MZOE-002

ANIMAL BEHAVIOUR AND

Indira Gandhi ANIMAL WELFARE ETHICS
National Open University
School of Sciences

Block

2
SYSTEMATIC APPROACH TO UNDERSTAND
IMPORTANT ANIMAL AND HUMAN
BEHAVIOUR
UNIT 6
Social Behaviour and Altruism 135
UNIT 7
Altruism 161
UNIT 8
Mating Behaviour 179
UNIT 9
Parental Care 211
UNIT 10
Human Behaviour and its Regulating Factors 249
COURSE NAME: ANIMAL BEHAVIOUR AND ANIMAL COURSE CODE: MZOE-002
WELFARE ETHICS

Course Design Committee

Prof. Sujatha Varma Prof. Bharati Chauhan Prof. Amrita Nigam (Retd.)
Former Director, School of Department of Zoology School of Sciences, IGNOU
Sciences, IGNOU, Maidan Garhi University of Rajasthan Maidan Garhi, New Delhi-110068
New Delhi-110068 Jaipur-302004
Prof. Neera Kapoor
Prof. Neeta Sehgal Prof. Devinder Kaur Kochar School of Sciences, IGNOU
Department of Zoology Department of Zoology, Punjab Maidan Garhi, New Delhi-110068
University of Delhi, Delhi-110007 Agricultural University
Ludhiana-141004 Dr. Siya Ram
Prof. Sarita Kumar School of Sciences, IGNOU
Department of Zoology, Acharya Prof. Varsha Wankhade Maidan Garhi, New Delhi-110068
Narender Dev College, University Department of Zoology
of Delhi, Delhi-110007 Savitribai Phule University Dr. Ravi Rajwanshi
Pune-411007 School of Sciences, IGNOU
Prof. S. Dinakaran Maidan Garhi, New Delhi-110068
Department of Zoology Prof. Sarita Sachdeva
Madurai Kamaraj University Department of Biotechnology
Tamil Nadu-625021 Manav Rachna International
University, Sector-43, Faridabad
Haryana-121004

Block Preparation Team

Prof. Kamal Kumar Gupta Dr. Aparna Sharma School of Sciences
Department of Zoology Department of Zoology
Prof. Neera Kapoor
Deshbandhu College, Kalkaji Acharya Narendra Dev College
School of Sciences, IGNOU
Main Road, Block H, Kalkaji University of Delhi, Delhi-110019
Maidan Garhi, New Delhi-110068
New Delhi-110019 (Units 6 and 8) (Unit 9)
(Units 6, 7, 8, 9 and 10)
Dr. Sweety Shrimali
Department of Zoology
Acharya Narendra Dev College
University of Delhi, Delhi-110019
(Units 7 and 10)

Programme Coordinators : Prof. Neera Kapoor and Dr. Siya Ram

Course Coordinator : Prof. Neera Kapoor

Course Editor : Prof. Rita Rath
Department of Zoology, Dyal Singh College
University of Delhi, Delhi-110003

Production : Mr. Hemant Kumar

SO(P), MPDD, IGNOU
Acknowledgement:
• Prof. Neera Kapoor and Mr. Ajit Kumar, Suggestions for figures and Cover Design.
• Mr. Vikas Kumar, JAT for word processing and CRC preparation.

April, 2024
Indira Gandhi National Open University, 2024
ISBN: ..................................
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means, without permission in writing from Indira Gandhi National Open University.
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Printed and published on behalf of Indira Gandhi National Open University, New Delhi by the Registrar,
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 BLOCK 2: SYSTEMATIC APPROACH TO
UNDERSTAND IMPORTANT ANIMAL AND
HUMAN BEHAVIOURS
In Block 1, you have studied about what animal behaviour is, mechanism of animal
behaviour and its categories. You also learnt about different learned behaviours, behavioural
homeostatsis and their hormonal regulation.

Block 2 consists of five units. Unit 6 “Social Behaviour and Altruism” speaks about social
organisation in animals with special reference to that of insects. It also tells about
cooperation and inclusive fitness. Cooperation is beneficial to all the individuals living in the
group. But behaviour, like altruism benefits the recipient at the cost of the individual who is
performing the behaviour. Group selection and kin selectional theories explain the evolution
of altruism. These concepts are discussed in details in Unit 7 “Altruism”.

Unit 8 “Mating Behaviour” explains about the asymmetry of sex and its role in the attraction
and selection behaviour of potential mates. It further describes the sexual dimorphism and its
adaptive significance. Evolutionary advantages of sexual selection are also discussed in this
Unit.

Unit 9 “Parental Care” will create an inquisitiveness among you regarding concept of parental
care among animals by discussing in a very interesting manner. Cost and benefits of
parental care and parental manipulation are discussed at length.

Unit 10 “Human Behaviour and Its Regulating Factors” will explain the basic tenets of human
behaviour. Regulation of human behaviour by environmental and genetic factors and
hormonal regulation are discussed at length.

Objectives
After studying this block, you should be able to:

• appreciate social organization in different animals with special reference insects,

• gain an insight into altruisms and compare with reciprocal altruism,

• narrate evolutionary advantages of sexual selection, and types of sexual selections,

• describe reproductive fires and factors governing it,

• elucidate numerous parental care examples across animal phyla,

• describe brood parasitism,

• appreciate the genic, environmental and hormonal control of human behaviour, and

• explain the relevance of altruism in family relationships of humans.

133
134
 UNIT 6
SOCIAL BEHAVIOUR AND
ALTRUISM

Structure
6.1 Introduction 6.7 Polyphenism

Objectives 6.8 Polyethism

6.2 An Overview of Sociality in Caste Polyethism

Animal Systems
Age Polyethism
Solitary vs Gregarious Animals
6.9 Swarming
Aggregation, Sub-Social and
6.10 Caste Determination in
Social Behaviour
Honey Bee
Characteristics of Social
6.11 Communication in Honey
Animals
Bee
Social Behaviour of Lion
Foraging in Honeybee
6.3 Cooperation and Conflict in
Round Dance
Animals
Waggle Dance
Cooperation
Chemical Communication
Conflicts
Trophallaxis in Honey Bees
6.4 Altruism and Inclusive
Fitness 6.12 Summary

6.5 Social Behaviour in Insects 6.13 Terminal Questions

6.6 Social Organization in 6.14 Answers

Honey Bees

Caste System

Beehive

6.1 INTRODUCTION
All animals adopt different lifestyles to maximize their survival and
reproduction. Solitary animals live in isolation without interacting except for
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
reproduction with the individuals of their species. Very often the animals live in
groups with different levels of organization. They may aggregate to avail better
life opportunities. In the subsocial hierarchy, the interactions are more intimate
as the young ones are cared for in the colony. Social organization is the
highest form of organization; eusocial organization involves a caste
system, division of labour and a greater degree of cooperation among
the individuals of the colony. The colony is perennial and cohesive. The
members communicate with each other in different ways. This unit speaks
about sociality in animal systems with special reference to social behaviour in
insects. It also elaborates upon cooperation, altruism and inclusive fitness in
animals.

Objectives
After studying this unit, you should be able to

 distinguish between solitary and gregarious animals and aggregation,

subsocial and social organization,

 describe cooperation, conflict and altruism, inclusive fitness,

 characteristic features of social organization in animals,

 appreciate social organization in lion,

 describe characteristic features of social organization in insects, and

 discuss social organization in honey bee.

6.2 AN OVERVIEW OF SOCIALITY IN ANIMAL

SYSTEMS
6.2.1 Solitary vs Gregarious Animals

Solitary animals spend the majority of their life in isolation without interacting
with other individuals of their species. The only group interaction is for
reproduction i.e. mating. Examples of solitary animals are bear, black
rhinoceros, platypus, skunk, leopard, mole, koala, sloth, etc.

Gregarious animals (grex, meaning "herd) live in groups, with the individuals
of their species. They are outgoing and sociable. Living in a group imparts
selective advantage to the individuals such as protection from physical factors;
protection against predators; selection of most suitable mates; foraging and
procuring food; “richer” learning environment for young; aiding (or receiving
aid) offspring or other relatives; modifying their environment. An advanced
form of gregarious habit leads to society formation with division of labour
among individuals.

Living in groups has certain disadvantages such as increased

competition for resources; increased chance of spread of diseases and
136 parasites; interference with reproduction; infanticide, etc.
Unit 6 Social Behaviour and Altruism

SAQ 1
Fill in the blanks:

i) ………………… is a temporary association of the animals in response to

an environmental stimulation.

ii) Common examples of pheromones are sex pheromones, alarm

pheromones and ………………… pheromones.

iii) ………………… male gains preferred positions in mating arenas and is

more likely to be chosen by females in a social group.

iv) Mating relationship of both sexes with multiple partners is called as

……………… .

v) The lion pride is ………………… .

6.2.2 Aggregation, Sub-Social and Social Behaviour

Aggregation Behaviour – is a behaviour in which a group of organisms of the
same or different species live closely together for feeding, protection or
reproduction. Aggregation is a chance gathering governed mainly by external
factors only. The group is formed solely in response to a common source
rather than the intrinsic factors of the individuals. For example, insects
attracted to a source of light for mating, woodlice gathered in humid areas,
collection of fruit flies at rotten fruit, a swarm of water fleas gathered in food-
rich areas, herds of grazing zebra and antelopes, mixed flocks of birds
foraging for food, etc. Slime moulds also form aggregations in which separate
cells aggregate in the process of constructing their reproductive structure.
Aggregation is a temporary association of the animals in response to an
environmental stimulus.

Sub-Social Behaviour – Such type of behaviour is observed in animals that

care for their young ones but show no other sociality trait. These animals live
in family groups consisting of parents and immature offspring and are
characterized by brood defence by parents.

Social behaviour refers to the type of behaviour in which a group of

individuals of the same species live in a closed community and cooperate. The
cooperation extends beyond sexual and parental behaviour. An animal
society is a relatively permanent association in which individuals live due to
intrinsic factors. Animal social behaviour involves interactions among
individuals of the same species to live in a group, cooperate in sexual or
parental behaviour, protect the territory, get access to mates, and
communicate. It may range from the release of a pheromone by a female moth
to attract a potential male mate, or loud roar by male red deer to signal
dominance and keep other males away to most complex societies
characterized by division of labour, cooperation and conflicts, altruism and
mating system. Social behaviour is exhibited by a wide variety of animals
including invertebrates, fish, birds and mammals. 137
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Human Behaviours
6.2.3 Characteristics of Social Animals
Some of the major characteristics of social animals are listed below:

Highest level of organization- In a social organization several animals of the

same species live and interact together. The level of interaction may vary in
different societies. It may be only during breeding season or lifelong. The
individuals have diverse roles to perform and show division of labour. In insect
societies, there is differentiation of individuals into caste; each caste has
morphological and functional distinction.

Overlapping generations – The social group in many animals consists of

many generations living together; it may be three or more generations at a
time. The parents defend and protect the young ones

Cohesion – Individuals in a society live together. Such as individuals in a herd

of deer, pride of lions or peck of wolves remain in close association in the
given home range.

Communication – Different organisms in a society communicate in different

ways to fulfil a wide range of purposes. In communication, the sender
produces a signal to convey some message. The signal is received by
receivers, who respond to it by modifying their behaviour. The main purposes
of communication are to maintain the cohesiveness of the colony, reproductive
behaviour, territoriality, parental care, dominance interactions, foraging,
defense, and cooperative interactions within the colony. Some of the common
ways of communication include visual, auditory, chemical and tactile. The
visual stimuli include dances, display of plumage, ornaments etc. The auditory
signals are mainly songs, notes and calls. The songs are very commonly used
as nuptial songs. A variety of chemical substances are used for
communication and group interaction. Pheromones are the chemicals
produced and released in the environment and perceived by other
members of society. Sex pheromones, alarm pheromones, and
aggregation pheromones are some of the common examples of
pheromones used in group interaction.

Territoriality in Social Organization – Territoriality is a defended space

where an individual or group shows exclusive monopolization. A
territory may be a short stretch of beach during high tides defended by
sanderlings for feeding, or small sections of ponds used by male white-
tailed skimmers as mating territories only for a few hours. A territory may
include sites of egg deposition, burrow entrances, nest sites, food plants,
feeding space, display sites, roosting sites, shelters, grazing areas, food
stores, foraging space, and even patches of sunlight in the forest. A territory
may be defended by songs or visual displays in birds; in mammals, it is often
marked by scent posts marked with urine, faeces, or special glandular
secretions. A territory is defended to varying degrees depending on the costs
and benefits of defence. Costs of defending territory depend upon the energy
required to keep out intruders and benefits include exclusive access to food,
mates, breeding sites, and shelter. The tendency to hold territories varies
depending on the distribution of resources, the competition and the individual’s
138 internal physiology.
Unit 6 Social Behaviour and Altruism
The costs and benefits of territoriality depend on the resources and
competitors. More clumped resources are easy to manage and defend but
have intense competition. In this case, it is too costly to maintain and
exclusivity is almost impossible. In contrast, the resources which are spread
out or with lesser quality, decline and the benefits of defending reduce.
Therefore, territoriality of intermediate resources quality is generally expected.
Timings of territoriality also correspond to the period of greatest benefit. For
example, in Yarrow’s spiny lizards (Sceloporus jarrovii) mating territories are
maintained only when the majority of females are receptive to mating.

Dominance in Social System – Individuals in a social group may compete for

social dominance to gain hierarchical advantages to access resources or
mates. Male red deer and elephant seals spend much of their time fighting
during the breeding season. A dominant male also called an alpha male
gains preferred positions in mating arenas and is more likely to be
chosen by females. The fight among males can be very damaging and may
cause the death of a fighting competitor. Dominant individuals are more
aggressive and successful in winning competitive interactions. Social
dominance may often be achieved by mutual display and signals. Animals
often learn the fighting ability of each other from past experience. They avoid
fighting by having a dominant hierarchy or ‘peck order’. Subordinates often
exhibit an array of tactics or behaviours that help them make the best of their
low status.

Mating Systems in Social Organization – Mating strategies in a social

organization are planned to maximize reproductive success. Mating strategies
are determined by the role of the two sexes in reproduction and subsequent
investment in parental care. Broadly mating groups are classified into.

Monogamy – It is an exclusive mating relationship between one male and one

female. The male and female join and rear at least one offspring. Monogamy
may last for one season or a lifetime. 90% of the birds are socially
monogamous. Otherwise, this phenomenon is rare.

Polygamy refers to multiple mating. It can be polygyny i.e. a single male

mates with multiple females. The male does not stay with the female to rear
the young. In polyandry, a single female is mated with many males.

Promiscuity refers to the mating relationship of both sexes with multiple

partners.

Wilson compiled a set of 10 essential characteristics of sociality

1. Group size

2. Distributions of different age and sex classes

3. Cohesiveness

4. Amount and pattern of connectedness

5. “Permeability,” or the degree to which societies interact with one another

6. “Compartmentalization,” or the extent to which subgroups operate as

discrete units 139
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
7. Differentiation of roles among group members

8. Integration of behaviours within groups

9. Communication and information flow

10. Fraction of time devoted to social behaviour as opposed to individual

maintenance.

6.2.4 Social Behaviour of Lion

Group of Lions living together is called pride. A pride typically contains five
or six females with their cubs. There may be only one or a maximum of two
males in a pride. However, only one is dominant and he has the right to mate
with the females. The pride is matriarchal i.e. the daughters remain in the
original pride while the males born in the pride leave it and occupy others.
Every member of the pride has a role. This can be shifted depending on the
needs and the size of the pride.

The pride can be taken over by an intruder male; he defeats the existing male
to take over the pride. The newly appointed male shows infanticide behaviour
i.e. killing of the newborn. This behaviour has a selective advantage to the
new male. After the death of infants, the females enter estrous and the male
gets an opportunity to mate and father the new cubs.

The social structure of the Lion also affects their eating patterns . When a kill
is made the female involved with it eats first. What is left has to go through the
hierarchy of pride.

Usually, all of the adults in the pride work as a team to protect each other and
the young from other predators. However, there is often a hierarchy that
determines who will be dominant in that fight. Some of the lions may cover the
outer perimeter with others remaining well protected inside. There can be
brutal battles between the entire pride of lions over territory. When they have
locations that overlap there may be a fight for who is going to keep it. The
male can take over other pride if he challenges the dominant male and wins.

