Lecture: Altruism: Answer Sheet

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Watch the following video: Richard Dawkins on Altruism and The Selfish Gene - YouTube

Introduction

1. Generate at least one hypothesis for why sentinel behavior has evolved in meerkats.
Hypothesis: The evolution of sentinel behaviour in meerkats is most likely due to the advantages
of altruistic behaviour and the kin selection theory.

Explanation: Survival in the harsh conditions where meerkats reside is difficult. Predators are a
constant concern, and food supplies can be limited. Sentinel behaviour, in which one meerkat
keeps guard while the others seek for food, can improve the group's survival rate. The sentinel
can warn the group of incoming predators, allowing them to flee and avoid being preyed upon.

This behaviour appears to be at the price of the sentinel, who sacrifices their own ability to
forage. However, the kin selection hypothesis suggests that this cost could be mitigated by the
enhanced survival rate of the sentinel's relatives inside the group. By assisting their kin in
surviving and reproducing, the sentinel indirectly passes on their own genes. This could explain
why sentinel behaviour arose and persists in meerkat groups.

Part 1: Paying the Price

1. What are some possible fitness costs of sentinel behavior for the sentinel?
Sentinel behaviour in social animals, in which specific individuals serve as guardians or lookouts
for the group, can have both advantages and disadvantages. Let's look at some of the possible
fitness costs connected with being a sentinel:

Sentinels are more vulnerable to predation as they survey the surroundings for the group. Their
elevated position makes them more conspicuous to predators, increasing their chances of being
targeted. This increased hazard may diminish the sentinel's chances of survival and reproductive
success.
Reduced Foraging Opportunities: Being a sentinel entails devoting time and energy to guard
tasks. Instead of actively seeking for food, sentinels must be alert. This shorter foraging time can
result in poorer energy intake and may have an influence on their overall fitness.

Energy Expenditure: Maintaining an attentive posture and scanning the surroundings requires
energy. Sentinels devote energy to staying watchful, which could otherwise be employed for
more important activities such as reproduction or self-maintenance.
Increased Stress: Sentinels may find it stressful to be always on watch. Chronic stress can harm
their health, immune system, and overall fitness.

Opportunity Costs: Time spent as a sentinel is time not spent on other useful tasks like grooming,
socializing, or mating. Missed opportunities might have an impact on an individual's fitness.

Reduced Reproductive Output: If sentinels devote a substantial amount of time to guarding, they
may have less opportunities to mate and care for their offspring. Reduced reproductive
production can have an impact on the group's genetic composition.

2. Now, analyze the data in Figure 3. They tested hypotheses in a wild population of meerkats;
half of helper meerkats were fed 25 g of hard-boiled egg for 30 successive days (fed), and half of
them were not (control). Their contributions to guarding behavior were documented during that
time period. GT refers to raised guarding, or sentinel behavior. Summarize the main
experimental findings shown in this figure.

Experimental setup: Helper meerkats were separated into two groups.

Fed Group: Received 25 g of hard-boiled egg everyday for 30 days.

The control group did not receive any additional food.

During this time, they were observed making contributions to guarding behaviour (particularly,
heightened guarding or sentinel behaviour).
The fed group of meerkats showed greater sentinel behaviour compared to the control group.
Specifically, meerkats given more food spent more time functioning as sentinels, assessing the
area for potential hazards.
The sentinel behaviour was distinguished by continual calls to alert other group members and
alarm sounds to signal the group to flee to their burrows if a predator was detected.
Interpretation: The fed group exhibited more sentinel behaviour, indicating a role for nutritional
status.
The enhanced sentinel behaviour in the fed group indicates that nutritional state influences
altruistic behaviour.
Meerkats given extra food were more likely to take up the sentinel position, maybe because they
had more energy to devote to guarding tasks.
Fitness implications:

While sentinel behaviour benefits the entire group by increasing collective safety, it imposes
fitness costs on individual sentinels.
Spending more time as a sentinel means less time foraging for food, potentially affecting their
general fitness.
However, the precise trade-offs between sentinel behaviour and individual fitness remain a
fascinating topic of research.

