenetic explanations
Adapted from Tinbergen (1963); Causal Relationships

Applied the to debate outlined above, it would seem that there is no apparent reason that a process of gene-environment interaction – including the cultural environment – can’t itself be subject to selection, or that developmental plasticity itself is not an adaptation in need of an ultimate explanation. It has long been the case that behaviour is no longer understood as either “nature” or “nurture”, but gene-environment interaction, with varying levels of heredity. The “reciprocal causation” suggested in Laland et al.’s paper, is (as they point out) very common in nature; feedback loops are uncontroversial proximate processes in biology. That a proximate process may give rise to a dominant variant of a trait in a population does not explain why it is adaptive, and this points to another problem with the proposing the abandonment of Mayr’s paradigm: a logical division of levels of explanation doesn’t seem to be the sort of thing that can be rendered outdated by empirical evidence. Indeed, claims about the particulars of traits and processes (and languages) themselves are a matter for empirical data – but the theoretical issue about the level of explanation that data is useful for does not itself seem to be subject to empirical findings.

The finding that a proximate process such as cultural transmission gives rise to a trait that is prolific in a population is itself exciting and surprising, and even shows us that the pressure for making language easier to learn gives us adaptive languages to learn; however, it could be argued that it is this process that is adaptive, and that the reason why humans so heavily rely on this process is an ultimate explanation.

One way of resolving these two perspectives may be to place cultural processes that give rise to variation at the level of what Tinbergen labels Phylogenetic (one subset of ultimate) explanation, as it concerns processes which produce some heightened frequency of traits over a language’s history. An explanation at the level of Phylogeny still must make recourse to natural selection at some point, since variants that result from mutation or drift are retained because of their adaptive value (or an adaptive trade-off). This approach may be a problem for the current understanding, which holds that the features resulting from cultural processes are themselves adaptive and therefore comparable to what Tinbergen labels Function.

The problem with this is that calling particular structures of language ‘adaptive’ obscures what it is about Language that is actually being selected for. To flesh out what I mean, I think it’s useful to consult Millikan’s (1993) distinction between Direct Proper Function and Derived Proper Function (… bear with me, it’ll be worth it, honest). The Direct Proper Function of a given trait T can be thought of as a “reproduction” of an item that has performed the exact same adaptive function F, and T exists because of these historical performances of F. Sperber and Origgi (2000) use the illustrative example of the heart, where the human heart has a bunch of properties (it pumps blood, makes a thumping noise, etc), but only its ability to pump blood is its Direct Proper Function. This is because even a heart that doesn’t work right or makes irregular thumping noises or whatever, still has the ability to pump blood. Hearts that pump blood have been “reproduced through organisms that, thanks in part to their owning a heart pumping blood, have had descendents similarly endowed with blood-pumping hearts”.

The Derived Proper Function, however, refers to a trait T that is the result of some device that, in some environment, has a Proper Function F. In that given environment, F is usually achieved by the production of something like T. If I unpack this idea and apply it to language, we can understand it as the acquisiton and production of a device that, in this environment, leads to, say, a particular SVO language, T. The Proper Function of adaptive communication is performed by T in this case, but could also be performed by any number of SOV, VSO, etc Ts in other cases. In other words, the Proper Function of this language is not the word order itself, but communication. The word order is the realisation of this device that is reproduced because of the performance of T in a particular environment, but does not necessarily lead to T in the next incarnation of that device (i.e. My child, if born and raised in Japan, will speak Japanese). We see, then, that a proximate process resulting in what a particular language spoken by a given population looks like does not necessarily speak to the evolutionary function. In other words, it is the device that allows the performance of Language that is adaptive, not the individual language itself.

One question being asked in the study of cultural transmission is why a particular language looks like it does, while we also know that there are 6000 different versions that perform the same (ultimate) function. I would even argue that asking how proximate processes shape languages is actually the most exciting and interesting avenue of inquiry precisely because it’s so blindingly obvious what the adaptive function of language is. But perhaps the value in this endeavour is somewhat neglected, in part, because of the same impression that Francis (1990) had: “the attitude, implicit in the term ultimate cause, [is] that these functional analyses are somehow superordinate to those involving proximate causes” which would be a shame. It seems to me that the coarse grain of ultimate vs proximate perhaps doesn’t do enough to help complex proximate study to position itself in the wider theoretical framework, and the best way to proceed from this might be to couch explanation in terms of Function, Phylogeny, Ontogeny and Mechanism. I think more fine-grained terminology grants us more explanatory power, in this case.