It is believed that males and females can be pushed out of pride if they can no
longer complete their roles. For example, if a female isn’t able to hunt any
more due to an injury or if a male is too old to be able to defend the pride.
Such changes in dynamics work to ensure those who are strong and healthy
can continue to thrive. Sometimes larger-sized prides push out members due
to overpopulation. A surge in the number of female offspring born and
surviving leads to the exclusion of some of the younger fully mature to leave
the pride.

6.3 COOPERATION AND CONFLICT IN ANIMALS

6.3.1 Cooperation
Cooperation is behaviour in the related animals in which the animals
work in a group for the benefit of each other. The cooperation increases
the fitness of an individual. The animals in the group cooperate for hunting,
140 defending territory, attracting mates, or forming coalitions.
Unit 6 Social Behaviour and Altruism
Cooperative foraging – Individuals in a group cooperate to get better access
to food. When in a group, the newly hatched insect larvae cope with the
physical defence mechanism of the host plant, and also they can penetrate
inside the leaf easily. Substances in the saliva overcome the chemical defence
of the plant and allow the plant to release more nutrients. The ants work
together to carry a larger insect to their nest. The honey bees collect nectar
and pollen by foraging in groups.

Cooperative hunting – Cooperative hunting is seen in many predatory

animals such as wolves, wild dogs, crocodiles, seals, walruses, etc.
Chimpanzees produce hunting bark during prey hunting. This helps to recruit
more group members and makes hunting easy. The jackals gather and
encircle a gazelle and her fawn at the time of hunting. The cooperation leads
to the separation of the fawn from the mother which is then prayed easily. The
pelicans form a semicircle around the fish to catch them. Hunting in groups by
the predators makes the prey confused; this makes it easy to capture the
predator.

Cooperative defence – In a group, the individual cooperates in defending

against a predator. When attacked, Australian sawfly larvae defend
themselves by aggregate regurgitation of noxious substances. The musk oxen
in the Arctic form a circle to defend against wolves. Groups of animals may
also confuse predators by looking larger than they are. These actions often
cause the predator to hesitate just long enough to permit the prey’s escape.

Communal nesting – Sociable weaver makes communal nest which looks

like a haystack hanging in a tree. This nest has multiple entrances, each
leading to different chambers. A single nest may have 5 to 100 nesting
chambers. This provides a home for 10 to 400 birds.

SAQ 2
Fill in the blanks:

i) Cooperative hunting is generally seen in ……………….. animals.

ii) Cooperative breeding is observed mostly in ……………….. .

iii) The queen termite that has exhausted eggs in the ovary is killed by
……………….. .

iv) ……………….. colonies face high risk of pathogen infection.

v) To attain territory ……………….. arises in the colony members.

141
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
Cooperative breeding/Cooperative brood caring – In cooperative breeding,
the young ones are cared for by more than two individuals. In many bird
species cooperative breeding involves ‘Helpers at the nest’. They are the
offspring from prior years that remain near their parents and help them rear
younger siblings. Species with helpers include common crows (Corvus
brachyrhynchos), and Florida scrub jays. Mexican jay lives in flocks of 8-20
individuals. The juveniles are fed by all the members of the group. Meerkats
and mongooses follow a cooperative approach to rear the young ones. In
meerkats, the young ones are raised and nurtured by many helpers. The
subordinate females lactate and feed during the first three weeks of life.
Subsequently, these females feed the young by sharing up to 40% of their
food for the next three months.

Cooperation in ants (social vaccines) – Ant colonies face a high risk of

pathogen infection. The invasive garden ant adopts a social immunization
strategy to counter this. The pathogen-exposed individuals are colour-marked
and healthy group members remove the pathogens by grooming. In this way,
they get exposed to the pathogen. However, the level of the pathogens in the
caregiver remains low which triggers an immune response resulting in a
protective immunization. Therefore, social contact with an infectious individual
results in 'social immunization' in its nest mates.

6.3.2 Conflicts
In a group, conflict arises in the individuals to attain territory, social
dominance, alpha status or mate.

Intra-sexual rivalry – Usually there is a male-male rivalry to acquire a mate. A

bower bird dismantles other bowers of other males; this reduces the
opportunities for competitors to copulate. Male-male fighting is seen commonly
in giraffes, red deer, and elephant seals. When male regularly fights for access
to mates, they tend to be larger than females of their species Males in animals
also evolved weapons in the form of horns, antlers, clubbed tails, and enlarged
spiny legs which they use when fighting with other males for females.

In Savanna baboon, male-male conflict is to attain dominant social hierarchy

or alpha status. The males compete intensely for high status. As a result of the
conflict individual males get bitten about once every six weeks; an injury rate
nearly four times greater than females. An alpha male generally mates more
often than subordinates; therefore contributes its genes to the progeny.

Infanticide – Infanticide is the killing of young offspring usually by a dominant

male of its own species. The infanticide behaviour increases the chances of
the male to mate with the female and attain fatherhood. This behaviour is seen
in hanuman langur, lions, dolphins, mice, etc.

Hanuman langurs live in groups called Band or Harem. It consists of several

females, their offspring and one or a few adult males. For the most part,
Hanuman langurs live a peaceful lifestyle. When a bachelor male overtakes
the leader of a harem, he targets the suckling infants for murder. This makes
the females stop lactating and enter estrous. Studies suggest a new ruling
male can increase his chances of having his offspring by killing the infants in a
142 troop.
Unit 6 Social Behaviour and Altruism
Conflicts in honey bees
In honey bees, the workers kill the drones after the mating is over. The queen
honey bee is monogamous i.e. it mates only once in her life. The only function
of the drones is to fertilize the queen. Therefore, after mating is over, they are
no more wanted in the beehive. Moreover, they are a liability to the worker as
they need honey for their survival.

The queen lays eggs in 5-7 queen cells before leaving for a new site. The
queen which emerges first visits the queen cells of the hive and stings to kill all
the developing queens.

Similarly, a queen termite that has exhausted eggs in the ovary is killed and
devoured by the workers. This ensures a perfect economy in the colony.

6.4 ALTRUISM AND INCLUSIVE FITNESS

Altruistic behaviour is a selfless form of cooperative behaviour for the
benefit of other individuals in the group; this behaviour may be
detrimental to the individual. In altruistic behaviour, an individual increases
the fitness of others at the cost of its own fitness. Altruistic behaviour is seen
as more common among closely related individuals than distantly related
individuals. This behaviour is often seen in the species with little dispersal as
they share kinship with their neighbours. The species showing altruism more
often discriminate against outsiders and favours insiders.

From evolutionary perspective, altruism poses a challenge to traditional

evolutionary theories centered on individual survival and reproduction.
However theories like kin selection and reciprocal altruism attempt to explain
its occurrence. Research suggests that empathy, compassion and a
sense of moral obligation are key psychological factors driving altruistic
behaviour. Altruism contributes to cohesion of social groups, fosters trust and
cooperation, and enhances overall well being within communities. Altruistic
behaviour may have limits particularly when individuals face competing
demands or risks to their own well being.

Relatedness is the probability of sharing an allele in the two individuals having

common descent. It is measured by the ‘Coefficient of relatedness’(r). The
value ranges from 1 (in identical twins) to zero in unrelated animals. Table 6.1
gives the value of ‘r’ in different relations.
Table 6.1: Coefficient of relatedness in different relations.
Relationship Coefficient of relatedness (r)

Identical twins 1

Parents progeny 0.5

Full siblings 0.5

Half siblings 0.25

Uncle/aunt-niece/nephew 0.25

Grandparents-grandchildren 0.25

First cousins 0.125

Unrelated individuals 0
143
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours

SAQ 3
Fill in the blanks:

i) Coefficient of relatedness (r) is ……………… in identical twins and

……………… in unrelated animals.

ii) Direct fitness + indirect fitness = ……………… .

iii) Altruism is favoured by natural selection when c< ……………… .

In honey bees the queen is diploid and produces two types of eggs while
males are haploid and produce only one type of sperms. Therefore the
coefficient of relatedness in the workers with the queen is 0.75, a value higher
than that of siblings in other diploid organisms. In queen and progeny, the
coefficient of relatedness is 0.5; this justifies being a worker in the bee
colony.

Inclusive fitness – inclusive fitness of an individual takes into consideration

the individual’s own reproductive success and the reproductive success of kin
along with their relatedness.

Inclusive fitness = direct fitness + indirect fitness

Inclusive fitness = individual’s reproductive success + (reproductive success of

kin x degree of relatedness)

Hamilton proposed that inclusive fitness offers a mechanism for the

evolution of altruism. He claimed that natural selection favours
maximizing their inclusive fitness. Hamilton's rule describes mathematically
whether or not a gene for altruistic behaviour will spread in a population.
Accordingly, altruism is favoured by natural selection when

c<rB where

c is the reproductive cost to the altruist; r is the degree of relatedness; and b is

the reproductive benefit to the recipient of the altruistic behaviour

Hamilton's rule supposes that an individual will be more altruistic for a

close relative rather than a distant relative.

6.5 SOCIAL BEHAVIOUR IN INSECTS

Eusocial insects are also called true social insects. The parental
generation of such insects lives together with many young generations of their
progeny. The individuals cooperate in caring for the progeny. In social insects,
a distinct caste system is seen. There is a division of labour; different castes
perform specific functions for the benefit of the colony. The individuals show
distinct morphological features suitable for the function to be performed.
Social behaviour is seen in termites (Order- Isoptera) and bees, wasps
and ants (Order- Hymenoptera).

144 Now we will explain the characteristics of social insects.

Unit 6 Social Behaviour and Altruism
Characteristics of social insects

• Large population – A nest of the social insect harbours a large

population. A honey bee hive may have 20,000-80,000 worker bees. In
ants, the number reaches up to 600,000; in a termitarium, the number of
termites reaches several million. Also, many generations live together in
an integrated manner in large groups in the colony. Colonies of social
insects are generally matriarchs i.e. members of the colony are offspring
of single females.

• Elaborate nest – Elaborated nests are constructed for the maintenance

of broods, and storage of food. The nests are perennial, initiated by the
reproductive caste but subsequently built and maintained by the
workers. It contains brood chambers for developing immature stages,
royal chambers for the reproductive caste, and storage cells. The nest
of termites also called termitaria is made up of an elaborate
architectural plan. It may contain a fungal garden where fungi are
cultivated. Bee hive is in the form of hexagonal chambers made up of
wax, secreted by the workers.

• Caste system – Members of the colony are morphologically

differentiated into different castes and each caste performs a distinct
function. Broadly the caste can be distinguished into reproductive
(queen and king) and sterile members (workers and soldiers). The
queen is the largest in size. It is the mother of the colony; the only
function performed by the queen is laying eggs. The reproductive caste
in termites can be macropterus, brachypterus or apterus. The sterile
caste is further differentiated into soldiers and workers. The function of
soldiers is to defend and protect the colony. The soldier caste can be
further distinguished into mandibulate and nasute based on the
morphological features. The ants have the most elaborate caste system.
The basis of caste determination is extrinsic in termites and genetic and
nutritional in honey bees.

• Protective devices – Different devices were developed in insects to

protect the individual and society from predators and prey. The worker
bees develop stings (modified ovipositor). In termites, soldiers are given
the task of protecting the colony. They are further differentiated into stout
mandibulate and individuals with chemical warfare- nasute. The
mandibulate forms have strong mandibles to seize the enemy; the
nasute forms eject chemicals from the rostrum to repel the enemy.

• Cohesiveness of the colony – The colonies are perennial. The

members of the colony are dependent on each other and cannot live
independently. They work in cooperation for the benefit of the colony.
Success is measured in terms of colony and by individual. Different
castes are bound by chemical and physiological mechanisms

• Parental care – It includes various activities such as provisioning of

food, feeding the young and queen, maintenance of hive temperature,
etc. Workers take charge of feeding the developing grubs and attending
to the queen. Some of the activities of the workers are cleaning and 145
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removing the debris from the cells, repairing and maintaining the hive,
foraging and provisioning food, cooling the chambers in the summer and
warming them in winter. Parental care is an instinctive behaviour.

• Progressive provisioning of food – Feeding of young ones

continuously from day to day until they metamorphosed into adults. This
is different from mass provisioning where one-time food is provided to
the developing stage. Usually, workers feed the developing grubs. The
nature of food offered to the immature stage may vary. Pastoral ants
carry ‘ant cows’ (aphids), keep them in overwintering quarters and feed
on honeydew produced by them. Harvester ants gather and store
seeds and pine needles for winter. The fungus-growing ants cultivate
pure crops of fungi in the underground gardens.

• Trophallaxis – Trophallaxis refers to mouth-to-mouth feeding in the

individuals of a colony. The young and reproductive castes are fed by
the workers. Some ants feed the nest guests and in return imbibe
nutritious secretions from them. In termites, trophallaxis is important in
caste determination. The reproductive caste secretes ectohormones;
during mutual feeding, they reach nymphs and prevent them from
developing into the same sex or caste.

• Swarming – Swarming is the marching of individuals in large numbers.

It occurs for feeding, migration, and mating. It also helps in reducing the
overcrowding of the colony and escapes the adverse conditions in the
inhabited area. A mating swarm is also called nuptial swarm. In this
queen is followed by drones for mating. In honey bee swarming is also
used for the multiplication of colonies. The honey bees also abscond the
preexisting site to tide over unfavourable conditions.

• Communication – Social insects communicate through tactile, chemical

(pheromones), visual, and auditory signals. Trail pheromones are
produced by ants and termites which help in foraging. Queen substance
is secreted by the queen honey bee which suppresses ovarian growth in
the workers. The honey bees perform dances which communicate
messages about the location of food.

SAQ 4
Give one word for the following:

i) Ants that gather and store seeds and pine needles for winter.

ii) Ants that cultivate pure crops of fungi in the underground.

iii) Ants that carry ant cows and keep them in overwintering quarters and
feed in honeydew produced by them.

iv) Termite soldiers with strong mandibles to size the enemy.

v) Termite soldiers that eject the chemicals from rostrum to repel enemy.

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6.6 SOCIAL ORGANIZATION IN HONEY BEE

6.6.1 Caste System
The caste system in the honey bee is well organized. The colony harbours
three types of morphologically distinct individuals. These are queens, drones
and workers (Fig. 6.1). All the individuals perform specific functions for the
benefit of the colony.

Fig. 6.1: Caste system in honey bee

Queen

The queen is the mother of the colony and the only fertile female. A colony can
have only one queen at a time. It is the largest in comparison to all other
members (Fig. 6.2). It has a long tapered abdomen and short legs and wings.
The brain and salivary glands of the queen are small. It cannot produce honey,
wax or collect pollen. It is diploid; and develops in special larger chambers of
the beehive called queen cell. The larvae which develop into queen are fed on
royal jelly throughout their development. The queen is attended by the
workers. The queen honey bee is strictly monogamous i.e. it mates only
once in its lifespan. It can lay thousands of eggs. Also, it can decide to
lay fertilized or unfertilized eggs. Post mating, the abdomen of the queen
gets greatly enlarged and membranous to accommodate large-sized
ovaries containing a large number of ova. This phenomenon is seen in
many other social insects such as termites and ants, known as physogastry.

SAQ 5
i) What is physogastry?

ii) What is the function of drones in bee hive?

Fig. 6.2: Queen and workers in a hive; Identify the queen 147
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Drones

Drones are sexually reproducing males in the bee hive. The wings are large
and extend beyond the tip of the abdomen in the folding state. They are
produced only during the reproductive season from unfertilized eggs and
develop parthenogenetically. The cells in which drones develop are large in
size and hexagonal in shape. The only function of the drones is to fertilize the
queen. They perform nuptial flight to mate. After mating the drones die or are
killed by the workers.

Workers

Workers are the smallest in size of all members and highest in number. Their
body is highly adapted to perform a variety of functions. The body is densely
covered with hairs. Mouth parts are biting and lapping types adapted for
foraging and industrious. The mandibles are spoon-shaped used to mould
wax. The legs are modified for pollen collection and storage. The forelegs
contain an eye brush and antenna cleaner. Mid legs have pollen spur and
hind legs with pollen basket. The legs have an important role in collecting
pollens and dislodging them in the hive. The ventral surface of the four
abdominal segments contains modified cells which act as wax glands; these
glands produce wax, which is used in making bee hives. Salivary glands in
workers are well developed. The salivary secretions make royal jelly which are
used to feed the grubs. Wings are well developed, the anterior margin of hind-
wings have hooklets called hamuli. The hamuli fit in the sockets present at the
posterior margins of the hind wing. This system makes the flight efficient and
effective. The ovipositor of the workers is modified into a sting apparatus. Its
function is in defence; the poison is mainly acidic. The workers perform all the
functions of the colony except reproduction. The functions range from feeding
and caring for young ones, attending queen, hive building and maintenance,
and foraging. The workers are sterilized females developed from fertilized
eggs. They develop in small hexagonal cells of the hive. The grubs which
develop into workers are fed on royal jelly for an initial few days but then fed
on bee bread- a diet made up of honey and pollen.

