UNIT 7

ALTRUISM

Structure
7.1 Introduction 7.5 Kin Selection
Objectives 7.6 Inclusive Fitness
7.2 Historical Background of 7.7 Reciprocal Altruism
Altruism 7.8 Summary
7.3 Examples of Altruistic 7.9 Terminal Questions
Behaviour
7.10 Answers
7.4 Group Selection

7.1 INTRODUCTION
In the previous unit, we discussed the social behaviour of animals. Animals
that live in groups show a great deal of cooperation amongst themselves.
Cooperation is beneficial to all the individuals living in the group. On the other
hand, there are many behaviours that benefit the recipient at the cost of the
individual who is performing the behaviour. This phenomenon, known as
altruism, is the topic of the present unit. We will begin with a brief historical
background of altruism. We will then proceed with a few interesting examples
of altruistic behaviours. A bit later in this unit, we will delve into the scientific
theories that seek to explain the evolution of altruism. With this aim, we will
first discuss the group selection theory, along with its merits and demerits.
Next, we will get acquainted with the postulates of the kin selection theory.
We will also take a closer look at some of the evidence that supports the kin
selection theory. This will be followed by a brief introduction to the concept of
inclusive fitness. Finally, we will deal with the concept of reciprocal altruism
which involves mutually beneficial interactions amongst unrelated individuals.

In the next unit of this course, we will learn about the wide variety of mating
behaviours among different animal species.

Objectives
After studying this unit, you would be able to:

 develop a better understanding of the term ‘altruism’,

 gain an insight into the historical context of altruism,

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 enumerate a few examples of altruistic behaviours,

 explain the postulates of group selection theory,

 differentiate between group selection and kin selection theories,

 comprehend the concept of inclusive fitness,

 describe why animals show altruistic behaviour towards others,

 appreciate the conditions that must be met for animals to show

reciprocal altruism towards unrelated individuals, and

 compare altruism with reciprocal altruism.

7.2 HISTORICAL BACKGROUD OF ALTRUISM

The term altruism was coined by Auguste Comte in 1858. It is derived from
the French word autrui which means other people. In a literal sense, altruism
refers to a selfless concern for the welfare of other people. Altruism can be
divided into three major categories: psychological, ethical and biological
altruism. Psychological altruism refers to intentionally helping someone.
Ethical altruism refers to the principle that one’s actions should be aimed at
bringing happiness to others and that individuals had a moral obligation to
renounce self interest and show love for all others. Biological altruism or
evolutionary altruism refers to a behaviour or an act that provides benefit to
the recipient(s) at a significant cost to the donor. In other words, biological
altruism involves the loss of direct fitness of the donor (or altruist). From the
perspective of animal behaviour, only biological altruism is relevant. Therefore,
we shall focus the rest of our discussion on biological altruism and for the sake
of simplification, we shall use the term altruism to refer to biological altruism.

In his famous book, On the Origin of Species, published in 1859, Charles

Darwin suggested that individuals must compete for the limited resources in
the environment. Therefore, the idea of helping someone else at a significant
cost to oneself clashed with the 19th century view of natural selection. Darwin
himself struggled with reconciling the idea of altruism with his theory of
evolution. Darwin reasoned that if a noble individual sacrificed his life for the
greater good of his fellow men, then he would not have any offspring and thus
he could not transfer this trait to his offspring. Thus, altruistic traits should
disappear over the course of evolution.

According to the classical approach to natural selection, the altruistic

phenotype seems to have a considerably reduced fitness, from an
evolutionary viewpoint. On the other hand, the selfish phenotype should have
a much higher fitness and thus should easily outcompete the altruistic
phenotype. Eventually, the altruistic gene should get eliminated over the
course of evolution. However, this does not happen. In fact, we frequently
come across examples of altruism in many animal groups. This scientific
puzzle came to be known as the altruism paradox. Ethologists have come up
with two main theories to resolve this paradox by explaining the evolution of
altruism. The first one is the group selection theory which was supported by
162 V.C. Wynne-Edwards, Konrad Lorenz and D.S. Wilson. The second theory is
Unit 7 Altruism
known as the kin selection theory which was supported by R.A. Fisher,
J.B.S. Haldane, W. D. Hamilton and J.M. Smith. In the later sections of this
unit, we will discuss both these theories pertaining to the evolution of altruism.

