Honest the possible mutations. This approach also

Honest signaling can
be seen in different types of scenarios such as the interactions between
predator and prey, mate selection between males and females, and offspring
fitness.  Many studies have looked into
the proximate and ultimate causes of behavior relating to honest signaling,
meaning if the causes of the individual animal’s behavior causes immediate
effects, or effects over time.  Honest
traits give off signals of an individual’s condition, such as strength, health,
and overall physical shape.  As
described, honest traits can either be true honest signaling, or deceptive
honest signaling.


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could potentially signal dishonestly to coerce others into behaving in ways
that benefit only signaler, not the receiver. 
Another study reflects on the conflict of interest and signal
interference and how they lead to the breakdown of honest signaling.  Honest signaling is said to be favored when
individuals share a common interest and signals carry reliable, accurate
information.  The method that was used to
test this was an alternative approach called quorum sensing, which involves
bacterial cells that release small diffusible signal molecules (Popat, 2015).  Cells respond to these signal molecules by
producing either more of the signal molecules that aid in cell growth.  Here, they started with a clonal population
of an opportunistic pathogen and examined how quorum sensing signaling evolves
through the spread of de novo mutations in response to variation in the
relatedness and signal reliability of signaling systems (Popat, 2015).  By using this approach, natural selection can
choose from all of the possible mutations. 
This approach also allows for identifying specific genes that can then
be sequenced and examined.  One theory
predicts that the extent of common interest between individuals can depend upon
their genetic relatedness.  Higher
genetic relatedness stabilizes honest signaling.  Another theory predicts that if signal
reliability is reduced by deceptive interference, this reduces the relative
benefit/cost ratio of responding to or producing a signal that can lead to the
breakdown of honest signaling (Popat, 2015). 
The prediction here is that because a higher relatedness favors quorum
sensing, this will allow the pathogen to better invade its host, which of
course, causes a higher virulence.   The
main conclusion from this study is that reduced relatedness, invasion of
mutants that have a reduced response to a signal, and interference by an
artificial signal all lead to the relative breakdown of signaling (Popat, 2015). 


research has been conducted on how communication relates to social or
physiological costs separately, but not much work has been done involving how
social and physiological costs interact with each other.  High quality individuals pay a much lower
cost for elaborate ornaments than lower quality individuals.  Models of social cost propose that aggressive
interactions with signal receivers provide the cost that maintains the signals
accuracy over evolutionary time (Tibbetts, 2014).  Two main types of
social cost have been suggested.  First,
is that receivers are more likely to attack when their competitor’s quality is
similar to their own.  The second type of
social cost is that a receiver may recognize a mismatch between the signal and a
specific behavior.  If this occurs, the
receiver may aggressively test the signaler to assess their true ability.  Physiological cost models propose that
production and maintenance of elaborate ornaments involve some type of physiological
cost that favors accuracy (Tibbetts, 2014). 
In this specific review, they looked at hormone-linked physiological
costs.  One of the best known models of
physiological cost is the immunocompetence-handicap hypothesis.  This hypothesis proposes that high
circulating levels of testosterone are required for the development of
elaborate ornaments (Tibbetts, 2014). 
Testosterone is extremely costly, as it may suppress immune
function.  This level of analysis focuses
on proximate causes of behavior, more specifically, sensory-motor
mechanisms.  The sensory-motor mechanism
described here involves hormone systems and how the individual adjusts their
responsiveness to environmental stimuli. 
As a result, social costs are very likely to have extensive physiological
consequences.  Social behavior could
maintain the link between elaboration of the ornament and amount of hormone
produced by the individual.   


