From Grooming to Speaking: Recent trends in social primatology and human ethology (Conference Announcement)

Should be of interest to some readers:

The Centre for Philosophy of Science of the Faculty of Science of the Portuguese University of Lisbon is organizing a 3-day international colloquium entitled “From Grooming to Speaking: recent trends in social primatology and human ethology”, on September 10-12th, 2012.

Conference website

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Observational learning in octopus vulgaris

A few months ago, a documentary I saw on the Discovery Channel covered some research by Graziano Fiorito and colleagues at the Stazione Zoologica  in Naples. They were investigating observational learning in wild Octopus vulgaris with a puzzle-box experiment similar to those demonstrating cultural transmission in chimpanzees.

It goes like this: there’s a tasty and terrified crustacean running around in a perspex box that has two possible ways of being opened by hungry octopuses. The experimenters capture a wild octopus (let’s call him Steve) from the harbour (which I’ll get back to in a minute), and they put it in a tank with the puzzle-box. After Steve stares hopelessly at the box for a while, it is then removed from the tank. Steve the kidnapped octopus then gets to watch a captive octopus in the next tank being presented with the same puzzle-box containing the delicious crab. Of course, the captive octopus has been confronted with the puzzle-box enough times that it has worked out a successful solution, and so opens the box like a pro. Steve is then presented with a crab in a puzzle-box again, except this time he goes straight for the crab using the same solution he just learned from the captive octopus. Here is a clip from the documentary, showing a trial of this experiment (NB: not the best quality).

It turns out that the papers on this go back as far as Fiorito & Scotto 1992, and it seems this was the first time observational learning had been demonstrated in invertebrates. The reason I’m interested in reporting this is because the documentary I watched explained another possible motivation/interpretation for Fiorito’s work that I can’t find in any of his actual papers. The octopuses used in the experiments were all caught from the harbour at Naples just before the experiments, which of course controlled for any prior experience with the puzzle boxes. But the results were reported as particularly interesting because the Naples harbour had been overfished and disrupted, resulting in an increase in marine predators that eat the small octopus vulgaris as well as fish that the octopuses themselves rely on. These harsher environmental conditions resulted in the octopuses being forced to inhabit a smaller space alongside each other. As a result, young octopuses were frequently exposed to, and even coexisted with, older octopuses. This is a weird situation for an octopus; they usually live solitary lives and never even meet their own mothers, who die of starvation while caring for the eggs (the fathers die within a few months of having mated). The only real interactions are mating, and conflicts between rival males while competing for a mate.

Toward the end of the documentary, the voiceover growled against some dramatic music about how the combination of observational learning capacities and increased predation pushing octopuses into groups meant that it was only a matter of time  before we’re overthrown by octopus vulgaris. This made me think of Dunbar’s “social brain hypothesis” for the emergence of language, and whether I really should prepare to welcome our new octopus overlords. Talking specifically about primates, Dunbar (1996) states that “[primates] in general exhibit two responses to increased predation: they grow physically bigger [or] they increase the size of their groups” (p.110). In order to maintain these groups, that are essential for survival in harsh ecological conditions, Dunbar suggests that standard primate grooming behaviour becomes too time consuming and costly in order to keep up with the rapidly expanding social group, creating a pressure for a more efficient method of bonding and communicating that allows the size of the group to continue increasing. Again with reference just to primate communication, Dunbar says “This [efficient mechanism] need not have involved any dramatic change, for as the studies by Seyfarth and Cheney have shown, primate vocalizations are already capable of conveying a great deal of social information and commentary.” (p.115) In addition to primates, and adding further credence to this idea, it’s been shown that the older Matriarchs of elephant groups make use of vocalisations to seemingly instruct their group on how to fend off lion attacks (McComb et al., 2011; previous Replicated Typo coverage here). It seems to me that the difference between the primates/elephants and the octopuses is that the former endeavour to actually enrich the environment from which their conspecifics extract information; they don’t just learn, they inform. As far as we can tell, there is no such communication like this – that is, enriching the environment in some way that helps other octopuses learn or survive – happening among the octopuses. That said, we know from mating displays and conflicts that cephalopods can communicate with chromatophore signalling. As an interesting aside that is reflective of their cognitive abilities and capacity for suffering, octopuses are treated as honorary vertebrates by UK animal testing laws.