7.3 EXAMPLES OF ALTRUISTIC BEHAVIOUR

In the previous section, you were introduced to the phenomenon called
altruism. The animal kingdom provides many fascinating examples of
behavioural traits that are altruistic in nature. Therefore, in order to achieve a
better grasp of this interesting phenomenon, let us look at a few examples of
altruism:

• One of the most convincing examples of altruism is provided by parental

care. Parental care refers to a wide variety of acts that are performed by
parents in order to ensure the offspring’s survival. This may include nest
building, feeding the offspring and defending the offspring from
predators. For example, sperm whales (Fig. 7.1) show extensive
parental care with a 14 to16 month long gestation period and a 14 to 42
month long lactation period. We will discuss parental care in greater
detail in unit 9 of this course.

Fig. 7.1: A mother sperm whale with her calf

• Another superb example of altruism is provided by social insects such as

Conspecific refers to
honey bees (Fig. 7.2), ants, termites and wasps. They show a special
the individual from the
kind of altruism in the sense that the worker castes are sterile and same species.
sacrifice their own reproduction (direct fitness) in favour of the
reproductive caste. This phenomenon of sacrificing one’s reproduction
for the benefit of conspecific individual(s) is known as reproductive
altruism. Besides foregoing their own reproduction and helping the
queen raise its offspring, the worker bees perform yet another altruistic
act. Upon seeing an intruder approach the hive, they sting the intruder
as a means of protecting the nest. The workers sacrifice their lives in this
process since they die after stinging an intruder.

Fig. 7.2: A beehive having a colony of honey bees 163

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• Around 200 species of birds, including pied kingfishers (Fig. 7.3), house
sparrow, acorn woodpecker, etc., have been known to show the
‘helpers at the nest’ phenomenon. In this type of breeding system, the
older siblings stay with their parents through at least one breeding cycle
and help them raise the younger siblings at a cost to the helper’s own
lifetime reproductive success and its compensation through benefits
from relatives other than their own offspring. These helpers invest as
much as the breeders with respect to guarding the nests and feeding the
young ones and this investment (measured in energy expenditure) affect
the helper birds’ future to such an extent that they have a lower chance
of survival and mating. This breeding system is also seen in a species of
Fig. 7.3: A female beavers (Castor canadensis), and some other mammalian species
pied kingfisher belonging to the families Canidae (dog family) and Callithridae
(marmosets and tamarins).

• A particularly interesting example of altruism is the alarm calling

behaviour of Belding’s ground squirrel. When a predator approaches
their group, the Belding’s ground squirrel is known to react by standing
on its hind legs and giving a high-pitched alarm call. This alerts the
group members about the approaching predator, thereby saving their
lives. However, this behaviour comes at a significant cost to the squirrel
who makes the alarm call because the high-pitched sound renders them
very conspicuous to the predator. Vervet monkeys also give out alarm
calls to alert others about an approaching predator. Surprisingly, they
have distinct alarm calls for different types of predators such as
leopards, eagles, pythons, baboons, etc.

• Meerkats live in packs (Fig. 7.4) and show guarding behaviour. One
meerkat stands guard against the predators while the rest of pack is
feeding. Another type of altruism exhibited by meerkats pertains to the
social hierarchy within the pack. The dominant meerkats reproduce while
the subordinate meerkats care for the pups.

Fig. 7.4: A pack of meerkats

• Humans reside in family groups and show a great deal of altruism

towards their kin. They also often behave altruistically towards strangers
(non kin) with no chance of reciprocation. Many people donate blood
and funds for the benefit of people they will never meet. (Please refer to
unit 10 of this module for a detailed account of altruism in the context of
164 human families).
Unit 7 Altruism

7.4 GROUP SELECTION

Group selection theory hypothesized that altruism evolved because natural
selection occurs at the level of the group rather than the individual. Thus, the
theory posits that the behaviours that benefit the whole group, are favored by
natural selection even if an individual must pay a heavy price for the same.
This theory argues that even though the altruist’s fitness gets reduced, the
fitness of the group with an altruist is higher than that of the group with no
altruists. Typically the group under selection is a small cohesive social unit and
members’ interactions are of altruistic nature.

