The Grand Slam: How Hard Should You Hit?

Today on the blog we're happy to have former Wilson Honour's student, Rebecca Wheatley, who describes her thesis work on gecko personality. Rebecca's currently working as a research associate in the Wilson lab, and plans to start a PhD next year. You can find out more about Rebecca on her science blog, The Adventures of the Integrative Ecologist. 

The trade-off between performance and accuracy is a problem faced by a lot of different animals in a variety of situations. For example, consider a squirrel running along a bare branch to get from one tree to another; the faster it runs, the less time it spends exposed to predators. However, as the squirrel runs faster, it also increases its chances of mis-stepping and falling to its potential doom. 

So, to get the best of both worlds, the squirrel needs to optimise its running speed depending on its chance of slipping (the width of the branch) and the cost of falling off (the height from the ground).

Squirrels know what’s going down (or do they)? Image source: Wikimedia commons.

These sort of performance/accuracy trade-offs are also commonplace in the human world. How fast should you smash out a text message to your supervisor asking him (politely) to email back your latest draft before the number of typos makes the whole thing unintelligible?  In particular, these trade-offs are of a great deal of interest in elite sports. An awesome example of a sport where this trade-off is of utmost importance is in singles tennis. 

Serving hard: Heather Watson, Roger Federer and David Ferrer. Image source: Wikimedia commons.

In tennis, it’s pretty well accepted that if you serve really hard, it’s more difficult for your opponent to return the ball. But the harder you serve, the more likely it is that you’ll miss the service area and fault. So, players will usually belt it out on their first serve, but if they miss the first serve they’ll hedge their bets and serve softer the second time round to make sure they don’t double fault.

A/Prof Robbie Wilson, Dr Chris Brown and I have been testing this idea about performance trade-offs and optimal strategies using data from the men’s singles in the 2013 Australian Open. We’ve found this observation to be generally true: the probability of winning the point increases as the serve speed approaches its maximum, but the probability of faulting increases as well (for most players – some players are really consistent at getting it in regardless of how fast they serve). This was reflected in the frequency of high serve speeds in the first and second serves.

Jérémy Chardy, Andy Murray and Janko Tipsarevic. Image source: Wikimedia commons.

We’ve also constructed an optimality model which predicts the optimal serve speed taking into account the probability of faulting and the cost of a fault. An optimality model is, in essence, a mathematical model where you input the risks and rewards of a specific situation for a given individual, and it will tell you the optimal response for that individual if it wants to both minimise the risks and maximise the rewards. 

Optimality modelling is useful because it allows us to calculate the optimal response of specific individuals to any situation. We are looking at whether their opponent’s world ranking (ability to return a fast serve) and the point they’re going for or defending against (normal, game, set or match) affects their serve speed in relation to their optimum, but more on those results later.

Rafael Nadal, Caroline Wozniacki and Jérémy Chardy. Image source: Wikimedia commons.

We hope that our research can teach us more about how animals optimise their behaviour and physical efforts to improve their chances of successfully performing a given task. Depending on what we find, we might even be able to offer specific recommendations to tennis players wanting to improve their service game – who knows what the future might hold!

Andrew Hunter, a PhD student in our lab, is looking at performance/accuracy trade-offs in soccer. Will the results be similar between an individual and a team sport? We don’t know yet, but it will be interesting to find out.

Novak Djokovic, Agnieszka Radwańska and Venus Williams. Image source: Wikimedia commons.

Can water dragons actually run on water?

Today's guest poster is Dr Christofer Clemente. After postdoctoral stints at Cambridge and Harvard, Chris obtained an ARC DECRA and joined the Wilson lab at UQ. You can keep up with Chris's adventures on his science blog, Biomechanics Downunder.

I have long been impressed with the ability of the South American Basilisk lizard to run on water. There are plenty of videos of it on youtube, for example a short one here. 

It's pretty amazing. Looking at this video alone shows 2 important aspects of its locomotion: 

1. it's able to lift the whole body out of the water, and 
2. it's able to do so for quite long distances (around 10-15m). 