6.6.2 Beehive

The beehive is built in a hanging-down position from trees, buildings or rocks.

The walls of the beehive are extremely thin and consist of two layers of
hexagonal chambers of cells made from bee wax called a honeycomb. These
cells are used to store food and house the brood. The wax used for building a
hive is secreted from the wax gland present in the abdomen of the worker's
bees. The wax is masticated and mixed with secretions of cephalic glands to
convert into plastic substances. Propolis prepared from pollen is used in
making the comb waterproof. The resins, and gums secreted from plants are
used for repairing hives.

Different types of cells are present in bee hive which serve different purposes
(Fig. 6.3).

Storage cells – These cells contain honey and pollen and are built at the
148 margin and top of the comb.
Unit 6 Social Behaviour and Altruism
Queen cells It is a special type of cell raised away from the rest of the cells.
The queen cells are peanut-like and larger. In this cell, the queen bee
develops. The larvae of the queen bee were fed on royal jelly by worker bees.
After pupation, the workers cap the cell by beeswax. There are three types of
queen cells-

a. Swarm cells: These cells are created to raise a second queen. When
the colony becomes overcrowded, the present queen flies away with the
half of colony and the other queen being raised in the swarm cells
emerges in the present hive. During swarm season multiple swarm cells
can be spotted in a hive. In a double brood hive, swarm cells hang
vertically and are lumpier peanut-shaped.

b. Supersedure cells: When the present queen is injured, sick or old, new
young larvae start feeding with royal jelly to create a new queen. To
protect this young larva a cell built around her and is called a
supersedure cell.

c. Emergency cells: These types of cells come into existence when the
queen dies suddenly. The bees get into emergency mode and start
preparation for the new queen by converting brood cells into
supersedure cells and feeding them with royal jelly. These cells are
partly horizontal and partly vertical with a right-angle bend in the middle.
These can be built anywhere in the hive.

Drone cells – Drone cells are larger than worker cells. These are formed
close to each other at the bottom of a hive. The cells are dome shaped, and
the cap is higher and rounded at the top. Drone cells are almost 30% in a
colony. The number of drone cells will vary depending on factors such as
season, genetics and weather conditions.

Worker cells – They are found in the centre of the frame and are slightly
domed. These are smaller in size. Worker cells which are present in the lower
part of the honeycomb are meant for rearing the worker. In the upper part,
they store pollen and honey. They are most common in number.

Fig. 6.3: Beehive showing the different types of cells

(Source: [Link]
bee-biology-for-beekeepers/) 149
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SAQ 6
Give one word for following:

i) Peanut like special type of cell raised away from the rest of the cells in
bee-hive.

ii) Dome shaped cells, formed close to each other at the bottom of a hive.

iii) Smaller sized cells found in the centre of the frame and are slightly
domed.

6.7 POLYPHENISM
Polyphenism is a phenomenon in which two or more distinct phenotypes
can arise from a single genotype because of differing environmental
conditions. It is a special case of phenotypic plasticity. Examples of
phenotypic plasticity are caste polyphenism in eusocial insects and seasonal
polyphenism in butterflies.

Caste Polyphenism

Morphological distinctions in the morphology of different castes have been

described earlier. It is a unique case of phenotypic plasticity where the
development exhibits reproducible bifurcations e.g. worker and queen bee.
The same genome gives rise to completely different morphologies and
behaviour. In honeybees, development nutrition and genetics regulate the
morphology of an individual. Males are haploid, developed from unfertilized
eggs through parthenogenesis while queens and workers are diploid
developed from fertilized eggs. The bee larvae which are destined to become
queen are fed on royal jelly throughout their development. The grubs
developed into workers are fed initially on royal jelly and subsequently on bee
bread only and develop into workers.

6.8 POLYETHISM
Polyethism is functional specialization in different members of a colony
of social insects which leads to a division of labor within the colony i.e.
different individuals to perform different tasks within the group.

6.8.1 Caste Polyethism

The various functions may be carried out by individuals of different
morphology. Honeybee colonies comprise a single queen bee, a hundred
drones and several thousands of workers. Queens are fertile. The queen
and workers are females but perform different functions.

The queen is the mother of the colony and the only function performed by the
queen is egg laying.

Drone bees (male bees) perform the function of mating with the queen and
150 fertilize her.
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Worker bees perform almost all the functions, except reproduction in the
colony. Different types of tasks performed are like building the honeycomb,
feeding and attending the queen and grubs, cleaning and ventilating the hive,
foraging etc. A specific function is performed by a worker of a specific age.

6.8.2 Age Polyethism

Age polytheism is also known as temporal polyethism. The various
functions may be carried out by individuals of the same caste of different ages.
Worker bees perform different function which changes according to age-

Newly emerged bees can’t fly or sting and are developmentally immature.
Therefore by staying at a place, they clean cells.

Nursing caste lasts for about 1 week from ages 4-12 days. They feed the older
brood and younger and older bees with bee bread for their development and
maintenance.

Worker bees of age around 2 weeks guard the hive and remove corpses.

Middle-aged bees perform various functions ranging from nest building,

maintenance, guarding the nest and nectar receiving and processing. At this
stage, the worker bees develop wax glands in their abdomen.

The workers develop hypopharyngeal and mandibular salivary glands which

produce royal jelly. At this stage, the workers feed the queen and the grubs.

Nearly after 3 weeks the worker bees start foraging for nectar, pollen and
water. Foraging is done via dance language. Bee remains forager until they
die.

6.9 SWARMING
Swarming is a means of reducing congestion in the overcrowded colony. It
usually takes place to divide the colony into two or more distinct colonies.
Changing weather conditions seem to stimulate swarming in a colony; mostly
it occurs in the spring season. While preparing for swarming the queen bee
lays eggs in the queen cell of the old hive. As the queen cells are covered and
capped, the hive starts swarming before the emergence of a new virgin queen.
The old gravid queen bee is heavy and will face difficulty in flight therefore, the
worker bee stops feeding her a few days before swarming takes place.
Swarms do not travel too far during their first flight, and they settle just 20-30
m away from the hive for a few days until some scout bees search for a new
site for the hive. Scout bees report the location of suitable nesting sites to
other bees through dancing. While swarming, the bees are dependent on
stored nectar in their stomach only.

Nuptial swarm: During the nuptial swarm, mating takes place. It starts with
the flying of virgin queens out of a hive. The queen then releases sex
pheromones to attract males. Guided by the pheromones, the drones start
following the queen bee. The queen escapes the male during flight and only
allows the fastest and fittest male to mate. Mating takes place in the air. The
queen bee has a storage pouch called a spermatheca in which the sperm, are 151
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stored and utilized for fertilizing the eggs in future. The queen bee is
monogamous, and mates only once in its lifespan. After the nuptial flight, the
queen bees return to existing hives and lay eggs when required.

Absconding refers to the movement of the entire bee colony along with the
queen bee due to adverse conditions. The scarcity of sufficient flora for the
collection of nectar and pollen, invasion of beehives by predators and adverse
weather conditions are some of the main causes of absconding in honey
bees. Bees start preparing in advance before they move out. The queen bee
stops laying eggs and reduces in size to prepare for flight. Only a small
amount of capped brood is left behind. Rest all the members fly away. During
absconding no queen cells are produced.

6.10 CASTE DETERMINATION IN HONEY BEES

In honey bees, caste is determined by genetic and nutritional factors.
Honeybees use a haplodiploid sex determination mechanism. Johann
Dzierzon was the first person to discover the method of sex
determination in honeybees. His key observation is that the virgin queen
who has not taken the nuptial flight produces only male progeny. From this,
he inferred that unfertilized eggs develop into males while fertilized eggs
develop into females. Further, the females are differentiated into queen and
worker bees by the role of nutrition provided to the developing grubs and the
effect of the queen substance.

The queen bee is responsible for the determination of the sex of offspring in
the hive. Queen bee stores all the sperm of a drone obtained during the
nuptial flight in the spermatheca. The spermatheca is guarded by a sphincter.
Therefore, a queen can lay an unfertilized egg by closing the sphincter or
a fertilized egg by opening the sphincter. The unfertilized eggs are when
developed into haploid males by parthenogenesis this process is called as
arrhenotoky. All the workers and the queen are females; they are diploid and
developed from fertilized eggs. The distinction between queen and worker
depends on the food provided to the developing grub. The queen-worker
developmental divide is controlled epigenetically by differential feeding. Royal
jelly which is a secretion from the hypopharynx of nurse bees is used in the
nutrition of larvae and adult queens. It comprises water, proteins, sugars, fatty
acids, minerals, vitamins, and 10-hydroxy-2-decenoic acid (10-HAD). All the
larvae, in the colony, irrespective of sex or caste, are fed on royal jelly for the
first 3 days. After that workers and drones are fed on beebread while the
queen cells are fed with copious amounts of royal jelly throughout their
developmental period, and this helps in the development of ovaries in the
queen bee. The workers distinguish the grubs to be developed into a queen or
workers by the shape of the brood cell. In addition, the secretion of queen
substance- a pheromone from the mandibular glands of the queen, inhibits
ovary development in workers, inhibits the rearing of new queens and also
calm the workers to improve their cooperation.

6.11 COMMUNICATION AMONG HONEY BEES

Bees communicate using various pheromones, including queen substance,
152 nasonov gland (Nasonov’s gland produces pheromone used in recruitment in
Unit 6 Social Behaviour and Altruism
worker honeybees. This gland is located on the dorsal side of the abdomen)
secretion, alarm pheromone emitted from sting and secretion of tarsal glands..
Bee communications include cognitive and sensory processes.

Karl von Frisch, an Austrian ethologist received the Nobel Prize in Physiology
or Medicine in 1973, along with Nikolas Tinbergen and Konrad Lorenz, first
demonstrated that honeybees have color vision; Secondly, he also concluded
that the dance of honey bees contained information about the distance and
direction to good foraging sites.

6.11.1 Foraging in Honeybee

Worker bees perform the foraging activities. Foraging is an interaction
between the bees and the ambient environment. It is the best example of
cooperative behaviour. Foraging consists of various behavioural components
that include-

• Search for food supply -Nectar-honey, pollen, water, balm, resins and
propolis for maintenance and repair of the hive

• Identification and memorization of food source location

• Carrying and storing food

• Interaction and communication

Worker bees perform dances when they return to the hive after finding the
food source. Karl von Frisch has correlated the runs and turns of the dance to
the distance and direction of the food source from the hive. Two types of
dances were performed by bees:

6.11.2 Round Dance

Round dance is performed to communicate the location of food sources close
to the colony. When the food source is located <50 meters from the hive, the
forager returns to the hive and performs a round dance. The dance is
performed in the form of a circle, while moving in a circle they switch directions
(Fig. 6.4). The dance does not give any information about the direction of the
food source. The scout bees distribute the newly found nectar to the waiting
bees and the odour is involved in guiding the bees to the site of food.

Fig. 6.4: Round dance 153

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6.11.3 Waggle dance (Tail Wagging Dance)
Waggle dance is performed when the food source is at a distance. The scout
bee dances in forming the digit 8. The bee communicates the distance and
direction of the food source. In waggle dance, two half circles are made in
opposite directions with a straight run in between (Fig. 6.5). Bee shakes its
abdomen from side to side while performing the waggle run portion. The
number of wags per unit of time is inversely proportional to the distance of the
food. The direction of food is predicted by measuring the waggle run angle
vertically and also by the dancer's facial direction during the waggle dance
(Fig. 6.6).

Fig. 6.5: Waggle dance

154 Fig. 6.6: Waggle dance communicating the direction of the food source
Unit 6 Social Behaviour and Altruism
If the scout bee waggles while facing directly towards the comb, indicates that
the food source will be found by flying directly towards the sun. While returning
to the hive after collecting pollen and nectar the time compensatory
mechanism helps the work to reach the hive.

Menotaxis – refers to an animal maintaining a constant angle to a

stimulus. The orientation of the long body axis is at a constant angle to the
source of the stimulus (Fig. 6.7). It is commonly known as ‘compass
orientation’. It is observed in honeybees. They use sunlight as a guiding light
to return to their hive from forage and to indicate the direction of food through
dance performed at a particular angle to the sun.

Fig. 6.7: Menotaxis

On a cloudy day the worker bees use the plane of polarization of light to
communicate the direction of food source.

6.11.4 Chemical Communication

Honeybees use chemical cues to interact with each other and to manage the
colony organization. Pheromones allow the communication in all the castes.
The pheromones can be broadly divided into two categories i.e.

Releaser Pheromones – They have a temporary but immediate effect on the

recipient’s behaviour Alarm Pheromones – These are released when a bee
stings another animal attracting bees to behave defensively. Two alarm
pheromones have been reported: one released from the koschevnikov gland
(near the sting shaft) and another one is released from the mandibular
glands. It is used to mark the depleting foraging sites for new foragers. It is
also used to paralyze the intruders.

i) Nasonov Pheromone – Used for aggregation during swarming or if the

bees are displaced from the colony

ii) Brood pheromone – It is produced by developing larvae which

stimulates feeding of the larvae, and capping of the cells before
pupation. 155
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iii) Primer Pheromones – They have long-term effects on the recipient’s
physiology. They trigger complex responses and generate both
developmental and behavioural changes.

iv) Queen Mandibular Pheromone – It is secreted from the mandibular

glands of the queen and helps in maintaining the social hierarchy,
maintenance of colony, homeostasis and queen reproductive
supremacy. Also known as queen substance (E)-9-Oxodec-2-enoic
acid or (9-ODA) (Fig. 6.8). It affects the social behaviour, swarming,
mating, and inhibition of ovary development in worker bees. This queen
substance maintains the colony structure as it is given to the worker
bees and prevents them from producing/ rearing new queens. It also
acts as a strong sexual attractant for drones when on a nuptial flight.

Fig. 6.8: Queen substance

v) Queen Retinue Pheromone elicits the workers from a short distance to

lick, antennate and groom their queen.

6.11.5 Trophallaxis in Honey Bees

Trophallaxis refers to the exchange of food between two individuals. The
food can be salivary secretions or regurgitated gut secretions. The transfer of
food or other fluids among members of a community takes place through
mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. It also
involves the transfer of molecules such as pheromones. It is highly developed
in social insects. Besides providing nourishment, it also helps in kin survival,
colony recognition, foraging communication, and maintaining a social insect
population.

In honey bees, trophallaxis is used by foragers in associative learning to form

long-term olfactory memories. Through this, the current foragers teach the
new foragers about foraging behaviour and where to search for food. In honey
bees, antennal movements of both donor and receiver are rapid during food
transfer vary according to the reward rate experienced by the food donor and
show a positive correlation between both trophallactic partners.

SAQ 7
Write 1-2 lines for each of the following in bee communication:

i) Nasonov pheromone

ii) Brood Pheromone

iii) Primer pheromone

iv) Queen mandibular pheromone

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6.12 SUMMARY
Let us summarise whatever you have learnt so far:

• Solitary animals spend the majority of their life in isolation without

interacting with other individuals of their species. On the other hand,
gregarious animals live in groups

• The grouping of animals can be aggregation, subsocial or social

depending on the interaction seen among the members in a group.

• Social organization is the highest grade of organization. It is seen in

diverse animal groups.

• The social set-up of a lion is called pride. It consists of 1-2 males, 5-6
females and their cubs. The pride is matriarchal i.e. the daughters
remain in the original pride while the males born in the pride leave it.

• The social behaviour in insects is characterized by a large population in

a colony, caste system, division of labour, cohesiveness of the colony,
parental care, progressive provisioning of food, trophallaxis, swarming,
protective devices, and communication. Termites, bees and ants show
social behaviour.

• In honey bees, the three castes are the queen- performs the function of
egg laying, drones- fertilize the queen and workers- perform all the
functions of the colony except reproduction.

• The phenomenon in which different individuals of a colony perform

different functions is called polyethism. The honey bee shows both caste
polyethism and age polyethism.