It appears that Charles Darwin was one of the earliest proponents of the group
selection theory. This is because in his book The Descent of Man, published in
1871, Charles Darwin presented a hypothetical scenario in which two tribes of
prehistoric humans were competing with each other. Darwin concluded that
the tribe which would emerge as a winner would be the one whose individuals
warned each other against danger and defended each other against the
enemy group. Thus, the group with altruistic individuals will win against the
other group. However, with the rise of neo Darwinism in the middle of 20th
century, in which Darwin’s theory was synthesized with genetics, the idea that
selection acted on groups was largely dismissed.

In 1962, group selection concept again reemerged when V. C. Wynne-

Edwards formally introduced the group selection theory. Subsequently, Konrad
Lorenz and D. S. Wilson also provided their support for the same. Wynne-
Edwards gave the example of red grouse birds (Fig. 7.5) to substantiate his
claims regarding group selection. He observed that when their population
density increases the birds curtail their reproduction. According to Wynne-
Edwards, the birds do this for the benefit of the entire group because
excessive population would exhaust their food supplies. However, later studies
revealed that the reduced reproduction rate at high densities was simply a
physiological response of each bird in response to the scarcity of food supply.

Fig. 7.5: A male red grouse

The major advantage of the group selection theory is that it could explain how
some of the most drastic transitions in the evolutionary history of life on the
planet were brought about. For example, this concept of ‘collective good’ could
explain why individual’s genes got together to form a chromosome. By the
same logic, it could also shed some light on the evolution of multicellularity
and social groups. 165
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The group selection theory does have merits but it also suffers from some
major flaws. For example, the basic premise of group selection is that altruism
evolved because it increases the fitness of the whole group. Now let us
assume that we have a population in which all the individuals are altruists.
This system will work quite smoothly for a while because it does seem to be
beneficial to all. However, such a system would crumble if a mutation occurred
which made the individual behave selfishly. Now, the altruists of the group will
keep sacrificing while the selfish individual will keep reaping the benefits. In
the long run, this would lead to a reduction in the fitness of altruists and a
concomitant increase in the fitness of the selfish individual. Thus, the
generous altruists would leave fewer offspring while the selfish individual
would leave much more offspring. Over a few generations, the selfish
individuals would outcompete the altruists, ultimately leading to the eradication
of altruism. In order to counteract this, the supporters of group selection
propose that this can be avoided if there are mechanisms within the group that
keep the selfish phenotype in check. For example, there could be rewards for
altruistic behaviour and punishments for selfish behaviour. However, one
cannot deny the possibility that even this system would be prone to cheaters
with the ‘selfish mutation’. This means that in the group selection model,
altruism is bound to be replaced by selfish mutations.

Unfortunately, at present, there is very little evidence that supports group

selection theory as a plausible explanation for evolution of altruism. However,
as inconsistent as group selection theory seems to be, it does have its merits.
Therefore, one must be cautious before disregarding it completely.

SAQ 1
a) Give any four examples of altruistic behaviours.

b) Briefly explain the limitations of the group selection theory.

7.5 KIN SELECTION

In the last section, we saw how group selection theory fails to answer many of
the questions regarding the evolution of altruism. This led to a quest for
alternative theories. Eventually, in 1964, John Maynard Smith coined the term
kin selection for R.A. Fisher and J.B.S. Haldane’s theory which offered one of
the best possible explanations for the evolution of altruism.

Kin selection posits that altruistic behaviours evolve because by helping its
relatives (or kin) increase their fitness (survival and/or reproduction), an
individual ascertains the transmission of the genes that it shares with kin to the
next generation. Therefore, the idea of kin selection is very different from that
of group selection. Contrary to group selection, kin selection seeks to explain
the evolution of altruism using a gene centric approach. Thus, in the
evolutionary context, selection occurs at the level of the gene. Therefore, any
behaviour that leads to an increase of frequency of genes that a particular
individual carries, will be selected even if this would lead to the sacrifice of the
166 individual in favour of its kin. Kin selection is a mechanism of natural selection
 Unit 7 Altruism
where instead of selecting the fittest indidividual to thrive and reproduce,
selects relatives of altruistic individuals to thrive and reproduce yet ensuring
that many of the genes of the altruistic individual still gets passed on to the
future generation through their relative.