A final question in this debate that came up too many times during discussions with the LEC is: what does keeping the traditional paradigm “buy us”? Well, the first answer to this is consilience with one of the most successful and robust theories in science. The same sentiment has been communicated by Pinker and Bloom (1990), who said: “If current theory of language is truly incompatible with the neo-Darwinian theory of evolution, one could hardly blame someone for concluding that it is not the theory of evolution that must be questioned, but the theory of language”. Part of the reason this debate may have arisen is that studies of cultural evolution have used evolutionary theory as an incredibly fruitful way of analysing cultural processes, but additional acknowledgement about how cultural adaptation is different to biological adaptation may be necessary. This difference is an aspect of Laland’s paper (shown in Fig 2) that I think is important, as it’s part of the reason that more nuanced frameworks for cultural evolution are now needed. Without this widespread acknowledgement, cultural evolution may be considered an extension of biological evolutionary theory instead of a successfully applied metaphor. It seems to me that the side of this debate one falls on is well predicted by whether one subscribes to the former interpretation of cultural evolution or the latter.

Knowing which level of explanation current work pertains to is a valuable part of evolutionary exploration, and abandoning this in favour of an approach where proximate processes are explanatory ends to themselves may mean the exploration of Function and Phylogeny may suffer. That said, it is telling, I think, that even in seeking to abandon the proximate/ultimate distinction, we must still exploit this existing terminology in order to explain such a position. That natural selection has explained countless adaptations in all living things is certainly not trivial, and to reject the theory giving rise to ultimate explanations as they’re currently defined is to reject this fundamental aspect of evolutionary theory. The big problem seems to be that we’re coming to understand proximate processes as so elaborate and complex, that a more nuanced framework is needed to deal with the dynamics of those processes. I reckon, however, that such a framework can be developed within the traditional paradigms of evolutionary theory.



Francis, R.C. (1990) – “Causes, Proximate and Ultimate” Biology and Philosophy 5(4) 401-415.

Laland, K., Sterelny, K., Odling-Smee, J., Hoppitt, W. & Uller, T. (2011) – “Cause and Effect in Biology Revisited: Is Mayr’s Proximate-Ultimate Distinction Still Useful?” Science 334, 1512-1516.

Mayr, E. (1993) – “Proximate and Ultimate Causations” Biology and Philosophy 8: 93-94.

Millikan, R. (1993) – White Queen Psychology and Other Essays for Alice, Cambridge, Mass: MIT Press.

Pinker, S. & Bloom, P. (1990) – “Natural language and natural selection” Behaviour and Brain Sciences 13, 707-784.

Scott-Phillips, T. Dickins, T. & West, S. (2011) – “Evolutionary Theory and the Ultimate-Proximate Distinction in the Human Behavioural Sciences” Perspectives on Psychological Science 6(1): 38-47.

Sperber, D. & Origgi, G. (2000) – “Evolution, communication and the proper function of language” In P. Carruthers and A. Chamberlain (Eds.) Evolution and the Human Mind: Language, Modularity and Social Cognition (pp.140-169) Cambridge: Cambridge University Press.

Tinbergen, N. (1963) “On Aims and Methods in Ethology,” Zeitschrift für Tierpsychologie, 20: 410–433.



Robustness, Evolvability, Degeneracy and stuff like that…

Much of the work I plan to do for this year involves integrating traditional and contemporary theories of language change within an evolutionary framework. In my previous post I introduced the concept of degeneracy, which, to briefly recap, refers to components that have a structure-to-function ratio of many-to-one, with a single degenerate structure being capable of performing distinct functions under different conditions (pluripotent). Whitcare (2010: 5) provides a case in point for biological systems: here, the adhesin gene family in A. Saccharomyces “ expresses proteins that typically play unique roles during development, yet can perform each other’s functions when expression levels are altered”.