• The body structures of different castes are modified according to the

functions they perform. The phenotypic plasticity of different castes is
called polyphenism.

• The bee colony thrives in a nest called a beehive. It is made up of wax

secreted by the workers. The beehive contains usually hexagonal
chambers for the development of brood and storage of honey and
pollen.

• The basis of caste determination in honey bees is both genetic and

nutritional. The drones are haploid and developed parthenogenetically
from unfertilized eggs. The workers and queen are diploid developed
from fertilized eggs. The difference between the workers and the queen
is made by the type of food offered to the developing grubs and the
effect of queen substance.

• Swarming is the movement of the queen followed by workers (a division

of colony or absconding) or with drones (nuptial). It is meant for
multiplying the colony or leaving and establishing a new colony at a
better place.

• Communication in honey bees is well studied. Karl von Frisch

deciphered the bee dances used to communicate about the food source.

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• Round dances are performed when the food source is close by. The
waggle dance is performed when the food source is at a larger distance.
It tells about the distance and direction of the food source.

• Chemical communication is done among the members of the colony with

the help of pheromones. Queen mandibular pheromone is secreted
from the mandibular glands of the queen and helps in maintaining the
social hierarchy, maintenance of colony homeostasis and queen
reproductive supremacy. It affects the inhibition of ovary development in
worker bees.

• Trophallaxis refers to the exchange of food between two individuals. The

food can be salivary secretions or regurgitated gut secretions.

6.13 TERMINAL QUESTIONS

1. i) Define the following terms:

a) Monogamy

b) Promiscuity

c) Communication

ii) Distinguish between Subsocial and Social behaviour.

iii) Explain the social behaviour of lion.

2. Define:

i) Infanticide

ii) Cooperative foraging

iii) Conflict

iv) Cooperative hunting

3. Explain the interrelationship of altruism and inclusive fitness.

4. Explain the important characteristics of social insects.

5. i) Define:

a) Waggle dance

b) Menotaxis

c) Swarming

d) Polyethism

e) Polyphenism

f) Royal jelly

ii) Explain the following:

a) Arrhenotoky
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b) Sociotomy

c) Division of labour

d) Queen substance

iii) Distinguish between the following:

a) Round dance and Tail wagging dance

b) Age and caste polyethism

iv) Draw a labeled diagram showing the waggle dance performed by a

scout honey bee in the following conditions.

6.14 ANSWERS
Self-Assessment Questions
1. i) Aggregation, ii) Aggregation, iii) Alpha, iv) Promiscuity,
v) Matriarchal.

2. i) Predatory, ii) Birds, iii) Workers, iv) Ant, v) Conflict.

3. i) One, Zero, ii) Inclusive, iii) rB.

4. i) Harvester ants, ii) Fungus growing ants, iii) Pastoral ants,

iv) Mandibulate, v) Nasute.

5. i) In social insects like ants and termites, post mating the abdomen
of the queen gets greatly enlarged and membranous to
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accommodate large sized ovaries containing a large number of
ova. This phenomenon is called as physogastry.

ii) Their only function is to fertilise the queen.

6. i) Queen cell, ii) Drone cells, iii) Worker cells.

7. i) Nasonov Pheromone – Used for aggregation during swarming or

if the bees are displaced from the colony

ii) Brood pheromone – It is produced by developing larvae which

stimulates feeding of the larvae, and capping of the cells before
pupation.

iii) Primer Pheromones – They have long-term effects on the

recipient’s physiology. They trigger complex responses and
generate both developmental and behavioural changes.

iv) Queen Mandibular Pheromone – It is secreted from the

mandibular glands of the queen and helps in maintaining the social
hierarchy, maintenance of colony, homeostasis and queen
reproductive supremacy.

Terminal Questions
1. i) Refer to Subsection 6.2.2 and 6.2.3.

ii) Refer to Subsection 6.2.2.

iii) Refer to Subsection 6.2.4.

2. i) Refer to Subsection 6.3.2.

ii) Refer to Subsection 6.3.1.

iii) Refer to Subsection 6.3.2.

iv) Refer to Subsection 6.3.1.

3. Refer to Section 6.4.

4. Refer to Section 6.5.

5. i) Refer to Sections 6.10 and 6.11.

ii) Refer to Sections 6.9, 6.10 and 6.11.

iii) Refer to Subsection 6.11.2 and Section 6.8.

iv) Refer to Subsection 6.11.3 (Fig. 6.6).

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 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
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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

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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
 UNIT 8

MATING BEHAVIOUR

Structure
8.1 Introduction 8.7 Courtship Behaviour in
Three-Spined Stickleback
Objectives
8.8 Sperm Competition and
8.2 Asymmetry of Sex
Post-Insemination Tactics
8.3 Reproductive Fitness
8.9 Summary
8.4 Sexual Dimorphism
8.10 Terminal Questions
8.5 Mating Systems
8.11 Answers
Monogamy

Polygamy

Promiscuity

8.6 Sexual Selection

Intra-Sexual Selection (Male-
Male Rivalry)

Inter-Sexual Selection (Female

Choice)

8.1 INTRODUCTION
It is an innate desire of all living organisms to reproduce and leave their own
type of progeny to future generations. Sexual reproduction involves the
participation of two sexes i.e. male and female. The two sexes differ in their
morphology, physiology, genetics and behaviour across the animal kingdom.
Sexual reproduction is a key process for bringing variation in the population
and therefore, helps in the process of evolution. On the other hand, evolution
by the process of sexual selection defines and optimizes the process of sexual
reproduction. All the individuals adopt strategies to maximize the contribution
of their genotype to the next generation. The key components of sexual
reproduction are premating behaviour, mating and insemination, and
postmating behaviour including parental care of the newborns. Premating
behaviour includes a sequence of events which bring the potential mates
together. Through the exchange of signals and behaviour, the mating partners
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exhibit mating receptivity, mate attraction, and mate choice and mate
selection. Usually, the male attracts and the female selects the potential mate
in different animals. This evolves sexual dimorphism and the evolution of
expansive ornaments and sexual displays in the males. The mating involves
the transfer of spermatozoa from the male to the female genital duct. The
duration of mating varies in different animals; it ensures insemination of the
females. The number of mating and the mating partners are also highly
variable in different animals. The mating system defines the number of mating
partners an individual has. Males in some animals show postmating
interactions and do not allow the females to mate with other males. Parental
care, although expensive, promotes the reproductive success of an individual.

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

 explain asymmetry of sex and its role in the attraction and selection
behaviour of potential mates,

 describe reproductive fitness and factors governing reproductive

fitness,

 appreciate sexual dimorphism and its adaptive significance,

 discuss the animal mating systems and difference among monogamy,

polygamy (polygyny and polyandry), and promiscuity,

 narrate evolutionary advantages of sexual selection; types of sexual

selection, and

 describe sperm competition and post-insemination tactics adopted by

the males to ensure paternity.

8.2 ASYMMETRY OF SEX

Males and females contribute differently in terms of the amount of energy,
time, and resources contributed towards parental investment in sexual
reproduction. This causes evolutionary differentiation of males and females in
terms of morphology, physiology and behaviour. The males produce many
Fig. 8.1: Fairy Wren sperms and invest little into each one. For example, Fairy wren has 8 billion
sperms in his testis (Fig. 8.1). Coho salmon releases 100 billion sperms on a
batch of 3500 eggs (Fig. 8.2). In human beings, a single ejaculate contains
350 million sperms. Theoretically, a single man can fertilize all the ova of the
females of the world.

On the other hand, egg production by females is a costly phenomenon; the

females produce significantly fewer eggs and invest heavily in each one.
Further, a female often spends time and energy in parenting the progeny.
These strong differences in the number of gametes formation and gamete
Fig. 8.2: Coho Salmon investment in the males and the females lead to the adoption of different
reproductive strategies of the two sexes. Male contribution of the genes to the
next generation increases with the number of females he mates, while, the
180 reproductive success of the female depends on the quality of her partner. This
Unit 8 Mating Behaviour
causes competition among the males to fertilize the limited supply of the eggs.
Due to the high investment in the production of their gametes, females often
take more care in selecting a mate than males. In a typical mating behaviour,
the males attract and the females select.

The operational sex ratio is the ratio of sexually receptive males and
sexually receptive females at a given time at a particular place. This ratio
decides the role of the two sexes in attraction or selection in mate finding.

Male-biased operational sex ratio (male-male rivalry, selective

females)

Usually, sexually active males are more in number compared to sexually

active females throughout the animal kingdom. This makes a ‘Male-Biased
Operational Sex Ratio’ and results in evolutionary pressure on the males to
attract females and win over other males. Under the influence of sexual
selection males evolved many structures for fighting and courting. The
availability of a large number of sexually mature males results in the evolution
of selection behaviour in the female (Fig. 8.3).

Fig. 8.3: Operational sex ratio

Female-biased operational sex ratio (sex role reversal) Fig. 8.4: Empid fly- The
male offers the female
Mating in many cases involves donations of expensive nuptial gifts by males. insect prey
In Empid flies, the male offers the female an insect prey (Fig. 8.4). Male
katydid transfers to the female with an edible spermatophore; this enhances
egg production by the female (Fig. 8.5). Male pipe fish Syngnathus offers the
female with brood pouch to lay eggs (Fig. 8.6). The pregnant males provide 181
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nutrition and oxygen to the fertilized eggs for several weeks. This incurs high
paternal investment by the males. Also, in the polyandrous system, females
mate with multiple males, resulting in female competition for males.
Consequently, the reproductive females are more in number and fewer males
are available for mating. This results in the ‘Female-Biased Operational Sex
Ratio’. Consequently, there is sex role reversal; males select and the females
attract (Fig. 8.3).

Fig. 8.5: Male Katydid with large spermatophore

Fig. 8.6: Pipe fish- Male with brood pouch

The operational sex ratio can change with the environmental conditions; this
changes the sexual tactics of the males and the females. In skinny Australian
katydid, the availability of food resources decides the operational sex ratio.
The food supply varies throughout the breeding season. In the scanty
availability of food resources, the male’s large spermatophore is difficult to
produce and valuable to the females. Therefore, only a few resourceful males
are available. This makes the operational sex ratio female-biased, the females
fight and the males are choosy. On the other hand, when pollen-rich grass
starts flowering, many males can produce large spermatophores more rapidly.
This makes the operational sex ratio male-biased. Therefore, many males are
available for mating; in these conditions, males attract and fight and the
182 females select.
Unit 8 Mating Behaviour

SAQ 1
i) Define (a) Asymmetry of sex (b) Operational sex ratio

ii) In a situation, the number of sexually active males is more than the
number of sexually active females; in this case

a) Both the males and females will be attractive

b) Both the males and females will be selective

c) The males will be attractive and the females will be selective

d) The males will be selective and the females will be attractive

iii) In which of the following situations the sex role reversal will not take
place?

a) The operational sex ratio is female-biased

b) The male offers expensive nuptial gifts to the female

c) The parental care is done by the male

d) The number of sexually active males is more than the number of

sexually active females

8.3 REPRODUCTIVE FITNESS

The reproduction fitness of an individual is the number of viable offspring
produced by it in a population. In this way, it passes its genes to the next
generation. In other words, it describes the genetic contribution of an individual
to the future generations. Reproductive success is not limited by the number of
offspring produced by one individual, but also by the reproductive success of
these offspring themselves.

The reproductive fitness of an individual can be measured in absolute or

relative terms. Absolute fitness measures the number of offspring contributed
to the next generation. Relative fitness indicates the fitness of an allele or
genotype in comparison to the maximum [Link], it measures the
differential reproductive success of a genotype. In statistical terms, it refers to
the comparison of reproduction rates between various individual members or
groups within generations. If there are two groups, each with a different
variation of a trait, then by determining the differential reproductive success
rate the “fittest” group can be found. For example, if X and Y, are two variants
of a trait. The X variant attains reproductive maturity more frequently and
produces more offspring than the Y variant. This suggests that the X variant
will have more genetic share in the next generation and future generations.
Also, a longer life span is favoured by natural selection, only if it brings a
higher reproductive success rate. For example, if a short-lived cat leaves more
progeny than a long-lived cat, its genes will spread faster and will be favoured
by natural selection. Therefore, any fitness has no meaning in evolutionary 183
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terms if it does not correlate with reproductive fitness. Parental investment is a
key factor in reproductive fitness; taking better care of progeny gives a fitness
advantage later in [Link] choice, sexual selection reproductive strategies,
and parental care are some of the important factors which decide the
reproductive success of an individual.

To maximize reproductive fitness different groups of animals adopt different

reproductive strategies. This includes premating, mating and postmating
strategies. Premating behaviour includes courtship behaviour and sexual
displays to attract mates, and mate selection behaviour. Premating behaviour
is important to transfer their genes to the next generation successfully; the
animals need to choose a suitable mate. Failure to do so leads to low or poor
reproductive success. Also, sexual displays and courtship behaviour present
health, skills and quality of mate. Mating behaviour includes the mating system
i.e. mating relationship of the partners, frequency of mating, and duration of
mating. This leads to a successful transfer of spermatozoa to the female.
Postmating behaviour includes mate-guarding behaviour and parental care. All
these behavioursare aimed to maximize the progeny output in terms of
number, health and reproductive abilities.

SAQ 2
i) Define reproductive fitness

ii) Differentiate between absolute fitness and relative fitness

iii) Fill in the blank:

Mate choice and mate selection are part of …………………………….

(premating/postmating) behaviour

8.4 SEXUAL DIMORPHISM

Sexual dimorphism refers to the morphological differences in the males and
the females of the same species. The males can be distinguished from the
females on the basis on morphological features such as colour pattern, shape,
size, and structure. The behavioural and cognitive differences may also be
present between males and females. The traits which bring sexual dimorphism
are heritable. The male is larger than the female in some species. While in
others, such as some spiders, the female is larger than the male. Other sex-
specific differences are the colour of plumage and songs in birds (Fig. 8.7),
structures such as horns, antlers, and tusks used in struggles for dominance,
and ornaments for display (Fig. 8.8 and Fig. 8.9). The size of eyes, possession
of stings in bees, and certain behaviours such as aggression, and infant care
are other traits that impart sexual dimorphism. The male baboon is large and
possesses huge canine teeth which offer better protection (Fig. 8.10). In the
case of herd animals such as deer, the antlers are a sexually dimorphic
weapon which helps the males fight and establish breeding rights. Although
the antlers are expensive in terms of survival, they offer the reproductive
184 advantage, and thereby inherited to the next generation.
Unit 8 Mating Behaviour
Colouration and ornamentation: Many birds show dimorphism in colour.
Usually, the females are cryptically coloured so that they remain concealed in
the nest. The males are more colourful which gives them a reproductive
advantage by display in courtship and territorial behaviours. The plumage of a
peacock is colourful and elaborate and used to attract peahen. Similarly,
male bird-of-paradise shows elaborate plumage in response to sexual
selection. In mallards, the male has a bright green plumage on its head
whereas the female has light plumage.

Fig .8.7: Plumage of Peacock

Fig .8.8: Antlers of deer 185

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Fig .8.9: Tusk of elephant

Fig .8.10: huge canine teeth of baboon

Size: Pronounced size differences may occur between the sexes. In humans
and many other mammals, the males are larger than their female
counterparts. Male baboons are more than twice as large as females. In
Steller sea lions (Eumetopias jubatus), a male weighs about 1,000 kg, roughly
three times the weight of a female (Fig. 8.11). In a few mammals, non-
mammalian vertebrates and many invertebrate species the females tend to be
larger than males.

Fig. 8.11: Steller sea lions (Eumetopias jubatus), a male is roughly three times
186 the weight of a female
Unit 8 Mating Behaviour
Song production: The males in many birds produce more elaborate and
complex songs than their female counterparts.

Courtship behavior: The male exhibits a variety of morphological and

behavioural displays to mate. The male behaviour is to attract the female; on
the other hand, the female selects.

Access to the opposite sex is not the only reason for sexual dimorphism. In the
predatory birds, the females are larger than the males. This allows the mated
pair to hunt a greater variety of prey for themselves and their juveniles. In
many insect species females are larger which enables them to produce and
lay more number of eggs. In some cases, sexual dimorphism enable males
and females to exploit different food resources, thus increasing their collective
ability to find food. The mountain spiny lizard is sexually dimorphic in feeding
habits; the equal-sized males and females look for different sizes of prey. In
some species of woodpecker, the shape and size of the beak differ; this
enables the two sexes to find insects in different layers of a tree's bark.