Some lizards' ability to run on water has been documented quite well by a series of papers by a group at Harvard University, particularly Tonia Hsieh. They've done some great work, including describing how smaller lizards are better able to support their body weight than are larger lizards, modelling 3D forces and recording 3D kinematics of the lizards' stride. Below is a gif showing some of the detailed kinematics of the lizard stride from George Lauders lab webpage. 

(NOTE: just click on the gifs if they are not running)

One other important point reported in these papers, based on the description given in Hsieh (2003), is it seems the lizards' kinematics change when running on water, such that the limb moves behind the hip, rather than being both in front and behind the hip.   

This is shown quite well in the gif above. So given this information on how Basilisk runs on water, we can then ask the question,  

Can the water dragon (Intellagama lesueurii) also run on water? 

I was led to believe it may be able to from two dominate and convincing lines of logic. 

1. they are water dragons! - it might behoove them to be able to do so and 
2. I heard reports of the juvenile lizard being observed doing so from a fellow researcher. 

So I modified the lizard racetrack which I have here at the University of Queensland, by placing a short, water-filled aquarium across the water dragons' path which they must cross to get to the other side. Then I sat back and filmed them using the fastec high speed camera system. And this is a typical (read: absolutely best) result below

Well the first thing I noticed is that they are no basilisk lizards. The body is not held out of the water and progress is significantly slowed. The first step seems hardly effective at all, and the second step is much deeper, and seems like a breaking step, with the foot held flat. However, the following step seem to have some similarity to those of the basilisk. From steps 3 onwards, the foot does not appear to be pushed as far forward, and much of the stroke seems to be posterior of the hip, as in basilisk. Secondly the trapped air bubbles on the foot are interesting, and these are also observed for basilisk, where they are thought to be the result of tiny fringes along the toes of the south american lizard. Such fringes however, are not obvious in the water dragons. Below is a snapshot of the bubble being dragged down on the trailing edge of the foot. 

So I'm unsure what to make of this all. It does look like they are capable of some run/swim locomotion, but it certainly falls short of the amazing prowess of the basilisk.

Here are some less impressive runs. Though notice that the right hind foot is actually brought out of the water - suggesting they could be using surface effects to give more downward force. 

And this one below shows a similar stroke. 

So that's as far as I've got. Let me know whether you think it is sufficiently interesting to warrant detailed kinematic analysis, or whether you think water dragons are just a little impaired when it comes to running on water. 

Finally, I leave you with what happens after several trials and the dragons know the water is coming up. It led me to believe, that for water dragons, they sure do not like water! 

The boldest gecko: personality in a reptile

Today on the blog we're happy to have former Wilson Honour's student, Rebecca Wheatley, who describes her thesis work on gecko personality. Rebecca's currently working as a research associate in the Wilson lab, and plans to start a PhD next year. You can find out more about Rebecca on her science blog, The Adventures of the Integrative Ecologist.

Animal behaviour is a big field - and it's constantly expanding as research reveals gaps in our understanding of why animals do the things they do. One topic in animal behaviour that holds a great deal of interest for me is that of animal personality. This is a relatively new concept and, frankly, it's a little bit controversial. 

The word "personality" conjures up a variety of mental images, most of which pertain to one animal in particular: us. It goes without saying that people have different personalities; we experience it every day. But do other animals have personalities as well?

Great tit (Parus major), beadlet sea anemone (Actinia equina) and pumpkinseed sunfish (Lepomis gibbosus); three species that display animal personality, from very different groups. Image source: Wikimedia commons.

In animal behaviour, the term "personality" is defined as consistent differences in behaviour displayed by individuals. An example of a personality trait is how an individual responds to a threatening situation, termed boldness or shyness. Bold individuals are undaunted by threatening situations and will approach the stimulus, while shy individuals will stay away or hide.

There are heaps of different personality traits that have been studied, including boldness, exploratory behaviour and aggression, amongst many others. Individuals' "personalities" are thought to range along a proactive-reactive continuum, where proactive individuals are aggressive and bold while reactive individuals are more passive and shy (sound familiar? It's not unlike a simplified version of the extroverted/introverted behaviour displayed by people).

There's growing evidence that "personality" is present within many groups of animals. Despite this, we don't really know much about what determines an animal's place along the proactive-reactive continuum or why this variation exists.

My study species: the Asian house gecko (Hemidactylus frenatus). Image credit: Wikimedia Commons (1 & 3) and Rebecca Wheatley (2).