For kin selection to work, it is imperative that the altruists be able to

differentiate between kin and non-kin. This poses two very pertinent questions.
How do the altruists make sure that their behaviour is directed specifically
towards the kin rather than an unrelated individual(s)? Also, do the altruists
need sophisticated neural networks and cognitive abilities to do so? William
Donald Hamilton provided answers to both these questions by positing that
there are two mechanisms which can ensure the preferential treatment of the
altruist’s kin. These mechanisms are:

(i) Kin discrimination: This is the ability to recognize relatives. It may

involve visual, auditory or olfactory senses. Therefore, kin recognition
requires a high degree of cognition and thus only those species that
have well developed cognitive abilities can use this mechanism. For
example, studies have indicated that humans have very elaborate
o kinship recognition circuits which are very effective at recognizing kin
and estimating the degrees of kinship (For details, please refer to Unit 10
of this course).

(ii) Population viscosity: This term refers to the fact that for many species,
the dispersal is very limited. Therefore, all the neighbours are their kin.
This mechanism does not require advanced cognitive abilities and thus
can be used by the simplest of organisms. Interestingly, it has been
observed that the species that show low mobility, and hence low
dispersal, show more altruism when compared to highly mobile species.

A rare but related phenomenon is called the green-beard effect, which posits
that if the carrier of a gene can recognize other individuals carrying the same
gene, then there is a high probability that an altruism might evolve in these
circumstances. Richard Dawkins tried to explain this effect in simpler terms in
his book entitled, The Selfish Gene, published in 1976. He used the analogy of
a green beard as a visible phenotype and proposed that altruism could evolve
if the individuals carrying the green beard phenotype are altruistic towards
other people with a green beard. In this case, phenotypic similarity between
individuals is more important than genetic relatedness. Many ethologists
consider the green-beard effect to be the third mechanism of kin selection (the
first two being kin discrimination and population viscosity). Although strictly
speaking, this is not selection of kin, but rather of a gene (or phenotype) only,
because in case of green-beard effect, the altruist need not be related to the
recipient.

Some of the most common examples of kin selection are provided by

behavioural traits such as parental care, helpers at the nest, suppressed
reproduction, alarm calling and guarding behaviour.

The proponents of kin selection claim that almost all types of altruistic
behaviours have kin selection at their core. Let us test this claim by revisiting
some examples of altruism that we discussed in Section 7.3 and determining
whether kin selection is involved in these cases. 167
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• Kin selection can very obviously account for all types of altruistic
behaviours pertaining to parental care.

• Consider the altruism shown by social insects like honeybees. Given that
the queen is the mother of the workers, and therefore, all its offspring are
sisters to the workers, this altruism can be very easily attributed to kin
selection. By helping their mother and sisters, the honeybees are
ensuring that more and more of their genes are passed on to the new
generation.

• You will recall from previous sections that the helpers at the nest
phenomenon is a special breeding system observed in some birds. The
older siblings forego their own reproduction for at least one breeding
season and help their parents take care of their younger siblings. This is
usually seen in monogamous birds. Kin selection can easily explain why
this phenomenon is correlated to monogamous species. Monogamy
ensures that the new brood will share both the parents with the older
siblings. On the other hand, in a polygamous system, there is a high
chance that one of parents would be different between successive
broods. Thus, the number of shared genes between the subsequent
broods would be reduced to half of that seen in monogamy. Hence, in a
monogamous mating system, the older siblings share more genes with
the newer siblings compared to a polygamous mating system. Therefore,
it makes sense that the helpers at the nest type of breeding system is
seen mostly in monogamous birds.

• Let us now take up the case of the alarm calling behaviour of Belding’s
ground squirrel (Fig. 7.6). The caller alerts the nearby squirrels against
the approaching predator at the cost of drawing the predator’s attention
to itself. Curiously, alarm calls are mostly given by female squirrels!
Even this seemingly bizarre observation can be explained by kin
selection. A closer examination of the social situation of the squirrels
reveals that related females occupy neighbouring burrows while the
males are more likely to move away to a different neighbourhood. This
means that for the females, most of the neighbours are their sisters,
daughters or nieces. Therefore, it makes more sense for it to risk its life
for its neighbours since they are its kin. On the other hand, the males of
the species are not closely related to their neighbours and thus they are
less likely to risk their lives by exhibiting the alarm calling behaviour.
Hence, kin selection can account for the fact that the female ground
squirrels show the alarm calling behaviour more often than the males of
the species.