Biological aspects of sexual dimorphism: The phenomenon of sexual

dimorphism is a direct product of evolution by sexual selection. Many male
and female organisms follow different evolutionary paths in response to
struggle for reproductive success. This results in evolution of many forms of
dimorphism which appear disadvantageous to organisms. For example, bright-
coloured male birds can easily be spotted by the predator and cost survival
fitness. Similarly, the antlers of deer and other forms of natural weaponry are
very expensive to grow and carry.
The explanation for this apparent paradox is that, at a biological level, the
reproductive success of an organism is often more important than its long-term
survival. This is particularly apparent in the case of game birds: A common
pheasant male in the wild lives about 10 months; in contrast, the females
survive twice as long as the males. However, a male pheasant's ability to
reproduce depends not on how long he lives but on whether females will
select him to be their mate. His bright colouration advertises his fitness and is
a good choiceto the female, to father her chicks.

8.5 MATING SYSTEMS

A mating system describes the mate pair relationships of the mating partners.
Three types of mating systems i.e. monogamous, polygamous (polygyny and
polyandrous), and promiscuous, are seen in different animals. Each type of
mating system is innate and evolutionarily selected (Fig. 8.12).

8.5.1 Monogamy
In monogamous mating systems, one male and one female pair remain in an
exclusive mating relationship for at least one breeding season. These mating
Fig. 8.12: Mating
alliances may be longer or last for a lifetime. Usually, in the monogamous
system in animals
system, both partners are involved in parental care and defence of offspring.

Evolutionary advantages of monogamy

Mate-guarding hypothesis states that males stay with the female to prevent
other males from mating with her. This behaviour is advantageous in an
environment where females are scarce. 187
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Male-assistance hypothesis: The males remain with a female and
participate in guarding and rearing their young. This produces more and
healthier offspring. It is seen that many monogamous species live in
environments with widely scattered resources; therefore, efforts of both the
parents are needed to forage for enough resources to rear the young.
Monogamy is observed in many bird populations where, both the female and
the male provide parental care for the young ones.

Female-enforcement hypothesis states the female ensures that the male

does not have other offspring that might compete with her own. In this case,
the female actively interferes with the male’s signalling to attract other mates.

Sexual monogamy: Sexual monogamy is true monogamy; both partners

mate only with each other. It is exceedingly rare.

Choosing an inappropriate mate in monogamous species has high fitness

costs. Monogamy also reduces the potential for genetic variation among a
female's offspring. By mating with more than one male throughout her lifetime,
a female gains higher genetic variation among her offspring. Therefore,
monogamous organisms adopt alternative forms such as serial monogamy or
social monogamy. In this way, the benefits of monogamy, such as shared
parental care and territorial resources, are maintained by having only one
mate at a time, or by concealing extra-pair partnerships.

Serial monogamy is a mating system in animals where they pair with a mate
for one mating season but change mates throughout a lifetime.

In social monogamy, two individuals are involved in rearing their offspring,

but they also engage in “extra-pair copulations,” with another individual. In
Starlings, both parents are engaged in caring for the young ones; however,
three to four males may be fathers in one brood. In many species, it is seen
that the female remains actively involved in copulation with outside males.
Social monogamy has both advantages and disadvantages for each partner. A
male mating with other females increases the total number of his offspring.
Same way a female may get genetically ‘better’ males’ in such mating which
helps the female in raising genetically superior progeny. In blue tits, a female if
mated to a lower-quality male more likely involves in extra-pair copulation.
This behaviour is also disadvantageous to males as he is most likely to render
help to raise offspring that are not their own. Same way the disadvantage to
the female is that the male may abandon her and her offspring if he detects
that she has mated with another male. Social monogamy is commonly seen in
songbirds; where up to 40% of the offspring in a mating pair’s nest were not
fathered by the male partner.

8.5.2 Polygamy
Polygamy refers to the mating system in which one partner mates with multiple
partners i.e. one male mating with many females (polygyny) or one female
mating with many males (polyandry).

A) Polygyny
In polygyny, one male mates with many females. In this system, usually, the
female takes responsibility for most of the parental care. Polygyny is a strategy
used by males to increase their reproductive fitness. This mating system is
188 most common in mammals but is also seen in a few birds and insects.
Unit 8 Mating Behaviour
Resource defence polygyny: It is a mating system in which males establish
a territory around the resources needed for mating success. A group of
females are attracted to a resource and join the male in his territory; the male
then mates with females. Therefore, the males compete for territorial
possession of the resource, which in return gets mating priority. The female is
benefited by mating with a dominant, genetically-fit male. In the yellow-rumped
honeyguide, the male defends the beehives. As the female approaches the
beehive to feed wax, the male defends the nest and mates with her.

Harems: It is a group of females associated with a single male. Typically the

male in the harem defends his group of females. The male also dominates
mating while controlling a territory with resources. Therefore, the males
compete for control of the groups. For example, in elephant seals, the alpha
male dominates the mating. Harems typically exhibit a dominance hierarchy
among the females in the group (Fig. 8.13).

Fig. 8.13: Harem of elephant seal

Leks: Lek is a communal courting area where several males perform

elaborate sexual displays to attract females; the females choose their mates
from the performing males. The leks attract more females than a single male
would attract. Aggregations of males may be near attractive females or in
areas where females are likely to travel. Possibly, the males from leks attract
more females than do isolated males and in this way increase their
reproductive success. Unlike most other mating systems, leks are not
associated with resources. This behaviour is observed in several bird species.
At the return of spring in the sagebrush steppe, greater sage-grouse begin
their spectacular mating displays (Fig. 8.14).

Fig. 8.14: Lek of Sage grouse 189

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B) Polyandry

In polyandrous mating systems, one female mates with many males. This type
of system is much rarer than monogamous and polygynous mating systems.
Polyandry is a reproductive strategy that promotes the reproductive fitness of
a female by providing her with multiple mating options. It involves sex role
reversal, where females invest less in offspring while males invest more.

In the spotted sandpiper, females possess resources, which in turn control

male mating associations. In pipefishes and seahorses, males receive the
eggs from the female, fertilize them, protect them within a pouch, and give
birth to the offspring. Pipefishes and sea horses differ in their mating
strategies. The seahorses are monogamous, while the pipefish are
polyandrous. This is due to ecological differences; the seahorses live in
habitats with widely distributed resources. This helps the seahorse population
to spread out and spread thin. In the scattered population it is difficult to find a
mating partner; therefore it is wise for the female to keep a partner for
reproductive assurance and natural selection favours monogamy. On the other
hand, the pipefish tends to live in very dense populations in resource-rich
environments. Here the male’s pouches, and not the female’s eggs, are the
limiting resource in reproduction. In these circumstances, the females compete
with each other for access to males; and natural selection favours polyandry.

Cooperative Polyandry: The Galapagos hawk exhibits cooperative

polyandry. In this case, all males in the group copulate with the female and all
participate in brood provisioning.

8.5.3 Promiscuity

In a ‘promiscuous mating system’ both the males and the females mate with
multiple partners. There exists no pair bonding and the mating takes place
randomly. In promiscuity, a single male fails to sexually monopolize a group of
females. This may be due to the wider range of females than the territory size
of a single male, so they interact with multiple males. When the males and the
females live together in large social groups, often all females are sexually
receptive at the same time. In this condition, he mates with one female, but
cannot prevent other males to mate with other females. In promiscuity each
female mates with multiple males, therefore, paternity is never certain. The
uncertainty selects males to avoid infanticide. Bonobos live in large social
groups where sexual activity plays a major role as a means of forming social
bonds and conflict resolution.

SAQ 3
i) Define (a) Sexual monogamy (b) Polygamy (c) Promiscuity

ii) Differentiate between

a) Polygamy and Promiscuity

b) Serial and social monogamy

190 c) Polyandry and Polygyny

Unit 8 Mating Behaviour
iii) Give one word which explains the following:

a) All the males of a group copulate with one female and all
participate in brood provisioning

b) Both the males and the females mate with multiple partners.

c) The monogamous female mates with multiple partners

d) A communal courting area where several males perform elaborate

sexual displays to attract females

e) A group of females associated with a single dominant male.

f) The males establish a territory around the resources needed for

mating success.

8.6 SEXUAL SELECTION

Sexual selection is a type of natural selection that favours individuals with
certain inherited characteristics to obtain mates more successfully than others.
Sexual selection has two components, intrasexual competition and
intersexual selection. The intrasexual selection consists of contests between
members of same sex, where the winner of the contest gains preferential
access to members of the opposite sex. Male-male rivalry and social
dominance are some of the main outcomes of intrasexual selection. The
intersexual selection operates between two sexes. In this one sex, usually
male, attracts the females and the female selects the best fit males. The
attracting sex evolves different strategies to win over the other males. They
may offer material benefits in the form of expensive nuptial gifts, or advertise
their fitness and resourcefulness by extravagant ornaments and courtship
displays.

SAQ 4
Fill in the blanks

i) Courtship behaviour is a type of ………………… intrasexual/intersexual

behaviour.

ii) Male-male rivalry is a type of ………………… intrasexual/intersexual

behaviour.

8.6.1 Intra-Sexual Selection (Male-Male Rivalry)

Intra-sexual selection is responsible for competition among the members of
the same sex for acquiring [Link], males attract; therefore, the males
compete to attract females for mating. This results in male-male rivalry and
fights. The female on the other hand is choosier sex. She contributes more
parental investment in the form of expensive egg production, sure maternity;
therefore, less number of females are available for mating. Hence females
select the males. 191
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Evolution of secondary sexual characters in males as a result of
intra-sexual selection

Sexual selection for fighting ability often leads to the evolution of larger body
size males. When male regularly fights for access to mates, they tend to be
larger than females of their species. Males in animals evolved weapons in the
form of horns, antlers, clubbed tails, and enlarged spiny legs which they use
when fighting with other males over females (Fig. 8.15).

Fig. 8.15: Horn and antlers in males

Darwin proposed the evolution of weapons in males due to sexual selection as

these are useful in fighting sexual rituals. However, biologists believe that
these weapons served to threaten and intimidate the rival. The antlers though
appear as a burden to the animal, provide the advantages of intrasexual
selection. This outweighs the pressure of natural selection. Competition
between two stags requires both fighting and assessment. Buffaloes and big-
horn sheep charge each other and clash head-on to get access to the female.
The two male giraffes hit each other violently with their heavy neck and
clubbed heads (Fig. 8.16). Therefore the neck of the giraffe is as much a
product of sexual selection as of natural selection.

Fig. 8.16: The two male giraffes hit each other violently with their heavy neck
192 and clubbed heads
Unit 8 Mating Behaviour
Male-male combat in Red deer (Cervus claphus)

In red deer, the intra-sexual selection is seen as male-male rivalry which

results in fights among the males. Stag grow antlers each year and in autumn
rutting season, directly challenge each other for ownership of females. The
females have no antlers and are herded into harems by successful males. The
females have little or no choice of sexual partner as male defends harems
against possible rivals (Fig. 8.17)

Fig. 8.17: Male- male fight in red deer

Tim Clutton and his coworkers made an intensive study on the male-male
rivalry and intra-sexual selection of red deer.

1. Males challenge each other initially by roaring, starting slowly and then
speeding up

2. Challenger usually retreats if the harem owner roars faster.

3. If the challenger matches the roaring of the harem owner, then the two
approache and walk parallel to each other more closely, particularly
concerning body size

4. Many contests end at this stage, but some escalate into proper fighting.

5. The stags interlock antlers and push against each other. Fighting is quite
fierce.

6. Usually the larger stag wins, other one which is handicapped by previous
injury or exhausted by the strenuous rutting season may be supplanted
by the young stag.

Another reason for male–male fighting is to attain dominance hierarchy. The

males of Savanna baboons compete intensely for high status. As a result,
individual males get bitten about once every six weeks. This injury rate in
males is almost four times greater than in females. As achieving high status is
a costly effort, high-ranking individuals are rewarded in terms of priority access
to useful resources and more importantly reproductive privilege. Alpha males
exhibit the social dominance in the hierarchy. Generally, they mate more often
than subordinates and get paternity benefits. Sometimes, it is seen that the
males of low and high status were equally likely to copulate. In these cases,
an alpha/dominant male mates with the fertile females (estrous cycles) and
subordinate males beta/satellite/subordinate males mate with infertile females. 193
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In the male elephant seal (Mirounga angusthrostris), two male- fighting (male
rivalry) is influenced by the behaviour of females (Fig. 8.18). The female
intensifies the competition among males by her protest and ensures herself a
dominant male. Dominant males may possess harems which have up to 40
females by the end of the season. Thus, females directly have no choice, their
behaviour indirectly has the effect.

Fig. 8.18: Male-male fight in elephant seal

Bower bird adopts a strategy to harm other males’ efforts to gain reproductive
advantage; they dismantle other bowers. In this way, they reduce the
opportunities for competitors to copulate.

In Iguanas, the dominant males are large in size; they remove small mounting
males from the back of the females. So female disapprove of mating by the
smaller males.

Alternative mating tactics in subordinate males

The subordinate males are at a reproductive disadvantage. To get the best out
of worst situation, they adopt alternating mating strategies.

Savanna baboons: The low-ranking males develop friendship with the female
by offering protection to her offspring. Male baboons also form friendship
alliances. This male-male cooperation is done to oppose alpha males
accompanying a fertile female. Sometimes, they collectively confront to
dominant male and force him to give up the mating partner.

Iguanas: In marine iguanas living in the Galapagos Islands, the males differ in
size; the large-sized male has reproductive advantage. When a small male
mounts a female the dominant big males remove him from the back of the
female. Normally, it takes three minutes for a male iguana to ejaculate.
Therefore, the chance of a small male inseminating a female is less as the
mating is forcefully terminated before ejaculation. To overcome this, the small
male Iguanas evolved an alternate strategy; to save time; they release sperms
and retain them before the copulation attempt. In the course of mounting, the
males do not have to wait for ejaculation; the smaller one can transfer the pre-
194 ejaculated sperm to the female and inseminate her.
Unit 8 Mating Behaviour
Horned scarab beetle Onthophagus nigriventris make both developmental
and behavioural decisions to get the best in the existing conditions. During
larval growth, if larval nourishment is poor the male shifts its nourishment
away from growing of horn to sperm-producing testis. The minor male enters
the burrow where the dominant male guards the female and tries to
inseminate the female by passing a large number of sperms (Fig. 8.19).

Fig. 8.19: Horned scarab beetle, Onthophagus nigriventris

Scorpion flies, Panorpa adopt a conditional strategy to optimize reproductive

efforts. The large alpha males offer dead insects as a nuptial gift highly
preferred by the receptive females. The beta males secrete saliva on the leaf
and wait for the occasional female to come and consume the nutritional gift;
the male mounts on the female when the female consumes the gift. The
gamma males are even smaller; they have nothing to offer, and they grab the
females for forced copulation. The three types of males adopt a strategy
based on their social standing.

Sandpiper-ruff shows three phenotypes i. e. territorial independents,

subordinates and female mimics. Territorial independent males have
predominantly dark ornamental plumage; they defend small mating courts on
leks. Non-territorial satellite males have predominantly white ornamental
plumage, join independents on courts, co-display to attract females and
compete for mating. Rare small, female-mimicking faeder males attend the
courts of independent males and attempt rapid copulations when females
solicit mating from ornamented-displaying males.

Similarly, marine isopods show three phenotypes concerning their

behavioural responses. These are large alpha, medium beta (mimics female)
and small gamma males. The behavioural phenotypes are under the influence
of a single gene represented by three alleles and are inherited. All three
phenotypes are equally fit.

SAQ 5
Distinguish between Alpha male and Subordinate male.

8.6.2 Intersexual Selection (Female Choice)

Intersexual selection involves both males and females. Mate choice is an
important aspect of mating systems. Usually, female is the choosier sex when 195
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making mating alliances with the males. This is for the reason that the females
make higher investment in each gamete than males. Additionally, in most
species, females provide parental care. Therefore, the females carefully select
their mates to remain safe as far as the risk of their reproductive investment is
concerned. The females prefer certain males having qualities such as "good
genes", to ensure better survivorship and reproductive potential of the
offspring, good parenting by the male, or possession of resources to support
the offspring during their growth and development. The males face a strong
selection for traits that are favoured by females. Most females look at these
traits as indicators of their partner's fitness. Elaborate ornamentations and
colourations of males are good indicators of male fitness; therefore, such
males win the chance to mate with a particular female.