It’s interesting enough that marine biologists (or at least those reporting on marine biologists) seem to have the same idea as Dunbar about the necessary preconditions for successful societies of animals, but why not let’s get wildly speculative? If (..!) existing in groups is in fact adaptive for these octopuses in the face of increased predation, and the competition between them for resources isn’t too great a counter-factor, it seems the only ingredient missing from an octopocalypse is the emergence of some cooperative behaviour. Someone should keep an eye on that harbour.



Dunbar, R. (1996) Grooming, Gossip, and the Evolution of Language. Harvard University Press: Cambridge, Massachusetts

Fiorito, G. & Scotto, P. (1992) “Observational learning in Octopus vulgaris” Science 256, 545-546.

McComb, K., Shannon, G., Durant, S., Sayialel, K., Slotow, R., Poole, J. & Moss, C. (2011) “Leadership in elephants: the adaptive value of age” Proceedings of the Royal Society B, published online.

Babies know who’s boss, whose boss, and who knows what else.

forthcoming paper (grateful nod to ICCI) in PNAS from Olivier Mascaro and Gergely Csibra presents a series of experiments investigating the representation of social dominance relations in human infants, and it’s excellent news: we’re special.

Social dominance can be inferred in a couple of ways. Causal cues such as age, physical aggression and size can tell us about the dominance status of an individual quite intuitively, so we can make a sensible decision about whether or not we get into a scrap with them. Another way we can establish this is to look for direct realisations of dominance, such as who gets the banana if two hungry chimps both want it; chances are, little Pan Pipsqueak isn’t going to get a look in. In order to be useful, we also have to use this information to expect certain things from the individuals around us, so those representations have some property of stability across time that allows us to have those expectations. The question being explored in this paper is whether the representations we have are about the relationship between the two agents who want the banana, or the individual properties each of them has.

In a series of experiments using preferential looking time as a dependent measure, human infants (9 and 12 month olds) were exposed to videos of geometric figures exhibiting similar goal-directed behaviour. Then they would watch, say, a dominant triangle picking up the last figurative banana when the nondominant pentagon also wanted it. For expository purposes and posterity’s sake, I have constructed an artist’s impression of a dominant triangle and a subordinate pentagon in MSPaint (below, right):

A dominant triangle and subordinate pentagon (artist’s impression).

I’m not just showing off my extraordinary artistic talent here; the good thing about these agents is that there are none of the cues like size or aggression that can give rise to the assignment of individual dominance properties. The task also doesn’t indicate anything similar; it’s just about who gets the desired object when there’s only one left. In other words, the goal-directed actions of two agents are in opposition. After seeing a triangle beat a pentagon to an object of ‘banana’ status, 12 month olds looked for longer when they were then presented with an incongruent trial where the pentagon gained over the triangle. 9 month olds (understandably?) couldn’t care less. So, on the basis of this social interaction alone, the 12 month olds were able to notice when something unexpected happened.

To rule out the possibility that this was just the result of some simple heuristic such as “when triangle and pentagon are present, triangle gets the object” and make sure the infants really were assigning some dominance, another experiment (with 12 and 15 month olds) showed the same test video of the two agents collecting little objects. This time, however, the preceding video was of the triangle dominating a little walled-in space that the pentagon also wanted to inhabit. The 12 month olds had no idea what was up, but the 15 month olds generalised from the first “get out of my room” interaction to the “I get the last banana” interaction. So, 15 month olds can extract, just from watching a social interaction, the dominance status of agents and can generalise that information to novel situations. So if a 15 month old watches you lose your favourite seat in front of the TV, they’ll also expect you to miss out on the last slice of pizza, because you’re a loser.