3. How might less time foraging translate to fitness costs for sentinels?
Reduced Energy Consumption: Spending more time as a sentinel results in less time actively
seeking for food. Meerkats rely on food to provide the energy they need for survival,
development, and reproduction. When sentinels spend a large amount of time on guard duty,
they may not ingest enough calories to maintain ideal health and energy levels. Reduced energy
intake can result in diminished fitness.
Nutritional deficiencies: Foraging gives vital nutrients, vitamins, and minerals required for good
health. Sentinels who prioritize guarding over foraging may not consume a well-balanced diet.
Over time, this insufficiency can have an influence on their immune system, reproductive
success, and overall health.

Meerkats with low physical condition are more vulnerable to illnesses, parasites, and
environmental disturbances. Reduced foraging time can lead to weight loss, muscular wastage,
and poor physical condition. A weak body state impairs their ability to survive and reproduce.

Every moment spent as a sentinel represents an opportunity cost. Instead of foraging, sentinels
devote time on vigilance. Missed foraging opportunities imply lost opportunity to obtain
necessary nutrients and energy. Over time, these lost opportunities add up and have an impact on
personal fitness.
Reproductive Trade-offs: Meerkats devote energy to several activities, including reproduction.
Sentinels may forego mating opportunities or parental care because of their sentinel job. Reduced
reproductive production has an influence on both genetic contribution to the group and overall
fitness.

Stress and Health: Constant vigilance can cause chronic stress and have a bad influence on
health. Stress hormones influence immunological function, reproductive hormones, and overall
health. Sentinels may face increased stress, which can contribute to decreased fitness.

4. Do these results support, refute, or have no relationship to the researchers’ hypothesis that
there is a fitness cost to sentinel behavior?

The findings back with the researchers' premise that there is a fitness penalty to sentinel
behaviour. Meerkats who received greater food (the "fed" group) displayed more sentinel
behaviour than the control group. However, increased guarding resulted in less foraging time,
perhaps affecting their overall fitness. Thus, the findings support the hypothesis that sentinel
behaviour in meerkat societies entails trade-offs and individual costs.

Part 2: For the Good of the Group

1. Imagine for a moment that sentinel behavior evolved by group selection, so meerkat groups
that had sentinels tended to have higher fitness than meerkat groups without sentinels, with
sentinel duty (and its associated costs) being shared among members of the group on an equal
rotation “for the good of the group.”

a. In a meerkat group with sentinels taking equal turns on duty, who is paying fewer fitness
costs: the foraging meerkats or the sentinel meerkat?
Fitness expenditures are likewise shared in a meerkat group where sentinel duty is evenly
distributed among all members. As a result, at any given time, the sentinel meerkat is paying
higher fitness costs because it is on duty and not foraging. However, as each meerkat takes its
turn as the sentinel, the costs eventually level out. So, in the long term, neither the foraging nor
the sentinel meerkats incur lower fitness costs. They all bear the costs evenly. This is the essence
of the group selection theory you cited, in which behaviours that are costly to an individual but
beneficial to the group might emerge.

b. Now imagine a meerkat with an allele for slightly more selfish behavior; this meerkat puts in
less time on sentinel duty than do other meerkats. Does this meerkat’s fitness increase or
decrease?
In the near run, a meerkat that spends less time on sentinel duty and more time foraging may
have a rise in fitness. This is because it will have more opportunities to acquire food, increasing
its chances of survival and reproduction.

However, if this behaviour becomes more widespread and fewer meerkats serve as sentinels, the
group's overall safety may suffer. This could result in increased predation rates and a decline in
fitness for all members of the group, even the more selfish meerkats. This is an example of a
contradiction between individual-level selection (which favours selfish behaviour) and group-
level selection (which promotes collaboration).

c. Does its allele for more selfish behavior increase or decrease in frequency in the population?
The frequency of the allele promoting more selfish behaviour in the population is determined by
the balance of individual and group selection.