this study, when the mother frog visits her tadpoles to provide them with eggs
for nutrition, they perform a begging signal by stiffening their bodies and vibrating
rapidly (Dugas, 2017).  Experimenters
performed a manipulation of the tadpole’s condition, which is classified as
need and quality. This manipulation also involved food deprivation, which is
classified as hunger.  If the tadpoles
begging signals need, the tadpoles in higher condition should beg less
intensely (Dugas, 2017).  On the other
hand, if the tadpole begging signals quality, high condition and more developed
tadpoles should instead beg more intensely (Dugas, 2017).  The results from this study revealed patterns
consistent with the signal of quality hypothesis and directly counter the
predictions of signal of need and signal of hunger (Dugas, 2017).  Begging effort was higher in more developed
and higher condition tadpoles and declined with food deprivation.  Mothers favor their high quality young.  This data suggests that maternal choices are
based at least in part on offspring signals, indicating that offspring begging
can evolve to signal high quality.   

signaling is not only relevant in sexual selection; it is also used to increase
an individual’s overall fitness.  Parental
and offspring fitness can be maximized at different levels of parental
investment.  Parental investment can be
defined as costly parental activities that increase the likelihood of survival
for some existing offspring but that reduce the parent’s chances of producing
offspring in the future (Alcock, 2009).  Parents
can be choosy and can often benefit themselves from distributing limited
resources to specific offspring.  One
study tested the predictions of three honest begging hypotheses; signal of
need, signal of quality and signal of hunger in tadpoles of a species of frog (Dugas,
2017).  The signal of need reflects the
assumption that the offspring with the lowest current reproductive values will beg
the most intensely (Dugas, 2017). 
Whereas the signal of quality hypothesis predicts the opposite, that
offspring in the best condition produce the most intense begging signals (Dugas,
2017).  This can help the individual by
making it known that their past accomplishments and achievements can be useful
in the future.  The signal of hunger
hypothesis predicts that signal intensity correlates with food deprivation,
with an individual signaling more intensely as its digestive tract empties and
it becomes hungrier (Dugas, 2017). 


prey often use warning signals, such as toxins or bright colors, to deter
potential predators away from them.  This
mechanism is called aposematism, which is mostly across many different animal
species, and it is perceived to be mostly honest.  Ultimately, an evolutionary simulated model
was utilized in which prey used some kind of warning display to their advantage.  This model assumed that there are two
possible outcomes.  One is that predators
are sensitive to the combined qualities of toxins and displays when they attack
prey.  It was shown that predators tend
to handle aposematic prey more carefully during attacks than non-aposematic
prey (Blount, 2009).  This could very well
be an evolutionary trait.  The second
assumption is that warning coloration and toxicity compete for the same
resources.  This assumption points to the
possibility that warning displays may be handicap signals (Blount, 2009).  This means that they are honest indicators of
defensive capability.  The big picture
showed that if warning displays and defenses compete for a shared resource,
warning signals can indeed be honest handicaps. 
However, when the availability of the key resource is not limiting, individuals
should be highly toxic and warning displays dishonest (Blount, 2009). 


using all of these different methods, this study looked at four different
pathways of color patterning in animals; predictable signals of social status,
developmental homeostasis, traits that increase somatic integrity, such as
elaborate feathers or scales, and traits of cost-added signals of maintenance
activities, such as grooming, that may link individual quality to the
expression of color patterns (Pérez-Rodríguez, 2017).  Certain traits that evolve are worn as badges
of social status.  It is typically
predicted that animals that hold a higher social status would have more simple
traits.  In this particular study, the
researchers looked at the more complex patterns of different animal species as
shown above.  The shape of certain
patterns has been shown to be influenced by body condition.  Also, uniformity, regularity and complexity
of color patterns are signals of developmental homeostasis (Pérez-Rodríguez,
2017).  The symmetry and uniformity of
the markings an animal possesses could indicate the stress levels suffered by
the individual.  Traits that increase physical
body condition, such as elaborate feathers or scales, play an important role as
well.  If an individual’s feathers or
scales appear damaged or are missing or broken, this in turn damages the
outward appearance of the individual. 
These outward damages could be caused by parasites, so grooming
activities are essential to maintain a pristine appearance.  Having a combination of clean markings and
beautiful contrasting colors may be a signal of good health and condition.  From the different models in this study, they
concluded that increasing pattern complexity requires controlling a higher
number of pathogens to achieve their target pattern (Pérez-Rodríguez, 2017).  The complexity in pattern can be used to
predict the evolutionary pathways in different animal species. 