What we still don’t know is whether they think your belly is inherently yellow, or if you’re just a pushover when interacting with a particular person. Is it the relationship between the triangle and pentagon that the babies are tracking, or do they just give each agent some sort of dominance score? This was addressed in experiment 4, where infants were presented with two interactions: one between A and B, where A wins, and then another between B and C, where B wins. If the babies are assigning an individual value to each agent, they should have some sort of linear, transitive representation of dominance like A > B > C. If they’re then presented with a novel interaction between A and C, they would have the expectation that A will beat C. So if they stare in surprise at a trial where C wins, we know it’s violated that kind of expectation, and that they’re representing this stuff linearly – I.E. each agent has a dominance value. In contrast, if the infant is tracking the relations between agents, they can’t really have an expectation of what will happen when A and C both want a banana, because they’ve never seen C before. The results find that the infants look preferentially when they get an incongruent trial using agent pairs they have seen before – as we’d expect from the previous experiment. When they’re presented with a new “I get the last banana” interaction between A and C, however, there’s nothing startling about it when C wins – which means their expectations are not based on something like A > B > C.

The only tiny little harrumph I have about this result is that all it does is falsify the linear representation account. Though I think their account is absolutely right, it’d be nice to see something more predictive come out of the relation-representation hypothesis that is a little more falsifiable. But this result is pretty huge, and stands in contrast with what we know about social cognition in other animals like baboons (Cheney et al, 1995; Bergman et al, 2003), lemurs (Maclean et al., 2008) and even pigeons (Lazareva & Wasserman, 2012), who seem to employ this sort of hierarchical, transitive inference when presented with novel interactions. It may also muddy the waters a little when we want to make the appealing claim that, since language surely emerged in order to enable communication as we navigated a social environment, hierarchical social cognition gives rise to the processing of languagey things like hierarchical syntax or our semantic representation (Hamilton, 2005), which can be characterised as hierarchical (e.g. hyperonym > hyponym). If we consider the nature of the human social environment, though, it should seem more intuitive that something more reliable than simple transitive inference is necessary in order to successfully navigate through our interactions. Due to our prolific production of (and reliance on) culture, humans have a much more diverse range of social currencies, which correspond to values for things like money, intelligence, blackmail information, who your friends are, ad infinitum. That means it’s pretty reasonable that our social cognition needs new strategies in order to get by; we have a little more to consider than just who’s big and angry enough to get all the bananas.


Bergman, T., Beehner, J., Cheney, D. & Seyfarth, R. (2003) “Hierarchical Classification by Rank and Kinship in Baboons” Science 14(302), 1234-1236.

Cheney, D., Seyfarth, R. & Silk, J. (1995) “The response of female baboons (Papio cynocephalus ursinus) to anomalous social interactions: evidence for causal reasoning?” Journal of Comparative Psychology 109(2), 134-141.

Hamilton, D.L. (2005) Social Cognition: Key Readings (p. 104) Psychology Press

Lazareva, O. & Wasserman, E. (2012) “Transitive inference in pigeons: measuring the associative values of stimulus B and D” Behavioural Process 89(3), 244-255.

Maclean, E., Merritt, D. & Brannon, E.M. (2008) “Social complexity predicts transitive reasoning in prosimian primates” Animal Behaviour 76(2), 479-486.

Mascaro, O. & Csibra, G. (forthcoming) “Representation of stable dominance relations by human infants” Proceedings of the National Academy of Sciences


3 PhD positions on the Evolution of Combinatorial Speech

Just a quick announcement that might very well be of interest to some readers of this blog: Bart de Boer (of the Vrije Universiteit Brussel in Brussels, Belgium) is looking for 3 PhD students for his ERC project ABACUS (Advancing Behavioral and Cognitive Understanding of Speech). The project is about investigating (the evolution of) cognitive adaptations for dealing with combinatorial speech. It uses a combination of iterated learning experiments, individual learning experiments and computational modeling.