Even though the theory of kin selection is very effective at providing

explanations for the evolution of altruism, it does have a few limitations. By
its very definition, it provides a strong basis for altruism that is exhibited by
an animal in favour of its own relatives. However, it fails to account for the
Fig. 7.6: A Belding's presence of some highly specialized animal societies which show a proper
ground squirrel division of labour and cooperation even though the individual members are
not necessarily related to each other. Moreover, the popularity of kin
selection sometimes biases ethologists towards overestimating the indirect
fitness benefits of a particular behaviour. In principle, this could lead to
overlooking of the direct fitness benefits, which would provide a much
168 simpler and accurate explanation for the behaviour.
Unit 7 Altruism

7.6 INCLUSIVE FITNESS

As discussed in the previous Section, kin selection posits that individuals can
assure the transmission of shared genes to the subsequent generation by
helping their kin. In 1964, William Donald Hamilton sought the mathematical
model to explain how helping its kin could potentially increase the altruist’s
own fitness. He proposed that an organism’s overall fitness, which he called
inclusive fitness, is much more relevant in the evolutionary context.
According to Hamilton:

Inclusive fitness = Direct fitness + Indirect fitness

Direct fitness of an individual is a measure of the direct transfer of its genes

to its own offspring. Indirect fitness of the individual is a measure of the
increase of the reproductive fitness of the recipient due to the donor’s altruism.

Natural selection favours an altruistic behaviour if it leads to a significant

increase in the indirect fitness of the individual. This is because by increasing
the indirect fitness, the altruistic behaviour is ensuring the transmission of the
shared genes to the next generation through the genetically related recipient.
Thus, Hamilton’s explanation for the evolution of altruism is gene centric since
the transmission of an organism’s genes to the next generation’s gene pool is
more important than the survival or reproduction of that particular organism.
Eventually, he proposed what is now known as Hamilton’s rule in his honor.
According to this rule, altruism evolves if the indirect fitness gain is greater
than the cost incurred by performing the altruistic act. Hamilton’s rule can
mathematically be represented as the inequality given below:

rB > C

Here, r represents the coefficient of relatedness which is a measure of the

genetic relatedness between the altruist and the recipient; B represents the
benefit or the gain in the recipient’s direct fitness; and C represents the cost or
the loss of the altruist’s direct fitness (survival &/or reproduction).

The left side of the above mentioned inequality is rB, which is the product of
genetic relatedness and the gain in the recipient’s direct fitness. It is thus a
measure of the total indirect fitness gain for the altruist. So, an altruistic
behaviour would evolve if the total indirect fitness gain (rB) due to the altruism
turns out to be more than the cost (C) incurred while performing it. Hamilton
argued that in such a scenario, the altruistic behaviour is increasing the overall
or inclusive fitness of the individual. Thus, he was able to demonstrate
mathematically how it is possible for an altruistic behaviour to be favored by
natural selection even though it leads to heavy losses in the individual’s direct
fitness.

A high coefficient of relatedness (r) between the altruist and the recipient,
means that the gain in the indirect fitness (rB) is higher too. Thus, Hamilton’s
rule provides a mathematical proof to the phenomenon that individuals are
more likely to be altruistic towards their kin rather than unrelated conspecific or
non-conspecific animals.

Now let us understand how the coefficient of relatedness (r) is calculated

between full siblings. In order to do so, we need to calculate the probability
that siblings 1 and 2 will inherit allele X from either their mother or father. 169
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Probability that sibling 1 will inherit allele X from father = ½

Probability that sibling 2 will inherit allele X from father = ½

Probability that siblings 1 & 2 will inherit allele X from father = ½ X ½ = ¼

Probability that sibling 1 will inherit allele X from mother = ½

Probability that sibling 2 will inherit allele X from mother = ½

Probability that siblings 1 & 2 will inherit allele X from mother = ½ X ½ = ¼

Probability that siblings 1 & 2 will inherit allele X from either parent = ¼ + ¼ =
½

Thus, r between siblings = ½ or 0.5

Similarly, we can calculate the coefficient of relatedness (r) for other relatives
too. Table 7.1 shows r of an individual with various relatives.

Table 7.1: Coefficients of relatedness (r) of an individual with various

relatives. Note that the r for self is 1.

Relative Coefficient of relatedness (r)

Identical twin 1.0

Parent 0.5

Offspring 0.5

Full Sibling 0.5

Half Sibling 0.25

Grandparents 0.25

Grandchild 0.25

Aunt/Uncle 0.25

Niece/Nephew 0.25

First Cousin 0.125

SAQ 2
a) What are two mechanisms which can ensure the preferential treatment
of the altruist’s kin?

b) What is the coefficient of relatedness of an individual with its (i) identical

twin, (ii) aunt, and (iii) nephew?