But what about degeneracy in language? For a start, we already know from basic linguistic theory forms (i.e. structures) are paired with meanings (i.e. functions). More recent work has expanded upon this notion, especially in developing the concept of constructions (Goldberg, 2003): “direct form-meaning pairings that range from the very specific (words or idioms) to the more general (passive constructions, ditranstive construction), and from very small units (words with affixes, walked) to clause-level or even discourse-level units” (Beckner et al., 2009: 5). When applied to constructions, degeneracy fits squarely with work identifying language as a Complex Adaptive System (see here) and as a culturally transmitted replicator (see here and here), which offers a link between the generation of first order synchronic variation – in the form of innovation (e.g. newly introduced linguistic material in the form of sounds, words, grammatical constructions etc) – and the selection, propagation and fixation of linguistic variants within a speaker community.

For the following example, I’m going to look at a specific type of discourse-pragmatic feature, or construction, which has undergone renewed interest over the last thirty-years. Known as General Extenders (GEs) – utterance- or clause-final discourse particles, such as and stuff and or something – researchers are realising that, far from being superfluous linguistic baggage, these features “carry social meaning, perform indispensible functions in social interaction, and constitute essential elements of sentence grammar” (Pichler, 2010: 582). Of specific relevance, GEs, and discourse-pragmatic particles more generally, are multifunctional: that is, they are not confined to a single communicative domain, and can even come to serve multiple roles within the same communicative context or utterance.

It is proposed the concept of degeneracy will allow us to explain how multifunctional discourse markers emerge from variation existent at structural components of linguistic organisation, such as the phonological and morphosyntactic components. If anything, I hope the post might serve as some food for thought, as I’m still grappling with the applications of the theory (and whether there’s anything useful to say!).

Continue reading “Robustness, Evolvability, Degeneracy and stuff like that…”

Linguistic diversity and traffic accidents

This post was chosen as an Editor's Selection for ResearchBlogging.orgI was thinking about Daniel Nettle’s model of linguistic diversity which showed that linguistic variation tends to decline even with a small amount of migration between communities.  I wondered if statistics about population movement would correlate with linguistic diversity, as measured by the Greenberg Diversity Index (GDI) for a country (see below).  However, this is a cautionary tale about obsession and use of statistics.  (See bottom of post for  link to data).

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Cognitivism and the Critic 2: Symbol Processing

It has long been obvious to me that the so-called cognitive revolution is what happened when computation – both the idea and the digital technology – hit the human sciences. But I’ve seen little reflection of that in the literary cognitivism of the last decade and a half. And that, I fear, is a mistake.

Thus, when I set out to write a long programmatic essay, Literary Morphology: Nine Propositions in a Naturalist Theory of Form, I argued that we think of literary text as a computational form. I submitted the essay and found that both reviewers were puzzled about what I meant by computation. While publication was not conditioned on providing such satisfaction, I did make some efforts to satisfy them, though I’d be surprised if they were completely satisfied by those efforts.

That was a few years ago.

Ever since then I pondered the issue: how do I talk about computation to a literary audience? You see, some of my graduate training was in computational linguistics, so I find it natural to think about language processing as entailing computation. As literature is constituted by language it too must involve computation. But without some background in computational linguistics or artificial intelligence, I’m not sure the notion is much more than a buzzword that’s been trendy for the last few decades – and that’s an awful long time for being trendy.

I’ve already written one post specifically on this issue: Cognitivism for the Critic, in Four & a Parable, where I write abstracts of four texts which, taken together, give a good feel for the computational side of cognitive science. Here’s another crack at it, from a different angle: symbol processing.

Operations on Symbols

I take it that ordinary arithmetic is most people’s ‘default’ case for what computation is. Not only have we all learned it, it’s fundamental to our knowledge, like reading and writing. Whatever we know, think, or intuit about computation is built on our practical knowledge of arithmetic.

As far as I can tell, we think of arithmetic as being about numbers. Numbers are different from words. And they’re different from literary texts. And not merely different. Some of us – many of whom study literature professionally – have learned that numbers and literature are deeply and utterly different to the point of being fundamentally in opposition to one another. From that point of view the notion that literary texts be understood computationally is little short of blasphemy.