Each PhD project focuses on one of the approaches:

  • iterated learning
  • individual learning
  • computational modeling

But it is envisaged that the PhD students interact in their research. The PhDs are paid a stipend of around 36.000 euros (before taxes) per year, and there is additional money for travel and research assistants.

More information, and instructions on how to apply can be found at under “vacancies”.

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. 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 to the 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.



Wild Replicator’s Got Funky Rhythm, Part 1

Now that the replicator meme is out and about I’ve got more to say. I’m going to republish two more posts from my 2010 cultural evolution series. These posts are about music. I have various reasons for using music as my cultural evolution conceptual sandbox. For one thing, it means that I don’t have to contend with semantic meanings arbitrarily associated with bits of music. In music, all we’ve got is the physical signal.

In these two posts I choose, not a simple musical example but, rather, a complex one, something jazz musicians know as Rhythm Changes. While I could talk about the four-note motif Beethoven used to construct the first movement of his Fifth Symphony, which is a memetic favorite, that’s too easy. Thinking about it won’t stretch our intuitions about the memetic properties of mere physical things. That motif has four notes, with specific durations and specific note-to-note pitch relationships.

Rhythm Changes isn’t like that. It’s an abstract property of a sound stream. There is now specific number of notes, no specific durations, and no specific note-to-note pitch relationships. Thousands upon thousands of specific musical streams, many quite different from one another, have exemplified the properties of Rhythm Changes.

In the previous post (in this series) I argued memes, the cultural parallel to the biological gene, are those physical properties of objects, events, and processes that allow different individuals to coordinate their participation in those things. In this view, memes are not physical objects, like genes, that spread through a population. Rather, memes are about sharability; they are physical properties that can easily be identified by human nervous systems and thus be the basis for shared (cultural) activity.

In that post I considered a very basic case, people making noise at regular intervals. In that case we have two memes, period (the interval between “hits”) and phase (the relationship between streams of hits by different individuals). Now I want to consider a considerably more complex case, the entity jazz musicians know as Rhythm Changes. This entity assumes that, for a given performance, period length and phase value are agreed upon. In fact it assumes a lot more. We’re dealing with a whole lot of memes here.

But I don’t want to get hung up in those details. I just want to characterize Rhythm Changes in a reasonable way and explain just why I insist that we regard Rhythm Changes as a structured collection of physical properties that can be ascribed to a stream of sound. While it would be nice to characterize Rhythm Changes using the language of acoustics, it’s not at all clear to me that we’ve got the necessary concepts. In any event, if we do, I don’t know them. Instead, I’ll couch my description in the schematic terms jazz musicians tend to use when talking about their craft; these terms are derived, in part, from descriptive and analytic concepts developed for European art music (i.e. classical music).

I’m going do this in two posts, the first will be confined to Rhythm Changes itself. The second will consider how Rhythm Changes came into being and how it functions in the popular music system. Continue reading “Wild Replicator’s Got Funky Rhythm, Part 1”

Evolang Previews: The Evolution of Morphological Agreement

Evolang is busy this year – 4 parallel sessions and over 50 posters.  We’ll be positing a series of previews to help you decide what to go and see.  If you’d like to post a preview of your work, get in touch and we’ll give you a guest slot.

Richard Littauer The Evolution of Morphological Agreement
Every lecture theatre but Lecture Theatre 1, all times except 14:45-15:05, and certainly not on Friday 16th

In this talk I’m basically outlining the debate as I see it about the evolution of morphology, focusing on agreement as a syntactic phenomenon. There are three main camps: Heine and Kuteva forward grammaticalisation, and say that agreement comes last in the process, and therefore probably last in evolution; Hurford basically states that agreement is part of semantic neural mapping in the brain, and evolved at the same time as protomorphology and protosyntax; and then there’s another, larger camp who basically view agreement as the detritus of complex language. I agree with Hurford, and I propose in this paper that it had to be there earlier, as, quite simply put, there is too much evidence for the use of agreement than for its lack. Unfortunately, I don’t have any empirical or experimental results or studies – this is all purely theoretical – and so this is both a work in progress and a call to arms. Which is to say: theoretical.