previously mentioned, male body size plays a significant role in determining
quality, health, and strength in individuals. 
Another study looked specifically at an animal’s color patterning and
its correlation to quality signaling despite size and color intensity.  Previously, analytical methods for determining
color patterning were very scarce, but more recently there has been a large
development of these tools.  In this
study, different animal species were sampled and their specific color patterns
were observed.  There were a few
different methods that were used to quantify the different color patterns in
this study.  For example, barred patterns,
which are classified as lines or stripes, were aligned through a specific
software and the deviation from the ideal pattern was then quantified (Pérez-Rodríguez,
2017).  Another method that was used was
a color adjacency method, which collected color characteristics by digital
photography and these photographs were then aligned to a grid of the animal’s
entire body and the indices of pattern elongation, regularity and complexity
were able to be provided (Pérez-Rodríguez, 2017).  A different way that color pattern
quantification was used throughout this study looked at the spotted patterns and
their quantifications in terms of contrast, spatial frequency, phase and
orientation by granularity analyses (Pérez-Rodríguez, 2017).  Also, fractal geometry can be useful in
measuring heterogeneous patterns by measuring the continuousness of a pattern
through scales (Pérez-Rodríguez, 2017). 
Lastly, geometric morphometrics was used which is the analysis of
morphological structures, using Cartesian geometric coordinates, which
specifies each point by using numerical coordinates rather than linear
variables.  This tool captures the shape
of an object independent of its size, position and orientation (Pérez-Rodríguez,


sexual signaling has also been experimentally tested in turtles.  One specific study shows that there is a
trade-off between immune response and the coloration in a turtle species of red-eared
sliders.  Researchers performed this experiment
by using a bacterial antigen, which doesn’t cause any pathogenic effects.  Once the turtle’s immune response was
activated by the introduction of this specific antigen, the visual colorful
ornaments located on the chin and head of these turtles were observed and it
was noted if the coloration was affected or not (Ibáñez, 2014).  Compared to the control animals that were not
injected with the antigen, the injected animals exhibited; reduced brightness,
lower long wavelength reflectance, and lower values for carotenoid chroma
(Ibáñez, 2014).  Also, the injected
animals showed darker and less “yellowish” chin straps and less “reddish”
patches on their heads at the end of the study (Ibáñez, 2014).  This trade-off between immune response and
coloration may allow turtles to honestly signal their individual quality
through their coloration.  Furthermore,
this trade-off could be beneficial for females in sexual selection in picking
the healthiest mates and again passing on the best genes to her offspring.  These turtles have evolved to signal their
colorful ornaments in hopes of gaining female mates and passing along their


study was performed by collecting a sample of male orchid bees and taking
measurements of their size.  An enzyme
that is imperative in the defense mechanisms of insects was used in this study
called phenoloxidase.  Since
phenoloxidase is commonly used as a measure of insect immune response, the
levels of phenoloxidase were also measured in the collected bee sample
(Arriaga-Osnaya, 2017).  It was concluded
from this study that male orchid bee’s body size and the amount of perfume each
bee had were positively correlated to that individual bee’s immune function
(Arriaga-Osnaya, 2017).  Female orchid
bees may use the honest signals of male orchid bee’s body size and scent blends
as an indicator that a potential male mate is healthier and able to ward off
infections.  If she mates with a
healthier male, those strong genes could then be passed on to her
offspring.  This study specifically has
looked at both the proximate and ultimate causes of the male orchid bees.  Since male orchid bees do not produce
pheromones, over time they have evolved a way to make up for that
deficiency.  From this, they have ultimately
increased their overall fitness. 