If the advantages of selfish behaviour (more time foraging and less time on sentinel duty) exceed
This is because meerkats with the selfish allele have higher fitness and thus more opportunity to
pass on their genes to future generations.

However, if the consequences of selfish behaviour are too great (for example, a drop in group
safety results in a considerable rise in predation), the allele's frequency may decline. The loss in
group safety would have a negative effect on meerkats with the selfish allele, perhaps lowering
their fitness and decreasing their chances of passing on their genes.

It's also worth mentioning that if selfish behaviour becomes too prevalent and group safety
suffers dramatically, the benefits of selfish behaviour may shrink. In such a case, the frequency
of the selfish allele may initially grow but then drop over time.

This is a difficult scenario that requires a delicate balance of individual-level selection (favouring
the selfish allele) and group-level selection (favouring cooperation). The real outcome would be
determined by the situation's specific features, such as the precise fitness costs and advantages of
selfish behaviour, as well as the strength of group selection.
d. Consider the definition of fitness you came to class with today. If there’s an allele for altruism,
or selfless behavior, what happens to it in an altruistic group?
An allele for altruism or selfless behaviour may become more common in an altruistic
population. This is because in such a group, the advantages of altruistic behaviour (such as
improved group survival or reproductive success) may outweigh the individual costs.

Altruistic behaviour can improve the overall fitness of a group by encouraging collaboration and
mutual aid, which increases the group's likelihood of survival and reproduction. If the group
members are closely related (kin), the altruistic individual may indirectly boost its genetic
representation in the population by assisting relatives (who share some of its genes) in surviving
and reproducing. This is referred to as kin selection.
However, it is important to note that the spread and maintenance of altruistic alleles in a
population can be complex and influenced by a variety of factors, including the group's degree of
relatedness, the specific costs and benefits of altruistic behaviour, and the possibility of
exploitation by selfish individuals. Thus, while the altruistic allele may grow in frequency in an
altruistic population, this is not guaranteed and is dependent on the individual circumstances.

2. If researchers have found that solitary meerkats spend as much time on raised guard as do
meerkats in groups, does this information support, refute, or have no relationship to the group
selection hypothesis for sentinel behavior?

The discovery that solitary meerkats spend as much time on elevated watch as meerkats in
groups may call into question the group selection hypothesis for sentinel behaviour.

According to the group selection hypothesis, sentinel behaviour evolved to benefit the group,
with the cost borne by all members. If solitary meerkats, who do not profit from sharing the
expense of sentinel conduct with a group, continue to participate in this behaviour to the same
level as meerkats in groups, it may indicate that sentinel behaviour is not entirely a result of
group selection.
It could imply that sentinel behaviour can also be advantageous on an individual level. For
example, a solitary meerkat may profit from spending time on high alert by lowering its own
danger of predation.

However, this does not necessarily invalidate the group selection hypothesis. Individual and
collective selection may have played roles in the evolution of sentinel behaviour. The particular
balance between these two levels of selection may differ depending on the social organization
and ecological surroundings of the meerkats.

Thus, while this evidence complicates our understanding of sentinel behaviour, it does not
clearly support or refute the group selection idea. More research is needed to properly understand
the evolutionary processes driving this behaviour.

3. Can group selection explain the evolution of meerkat sentinel behavior? Explain.
Yes, group selection may have contributed to the evolution of meerkat sentinel behaviour, but it
is unlikely to be the main driver.

Group selection is the concept that natural selection can act at the group level, favouring features
that may be expensive to individuals but beneficial to the group as a whole. In the instance of
meerkat sentinel behaviour, the individual meerkat may incur a cost (such as lost foraging time),
but the group benefits from enhanced vigilance and lower predation risk.