Not so. Not quite.

The question of just what numbers are – metaphysically, ontologically – is well beyond the scope of this post. But what they are in arithmetic, that’s simple; they’re symbols. Words too are symbols; and literary texts are constituted of words. In this sense, perhaps superficial, but nonetheless real, the reading of literary texts and making arithmetic calculations are the same thing, operations on symbols. Continue reading “Cognitivism and the Critic 2: Symbol Processing”

Laryngeal Air Sacs

So, I got a request from a friend of mine to make an abstract on the fly for a poster for Friday. I stayed up until 3am and banged this out. Tonight, I hope to write the poster justifying it into being. A lot of the work here builds on Bart de Boer’s work, with which I am pretty familiar, but much of it also started with a wonderful series of posts over on Tetrapod Zoology. Rather than describe air sacs here, I’m just going to link to that – I highly suggest the series!

Here’s the abstract I wrote up, once you’ve read that article on air sacs in primates. Any feedback would be greatly appreciated – I’ll try to make a follow-up post with the information that I gather tonight and tomorrow morning on the poster, as well.

Re-dating the loss of laryngeal air sacs in hominins

Laryngeal air sacs are a product of convergent evolution in many different species of primates, cervids, bats, and other mammals. In the case of Homo sapiens, their presence has been lost. This has been argued to have happened before Homo heidelbergensis, due to a loss of the bulla in the hyoid bone from Austrolopithecus afarensis (Martinez, 2008), at a range of 500kya to 3.3mya. (de Boer, to appear). Justifications for the loss of laryngeal air sacs include infection, the ability to modify breathing patterns and reduce need for an anti-hyperventilating device (Hewitt et al, 2002), and the selection against air sacs as they are disadvantageous for subtle, timed, and distinct sounds (de Boer, to appear). Further, it has been suggested that the loss goes against the significant correlation of air sac retention to evolutionary growth in body mass (Hewitt et al., 2002).

I argue that the loss of air sacs may have occurred more recently (less than 600kya), as the loss of the bulla in the hyoid does not exclude the possibility of airs sacs, as in cervids, where laryngeal air sacs can herniate between two muscles (Frey et al., 2007).  Further, the weight measurements of living species as a justification for the loss of air sacs despite a gain in body mass I argue to be unfounded given archaeological evidence, which suggests that the laryngeal air sacs may have been lost only after size reduction in Homo sapiens from Homo heidelbergensis.

Finally, I suggest two further justifications for loss of the laryngeal air sacs in homo sapiens. First, the linguistic niche of hunting in the environment in which early hominin hunters have been posited to exist – the savannah – would have been better suited to higher frequency, directional calls as opposed to lower frequency, multidirectional calls. The loss of air sacs would have then been directly advantageous, as lower frequencies produced by air sac vocalisations over bare ground have been shown to favour multidirectional over targeted utterances (Frey and Gebler, 2003). Secondly, the reuse of air stored in air sacs could have possibly been disadvantageous toward sustained, regular heavy breathing, as would occur in a similar hunting environment.


Boer, B. de. (to appear). Air sacs and vocal fold vibration: Implications for evolution of speech.

Fitch, T. (2006). Production of Vocalizations in Mammals. Encyclopedia of Language and Linguistics. Elsevier.

Frey, R, & Gebler, A. (2003). The highly specialized vocal tract of the male Mongolian gazelle (Procapra gutturosa Pallas, 1777–Mammalia, Bovidae). Journal of anatomy, 203(5), 451-71. Retrieved June 1, 2011, from

Frey, Roland, Gebler, Alban, Fritsch, G., Nygrén, K., & Weissengruber, G. E. (2007). Nordic rattle: the hoarse vocalization and the inflatable laryngeal air sac of reindeer (Rangifer tarandus). Journal of Anatomy, 210(2), 131-159. doi: 10.1111/j.1469-7580.2006.00684.x.

Martínez, I., Arsuaga, J. L., Quam, R., Carretero, J. M., Gracia, a, & Rodríguez, L. (2008). Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain). Journal of human evolution, 54(1), 118-24. doi: 10.1016/j.jhevol.2007.07.006.