most of the animal kingdom, regardless of species, larger males have a higher
mating success because they are considered more dominant and in better physical
condition.  In one study, researchers tested
the hypothesis that a male’s body size and the scents that they emit are honest
signals of their overall immune response (Arriaga-Osnaya, 2017).  Interestingly, male orchid bees don’t produce
pheromones so they resort to having to collect different blends of scents from
objects such as flowers, fungi, sap, and resins within their environment
(Arriaga-Osnaya, 2017).  These scent
blends that are collected are then displayed resembling perfumes during
courtship with female orchid bees. 


researchers have looked into how honest signaling fits into the different
levels of analysis in the fascinating study of animal behavior.  These different levels of analysis include
proximate and ultimate causes of behavior. 
Proximate causes of behavior refer to the immediate effects of the
individual animal’s internal developmental and physiological causes that occur
during that animal’s lifetime (Alcock, 2009). 
Ultimate causes of behavior refer to the evolution of the individual
animal’s historical events that happened in previous generations (Alcock, 2009).  From these analyses, there are four main
questions that behavioral researchers ask:  How does the behavior promote an animal’s
ability to survive?  How does an animal
use its sensory and motor abilities to modify its behavior patterns?  How does an animal’s behavior change in
response to the experiences that it has while maturing?  And what does an animal’s behavior tell us
about the origins of its behavior and the changes that have occurred during the
history of the species (Alcock, 2009)?   Here, a few of those questions will be
answered in regards to honest signaling within different species.

fitness refers to the measure of the genes contributed to the next generation
by an individual, often stated in terms of the number of surviving offspring produced
by the individual (Alcock, 2009).  An
example relating to how honest signaling is seen in overall fitness would be looking
at the question as to why baby birds beg so loudly.  Baby birds tend to do this because it then results
in competition between siblings that are in the nest.  Each individual baby bird uses its own
vocalization skills and begging behaviors and specific calls in hopes to get
its parents to deliver the ideal amount of food needed to maximize that individual
baby bird’s own fitness (Alcock, 2009).  The
most successful nestling’s genes will have the greatest probability of being
passed on in future generations. 

signaling is especially relevant in sexual selection and to overall fitness of
an individual.  Sexual selection is a
form of natural selection that occurs when individuals vary in their ability to
compete with others for mates or to attract members of the opposite sex
(Alcock, 2009).  A simple example of
honest signaling relating to sexual selection would be two male elks engaging
in a roaring battle to see who is in better overall physical condition.  The long duration of roaring expends a lot of
energy and only the male in the most top shape will be able to roar for many
minutes.  Weak male elk or elk that are
not in the best physical condition would in turn not be considered by females
and there genes may not be passed on to future generations.  Female elk would then benefit from this
honest signaling because they would be able to pick out a potential dominant
mate based on this honest signaling battle and the dominant males genes could
then be passed on in future generations.

The many
different behaviors that animals possess and the specific reasons as to why
they behave in these ways is a very interesting phenomenon.  Animal behavior relates to how animals interact
with each other and their environment.  More
specifically, the interactions between predator and prey, mate selection
between males and females, and offspring fitness use what is called “honest
signaling.”  Honest signaling refers to true
traits that can be either physical structures or behaviors of the
individual.  Honest traits also give off
signals that reflect the actual condition of the individual.  These specific traits or actions in turn end
up mutually benefiting the signaler of these traits and also the receiver of
the traits.

behavior relates to how animals interact with each other and their
environment.  More specifically, the
interactions between predator and prey, mate selection between males and
females, and offspring fitness use what is called “honest signaling.”  Honest signaling refers to true traits that
can be either physical structures or behaviors of the individual.  Honest traits give off signals of an
individual’s condition, such as quality, health, and strength.  It has been shown that this can either be
true honest signaling, or deceptive honest signaling.  Many studies have looked into the proximate
and ultimate causes of behavior relating to honest signaling, meaning if the
causes of the individual animal’s behavior causes immediate effects, or
effects over time. 



Honest Signaling in
Animal Behavior



Brittny Schnur


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