Sunday, November 17, 2013

Discussion Leading Tuesday November 12- Handicaps, deception, and manipulation






We began with “Is honesty the best policy? Testing signal reliability in fiddler crabs when receiver-dependent costs are high” (Bywater, C., et al, 2012).  The authors of this paper took notice that signal reliability has only been manipulated in the lab.  While it has been proven costly to the unreliable signaler, the writers pointed out that no studies have been done in the natural habitat.  This paper was one of the first to examine signaling and receiver-dependent costs in the wild, and their findings advanced the field of Animal Behavior.  They showed a correlation between signal reliability and population density.  A signal was more likely to be unreliable when the signaler was in a low-density population.  I began by asking why this is.  Through their study Bywater, et al found that in low population density, individuals were less likely to come into contact with a competitor.  Because of this, they allocated less energy to developing muscle, yet retained large claw size.  This leads to an unreliable signal.  Crabs first present their claw to any competitor or potential mate.  A large claw size is more likely to deter a competitor and attract a female.  However, if a competitor wished to engage in physical contact, they might find that claw size is not representative of claw strength.  This is an unreliable signal.
                It was often mistaken during the discussion that claw strength is dependent on claw size.  This is not what Bywater, et al found.  In actuality, it was the shape of the claw that determined strength.  We then postulated why a large, shapely claw might not be representative of strength.  It was brought up that resource availability might be an effect on claw strength, a factor Bywater, et al admittedly did not take into account.  A rocky terrain vs. a smooth terrain would definitely affect population density and potential competitor interaction.  We further discussed the ability of the crab to better the representation of its claw strength after molting.  It is possible that the claw could get bigger after a molting season, and using information from a previous season, it could allocate more muscle to the claw.  In this way, the crab could become a more reliable individual.  We also agreed during discussion that Bywater, et al missed out on some great data by leaving out crabs that lost a claw and then replaced it.  They assumed that because they are generally weaker, they are automatically unreliable signals.  But if they had included these crabs as isolated events, their findings might have been surprising and the discussion could have delved deeper in what it means to develop a stronger claw.
                The discussion on the second paper “A mechanism of extreme growth and reliable signaling in sexually selected ornaments and weapons” (Elmen, D., et al) was admittedly better.  This paper examined horn growth in rhinoceros beetles.  They examined its dependence on the insulin/IFG pathway in comparison to genital and wing size.  When insulin receptor gene was knocked out, they found that genital size was not affected significantly; wing size was affected, but not as drastically as horn size.  This is thought to be controlled by the intracellular signaling as a product of blocking insulin receptors.  From this we can define two terms: handicaps vs. indices.  Handicaps are those signals that are unfaithful and incur a high cost on the signaler.  Indices are those that are impossible to replicate.  The insulin/IFG pathway is an example of indices.  This supports Condition Dependence.  This is a process defined by Animal Behaviorists as one that keeps a signal honest through many pathways.  This paper supports that horn size in rhinoceros beetles is a reflection of cellular processes (insulin/IFG pathway).
                Now that we know the trait is dependent on insulin, we can look at the proximate question (ultimate was assumed) on the trajectory of evolutionary change.  If there is an increase in sensitivity to insulin, then selection pressure could select for increased insulin sensitivity.  This brings up the question, what came first, the chicken or the egg?  In other words, did selection lead to sensitivity or did sensitivity lead to selection?  Either answer could be correct, but could be determined by looking at phylogenetic similarities to insulin sensitivity.  If it arose early in rhinoceros beetle evolution, the former can be assumed.  If sensitivity arose late in their evolutionary lineage, then the latter can be assumed.  Since we did not have a rhinoceros beetle phylogeny, we moved onto another question.  One that was even more complicated, but proved fun to speculate on.
Someone brought up the question, what is quality?  Elmen, et al repeatedly defined beetles as high-quality individuals and low-quality individuals, but failed to define quality.  It is assumed that quality is a mark of some previous defined standard, but that remains unclear throughout the paper.  We assume that quality is reflective of horn size, but what does horn size say about the individual.  Are they more likely to reproduce?  Are they more competitive?   Do they gain more resources?  Is there a top barrier to horn size, and how does that affect the individual?  Ultimately, this part of the discussion led to more questions than answers.  It felt like we were going in a circle when it was brought to our attention that, indeed, we were.  Someone in the group mentioned a certain trait controlling quality.  From this I drew a little diagram that encapsulated our discussion of the term quality.  Trait→ Reproductive Success→ Quality→ Trait.  This was just too mind-boggling to handle so I diverted our attention back to the paper.
Figure 2 D, E, and F plotted body size (g) against Aedeagus (genital) length (mm), wing length (mm), and horn length (mm) respectively.  These can be seen above, courtesy of Elmen et al.  These graphs show a simple linear regression for control animals vs. knockout individuals (those that received dsln-R injections to knockout insulin receptors).  These graphs were simple, yet showed something very interesting and significant.  The slope of the line did not change for each graph while the y-intercept did.  I brought this to the attention of the class and asked why this was important and what it says.  Put simply, y=mx+b.  Body size remains proportional to the three measurements taken (slope).  This shows that insulin affects growth proportionally.  The only thing that changed was the base line (y-intercept) at which the control and knockout individuals began to develop the three measurements taken.  Because there was only one variable, a goal of every experiment, Elmen et al were able to measure the magnitude of difference between the control and knockout individuals.  This is where their significant data came from and how they were effectively able to support their hypothesis that sexual selected ornaments and weapons are highly controlled.  When looking at these graphs, we see that genital length differed between the two groups by 0%.  They were able to conclude that genital formation is independent of insulin/IFG.  Wing size differed slightly (2%) showing a slight dependence on insulin/IFG pathway.  They were probably very excited when they analyzed horn length.  The knockouts showed a 16% change in length when compared to those of the control group.  This shows an astounding dependence on insulin/IFG.  It remains to be seen as to why such an important ornament is dependent on insulin, but data shows time and time again that we have a pretty clear idea of how.

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