Hewitt, G., MacLarnon, A., & Jones, K. E. (2002). The functions of laryngeal air sacs in primates: a new hypothesis. Folia primatologica international journal of primatology, 73(2-3), 70-94. Retrieved from

Sound good? I hope so! That’s all for now.

The Return of the Phoneme Inventories

Right, I already referred to Atkinson’s paper in a previous post, and much of the work he’s presented is essentially part of a potential PhD project I’m hoping to do. Much of this stems back to last summer, where I mentioned how the phoneme inventory size correlates with certain demographic features, such as population size and population density. Using the the UPSID data I generated a generalised additive model to demonstrate how area and population size interact in determining the phoneme inventory size:

Interestingly, Atkinson seems to derive much of his thinking, at least in his choice of demographic variables, from work into the transmission of cultural artefacts (see here and here). For me, there are clear uses for these demographic models in testing hypotheses for linguistic transmission and change, as I see language as a cultural product. It appears Atkinson reached the same conclusion. Where we depart, however, is in our overall explanations of the data. My major problem with the claim is theoretical: he hasn’t ruled out other historical-evolutionary explanations for these patterns.

Before we get into the bulk of my criticism, I’ll provide a very brief overview of the paper.

Continue reading “The Return of the Phoneme Inventories”

The Parental Antagonism Theory of Language Evolution

Human Biology are publishing a special issue on “Integrating genetic and Cultural Evolutionary Approaches to Language” this month! Abstracts for all of the papers can be found here.

William Brown‘s paper has been published on his blog ahead of the boat today. The Abstract is below and there is a link to the paper at the bottom.

Language—as with most communication systems—likely evolved by means of natural selection. Accounts for the natural selection of language can usually be divided into two scenarios, either of which used in isolation of the other are insufficient to explain the phenomena: (1) there are group benefits from communicating, and (2) there are individual benefits from being a better communicator. In contrast, this paper argues that language emerged during a coevolutionary struggle between parental genomes via genomic imprinting, which is differential gene expression depending on parental origin of the genetic element. It is hypothesized that relatedness asymmetries differentially selected for patrigene-caused language phenotypes (e.g., signals of need) to extract resources from mother early in child development and matrigene-caused language phenotypes (e.g.,  socially transmitted norms) to influence degree of cooperativeness  among kin later in development. Unlike previous theories for language evolution, parental antagonism theory generates testable predictions at the proximate (e.g., neurocognitive areas important for social transmission and language capacities), ontogenetic (e.g., the function of language at different points of development), ultimate (e.g., inclusive fitness), and phylogenetic levels (e.g., the spread of maternally derived brain components in mammals, particularly in the hominin lineage), thus making human capacities for culture more tractable than previously thought.

Brown, W.M. (2011). The parental antagonism theory of language evolution: Preliminary evidence for the proposal. Human Biology, 83 (2)

Animal Signalling Theory 101: Handicap, Index… or even a signal? The Case of Fluctuating Asymmetry

The differences between handicaps and indices are usually distinguishable in formal mathematical models or in unambiguous real-world cases. Often though, classifying a trait as a handicap, an index, or even a signal at all, can be quite a difficult task.

For the purposes of illustration I will use Fluctuating Asymmetry (FA for short) as an example.  Fluctuating asymmetry is the term used to refer to deviation from symmetry in paired morphological structures (ranging from birds’ tails to human faces) that should be, all being well, bilaterally symmetric. Deviations from the ideal symmetrical phenotype are caused by inherent genetic perturbations and exposure to environmental disturbances occurring in early development.

Is FA a signal?

In their 2005 book Animal Signals, Maynard-Smith and Harper define a signal as:

‘Any act or structure which alters the behaviour of other organisms, which evolved because of that effect, and which is effective because the receiver’s response has also evolved’

They then argue that FA is unlikely to function as a signal because it is difficult to discern whether receivers respond directly to FA and because there appear to be few examples of displays in which signallers actively advertise their symmetry to receivers.