The male mate-choice is less common in animals. However, it is seen in the

cases where male reproduction contribution is significantly high i.e. the males
offer expensive nuptial gifts to the mating partner, are involved in parental
care, or when there is great variation in the quality of the females as mates
within a population. In these cases, females may evolve ornamentation or
coloration in response to sexual selection.

Nuptial gifts

The female choice of the male may be influenced by the nuptial gifts offered
by him. Males of many other animals present and provide food before or
during copulation. In chimpanzees, the male hunts and offers the meat to the
females; the female selects the successful hunter for mating. The male dung
beetles make a dung ball from cowpat and roll it to the burrow (Fig. 8.20); the
female accompanies him to the burrow. In this way, he is awarded with the
mating in return for the nuptial gift. In black-tipped hanging fly both the
quality and quantity of the nuptial gift decide the selection behaviour of the
female (Fig. 8.21). The male offers insect food to the female; if the food is
unpalatable, the female rejects the male. The size of the nuptial gift is also
important as the male is allowed to copulate only till the meal lasts. For
example, if it is consumed within five minutes the female separates from the
male without accepting any sperm. If the feeding on the nuptial gift continues
for twenty minutes, the female receives a complete sperm complement. A
male firefly adds a packet of protein to the ejaculate which adds to the
nutrition of the females. In some species of the firefly, the female evaluates the
male based on the duration of light flashes produced by them. It is seen that
there is a direct correlation between the duration of light flesh and the size of
spermatophore transferred by the males.

196 Fig. 8.20: Male dung beetles carrying dung ball, nuptial gift for the female
Unit 8 Mating Behaviour

Fig. 8.21: black-tipped hanging fly

In the case of the praying mantis and the spiders, the female feeds the
sexual partner while copulation is still going on (Fig. 8.22). In some cases, it
has been seen that the male after wrapping up copulation, turns himself into a
meal for his sexual partner. The male after inseminating the female, blocks the
female’s sperm receptacles with his two sperm-transferring organs, performs a
somersault, and throws his body to the partner's jaws; the female accepts this
sacrifice of the male and devours the sexual mate (Fig. 8.23). The deceased
male redback spider males get a higher reproductive advantage. It is seen that
in the cases where the male is cannibalized by the female, she uses his sperm
for egg fertilization; a fed female is less likely to remate. On the other hand, if
the male is not cannibalized by the female, chances of egg fertilization by his
sperm remain less; in this case, the female goes for remating. Moreover, the
cost of postcopulatory death is very low for the male, as the chances of the
male finding a second mate are very low as while searching mate it gets
preyed to the predatory ants.

Fig. 8.22: Female praying mantis feeds the sexual partner while
copulation is still going on 197
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Fig. 8.23: Redback spider, the male after mating offers himself to the
cannibalistic female partner

Female mate choice for paternal males

In many animals the males contribute to the parental care, therefore, the
females in these cases select the males that can provide better parental care
to the developing progeny than average. In fifteen-spined stickleback, male
shaking their body during courtship is preferred. Such males can also perform
nest fanning after courtship. Therefore, they are useful for sending oxygenated
water into the nest, which increases gas exchange and egg-hatching success.
Males of three-spined sticklebacks exhibit their parental helpfulness by
displaying their red colourful bellies; this behaviour helps them to be selected
by the female. A male's redder belly reflects their carotenoid content, a
reflection of their health. The males with redder bellies can fan the eggs for
longer periods in low oxygen conditions than the males with paler bellies. This
appearance advertises his ability to supply oxygen for the developing brood.

Many birds use the colour of plumage or bill to reflect and advertise their
health and reproductive fitness. These males show better parental behaviour.
Also, the quality of an individual immune system is enhanced by a diet rich in
carotenoids. Males with bright plumage are, therefore, preferred by the female.
In Zebra finches, the males when fed on a diet containing extra carotenoids,
contain more carotenoids in their blood, have brighter red beaks and stronger
immune systems (Fig. 8.24). Therefore, they are more attractive and preferred
by the females for mating. Also, the female prefers the males given the red leg
bands and lays large-sized egg. This means that in the presence of a male
who can support parenting, a female is benefitted by investing more in egg
production. Blue tit, a songbird, shows carotenoid-based plumage ornament
and bright yellow breast. The male collects and delivers food in the form of a
carotenoid-rich caterpillar to the nestlings. It has been seen that the offspring
of brighter parents are healthier with a stronger immune system than the ones
198 which are fed by less yellow parents.
Unit 8 Mating Behaviour

Fig. 8.24: Male Zebra Finch with red bright beak

Female mate choice without material benefit

In many animals, the choosy female does not receive material gains from the
male partner. For example, the Japanese damselfly evaluates the males
based on their body temperature. The thorax of the male is warmer in the
males, who occupy streamside territory exposed to the sun. These males also
beat their wings faster at the time of courtship flight. A female prefers to
oviposit in relatively sunny and warm locations, which allows better
development of the young ones. Therefore, selecting and mating with a male
with a high body temperature provides a reproductive advantage to the
female.

Sensory stimulus provided by the male at the time of courtship, makes him
more attractive to the female. In the bower bird, a female prefers the male to
have more ornaments in the bower and display sexual display in the form of
courtship dance (Fig. 8.25). A female canary bird chooses and selects the
male based on their ability to sing and certain portion of the song. The golden-
collared mankin, a tropical bird exhibits complex courtship behaviour in the
form of a jump-snap display. The faster display makes the male more
attractive and more likely to acquire mates.

Fig. 8.25: Ornamented bower of Bower bird 199

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In some animals, male performance during or even immediately after the
copulation suggests his quality. The tactile signals provided by the male
dagger bee mounted on the already inseminated female affects her readiness
to mate again.

Evolution of extreme male ornamentation and striking courtship

displays
In the absence of any material benefits to the female, what makes the
evolution of extreme male ornamentation and striking courtship displays?
Males benefit by acquiring attractive features, but why do females choose
males with features that have no survival value? E.g. the enormous tail of a
peacock (Pavo cristatus) is costly to produce, impairs motility and is likely to
hinder escape from predators. Then why these traits are favoured by
evolution? Four theories are proposed to explain this.

1. Healthy mates: The courtship behaviour is indicative of male health.

Unhealthy males can transmit various diseases and parasites to the
female and her offspring. The female judges the health of the male by
his courtship behaviour. For example:a male bower bird with high-quality
bowers is less likely to carry ectoparasite in their feathers.

2. Good genes: The good gene theory proposes that the female choice of
partner with ornaments and showing sexual displays is indicative of good
genes for its viability. In this way, she ensures that her offspring inherit
the viability advantage by inheriting such genes from the father.

3. Runaway selection: It explains that secondary sexual characters of

males are evolved in response to the sexual preference of the females.
The female chooses a sexually attractive male. In this way, the sons
inherit the traits that make them sexually attractive. Ronald Fischer
proposed that females mate with attractive males and have attractive
sons with attractive inherited features. These sons, in turn, will be
successful in attracting females and in reproducing themselves. In this
way, the females who choose mates based on sexual attractiveness are
likely to have sexy sons and more grandchildren. The daughters inherit
the majority mate preference from the mother. In this way, mate choice
genes and genes for preferred male attributes are inherited together.
This could generate a runaway process in which extreme female
preferences and male attributes evolved. In runaway selection, the male
character is more exaggerated and female preference of the trait
increases. It is mutualistic coevolution of trait and preference of trait.

4. Chase away selection: In chase away selection, the female fitness

decreases with increase in the male fitness or vice versa. A male as
a result of mutation gets a novel display trait. This taps into preexisting
sensory bias. Such a male induces the female to mate with him. This
male may not provide any material or genetic benefits to the female.

8.7 COURTSHIP BEHAVIOUR OF THREE-SPINED

STICKLEBACK GASTERSTEUS ACULEATUS
Three-spined stickleback, Gastersteus aculeatus is a small freshwater fish of
greyish green colour. They live peacefully in schools having no leader. The
200 day length acts as a stimulus; sex hormones are secreted in response to
Unit 8 Mating Behaviour
longer day length which promotes fish migration from deep to shallow water
resulting in a change in the ambient temperature. At this time, the males
exhibit a change in appearance and behaviour. The male’s belly becomes red
colour, the sides iridescent blue and the eyes emerald green. No colour
change is observed in the females; however, the belly gets enlarged and fully
loaded with eggs. At this time, the males leave the school and set up a
territory in an area of sandy bottom. The territory is extended upwards and
outwards as well. The male builds a nest in a sandy pit with the help of algae
and weeds; the sticky secretions from the kidney act as glue. The nest is given
a tunnel shape. The nest serves as a focus for courtship and a receptacle for
eggs spawned by females. The nest site choice, composition and construction
all affect the quality of the nest and its efficacy. The territory is guarded by the
male by vertical position, a threat posture, and exposing red colour to the
intruder.

After the nest building is complete, another colour changes in the male i.e. the
upper portion of the body changes to blue. This colour attracts the female. If
the nest is not ready, the male is too aggressive and frightens the female.

The ripe females are bold intruders, they do not go away even when butted
and bitten by the male. Rather they adopt an appeasement posture - hanging
obliquely upwards in the water. This posture displays the belly of the female.
The egg-filled belly of the female stimulates the male to display courtship
behaviour in the form of a zigzag dance.

Fig. 8.26: Courtship behaviour of three-spined stickleback

During courtship, the male swims jerkily with alternate leaping movements
towards the female; sideways, jumps away from her, towards the nest, and
circles around her. In this way, he leads the female towards the nest. The 201
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male performs a zigzag dance till the female follows to the nest. An unripe
female fails to respond to the sequences, then the male chases her away. The
male shows her entry into the nest by thrusting his snout into the nest and the
female pushes into the nest. In case, the female does not, the male starts
gently tapping with his head against her tail. The female after entering the nest
spawns and swims out from the other side. The male follows the female in the
nest and releases the sperm over the eggs to fertilize. The male also repairs
the nest and invites other females to spawn. Sometimes when one female is
spawning, and the male observes another female in the territory, he rushes for
the second female; bringing her to the nest without fertilizing the eggs of the
first.

After fertilizing the male begins parental care, comes to its original colour, and
aerates the eggs by fanning movements. After the eggs hatch about a week,
the male guards the young one. The male carries hatchlings in the mouth and
spits back into the nest (Fig. 8.26).

8.8 SPERM COMPETITION AND POST-

INSEMINATION TACTICS
Sperm Competition: Intrasexual selection decides the mating rights of sexual
partners. Postmating, a sperm war, is seen in many animal groups including
insects. If more than one male mates with a female in a short period,
competition can occur among sperms for the fertilization. The male tends to
ensure that the female uses his sperm for fertilization and displaces the sperm
of the rival males. Sperm competition is a form of male-male competition that
plays an important role in mating systems. Several mechanisms have evolved
to facilitate a male's reproductive success with females that have multiple
mates.

Sperm displacement: Male dragonflies and damselflies genitalia have

lateral horns and spines that enable them to draw out sperms and displace
sperms of other males before mating. In Calopteryx maculata, the black-
winged damselfly male tries to win the war by physically removing rival
gametes from his mate’s body before transferring his own. Males of black-
winged damselflies have an extraordinary penis. A spiky penis acts as a scrub
brush, catching and drawing out any sperm stored in the female sperm
storage system.

Sperm precedence or last male advantage: In case female mates with more
than one male, then the females decide which partner’s sperm will fertilize her
eggs. The female may remove the sperm of a male, whom she doesn’t favour
by copulating with another male. It was seen that the male who mates last has
an edge over other males in fertilizing the ova. The females store sperm in
tubules that are filled up from the back; therefore the sperm to enter last are
first to be released. Female bird lives and mates with a social partner but they
may also engage in extra-pair mating with other males. Females of the
collared flycatcher - use their sperm storage site to bias male fertilization
chances by copulating in ways that give one male sperm an advantage over
202 another.
Unit 8 Mating Behaviour
Post-insemination tactics: After the completion of mating and copulation, the
male ensures that the female does not mate with another male and only his
sperms are utilized for fertilization. In this way, he contributes his genotype to
the progeny. The male adopts different post-insemination tactics to achieve
this. The strategies are costly to the males in terms of time energy and
resources; however, benefit the male by reducing the probability of a female
copulating with other males which dilutes or removes the sperms of the
guarding male.

Mate guarding: Mate guarding is adopted by males to reduce the chances of

a female copulating with another male. The males of the Mexican whiptail
lizard were reported to be engaged more in fighting than foraging after mating.
The male New Zealand stitch bird chases other males away from their
previous sexual partner; in this process, they lose about 4% of their body
weight. In a baboon, the female attracts additional sex partners. The dominant
males must guard the fertile partners to maintain exclusive reproductive
excess.

Remain in company or mating position till oviposition: The males may

remain in the company of the mated female constantly to ensure that other
males do not visit her and she does not mate a second time at least before
oviposition. The male blue band goby fish in the Indonesian reef closely
accompanies his mate and follows her wherever she goes. Males of the bee,
Centris pallida inseminate the female quickly but post-mating remains with her.
During this time the pair flies to the nearby bush. The blue milkweed beetle
remains mounted on the female back for some time after copulation. Similarly,
in cotton stainer, Dysdercuskoenigii the male remains in the mating posture
far beyond the time needed for insemination (Fig. 8.27). The male of red
damselflies grasps its mate in the tandem position so that she cannot mate
with another mate (Fig. 8.28).

Fig. 8.27: Male cotton

stainer, Dysdercus
koenigii, the male
remains in the mating
posture far beyond
the time needed for
insemination

Fig. 8.28: The male of red damselflies grasps its mate in the tandem position so
that she cannot mate with another mate

Blocking of genital ducts: The males in many species of crabs and

butterflies block the genital ducts of the females by introducing mating plug.
In about 1% of butterfly species, the male deposits sphragis, an external
mating plugs to block the genital ducts of the female. The sphragis can be
simple in the form of a scab or a blob of petroleum jelly or with distinct
architectural forms (Fig. 8.29). 203
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In some spider species, the male amputates his pedipalp, the sperm-
transferring appendage during copulation and blocks the genital openings of
the females. This sperm-containing appendage acts as a mating plug and also
continues to pump sperm into the female duct, even after the male is eaten
away by the cannibalistic mate. An extreme case is seen in Orb weaving
spiders; the male expires after inserting the pedipalp, the sperm transferring
appendages into paired genital openings of the female.

Induction of monogamy: In Drosophila, the male transfers, paragonial

substances, secretion of male accessory glands to the female. The paragonial
substance suppresses the mating receptivity of the female; consequently, the
mated females exhibit mating refractory behaviour. Induction of monogamy in
the females by the male accessory gland secretions has also been reported in
the mosquito Aedes aegypti.

Fig. 8.29: In butterfly, the male deposits sphragis, an external mating plug, to
block the genital ducts of the female

SAQ 6
i) Define (a) Sperm precedence (b) Sperm competition

ii) Match the items given in Column A with the items in Column B:

Column A Column B

a) Drosophila I. Remain in company of female

after mating

b) Butterflies II. Sperm displacement

c) Blue band goby fish III. Blocking of genital duct

d) Dragon flies IV. Induction of monogamy

204
Unit 8 Mating Behaviour

8.9 SUMMARY
Let us summarise what you have learnt so far:

• In animals, the male and female differ in their morphology, physiology

and behaviour on account of their differential contribution to the progeny
in terms of time, energy and resources; this brings asymmetry of the sex.

• Operational sex ratio is the ratio of sexually active males and females at
a given time. The males produce a large number of sperms, with little
investment, therefore, at a given time more sexually active males are
available. On the other hand, the number of eggs produced by the
female is limited and investment is high. This makes the less availability
of sexually active females. As a result, the operational sex ratio is biased
towards males. The male is an attractive sex and the female is a
selective sex.

• When the male has to offer expensive nuptial gifts to the female; the
availability of males decreases. The operational sex ratio is female-
biased; this causes sex role reversal. The females are attractive and the
males are selective. The operational sex ratio may also be dependent on
the availability of food i.e. in one set of environmental conditions it is
male-biased while at other times it may be female-biased.

• Reproductive fitness of an individual refers to the number of viable

offspring contributed by it. In this way, an individual represents his genes
in the next generation.

• Absolute fitness measures the number of offspring contributed to the

next generation. Relative fitness indicates the fitness of an allele or
genotype in comparison to the maximum fitness.

• Mate choice, sexual selection, reproductive strategies and parental care

are some of the important factors which decide the reproductive fitness
of an individual.

