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.

 

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Phonology and Phonetics 101: Vowels pt 1

In phonetics and phonology there is an important distinction to be made between sounds that can be broadly categorised into two divisions: consonants and vowels. For this post, however, I will be focusing on the second, and considered by some to be the more problematic, division. So, what are vowels? For one, they probably aren’t just the vowels (a, e, i, o, u) you were taught in during school. This is one of the big problems when teaching the sounds systems of a language with such an entrenched writing system, as in English, especially when there is a big disconnect between the sounds you make in speech and the representation of sound in orthography. To give a simple example: how many different vowels are there in bat, bet, arm, and say? Well, if you were in school, then a typical answer would be two: a and e. In truth, from a phonological standpoint, there are four different vowels: [æ], [e], [ɑː], [eɪ]. The point that vowel-sounds are different from vowel-letters is an easy one to get across. The difficultly arises in actually providing a working definition. So, again, I ask:

What are vowels?

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Python: Your one stop shop for social science research.

Python is easy to use and pretty solid across platforms.  Drew Conway has written an essential list of python tools for social science researchers.  From running experiments to analysis and modelling, Python can do practically anything you’d ever want, mostly in two or three lines.  Hooray!

http://www.drewconway.com/zia/?p=204

Mathematical Modelling 101 – The Price Equation

So in this post I’m going to assume you know absolutely nothing about anything. If you know something about something this probably isn’t what you’re looking for. If you’re looking for something which will go into depth on how the price equation is derived this probably isn’t what you’re looking for either. If you simply want to know what the price equation does and how to use it at face value then welcome! You’ve found the right place.

The price equation is used to calculate how the average value of any variant can change within a population from generation to generation.

Here I will cover everything you need to know to understand the equation and slot in the right values:

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Animal Signalling Theory 101 – The Handicap Principle

One of the most important concepts in animal signalling theory, proposed by Amotz Zahavi in a seminal 1975 paper and in later works (Zahavi 1977; Zahavi & Zahavi 1997), is the handicap principle. A general definition is that females have evolved mating preferences for males who display exaggerated ornaments or behaviours that are costly to maintain and develop, and that this cost ensures an ‘honest’ signal of male genetic quality.

As a student I found it quite difficult to identify a working definition for this important type of signal mainly due to the apparent ‘coining fest’ that has taken place over the years since Zahavi outlined his original idea in 1975. For this reason, I have decided to provide a brief outline of the terminological and conceptual differences that exist in relation to the handicap principle in an attempt to help anyone who might be struggling to navigate the literature.

As Zahavi did not define the handicap principle mathematically, a number of interpretations can be found in the key literature due to scholars disagreeing as to the true nature of his original idea. Until John Maynard Smith and Harper simplified and clarified things wonderfully in their 2003 publication Animal Signals, to my knowledge at least four different interpretations of the handicap were being used and explored empirically and through mathematical modelling, each with distinct differences that aren’t all that obvious to grasp without delving into the maths.

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Phonology and Phonetics 101

What I’m going to try and do in this series of posts is follow my phonology module at Cardiff. As such, these posts are essentially my notes on the topic, and may not always come across too clearly. First, I thought it would be useful to give a quick definition of both phonology and phonetics, before moving on to discuss the anatomical organisation of our vocal organs.

Phonetics and Phonology

To begin, phonetics, often referred to as the science of speech sound, is concerned with the physical production, acoustic transmission and perception of human speech sounds (see: phone). One key element of phonetics is the use of transcription to provide a one-to-one mapping between phones and written symbols (something I’ll come back to in a later post). In contrast, phonology focuses on the systematic use of sound in language to encode meaning. So, whereas phonetics is specifically concerned with human speech sounds, phonology, despite having a grounding in phonetics, links in with other levels of language through abstract sound systems and gestures. SIL provides a useful little diagram showing where phonetics and phonology lie in relation to other linguistic disciplines:

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Mathematical Modelling 101 – Evolutionary Game Theory

Game Theory was fist applied to evolution by John Maynard-Smith and George Price in 1973. It differs from traditional game theory is that it focusses on dynamics of strategy change more than the properties of strategy equilibria, although equilibria still exist within EGT but are know as Evolutionary Stable Strategies as opposed to Nash Equilibria.

Dove-Hawk

Imagine a situation in which 2 members of a species come into conflict over a resource. Within this conflict each animal has the optional to ‘fight’, ‘display’ or ‘run away’. There are 2 strategies within this species, either the Dove strategy or the Hawk strategy. In the Dave strategy, upon meeting someone also adopting the Dove strategy both “Doves” display and share the resource or upon meeting a “Hawk” the Dove runs away. Adopting the Hawk strategy entails always fighting. So upon meeting a Dove the Hawk will fight and the Dove will run away and the Hawk will take all of the resource, and upon meeting another Hawk, both will fight and one will win out. On average across many interactions with other Hawks, the payoff gained ends up being (v/2)-c where v=value of resource and c=cost.