During my honours project, I investigated "personality" in male Asian house geckos (Hemidactylus frenatus). I measured the anti-predator behaviour (a proxy for boldness) of 100 geckos by filming each gecko for one hour and then by calculating the proportion of time it spent inside the shelter in its terraruim. 

Each gecko was measured under three different treatments:

1. "empty terrarium": where nothing (aside from the shelter) was added to the terrarium, to give me a measure of each gecko’s normal amount of anti-predator behaviour
2. "terrarium with novel object": where I added a novel object to the terrarium, to see what happened to their anti-predator behaviour when something new was added to the environment
3. "terrarium with threatening stimulus": where I added a threatening stimulus, to see how their anti-predator behaviour changed when something scary was added to their environment

I found that different individuals reacted to the treatments in different ways, but the overall trend looked like this:

 

We can see that when a novel object was added to the environment, the geckos' anti-predator behaviour generally decreased when compared to their standard level of anti-predator behaviour. This might be because they wanted to check out the new object to make sure it wasn't food or some other valuable resource. 

However, when I added a threatening stimulus, their anti-predator behaviour jumped back up again to around the same as its standard level. So it seems that the threatening stimulus effectively cancelled out the novel object effect.

How do we know if these behaviours constitute as "personality"? Well, I found that while different individuals displayed consistent anti-predator behaviour within treatments, they also responded to the treatments in different ways. Some displayed more anti-predator behaviour when the environment was altered (were "shyer"), while others displayed less (were "bolder").  Therefore, from our definition, we can see that their anti-predator behaviour is a personality trait: they display consistent differences in behaviour that are context-specific.

Checking on my gecko housing set-up. Image credit: Amanda Niehaus.

But why do individuals have different personalities? Previous research has found that a few things can be associated with an animals' boldness or shyness. A large body mass is often associated with a bold personality, which is probably because heavier individuals are usually larger and more likely to win in a fight (so they have a good reason to be bold). Similarly, individuals with a hard bite force, a strong claw pinch or any other performance trait which would give them an advantage in a contest are usually bolder as well.  

The possession of traits that might make it easier for them to escape from a predator in a pinch, like fast running speed, have also been associated with boldness. In addition, resting (or "standard" for reptiles) metabolic rate has been linked to animal personality; it's thought that bolder, more aggressive individuals need a higher metabolic rate to keep up with their energetic demands.

I investigated how some of these traits interact to effect boldness in my geckos. I measured each gecko's mass, standard metabolic rate, maximum running speed and maximum bite force and analysed their interactive effects on anti-predator behaviour. Contrary to what I expected (and to what the literature would lead us to predict), I found that none of these traits affected anti-predator behaviour. This could be due to a few different things: one possibility is that boldness and shyness in Asian house geckos has a hormonal basis. It could also be that "personality" in geckos develops based on experiences rather than any specific physiological or performance trait. To discover the answer to this question, further research into the interactive effects of such traits on personality needs to be done.

One of my geckos in his metabolic chamber. I did all my metabolic tests during the day (when they are least active, being nocturnal animals) so I could get an accurate estimate of their resting (standard) metabolic rate. Image credit: Amanda Niehaus.

Anyway, why does it all matter – why does "personality" even exist? The fact is there are costs and benefits to being both proactive and reactive. Proactive individuals are bolder and more aggressive, so they are usually better at holding territories and getting laid – but they're also a lot more conspicuous to predators, so they tend to "live hard, die young". Reactive individuals, on the other hand, might not have the best real estate or as many mates at any given time, but their shy behaviour means they usually live longer. So, if we imagine an ecosystem where predation is low, it's better to be proactive and reap the benefits without the risk of being eaten. But if the ecosystem changes (for example, a bunch of predators move into the neighbourhood) and all the proactive guys die off – who is left? This is the most popular theory as to why different personalities exist; so that if conditions change quickly, some individuals survive and the population continues.

Although extremely interesting, these personality experiments were only one small aspect of my honours project, which aimed to answer questions about fighting ability (resource-holding potential) and fighting strategies. More on that later!

- by Rebecca Wheatley

Bit of a teaser for the rest of my project. Image credit: Amanda Niehaus.