However, it is vital to recognize that group selection is not always the sole evolutionary factor at
work. Individual selection may also be crucial. For example, a meerkat working as a sentinel
may obtain individual benefits like as improved social standing within the group or increased
survival probability as a result of being more watchful.

Furthermore, kin selection, a type of natural selection in which individuals can improve their
genetic success by assisting relatives, could have a role in the evolution of sentinel behaviour in
meerkats, as meerkat groups are frequently made up of closely related individuals.
As a result, while group selection may play a role in the development of sentinel behaviour in
meerkats, it is more likely that a combination of group selection, individual selection, and kin
selection is involved. The particular balance between these multiple forms of selection can be
influenced by a number of factors, including the meerkats' social structure, the specific costs and
benefits of sentinel behaviour, and the degree of relatedness within the group.

Part 3: Benefits if Being Sentinel

1. What are some possible individual fitness benefits of sentinel behavior for the sentinel?

Increased Survival Probability: Being more alert while on sentinel duty may assist the sentinel to
detect predators earlier, boosting its odds of survival.
Social Status and Mating Opportunities: If sentinel behaviour is regarded as favourable, it may
improve the sentinel's social standing within the group. This could lead to more mating
opportunities and consequently better reproductive success.
Kin Selection: If the sentinel is closely connected to other members of the group, it may improve
its indirect fitness by assisting its relatives in survival and reproduction. This is referred to as kin
selection.
Reciprocal Altruism: If sentinel duty is shared among group members, a meerkat may profit
from acting as a sentinel now in exchange for future protection from others' sentinel duty. This is
referred to as reciprocal altruism.

2. Now, analyze some data as you did earlier. As before, these data result from a study of wild
meerkats in the Kalahari. Summarize the main findings shown in Figure 4.
Direction of Attention (Graph A): When on elevated guard, meerkats spent more time looking
away from the group than towards it. This shows that the sentinel meerkats are primarily
concerned with detecting potential hazards in the environment rather than monitoring the group's
activities.
Distance to Nearest Bolt-Hole (Graph B): Foraging meerkats had a longer average distance to the
nearest bolt-hole (a place to escape predators) than guard meerkats. This suggests that meerkats
on guard duty relocate themselves closer to safety, presumably to offset the greater risk
associated with their exposed position.

3. Does sentinel behavior benefit the group or the sentinel? Who benefits most?

The biggest benefit for the group is greater safety. The sentinel acts as an early warning system,
informing the group of the presence of predators. This enables the group to act swiftly,
improving their chances of survival.
Benefit to the Sentinel: Being the first to see a predator allows the sentinel to initiate the escape
response, potentially enhancing its own chances of survival. Furthermore, if sentinel behaviour is
connected with higher social rank within the group, it may lead to more mating opportunities.

4. It was also reported that “during over 2000 hours of observation, no raised guards were
attacked or killed by predators, probably because raised guards were usually the first to detect
them.” Summarize the benefits and costs of sentinel behavior to the sentinel.

Benefits:

Increased Survival: The fact that no raised guards were attacked or killed by predators implies
that being a sentinel may improve an individual's survival rate. Sentinels typically notice
predators early, giving them the benefit of initiating the escape response, potentially enhancing
their own odds of survival.
Potential Social Benefits: If sentinel behaviour is connected with higher social rank within the
group, it may lead to more mating opportunities.

Costs:

Opportunity Cost: The time a meerkat spends on sentinel duty is time it cannot spend doing other
things, like scouting for food or relaxing. This could affect its energy levels and overall health.
While the sentinel may be the first to detect a predator, its elevated posture may make it more
noticeable to predators. However, the fact that no sentinels were assaulted or killed implies that
this risk may not be as high as previously believed.