First, I define agreement and explain that I’m using Corbett’s (who wrote the book Agreement) terms. This becomes relevant up later. Then I go on to cite Carstairs-McCarthy, saying that basically there’s no point looking at a single language’s agreement for functions, as it is such varied functions. It is best to look at all languages. So, I go on to talk about various functions: pro-drop, redundancy, as an aid in parsing, and syntactic marking, etc.

Carstairs-McCarthy is also important for my talk in that he states that historical analyses of agreement can only go so far, because grammaticalisation basically means that we have to show the roots of agreement in modern languages in the phonology and syntax, as this is where they culturally evolve from in today’s languages. I agree with this – and I also think that basically this means that we can’t use modern diachronic analyses to look at proto-agreement. I think this is the case mainly due to Lupyan and Dale, and other such researchers like Wray, who talk about esoteric and exoteric languages. Essentially, smaller communities tend to have larger morphological inventories. Why is this the case? Because they don’t have as many second language learners, for one, and there’s less dialectical variation. I think that today we’ve crossed a kind of Fosbury Flop in languages that are too large, and I think that this is shown in the theories of syntacticians, who tend to delegate morphology wherever it can’t bother their theories. I’m aware I’m using a lot of ‘I think’ statements – in the talk, I’ll do my best to back these up with actual research.

Now, why is it that morphology, and in particular agreement morphology, which is incredibly redundant and helpful for learners, is pushed back after syntax? Well, part of the reason is that pidgins and creoles don’t really have any. I argue that this doesn’t reflect early languages, which always had an n-1 generation (I’m a creeper, not a jerk*), as opposed to pidgins. I also quote some child studies which show that kids can pick up morphology just as fast as syntax, nullifying that argument. There’s also a pathological case that supports my position on this.

Going back to Corbett, I try to show that canonical agreement – the clearest cases, but not necessarily the most distributed cases – would be helpful on all counts for the hearer. I’d really like to back this up with empirical studies, and perhaps in the future I’ll be able to. I got through some of the key points of Corbett’s hierarchy, and even give my one morphological trilinear gloss example (I’m short on time, and remember, historical analyses don’t help us much here.)  I briefly mention Daumé and Campbell, as well, and Greenberg, to mention typological distribution – but I discount this as good evidence, given the exoteric languages that are most common, and complexity and cultural byproducts that would muddy up the data. I actually make an error in my abstract about this, so here’s my first apology for that (I made one in my laryngeal abstract, as well, misusing the word opaque. One could argue Sean isn’t really losing his edge.)

So, after all that, what are we left with? No solid proof against the evolution of morphology early, but a lot of functions that would seem to stand it firmly in the semantic neural mapping phase, tied to proto-morphology, which would have to be simultaneous to protosyntax. What I would love to do after this is run simulations about using invisible syntax versus visible morphological agreement for marking grammatical relations. The problem, of course, is that simulations probably won’t help for long distance dependencies, I don’t know how I’d frame that experiment, and using human subjects would be biased towards syntax, as we all are used to using that more than morphology, now, anyway. It’s a tough pickle to crack (I’m mixing my metaphors, aren’t I?)

And that sums up what my talk will be doing. Comments welcomed, particularly before the talk, as I can then adjust accordingly. I wrote this fast, because I’ll probably misspeak during the talk as well – so if you see anything weird here, call me out on it. Cheers.

*I believe in a gradual evolution of language, not a catastrophic one. Thank Hurford for this awesome phrase.

Selected References

Carstairs-McCarthy, A. (2010). The Evolution of Morphology. Oxford, UK: Oxford University Press.
Corbett, G. (2006). Agreement. Cambridge, UK: Cambridge University Press.
Heine, B., and Kuteva, T. (2007). The Genesis of Grammar. Oxford, UK: Oxford University Press.
Hurford, J.R. (2002). The Roles of Expression and Representation in Language Evolution. In A. Wray (Ed.) The Transition to Language (pp. 311–334). Oxford, UK: Oxford University Press.
Lupan, G. & Dale, R (2009). Language structure is partly determined by social structure Social and Linguistic Structure.