Dove Hawk
Dove v/2, v/2 v, 0
Hawk 0, v (v-c)/2, (v-c)/2

The question to ask of this game is, given values v and c, which strategy will evolutionarily win out?

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Mathematical Modelling 101 – Intro to Game Theory

This post is going to just be a very brief introduction to what Game Theory is, how it works and some basic terminology. In later posts I will get more advanced and cover how it can be applied to Cultural Evolution.

What is Game Theory?

Game theory is a branch of applied mathematics most commonly used in Economics. However, it can be very successfully applied to other social sciences as well as Evolutionary Biology. It gives both descriptive answers (what people do) and prescriptive answers (what people should do) in a given game.

Why is this relevant?

Game Theory is a very good tool in predicting outcomes, not only in the very simple games covered in this post, but also in predicting the outcomes of evolutionary strategies and of predicting outcomes for signalling games which can inform us on human and animal communicative strategies.  Running iterated games over populations can introduce interesting qualifications to these very simple ideas as well and explain some things which may, at first, appear maladaptive, as such is a very useful tool in bypassing our intuitions. It is of these things that the next few posts will explore.

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That’s Linguistics (Not logistics)


Linguists really need a catchy tune to match those in logistics. Any takers?

I always remember when one of my former lecturers said he was surprised by how little the average person will know about linguistics. For me, this was best exemplified when, upon enquiring about my degree, my friend paused for a brief moment and said: “Linguistics. That’s like logistics, right?” Indeed. Not really being in the mood to bash my friend’s ignorance into a bloody pulp of understanding, I decided to take a swig of my beer and simply replied: “No, not really. But it doesn’t matter.” Feeling guilty for not gathering the entire congregation of party-goers, sitting them down and proceeding to explain the fundamentals of linguistics, I have instead decided to write a series of 101 posts.

With that said, a good place to start is by providing some dictionary definitions highlighting the difference between linguistics and logistics:

Linguistics /lɪŋˈgwɪs.tɪks/ noun

the systematic study of the structure and development of language in general or of particular languages.

Logistics /ləˈdʒɪs.tɪks/ plural noun

the careful organization of a complicated activity so that it happens in a successful and effective way.

Arguably, linguistics is a logistical solution for successfully, and rigorously, studying language through the scientific method, but to avoid further confusion this is the last time you’ll see logistics in these posts. So, as you can probably infer, linguistics is a fairly broad term that, for all intensive purposes, simply means it’s a discipline for studying language. Those who partake in the study of language are known as linguists. This leads me to another point of contention: a linguist isn’t synonymous with a polyglot. Although there are plenty of linguists who do speak more than one language, many of them are quite content just sticking to their native language. It is, after all, possible for linguists to study many aspects of a language without necessarily having anything like native-level competency. In fact, other than occasionally shouting pourquoi when (drunkly) reflecting on my life choices, or ach-y-fi when a Brussels sprout somehow manages to make its way near my plate, I’m mainly monolingual.

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Language, Thought, and Space (IV): Comparing Different Cultures

In my last post on the relationship between language, thought and (thinking and talking about) space I wrote that one of the most interesting, but also one of the most difficult questions is to what extent linguistic differences in talking about space reflect conceptual and perceptual differences.
Researchers at the Max-Planck-Institute for Psycholinguistics (Nijmegen, Netherlands) and at the Max-Planck-Institute for Evolutionary Anthropology (Leipzig, Germany) have done very interesting experiments that shed light on this question
As I mentioned in a previous posts (see here) work done by Stephen Levinson and others on how different cultures talk about and conceptualize space has shown that not all of them employ a bodily, egocentric frame of reference or coordinate system as their dominant organizing principle for their experiences and thoughts. Speakers of “several indigenous languages of Australia, Papua New Guinea, Mexico, Nepal, and south West Africa,” in contrast, organize the axes of their dominant coordinate system by absolute principles such as fixed landmarks (e.g. uphill vs. downhill) or cardinal directions (e.g. move the chair to the north). In addition, there are also langouages that primarily use “intrinsic,” object-centred Frames of Reference, such as in “The dog is at the front of the library.”
Differences between Relative and Absolute Speakers in Non-linguistic Spatial Tasks
In a set of clever experiments Levinson and his colleagues have also shown that speakers of relative and absolute languages differ in how they solve non-linguistic spatial tasks.