7.7 RECIPROCAL ALTRUISM

The previous sections of this unit have convinced us that kin selection offers a
170 very plausible explanation for the evolution of altruism between conspecific
Unit 7 Altruism
animals. But if kin selection was the only mechanism for altruism to have
evolved, then we should never observe altruism between unrelated animals let
alone animals belonging to different species. This is because animals from
different species are not kin to each other. In order to explain altruism between
unrelated animals, Robert Trivers introduced the theory of reciprocal altruism
in 1971.

Reciprocal altruism can be defined as a behaviour through which the donor

increases the recipient’s fitness at the cost of its own fitness, all the while
expecting to get a subsequent payback or repayment from the recipient in the
future. Thus, reciprocal altruism differs from altruism in the sense that the
recipient is expected to reciprocate later. In other words, the recipient must
return the favour to the donor whose behaviour is altruistic in the short-term
only. Therefore, in contrast to altruism wherein the altruist’s reduction in fitness
is permanent, it is temporary in case of reciprocal altruism. Hence, Trivers
argued that since both the recipient, and subsequently the donor, get
benefitted in this process, their individual fitness increases. Over many
subsequent generations, such a reciprocity would be favored by natural
selection. This, in turn, could explain why we see altruism between two
unrelated or non-conspecific animals.

It is noteworthy that reciprocal altruism does not evolve randomly between

unrelated individuals. For reciprocal altruism to evolve between two animals,
certain conditions need to be met. Ideal conditions increase the probability of
evolution of reciprocal altruism. These conditions are:

a) The prospective reciprocal altruists must have many opportunities for

reciprocation. The association between the prospective reciprocal
altruists must be long or should involve repeated encounters between
them. This would increase the possibility that the donor will receive the
benefit when the recipient reciprocates the favour. This is an essential
precondition for evolution of reciprocal altruism as the association needs
to be beneficial for both the animals in the long term. On the contrary,
reciprocal altruism is very unlikely to evolve between individuals whose
associations with each other are short or involve very infrequent contact
with each other. This makes perfect logical sense. If the prospective
altruists are only going to meet once in their lifetime, then none of them
has the incentive to help the other and therefore reciprocal altruism
would not evolve between them.

b) The prospective reciprocal altruists must have sufficient cognitive

abilities to recognize each other at an individual level. This is important
because in order to return the favour, the recipient needs to be able to
identify the donor as a specific individual. This will ensure that the favour
is returned to the correct individual. Moreover, if this reciprocity is to be
extended to multiple encounters, then the initial donor must also
recognize the initial recipient who has reciprocated the favour during
their second encounter. This would ensure long term associations which
are crucial for the evolution of reciprocal altruism.

c) The prospective reciprocal altruists must remember the favour and their
obligations in relation to that. Just as in previous case, this requires 171
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sufficient memory in both the animals, so that they can have multiple
encounters. The long-term associations thus produced will eventually
serve to increase the fitness of both the animals. Consequently, there
will be a high probability of evolution of reciprocal altruism.

d) The prospective reciprocal altruists must have a mechanism to detect

and possibly punish the ‘cheaters.’ This would ensure that the cheaters
(or non-altruists) are not able to exploit the altruists and increase their
own fitness at the cost of the altruists. This mechanism is essential for
the behaviour to evolve into reciprocal altruism. However, in some cases
of reciprocal altruism, the exact mechanism through which cheaters are
detected and punished has not been discovered yet.

Now that we have developed a clear understanding of the concept of

reciprocal altruism, let us enumerate a few examples of reciprocal altruism.
Humans show a lot of behaviours that would qualify as reciprocal altruism.
However, the examples for reciprocal altruism are rarer in case of other
animals. Some of these examples are:

• Birds give an alarm call to alert all the nearby birds about an
approaching predator. They do so at a great cost to themselves. By
raising the alarm, they catch the predator’s attention and may even get
attacked by it, thereby decreasing their own fitness in the process.
However, this risky behaviour is useful for all the nearby birds many of
whom may be unrelated or even non-conspecific to the bird giving the
alarm call. It is easy to see how this behaviour qualifies as reciprocal
altruism since there may be another instance where our altruistic bird
gets benefitted because yet another bird raised a call upon seeing a
predator and alerted the initial altruist about the impending danger.