Part 4: Summary

1. Given the data you just analyzed, provide an explanation for the evolution of sentinel behavior
in meerkats, being careful to avoid explanations that require altruism or group selection.
Individual selection, rather than altruism or group selection, can explain the evolution of sentinel
behaviour in meerkats. Here's how.

Sentinel behaviour improves the individual's survival rate. As the first to detect predators, the
sentinel has the benefit of initiating the escape response, potentially enhancing its odds of
survival.
Reduced Predation danger: While on watch duty, a meerkat can stay closer to bolt-holes (escape
routes), lowering its danger of predation. This provides a direct benefit to the individual meerkat,
regardless of the group.

Potential for Increased Reproductive Success: If sentinel behaviour is connected with higher
social standing within the group, it may result in more mating opportunities for the sentinel,
enhancing its individual fitness.
Reciprocal Altruism: Even in the absence of group selection or altruism, reciprocal altruism can
play a role. If sentinel duty is divided among group members, a meerkat may benefit from
serving as a sentinel now in exchange for future safety offered by the sentinel duty of others.
This is an example of direct reciprocity, in which people trade positive acts.
In conclusion, while sentinel behaviour benefits the group, it can also bring direct benefits to the
individual meerkat performing the function. These individual-level advantages can drive the
evolution of sentinel behaviour via individual selection.

2. Reciprocal altruism is a type of altruism where individuals take turns helping each other, even
if one individual has to wait for the payoff (e.g., I’ll scratch your back today if you’ll scratch
mine tomorrow). It tends to occur mostly in groups of social animals that continue to interact
with each other over time.

a. Considering what you have learned today, why do you think that repeated future interactions
are key to the evolution of reciprocal altruism?
Repeated contacts allow people to build trust and reliability. Trust is formed when one person
assists another and then receives support in return in subsequent exchanges. This mutual trust
promotes the continuation of reciprocal charity.
Cost-Benefit Balance: In reciprocal altruism, one person pays to help another in exchange for a
future benefit. Repeated encounters guarantee that individuals have the opportunity to reap the
anticipated future advantages while balancing the original investment.
Cheaters' Punishment: In a reciprocal altruism system, people who frequently accept help but do
not reciprocate can be detected and perhaps removed from future interactions. This discourages
'cheating' and encourages the continuation of reciprocal generosity.
Stability of Social Structure: Reciprocal altruism is more stable in communities where people
interact repeatedly throughout time. It enables the formation of societal norms and structures that
encourage the practice of providing and receiving assistance.

b. Do you consider reciprocal altruism to be a selfless behavior? Why or why not?

In reciprocal altruism, one person assists another in the idea that the favour would be
reciprocated in the future. While the act of helping may appear altruistic in the near term, it is
done with the anticipation of future benefits. The anticipation of reciprocation infuses self-
interest into the behaviour.

As a result, while reciprocal altruism contains behaviours that benefit others, it is not fully
selfless because it includes a component of self-benefit. The individual is willing to pay a current
expenditure in exchange for a future reward. This is what distinguishes reciprocal altruism from
pure altruism, in which an individual pays a cost to help others with no prospect of direct
reciprocation.

3. What new questions do you have now about apparently altruistic behavior in animals?

How does the balance of individual and group advantages affect the evolution of altruistic
behaviours across species?
How do environmental factors like predation pressure and resource availability influence the
evolution and manifestation of altruistic behaviours?

Part 5: Assessment

1. Imagine that you’re talking with a friend who has just watched a nature documentary about
meerkats. Your friend says, “Meerkats are so cute! They even protect each other by taking turns
looking out for predators so that the whole group stays safe!” Respond to your friend by
explaining the problem with this summary, based on what you’ve learned in the meerkat lesson.

Meerkats are wonderful creatures, and their sentinel behaviour is very astonishing! However,
while this behaviour benefits the group, it is not fully correct to claim that meerkats take turns
seeking for predators only for the group's safety.