Evolving Linguistic Replicators: Major Transitions and Grammaticalisation

ResearchBlogging.orgJust before Christmas I found myself in the pub speaking to Sean about his work on bilingualism, major transitions and the contrast between language change and the cultural evolution of language. Now, other than revealing that our social time is spent discussing our university work, the conversation brought up a distinction not often made: whilst language change is part of language evolution, the latter is also what we consider to be a major transition. As you evolutionary biologists will know, this concept is perhaps best examined and almost certainly popularised in Maynard Smith & Szathmáry’s (1995) The Major Transitions in Evolution. Here, the authors are not promoting the fallacy of guided evolution, where the inevitable consequence is increased and universal complexity. Their thesis is more subtle: that some lineages become more complex over time, with this increase being attributable to the way in which genetic information is transmitted between generations. In particular, they note eight transitions in the evolution of life:

What’s notable about these transitions, and why they aren’t necessarily an arbitrary list, is that all of them share some broad commonalities, namely:

Continue reading “Evolving Linguistic Replicators: Major Transitions and Grammaticalisation”

Fungus, -i. 2nd Decl. N. Masculine – or is it?: On Gender

ResearchBlogging.orgIn an attempt to write out my thoughts for others instead of continually building them up in saved stickies, folders full of .pdfs, and hastily scribbled lecture notes, as if waiting for the spontaneous incarnation of what looks increasingly like a dissertation, I’m going to give a glimpse today of what I’ve been looking into recently. (Full disclosure: I am not a biologist, and was told specifically by my High School teacher that it would be best if I didn’t do another science class. Also, I liked Latin too much, which explains the title.)

It all started, really, with trying to get my flatmate Jamie into research blogging. His intended career path is mycology, where there are apparently fewer posts available for graduate study than in Old English syntax. As he was setting up the since-neglected Fungi Imperfecti, he pointed this article out to me: A Fungus Walks Into A Singles Bar. The post explains briefly how fungi have a very complicated sexual reproduction system.

Fungi are eukaryotes, the same as all other complex organisms with complicated cell structures. However, they are in their own kingdom, for a variety of reasons. Fungi are not the same as mushrooms, which are only the fruiting bodies of certain fungi. Their reproductive mechanisms is rather unexpectedly complex, in that the normal conventions of sex do not apply. Not all fungi reproduce sexually, and many are isogamous, meaning that their gametes look the same and differ only in certain alleles in certain areas called mating-type regions. Some fungi only have two mating types, which would give the illusion of being like animal genders. However, others, like Schizophyllum commune, have over ten thousand (although these interact in an odd way, such that they’re only productive if the mating regions are highly compatible (Uyenoyama 2005)).

Some fungi are homothallic, meaning that self-mating and reproduction is possible. This means that a spore has within it two dissimilar nuclei, ready to mate – the button mushroom apparently does this (yes, the kind you buy in a supermarket.) Heterothallic fungi, on the other hand, merely needs to find another fungi that isn’t the same mating type – which is pretty easy, if there are hundreds of options. Other types of fungi can’t reproduce together, but can vegetatively blend together to share resources, interestingly enough. Think of mind-melding, like Spock. Alternatively, think of mycelia fusing together to share resources.

In short, the system is ridiculously confusing, and not at all like the simple bipolar genders of, say, humans (if we take the canonical view of human gender, meaning only two.) I’m still trying to find adequate research on the origins of this sort of system. Understandably, it’s difficult. Mycologists agree:

“The molecular genetical studies of the past ten years have revealed a genetic fluidity in fungi that could never have been imagined. Transposons and other mobile elements can switch the mating types of fungi and cause chromosonal rearrangements.Deletions of mitochondrial genes can accumulate as either symptomless plasmids or as disruptive elements leading to cellular senescence…[in summary,] many aspects of the genetic fluidity of fungi remain to be resolved, and probably many more remain to be discovered.” (Deacon, 1997: pg. 157)