• Cleaner fish have a cleaning symbiosis with the host fish (Fig. 7.7).
However, this interaction also has elements of reciprocal altruism in it.
For example, the host fish allows the cleaner fish safe entry into its
mouth. Even after the cleaning has been done, the host fish allows the
cleaner to exit its mouth rather than swallowing it. Moreover, the host
fish has been shown to defend the cleaner fish against predators.
Interestingly, there are repeated interactions between both the fish.

Fig. 7.7: A white-spotted pufferfish (host) getting cleaned by a Hawaiian cleaner

172 wrasse
Unit 7 Altruism
• Red winged blackbird males (Fig. 7.8) perform cooperative nest defence
by mobbing predators such as crows. However, the male blackbirds only
cooperate with those males who have defended their nests in the past.
Thus, if the neighbouring male turns out to be a cheater and does not
help them defend their nest, then in the future, they will also not defend
the cheating neighbour’s nest.

Fig. 7.8: A male red winged blackbird

• Grooming in primates is another example of reciprocal altruism. A

particularly interesting example is presented by vervet monkeys (Fig.
7.9). These monkeys groom their kin as well as unrelated conspecifics.
Moreover, in case a monkey calls for help, the other monkeys are more
likely to help if they share a grooming based reciprocal altruistic
relationship with them.

Fig. 7.9: Two vervet monkeys. The one on the left is grooming the one on the
right

• The common vampire bat feeds on mammalian blood and lives in

colonies (Fig. 7.10). It can only survive for two nights without a blood
meal. Unfortunately, there is always a chance that it may not have been
able to get any feeding throughout the night. Each night that it does not
feed, it loses quite a bit of its body weight. In such a scenario, the other
bats of the colony will feed it by regurgitating blood, thereby saving its
life. Interestingly, studies have indicated that the donors are more likely
to share blood with those recipients who had shared with them on a
previous occasion! 173
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Fig. 7.10: A colony of vampire bats

Although the examples given above have been cited as examples of reciprocal
altruism, we must remember that kin selection is still a key component here.
All the animals in the aforementioned examples are much more likely to help
their own kin before helping an unrelated individual.

Reciprocal altruism involves short-term loss and long-term gain, thereby

increasing the direct fitness of the altruists too. Moreover, since reciprocal
altruism is a kind of win-win mutualism wherein all the participants benefit, this
is a selfish cooperation. As a result, it is effectively long-term self-interest
rather than genuine sacrifice. Therefore, many scientists including Hamilton
believe that reciprocal altruism is not true altruism but Delayed Return
Altruism. Rothstein and Pierotti in 1988 proposed the term pseudo reciprocity
in place of Delayed Return Altruism. Instead, they argue that this ‘reciprocity’
ultimately results in the evolution of non-altruistic cooperation wherein both the
donor and the recipient get benefitted.

SAQ 3
a) What are four conditions that increase the probability of evolution of
reciprocal altruism?

b) Give any three examples of reciprocal altruism.

7.8 SUMMARY
In this Unit you have learnt that:

• Biological altruism or evolutionary altruism refers to a behaviour or an

act that provides benefit to the recipient(s) at a significant cost to the
donor.

• Examples of altruistic behaviours include parental care, reproductive

altruism in honeybees, alarm calling behaviour, helpers at the nest,
guarding behaviour, etc.

• The phenomenon of sacrificing one’s reproduction for the benefit of

174 conspecific individual(s) is known as reproductive altruism.
Unit 7 Altruism
• In the ‘helpers at the nest’ type of breeding system, the older siblings
stay with their parents through at least one breeding cycle and help them
raise the younger siblings.

• Group selection theory hypothesized that altruism evolved because

natural selection occurs at the level of the group rather than the
individual.

• Kin selection posits that individuals can assure the transmission of

shared genes to the subsequent generation by helping their kin.

• Kin discrimination and population viscosity are two main mechanisms

which can ensure the preferential treatment of the altruist’s kin.

• An organism’s overall fitness or inclusive fitness is calculated by adding

its indirect fitness to its direct fitness.

• According to Hamilton’s rule, altruism evolves if the indirect fitness gain

is greater than the cost incurred upon performing the altruism.