The sentinel behaviour can also bring direct benefits to the individual meerkat that plays the
function. For example, being the first to see a predator gives the sentinel the benefit of
commencing the escape response, potentially enhancing its odds of survival. Furthermore, if
sentinel behaviour is connected with higher social rank within the group, it may lead to more
mating opportunities.

Furthermore, the emergence of such behaviours is frequently fueled by a combination of

individual and collective benefits, rather than only group benefits. This is a typical
misunderstanding when reading animal behaviour.
2. Use a key piece of evidence from the figures in the meerkat case to justify the role of natural
selection in the evolution of meerkat sentinel behavior.
One crucial piece of evidence from the figures in the meerkat case is the discovery that meerkats
on guard duty were discovered to be closer to bolt-holes (escape routes) than those foraging. This
shows that sentinel behaviour may have originated as a survival strategy through natural
selection.

Being closer to a bolthole minimizes a meerkat's predation risk, giving the sentinel a direct
survival advantage. Meerkats who exhibit this behaviour would have a greater survival rate over
time, making them more likely to pass on their genes to the next generation, resulting in the
evolution of sentinel behaviour.

This research backs up the role of natural selection at the individual level in the evolution of
meerkat sentinel behaviour. It demonstrates that, while sentinel behaviour benefits the group, it
can also provide direct benefits to the individual meerkat that plays the position, which is a
crucial driver of natural selection.

3. In meerkats, another behavior that they exhibit is helping care for young. Adult helpers remain
in the colony for a few years, feeding and caring for the young of the dominant female before
having any of their own offspring. Highlight the letter below (i.e., a, b, c, d) of any hypotheses
about helping behavior that are based on group selection.

a. Helping allows the non-breeding adults to learn how to care for young before having their
own.

b. Caring for the young of the dominant female ensures that more young survive, improving the
genetic quality and size of the colony.

c. Non-breeding adults spend more time foraging if they are collecting food for young, and
occasionally when the young are full the helpers get to eat the excess food themselves.

d. Adult helpers are treated less aggressively than non helpers by the dominant breeders in the
colony.

4. In another study, biologists studied how adult helper behavior differed depending on whether
pups of the dominant female were present in the foraging group or stayed behind at the burrow.
Bipedal vigilance refers to helpers standing on their hind legs to look for predators.

a. Summarize the findings presented in Figure 5.

Bipedal Vigilance (Graph A): Adult helpers were found to execute bipedal vigilance much more
frequently when pups were present in the foraging group than when pups were at the burrow.
This shows that having pups in the group alters the helpers' attentiveness behaviour.
Time Foraging (Graph C): Adult helpers' time spent foraging was not significantly different
whether the pups were in the group or at the burrow. This suggests that the presence of pups has
no substantial effect on the helpers' foraging behaviour.

Amount of Food discovered (Graph E): Helpers discovered somewhat more food when pups
were in the burrow, but the difference was not statistically significant. This shows that the
presence or absence of pups in the foraging group has no meaningful effect on how much food
the helpers find.

b. Based on what you see here, and what you know about meerkat fitness and selection, describe
what must be true about the fitness costs and benefits of guarding behavior for female helper
meerkats.
Fitness Benefits:

Increased Survival: The fact that helpers exhibit greater bipedal alertness when pups are present
shows that this behaviour may increase the helpers' survival rate, as they are more watchful and
hence more likely to detect predators early.
Potential Reproductive Benefits: If the helpers are related to the pups they are guarding (as is
frequently the case in meerkat groups), assisting in the survival of these pups can result in
indirect fitness benefits via kin selection. Helping their relatives' offspring survive allows them
to pass on shared genes to the next generation.

Fitness Costs:

Energy Expenditure: Protective behaviour, particularly bipedal vigilance, requires energy. This
may reduce the energy available for other activities, like as foraging or reproduction.
Reduced Foraging Time: The time spent on guard duty cannot be used to forage for food.
However, the data imply that the presence of pups has no major impact on the helpers' foraging
time.