Continue reading “Animal Signalling Theory 101: Handicap, Index… or even a signal? The Case of Fluctuating Asymmetry”

Intelligence: Darwin vs. Wallace

It’s Charles Darwin’s birthday today! He’s 202. So in celebration I’ve written a post on the still ongoing controversy which the theory of evolution by natural selection caused and is causing, specifically with regards to the emergence of human intelligence.

Alfred Russel Wallace is widely seen as the co-discoverer of the theory of evolution by natural selection. While Darwin had been formulating his theory from as early as the late 1830s, he kept quite about it for more than twenty years while he amassed evidence to support it. In 1858 Alfred Russell Wallace, a naturalist of the same time, sent Darwin a letter outlining for him a theory of evolution which very closely mirrored Darwin’s own. The pair co-presented their theory to the Linnaean Society in 1858 but due to Darwin’s long time amassing evidence and refining his ideas, it was his book, On The Origin of Species, which was published in 1859 and set Darwin’s name firmly in the history books as the discoverer of natural selection.

While Wallace’s part in the discovery of natural selection is far from undocumented or unknown, it is largely for presenting ‘the same ideas’ as Darwin for which he is known and what is rarely discussed in the differences in their ideas. In this post I will briefly discuss a new(ish) paper by Steven Pinker on the evolution of human intelligence and some the differences between the thinking of Darwin and Wallace on the subject.

Darwin, unsurprisingly, asserted that the abstract nature of human intelligence can be fully explained by natural selection. In opposition to this Wallace claimed that it was of no use to ancestral humans and therefore could only be explained by intelligent design:

“Natural selection could only have endowed savage man with a brain a few degrees superior to that of an ape, whereas he actually possesses one very little inferior to that of a philosopher.”(Wallace, 1870:343)

Unsurprisingly most scientists these days do not agree with Wallace on either the point that the human brain could not be the result of natural selection or that as a result of this problem it must have been a product of design by a higher being. It would be both dismissive and dull to leave the discussion at that however, which is where Pinker comes in. Despite Wallace’s argument probably coming to the wrong conclusion he does bring up some very interesting questions which need answering, namely that of; “why do humans have the ability to pursue abstract intellectual feats such as science, mathematics, philosophy, and law, given that opportunities to exercise these talents did not exist in the foraging lifestyle in which humans evolved and would not have parlayed themselves into advantages in survival and reproduction even if they did?” (Pinker, 2010:8993)

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From Natyural to Nacheruhl: Utterance Selection and Language Change

Most of us should know by now that language changes. It’s why the 14th Century prose of Geoffrey Chaucer is nearly impenetrable to modern day speakers of English. It is also why Benjamin Franklin’s phonetically transcribed pronunciation of the English word natural sounded like natyural (phonetically [nætjuɹəl]) rather than our modern variant with a ch sound (phonetically [nætʃəɹəl]). However, it is often taken for granted on this blog that language change can be understood as an evolutionary process. Many people might not see the utility of such thinking outside the realm of biology. That is, evolutionary theory is strictly the preserve of describing biological change, and is less useful as a generalisable concept. A relatively recent group of papers, however, have taken the conceptual machinery of evolutionary theory (see Hull, 2001) and applied it to language.

It's all natyural, yo!

Broadly speaking, these utterance selection models highlight that language change occurs across two steps, each corresponding to an evolutionary process: (1) the production of an utterance, and (2) the propagation of linguistic variants within a speech community. The first of these, the production of an utterance, takes place across an extremely short timescale: we will replicate particular sounds, words, and constructions millions of times across our production lifetime. It is as this step where variation is generated: phonetic variation, for instance, is not only generated through different speakers having different phonetic values for a single phoneme — the same speaker will produce different phonetic values for a single phoneme based on the context. Through variation comes the possibility of selection within a speech community. This leads us to our second timescale, which sees the selection and propagation of these variants — a process that may “take many generations of the replication of the word, which may–or may not–extend beyond the lifetime of an individual speaker.” (Croft, in press).

Recent mathematical work in this area has highlighted four selection mechanisms: replicator selection, neutral evolution, neutral interactor selection, and weighted interactor selection. I’ll now provide a brief overview of each of these mechanisms in relation to language change.

Continue reading “From Natyural to Nacheruhl: Utterance Selection and Language Change”