• The difference in the morphology of the males and the females is termed
as sexual dimorphism. Males are usually large; bright in colour and bear
costly ornaments in the form of plumage, horns, antlers and tusks.
Besides, the males produce elaborate songs and exhibit courtship
behaviour and attractive sexual displays.

• The characteristics of males are expansive in terms of investment and

disadvantageous in terms of survival. But they are selected by evolution
as they offer reproductive advantage to the male.

• Three general mating systems are seen in animals i.e. monogamy,

polygamy (polygyny and polyandry) and promiscuity.

• In monogamous systems, one male and one female are paired for at
least one breeding season. These partnerships may last even longer,
sometimes an entire lifetime. In the monogamous mating system, the
males provide substantial parental care. 205
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• In a polygynous mating system, one male mates with multiple females.
The female contributes to most of the parental care. In a polyandrous
mating system, one female mates with many males.

• In a ‘promiscuous mating system’ both the males and the females mate
with multiple partners. There exists no pair bonding and the mating takes
place randomly.

• Sexual selection is a type of natural selection that favours individuals

with certain inherited characteristics, to obtain mates more successfully
than others. Sexual selection has two components, intrasexual
competition and intersexual selection.

• Intrasexual selection is responsible for competition among the members

of the same sex for acquiring mates. Usually, the males compete to
attract females for mating. This results in male-male rivalry and fighting;
the male evolved weapons in the form of horns, antlers, clubbed tails,
and enlarged spiny legs.

• The males also fight to attain dominance or alpha status. The alpha male
gets the privilege to mate with the reproductively active male. The beta
or subordinate males adopt alternative tactics to get the best out of the
worst situations.

• Intersexual selection involves both males and females. Usually, female

is the choosier sex when making mating alliances with the males.

• The male may offer nuptial gifts in the form of food to the female. He
may advertise his fitness in terms of his health or display parenting skills.

• The male, in many species, exhibits courtship behaviour to mate. Three-

spined stickleback,Gastersteus aculeatus a small freshwater fish exhibits
an elaborate courtship behaviour for mating.

• Sperm competition is seen in the female mated with more than one male
in a short interval of time. The male tends to ensure that the female uses
his sperm for fertilization and displaces the sperm of the rival males. For
this different animals adopt different strategies.

• Sperm precedence is a phenomenon in which the sperms of the last

male in a mating sequence are used for the fertilization of eggs.

• Male dragonflies and damselflies remove the sperm of previous mating

before transferring their sperms to the female genital ducts.

• The males after insemination, attempt to prevent the female to mate with
other males. Territory guarding, constant companionship, blocking the
genital ducts of the mated female, and induction of monogamy in the
female are some of the post-insemination tactics adopted by the males.

8.10 TERMINAL QUESTIONS

1. Describe the asymmetry of the sex. Explain morphological, physiological
and behavioural differences observed in the two sexes to maximize their
206 reproductive contribution to the next generation.
Unit 8 Mating Behaviour
2. Explain the role of the operational sex ratio in mate attraction and mate
selection behaviour.

3. Explain key features of different types of mating systems seen in animal

kingdom.

4. Describe intrasexual selection. With suitable examples explain the

strategies adopted by the males to maximize their reproductive
contribution to the next generation.

5. Describe intersexual selection. Explain various tactics adopted by males

in response to intersexual selection.

6. Describe sperm competition and post-insemination tactics followed by

the males to ensure their paternity in the progeny.

7. Describe courtship behaviour in a 3-spined stickleback.

8. Write short notes on:

i) Post-insemination mate guarding

ii) Evolutionary importance of sexual dimorphism

iii) Reproductive fitness

8.11 ANSWERS
Self-Assessment Questions
1. i) (a) Asymmetry of sex: Males and females contribute differently
in terms of the amount of energy, time, and resources
contributed towards parental investment in sexual
reproduction. This causes evolutionary differentiation of
males and females in terms of morphology, physiology and
behaviour

(b) Operational sex ratio: The operational sex ratio is the ratio
of sexually receptive males and sexually receptive females at
a given time at a particular place. This ratio decides the role
of the two sexes in attraction or selection in mate finding.

ii) (c) The males will be attractive and the females will be selective

iii) (d) The number of sexually active males is more than the
number of sexually active females.

2. i) Reproductive fitness: The reproduction fitness of an individual is

the number of viable offspring produced by it in a population. In
this way, it passes its genes to the next generation

ii) Absolute fitness measures the number of offspring contributed to

the next generation. Relative fitness indicates the fitness of an
allele or genotype in comparison to the maximum fitness.

iii) Premating 207

.

Block 2 Systematic Approach to Understand Important Animal and

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3. i) (a) Sexual monogamy: Sexual monogamy is true monogamy;
both partners mate only with each other. It is exceedingly
rare.
(b) Polygamy: Polygamy refers to the mating system in which
one partner mates with multiple partners i.e. one male
mating with many females (polygyny) or one female mating
with many males (polyandry).
(c) Promiscuity: In a ‘promiscuous mating system’ both the
males and the females mate with multiple partners.
ii) (a) Polygamy- Polygamy refers to the mating system in which
one partner mates with multiple partners i.e. one male
mating with many females (polygyny) or one female mating
with many males (polyandry).

Promiscuity: In a ‘promiscuous mating system’ both the

males and the females mate with multiple partners.
(b) Serial monogamy is a mating system in animals where they
pair with a mate for one mating season but change mates
throughout a lifetime.
In Social monogamy, two individuals are involved in rearing
their offspring, but they also engage in “extra-pair
copulations,” with another individual.
(c) Polyandry: In polyandrous mating systems, one female
mates with many males.

Polygyny: In polygyny, one male mates with many females.

In this system, usually, the female takes responsibility for
most of the parental care.

iii) a) Cooperative polyandry

b) Promiscuity

c) Social monogamy

d) Lek

e) Harem
f) Resource defence polygyny

4. i) intersexual; ii) intrasexual

5. Alpha males are the social dominance in the hierarchy. Generally, they
mate more often than subordinates and get paternity benefits.

6. i) a) Sperm precedence: Last male advantage-It was seen that

the male who mates last has an edge over other males in
fertilizing the ova.
b) Sperm competitopn: If more than one male mate with a
female in a short period, competition can occur among
sperm for the fertilization.

208 ii) a) IV, b) III, c) I, d) II.

Unit 8 Mating Behaviour
Terminal Questions
1. Refer to Section 8.2.

2. Refer to Section 8.2.

3. Refer to Section 8.5.

4. Refer to Subsection 8.6.1.

5. Refer to Subsection 8.6.2.

6. Refer to Section 8.8.

7. Refer to Section 8.7.

8. a) Refer to Section 8.8.

b) Refer to Section 8.4.

c) Refer to Section 8.3.

Acknowledgement of Figures
Fig. 8.1: [Link]
_-_cropped.jpg

Fig. 8.2 [Link]

m/

Fig. 8.4 [Link]

Fig. 8.5 [Link]

17/issue-2/1082-6467-17.2.231/Understanding-nuptial-gift-size-in-
bush-crickets--an-analysis/10.1665/[Link]

Fig. 13.5: [Link]

Fig. 8.6: [Link]

great-giving-embryos-oxygen

Fig. 8.7: [Link]

k_courting_peahen.jp

Fig. 8.8: [Link]

er_stag_2009_denmark.jpg

Fig. 8.9 [Link]

africa-tusks-african-elephant-wild-wildlife

Fig. 8.10 [Link]

Fig. 8.11: [Link]

sciences/steller-sea-lion

Fig. 8.13: [Link]

Fig. 8.14: [Link]

_Colorado_-_USA_99_0004_%2815519986626%[Link] 209
.

Block 2 Systematic Approach to Understand Important Animal and

Human Behaviours
Fig. 8.15: [Link]

[Link]
wildlife-male-antlers-horns

Fig. 8.15: [Link]

Fig. 8.16: [Link]

Fig. 8.17: [Link]

Fig. 8.18: [Link]

[Link]

Fig. 8.19: [Link]

Fig. 8.20: [Link]

e_%28Pachylomerus_femoralis%29_rolling_a_ball_of_elephant_dun
g_..._%2846628617921%[Link]

Fig. 8.21: [Link]

Fig. 8.22: [Link]

[Link]

Fig. 8.23: [Link]

Fig. 8.24: [Link]

Fig. 8.25: [Link]

ptilonorhynchus-violaceus

Fig. 8.26: [Link]

spined_stickleback#/media/File:Gasterosteus_aculeatus.jpg

[Link]
spined_stickleback#/media/File:[Link]
G

[Link]
spined_stickleback#/media/File:Unsere_S%C3%BC%C3%9Fwasserfi
sche_(Tafel_4)_(6102595905).jpg

Fig. 8.27: [Link]

Fig. 8.28: [Link]

Fig. 8.29: [Link]

_sphragis_03_%28HS%[Link]

210
 UNIT 9
PARENTAL CARE

Structures
9.1 Introduction Parental Care in Amphibians
Objectives Parental Care in Reptiles
9.2 Concept of Parental Care Parental Care in Birds
Types of Parental Care Parental Care in Mammals
Provided by the Parents 9.7 Cost Benefit Analysis of
9.3 Types of Offspring Parental Care
Altricial The Cost of Parenting
Precocial The Benefits of Parenting
9.4 Parental Investment 9.8 Evolutionary Stable
Theories Strategy
Triver’s Theory 9.9 Familial Conflicts
Dawkins and Carlisle’s Theory Parents-Offspring Conflict
Williams’s Hypothesis Sibling-Sibling Conflict
Certainty of Paternity Parental Favoritism
Hypothesis 9.10 Parental Manipulation
9.5 Types of Parental Care Maternal manipulation in
Maternal Parental Care Eusocial Society
Paternal Parental Care Brood Adjustment
Bi-Parental Care 9.11 Brood Parasitism
Allo-Parental Care Intra-species Brood Parasitism
9.6 Parental Care Across Inter-species Brood Parasitism
Various Phyla 9.12 Summary
Parental Care in Insects 9.13 Terminal Questions
Parental Care in Fishes

9.1 INTRODUCTION
One of the most intriguing and patient behaviours shown by an animal is
taking care of their eggs or young ones. Ask any human parent and they will
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
let you know although it is rewarding but parenting is the one of most difficult
task handled by them and it is same for animal parents also. Parenting in
animal world comes in various shapes and forms. From a male giant water
bug carrying its eggs on back to tigress teaching its cubs to hunt to mother
monkey carrying, feeding and grooming its baby all exhibit some or other form
of parenting. In the last Unit we learned about intra-sexual and inter-sexual
selection.

This unit takes you on a journey how post mating an embryo or an offspring is
taken care of. Here we also observe the differential investment of each sex in
parenting duties depending on their phyla. Here various theories will be
discussed that tries to explain differential investment of male and female
member of species in parental care. The key to explaining variety in parental
care lies with the cost benefit approach. The benefits of parental care is the
improved survival of the assisted offspring and costs are reduced fecundity in
future and fewer mating opportunities in the present. This unit will discern
various types of parental care including maternal, paternal, bi-parental and
allo-parental and what’s the evolutionary cause behind it. In this unit,
evolutionary stable strategies and brood parasitism will be discussed to have
better understanding of this complex behaviour. Once finished with this part
we can progress into similarities and dissimilarities between human
behaviours in comparison to other animals. The next unit will showcase the
human behaviour and how it is being regulated.

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

 explain the concept of parental care,

 describe various types of parental care in nature,

 give reasons for differential parental investment by male and female of

a species,

 elucidate numerous parental care examples across phyla,

 determine the cost and benefits of parental care,

 define what is Evolutionary Stable Strategy,

 define parental manipulation,

 explain about the various familial conflicts and how they are resolved,
and

 describe brood parasitism and its sub-types.

9.2 CONCEPT OF PARENTAL CARE

Any behaviour on the part of either one or both parent that enhances the
chances of survival of offspring is termed as parental care. Sometimes
besides parents, other members of the family also help in taking care of the
212 offspring.
Unit 9 Parental Care
In animal world, most species do not provide any kind of parental care. The
gametes are released in many species outside the body (mostly in water) in
case of external fertilization. In these cases eggs are released outside and
sperms fertilize them with no-body taking care of zygote or embryo. In case of
internal fertilization, post fertilization zygote is abandoned. Most species that
do not provide parental care produce large number of gametes and thus large
number of uncared offspring.

Those species that do provide parental care do so at the cost of having more
progenies. So they will have less number of progenies, but chances of survival
of these offspring are much higher compared to uncared one. Parental care
increases inclusive fitness in a species and usually conflicts with sexual
selection pressure.

9.2.1 Types of Care Provided by the Parents

Nest/Home Building: Providing shelter or safe space for the eggs or the
offspring.

Brooding: Providing proper temperature to eggs or offspring.

Feeding: Taking care of the nutritional needs of an offspring (Fig. 9.1).

Defense: Providing protection from predators.

Teaching Skills: Teaching various survival and behavioural skills to offspring.

Fig. 9.1: A female calliope hummingbird feeding fully grown chicks

9.3 TYPES OF OFFSPRING

In this Section, you will study about types of offspring of different species and
the type of parental care needed by them.

9.3.1 Altricial Species

Species whose offspring are born completely helpless and require parents to
feed and take care of them during initial stages of life are called altricial
species (Fig. 9.2). Altricial word comes from a Latin root meaning “to nourish”
in reference to the need for extensive parental care.

Examples: Hawks, woodpecker, owls, swallows, marsupials, domestic dogs,

cats, human, etc. 213
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Fig. 9.2: Altricial chicks of white-breasted wood swallow

9.3.2 Precocial Species

Species whose offspring are born relatively mature and can move around from
the moment they are born or hatch are called precocial species (Fig. 9.3).
The word comes from the same Latin root as ‘precocious’.

Examples: Domestic chicken, ducks, geese, guinea pig, most ungulates etc.

Fig. 9.3: A precocial Muscovy duckling

SAQ 1
a) Define parental care in animals with suitable examples.

b) Differentiate between altricial and precocial species.

9.4 PARENTAL INVESTMENT THEORIES

This section will discuss about various theories of parental investment.

9.4.1 Trivers’s Theory

This term was coined by Robert Trivers in 1972. It hypothesizes which sex will
be more inclined to provide parental care. It connects sexual selection with
parental care. According to this the sex that invests less in parental care will
214 invest more in trait that help them getting chosen by mate (intra-sexual
Unit 9 Parental Care
competition) and the sex that will invest more in parental care will be more
choosy and will be more selective of the mate (inter-sexual selection). This
suggests, since males invest in trait that help them outcompete fellow males of
same population, they are less invested in parental care and since females
usually invest less in intra-sexual competition and choose mate very carefully,
they are usually more involved in parental care giving.

9.4.2 Dawkins and Carlisle’s Theory

This theory given in 1976 by R. Dawkins and T.R. Carlisle, suggests that
parental care depends upon the order of release of gametes and the chance
of mate desertion. In case of fishes, female lays eggs before fertilization by
male, giving them chance to desert first and leaving fertilized eggs to be taken
care by male. On the other hand, internal fertilization provide males the
chance to desert first and leave the offspring to be taken care by the mother.
This tries to explain mostly paternal care in fishes and maternal care in
mammals.

9.4.3 Williams’ Hypothesis

Hypothesized by George William in 1975, this correlates association between
embryo and parental care. According to this, in case of external fertilization,
female lays eggs in male territories that are fiercely guarded by males. Now
males will fertilize them and remain in close proximity and create an
association that causes males to take care of them. In case of internal
fertilization, it's the female who are closely associated with the zygote and take
care of the growing embryo and infant.

9.4.4 Certainty of Paternity Hypothesis

This hypothesis states that males invest in parental care when they are sure of
the paternity. In case of fishes and amphibians, external fertilization takes
place in aquatic environment. In external fertilization, female lays eggs first as
they are larger in size and have less chances of being swept away by water
current. Male later fertilizes the eggs as sperm are smaller and can get lost if
released first. So in external fertilization males are sure of paternity and
paternal parental care is higher in fishes and amphibians. On the other hand in
internal fertilization males are not sure of paternity and do not invest in
parental care. In most cases of internal fertilization maternal care is seen.
Male members of the species contribute along with female in bi-parental care
mostly in monogamous relation as seen in the birds.

SAQ 2
a) Explain William’s Hypothesis of parental Investment?

b) How Trivers’s theory of parental investment is different from Dawkins

and Carlisle’s theory?

c) Define Certainty of paternity hypothesis.