At this point you’re probably asking why I’ve posted this here. Well, perhaps understandably, I started drawing comparisons between mycologic mating types and linguistic agreement immediately. First, mating-type isn’t limited to bipolarity – neither is grammatical gender. Nearly 10% of the 257 languages noted for number of genders on WALS have more than five genders. Ngan’gityemerri seems to be winning, with 15 different genders (Reid, 1997). Gender distinctions generally have to do with a semantic core – one which need not be based on sex, either, but can cover categories like animacy. Gender can normally be diagnosed by agreement marking, which, taking out genetic analysis of the parent, could be analogic to fungi offspring. Gender can be a fluid system, susceptible to decay, mostly through attrition, but also to reformation and realignment – the same is true of mating types. (For more, see Corbett, 1991)

As with all biologic to linguistic analogues, the connections are a bit tenuous. I’ve been researching fungal replication partly for the sake of dispelling the old “that’s too complex to have evolved” argument, which is probably the most fun point to argue against creationists with. However, I’ve mostly been doing this because fungi and linguistic gender distinctions are just so damn interesting.

If anyone likes, I’ll keep you updated on mycologic evolution and the linguistic analogues I can tentatively draw. For now, though, I’ve really got to get back to studying for my examination in two days. Which means I’ve got to stop thinking about a future post involving detailing why “Prokaryotic evolution and the tree of life are two different things” (Baptiste et al., 2009)…


  • Corbett, G. Gender. Cambridge University Press, Cambridge: 1991.
  • Deacon, JW. Modern Mycology. Blackwell Science, Oxford: 1997.
  • Reid, Nicholas. and Harvey, Mark David,  Nominal classification in aboriginal Australia / edited by Mark Harvey, Nicholas Reid John Benjamins Pub., Philadelphia, PA :  1997.

Uyenoyama, M. (2004). Evolution under tight linkage to mating type New Phytologist, 165 (1), 63-70 DOI: 10.1111/j.1469-8137.2004.01246.x
Bapteste E, O’Malley MA, Beiko RG, Ereshefsky M, Gogarten JP, Franklin-Hall L, Lapointe FJ, Dupré J, Dagan T, Boucher Y, & Martin W (2009). Prokaryotic evolution and the tree of life are two different things. Biology direct, 4 PMID: 19788731

Domain-General Regions and Domain-Specific Networks

The notion of a domain-specific, language acquisition device is something that still divides linguists. Yet, in an ongoing debate spanning at least several decades, there is still no evidence, at least to my knowledge, for the existence of a Universal Grammar. Although, you’d be forgiven for thinking that the problem was solved many years ago, especially if you were to believe the now  sixteen-year old words of Massimo Piattelli-Palmarini (1994):

The extreme specificity of the language system, indeed, is a fact, not just a working hypothesis, even less a heuristically convenient postulation. Doubting that there are language-specific, innate computational capacities today is a bit like being still dubious about the very existence of molecules, in spite of the awesome progress of molecular biology.

Suffice to say, the analogy between applying scepticism of molecules and scepticism of Universal Grammar is a dud, even if it does turn out that the latter does exist. Why? Well, as stated above: we still don’t know if humans have, or for that matter, even require, an innate ability to process certain grammatical principles. The rationale for thinking that we have some innate capacity for acquiring language can be delineated into a twofold argument: first, children seem adept at rapidly learning a language, even though they aren’t exposed to all of the data; and second, cognitive science told us that our brains are massively modular, or at the very least, should entail some aspect that is domain specific to language (see FLB/FLN distinction in Hauser, Chomsky & Fitch, 2002). I think the first point has been done to death on this blog: cultural evolution can provide an alternative explanation as to how children successfully learn language (see here and here and Smith & Kirby, 2008). What I haven’t really spoken about is the mechanism behind our ability to process language, or to put it differently: how are our brains organised to process language?

Continue reading “Domain-General Regions and Domain-Specific Networks”