• Reciprocal altruism can be defined as a behaviour through which the

donor increases the recipient’s fitness at the cost of its own fitness, all
the while expecting to get a subsequent payback from the recipient.

• In case of altruism, the actor’s reduction in fitness is permanent, while in

case of reciprocal altruism, it is temporary.

7.9 TERMINAL QUESTIONS

1. Define the following terms:

a) Biological altruism

b) Green-beard effect

c) Coefficient of relatedness

d) Reciprocal altruism

2. Indicate whether the following statements are ‘True’ or ‘False’:

a) In a polygamous mating system, the older siblings share more

genes with the newer siblings compared to a monogamous mating
system.

b) Direct fitness of an individual is the direct transfer of its genes to its

own offspring.

c) Coefficient of relatedness (r) of an individual with self is 1.

d) An organism’s inclusive fitness is the measure of its indirect fitness

only.

e) In case of reciprocal altruism, the actor’s reduction in fitness is

permanent, while in case of altruism, it is temporary. 175
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
3. Fill in the blanks.

a) According to ...................... rule, altruism evolves if the indirect

fitness gain is greater than the cost incurred by performing the
altruistic act.

b) ...................... fitness of the individual is a measure of the increase

of the reproductive fitness of the recipient due to the altruistic act.

c) The phenomenon of sacrificing one’s reproduction for the benefit

of conspecific individual(s) is known as ...................... .

d) In the ...................... type of breeding system, the older siblings

stay with their parents through at least one breeding cycle and
help them raise the younger siblings.

Activity

Search the internet for videos with animals displaying altruistic behaviour. Pick
your favourite one and write a short note on it.

……………………………………………………………………………..……………
…..………………………………………………………………………………..…..…
……..………………………………………………………………..……………..……
………..…………………………………………………………..…………..…………
…………..……………………………………………………..……………...………...
……………………………………………………………..……………………………

7.10 ANSWERS
Self-Assessment Questions
1. a) Examples of altruistic behaviours include parental care,
reproductive altruism in honeybees, alarm calling behaviour,
helpers at the nest, guarding behaviour, etc (any four).

b) A major limitation of group selection theory is that in this model,

altruism is bound to be replaced by selfish mutations. Moreover, at
present, there is very little evidence that supports the group
selection theory.

2. a) Kin discrimination and population viscosity.

b) Coefficient of relatedness (r) for:

(i) identical twin = 1

(ii) aunt = 0.25

(iii) nephew = 0.25

3. a) The prospective reciprocal altruists must

176 (i) have many opportunities for reciprocation.

Unit 7 Altruism
(ii) have sufficient cognitive abilities to recognize each other at
an individual level.

(iii) remember the favour and their obligations in relation to that.

(iv) have a mechanism to detect and possibly punish the

cheaters.

b) Examples of reciprocal altruism (any three):

(i) Birds give an alarm call to alert all the nearby birds about an
approaching predator.

(ii) The host fish defends the cleaner fish against predators.

(iii) Cooperative nest defence by male red winged blackbirds.

(iv) Grooming in primates.

(v) Blood meal sharing in vampire bats.

Terminal Questions
1. a) Biological altruism refers to a behaviour or an act that provides
benefit to the recipient(s) at a significant cost to the donor.

b) Green-beard effect is a theory which posits that if the carrier of a

gene can recognize other individuals carrying the same gene, then
there is a high probability that an altruism might evolve in these
circumstances.

c) Coefficient of relatedness is the proportion of shared genes

between the altruistic individual and the recipient.

d) Reciprocal altruism can be defined as a behaviour through which

the donor increases the recipient’s fitness at the cost of its own
fitness, all the while expecting to get a subsequent payback from
the recipient.

2. a) False, b) True, c) True, d) False, e) False.

3. a) Hamilton’s, b) Indirect, c) Reproductive altruism,

d) Helpers at the nest.

Acknowledgement of Figures
Fig. 7.1: [Link]

Fig. 7.2: [Link]

Fig. 7.3: [Link]

Fig. 7.4: [Link]

Fig. 7.5: [Link] 177

Block 2 Systematic Approach to Understand Important Animal and
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Fig. 7.6: [Link]

Table 7.1: Author

Fig. 7.7: [Link]

Fig. 7.8: [Link]

Fig. 7.9: [Link]

Fig. 7.10: [Link]

178