215
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9.5 TYPES OF PARENTAL CARE
9.5.1 Maternal Care
The animal where parental caregiver is female of the species comes under
maternal care. This is most common among all types of parental care.
Females either alone or with the help from father or other family members take
care of the young ones.

In viviparous animal, since zygote develops inside the body of mother, the
gestation and lactation (in case of mammals) brings hormonal changes in the
body of female making them predisposed to parental care. Many mammals
like tiger, leopard, and polar bears show parental care by females only.

In case of orangutans, the babies are entirely dependent on their mother and
breastfeed up to eight years in some cases. Young orangutans learn survival
skills from mother like searching for food, differentiating between good and
bad food and remaining steer clear of predators. The maternal bond is so
strong in orangutans that even 15years after birth the adult offspring visits
mother from time to time (Fig. 9.4).

In case of oviparous animals, females are seen to take care of the clutch and
also providng care post hatching. Many females of crocodilian species provide
parental care for months to its offspring. In invertebrates, maternal care forms
the core of eusocial groups like honeybees.

The hypothesis for female as primary or sole caregiver is explained on the

basis of gamete size. Female produces large sized and fewer gametes,
whereas male produces gametes small in size, but in large number. This way,
females invest more in gametes and are more invested in the survival and
wellbeing of her offspring.

Fig. 9.4: Mother Orangutan with her baby

9.5.2 Paternal Care

Paternal care is the care provided by the male member of the animal species.
Male-only parental care is more common in species with external fertilization.
Many species of fishes and amphibians provide male-only parental care to
their offspring. Various species of sea horse are most well studied example of
216 paternal care.
Unit 9 Parental Care
The scientific explanation given for this behaviour is surety of paternity. In
case of external fertilization, mostly females lay eggs first that are later
fertilized by males. In these cases, males are sure that the growing offspring
are theirs and provide them with parental care. In case of polygynous system,
males usually skip the parental caring e.g. Spotted hyena, red winged prinia,
red deer, Bengal tiger, house wren, etc, where as in case of polyandrous
species many times males are the sole caregivers to offspring e.g. as in case
of rhacophorid frog (Kurixalus eiffingeri), acorn woodpecker, etc.

Males in case of internal fertilization are susceptible of raising young ones that
are genetically not their offspring and have reduced fitness due to it. This
makes them to invest less or not at all in parental care, when they are not sure
of paternity. During internal fertilization, male provides parental care in only
monogamous systems. For Example, male emperor penguins incubate the
egg for almost 2 months in harsh winter. Male giant water bug carries all eggs
on his back to provide aeration (Fig. 9.5).

Paternal care is seen rarely in non-monogamous systems and in case of

polygamy it’s the females who are the main or sole caregiver. Exception to this
rule is seen in mountain gorillas. Those gorilla who irrespective of offspring
parentage, provided parental care, are able to attract more females and have
5 times higher chances of fathering a child than males who do not engage in
any parental care.

There are two hypothesis given for male investing in parental care:

1) The mating effort hypothesis: This suggest that those males who
engage in parental care have better chances for attracting females in
next reproductive season and indirectly it increases their reproductive
success.

2) The maternal Relief hypothesis: This hypothesis suggests that male

contributing to parental care duties reduces burden on females. This
helps females to be ready for next mating cycle faster and reduces the
inter-birth gaps. Indirectly this will contribute to more mating chances
and enhances the reproductive success of caregiver males.

9.5.3 Bi-parental Care

When both parents work together to take care of offspring, such system is
called bi-parental care. It is mostly seen in species, where sex ratio in Fig. 9.5: Male giant
water bug
reproductively mature individuals is well balanced. This means, species whose Abedusindentatus
population has almost similar number of sexually mature males and females with eggs on his back
are most likely to engage in bi-parental care.

This type of care involves complete cooperation and coordination among the
parents. Bi-parental care is one of the key factors that lead individuals of a
species to be in a monogamous relationship, at least for the season.

There is no evidence that suggests that bi-parental care is better or worse

than single parent care giving system. In most cases both parents share
workload of parenthood almost equally, providing offspring with excellent
resources. Offspring gets highly benefitted from such arrangement, with
increased survival and fitness. 217
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But in few cases, there is evidence of parent of either sex withholding the
amount of parental care, making conditions worse for offspring. This usually
happens due to high sexual conflict and lack of coordination among parents.

Bi-parental care is most common in birds with 90% of them engaging in it,
followed by mammals. Example of bi-parental care include Indian striped
hyenas, wire-tailed swallow, etc.

9.5.4 Allo-parental Care

When other members of the family besides parents help in taking care of
the young ones, such a system is termed as allo-parental care. Many
times, members of same gender come together to take care of the offspring as
in honeybees, but it can have both parents and other members involved in
parenting as seen in wolves. This system is product of altruism and
development of eusociality. For mammalian mothers allo-parenting helps in
multiple ways. The infant weans earlier with allo-parenting shortening the inter-
birth interval. This improves reproductive success among the members of
species. With outside help, mammalian mother now will have more time to
feed young ones and more energy to produce rich milk. This improves survival
rate and fitness of offspring.

Example of allo-parenting include meerkats, that have one alpha male and
female who do most of the breeding. The rest of the members take care of
offspring and protection of burrow, while parents are out foraging (Fig. 9.6 &
Fig. 9.7).

In invertebrates, example of allo-parenting is seen in eusocial insects like

honeybees and termites. To maintain coordination among members, a proper
caste system is developed. In this single female (queen) and in case of
termites also male (king) are the only reproductive individuals and the rest of
the members sacrifice their reproductive fitness for them. The sterile castes
are responsible for taking care of all the duties of eusocial structure including
caring for reproductive caste members and their offspring.

Fig. 9.6: Meerkat Pups Fig. 9.7: Meerkat family

SAQ 3
a) What is allo-parental care?

b) Differentiate between maternal and paternal care.

c) With suitable example explain bi-parental care.

218
Unit 9 Parental Care

9.6 PARENTAL CARE ACROSS VARIOUS PHYLA

Here we will discuss parental care in invertebrates (insects) and vertebrates
(fish, amphibians, birds, mammals)

9.6.1 Parental Care in Insects

Parental care is not prevalent among the invertebrates with the exception of
insects. Insects not only engage in parental care, but also show variety of
social structure. You may find from solitary social insects, to semi-social and
eusocial ones. The more complex the society, the better parental care is
observed in that species. In hymenopteran insects, bees, ants and wasps are
known to invest heavily in parental care. The female potter wasp, which is
solitary insect is known to build nest for the young ones and stores food inside
for their growth (Fig. 9.8).

Fig. 9.8: Female potter wasp building nest for her offspring

Honeybees, ants and termites are eusocial insects that have complex social
structure with army of workers to take care of all parental requirements of
offspring from egg till adult stage. Paternal care is observed in giant water
bugs. Giant water bug’s eggs are large in size and need proper aeration for
breathing and growth, so male carries eggs on its back till they hatch.

9.6.2 Parental Care in Fishes

Most of the fishes do not show any kind of parental care. Only about 17% of
the fishes are known to show some kind of parental care out of which most
exhibit paternal care. Beside paternal care, some fishes also exhibit bi-
parental and in rare cases maternal care.

Few Examples

Nest Building: Nest Building is used as a tool to attract females and to take
care of fertilized eggs. Nest in case of fishes can be simple depression at the
floor of aquatic body as in case of lungfishes or it can be complex structure.
Male Stickleback fish makes the nest using rootlet and other fragments from
plants and it glues the material together with adhesive secretion from his
kidney (Fig. 9.9). The nest in this case is spherical in shape with a tunnel for
the deposition of eggs. Male attracts female into his nest, and once she lays
eggs inside the nest, it chases her away. The usual clutch size varies from
50-100 eggs. Male fertilizes the eggs deposited in his nest and guards them till
they hatch. Males are also known to fan the eggs with their pectoral fin to keep
them aerated. 219
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Fig. 9.9: Nest of Three-spined Stickleback

Bubble Nest/Foam Nest: Male fish build these nests by producing bubbles
from mouth along with saliva which float at the surface of aquatic body. These
types of nest are usually built in oxygen depleted stand still water bodies and
fishes that build them are called aphrophils. Examples of fishes that build
bubble nest are Gourami (Fig 9.10), eel and Betta fish.

Fig. 9.10: Bubble nest of dwarf Gourami fish

Coiling around Eggs: In this case fish coils itself along the mass of eggs till
they hatch. The Pholis gunnellus, butter fish is known to show such parental
care (Fig. 9.11). This is also an example of bi-parental care as both male and
female take turns guarding eggs.

Fig. 9.11: Butterfish coiled around mass of eggs

Mouth Brooding: This type of parenting involves carrying fertilized eggs in

the mouth of parent fish and incubating it till they hatch. Depending on species
both male and female fish exhibit mouth brooding, most time alone and
sometimes together. Examples of male mouth brooder are American sea
catfish and female are African and South American cichlids (Fig. 9.12). A good
220 example of bi-parental care in fishes is seen in Spatula barbed catfish.
Unit 9 Parental Care

Fig. 9.12: Mouth brooding Female Cyphotilapia frontosa

Egg attached to body parts: In nursery fish, mass of fertilized eggs are
attached to cephalic hook of male fish (Fig. 9.13). They remain attached till
they are hatched.

Brooding Pouch: The most common example of paternal care in fishes is

seen in male sea horse (Hippocampus) (Fig. 9.14). It carries the fertilized eggs
in the brood pouch till they are hatched. Female transfers the fertilized eggs
Fig. 9.13: Nursery
into the pouch present on the belly of the male. The pouch has multiple folds
fish male carrying
into which each egg attaches and is able to respire due to exchange of gases.
eggs

Fig. 9.14: Male Hippocampus with Brood pouch

Male Pipefish also has a brood pouch made up of two flaps at its underbelly
(Fig. 9.15). It protects its fertilized eggs till giving birth to young ones.

Fig. 9.15: Beady Pipefish 221

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Integumentary Cup: In case of fish, Platystacus (Fig. 9.16) and Aspredo
fertilized eggs are carried by females in soft folds in skin known as
integumentary cup till hatching.

Fig. 9.16: Female Platystacus carrying eggs in Integumentary cup

Viviparity: In this condition, the embryo develops inside the body of mother
and takes nourishment from it till it is fully developed. In case of
elasmobranchs many families of sharks exhibit viviparity. Female shark’s
uterus is lined with mucus that forms fluid filled compartments. Each embryo
grows in separate compartment and receives nourishment from uterine wall
via yolk sac placenta.

Some species of teleosts such as Poicilia and Zoarces how viviparity. Here
embryo grows in the ovary, but it is not attached to it. The discharged ovarian
follicle nourishes growing embryo in such species.

Ovoviviparous condition: This condition is seen in nurse shark (Fig. 9.17)

that lays eggs covered with horny case. The eggs are fertilized and then
develop inside the body of the mother shark. The egg hatches out of the shell
inside the uterus and then pups are born after 6 months of gestation period.

222 Fig. 9.17: Nurse Shark

Unit 9 Parental Care
9.6.3 Parental Care in Amphibians
Most amphibians show some or other kind of parental care. Both maternal and
paternal parental cares are almost equally seen across the phylum, but bi-
parental care is rare. This is the group where you find most diverse
parenting behaviour.

It includes taking care of egg clutch, protection from predator, keeping eggs
moistened and even eating fungi infected eggs to safeguard rest of the clutch.

Nest Building: Amphibian showcase variety of nest

Foam Nest: Japanese tree frog (Rhacophorus schlegelii) covers their fertilized
eggs with mucus secretion. Both male and female dig a tunnel together and
leave their mucus-enveloped eggs in there. The mucus keeps eggs hydrated
and prevents their desiccation. Tadpoles hatch during rainy season and
rainwater takes them to nearby water body (a pond or river).

Malabar gliding frog (Rhacophorus malabaricus) produces foam nest on the

trees, where female accumulates her eggs (Fig. 9.18).

The female South American tree frog (Leptodactylus mystacinus) produces

foam nest in a hole near water body. In this case tadpoles directly enter the
water as soon they are hatched.

In case of Adelotus brevis males produces foam nest and also defend the
fertilized eggs actively till they are hatched.

Fig. 9.18: Foam nest of Rhacophorus malabaricus mating pair

Mud Nest: Mud nest are prepared by male Brazilian tree frog (Hyla fabre) by
digging a hole or making a nursery in the mud of shallow water (Fig 9.19).
Female lays her eggs in the mud nest and the tadpoles after hatching move
directly into the water body. 223
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Fig. 9.19: Mud nest of Brazilian tree frog

Leaf Nest: The tiger-striped leaf frog (Phyllomedusa tomopterna) lays its eggs
in gelatinous mass on the leaf above a pond (Fig. 9.20). It then folds the leaf
and seals it to give the shape of a nest. When eggs hatch, tadpoles directly fall
into the water below.

Fig. 9.20: Leaf Nest of tiger-striped leaf frog

Tree Nest: Tree frog, Hyla resinfictrix uses shallow tree cavity as a nest. They
line this cavity with beeswax obtained from the hives of stingless bees. When
these cavities are filled with rainwater, female lays eggs in it and their offspring
develop in a protected environment.

Gelatinous bags: Salamandrella keyserlingi secretes gelatinous bags that

contain 50-60 eggs each. These bags are attached to aquatic plants and when
eggs mature, they hatch directly into water.

Communal Nest: The toad, (Nectophyrynoides malcolmi) prepare communal

nest close to one another. Several females lay eggs in that communal nest,
which are fertilized by a single male. This male will guard the nest until eggs
224 are hatched.
Unit 9 Parental Care
Direct Caring by Parent

Coiling around eggs. Female Ichthyophis glutinosa digs a shallow hole near
the water body. She lays around 30 eggs coated with gelatin and coils around
this fertilized egg mass till they are hatched (Fig. 9.21). Similarly, female
salamander coils around her fertilized eggs that are deposited beneath a rock
or a cavity of rotten wood.

Fig. 9.21: Female Ichthyophis glutinosa with her egg clutch

Carrying eggs on the body: Many amphibians carry their eggs attached to
the body parts till they are hatched. Female salamander (Desmognathus
fuscus) carries eggs attached to her neck and keeps them safe from
predators by attacking them if needed. In common midwife toad (Alytes
obstetricans) fertilized eggs are glued to the hind leg of male (Fig. 9.22),
whereas in Sri Lankan tree frog they are attached to the belly of female.

Fig. 9.22: Male common midwife toad with eggs

Eggs in back pouches: Back pouches are observed in certain frogs and
toads and these are called marsupial frogs or toads. The females have oval
depressions or closed individual pouches in their back, which are used to carry
eggs. Eggs develop into mini-frog before they leave back pouches.

In Brazilian tree frog, female brood pouch is a groove, where eggs remain
exposed where as in Nototrema marsupium they are covered with a layer of
skin. In Pipa pipa skin of female frog during breeding season changes to thick
gelatinous mass, on to which male frog inserts fertilized eggs. In case of
marsupial frog Fritziana goeldii, they carry eggs in back brood pouch and
deposit tadpoles in water (Fig. 9.23). 225
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Fig. 9.23: Marsupial frog Fritziana goeldii carrying eggs in back brood pouch

Organ used as brooding pouch: In case of South American Darwin frog

Rhinoderma darwini, males have large vocal sac. They use it as brooding
pouch during mating season and keep up to at least two fertilized eggs in it.
Post metamorphosis, mini frogs come out of the vocal cord. In Hylambates
breviceps, the females carry the eggs and in Artholeptis, the males carry
larvae in buccal cavity. In Rheobatrachussilus female keeps her eggs in
stomach and metamorphosed tadpoles come out of mouth.

Feeding Larvae: Dendrobates feeds their offspring.

Ovoviviparity: Some anurans keep their fertilized eggs in oviduct for

maturation and give birth to tadpoles.

9.6.4 Parental Care in Reptile

Reptiles have internal fertilization and many leave their fertilized eggs at safe
site for embryo development. Most eggs hatch on their own and young ones
find their way out themselves. Most reptiles do not provide parental care to
their offspring except crocodiles.

Maternal care is very common among crocodilians that includes making nest,
protecting eggs from predators and carrying them in her mouth, post hatching,
to water from the nest. Mother crocodiles are also known to stay with their
young ones for the first few months of their life (Fig. 9.24).

Fig. 9.24: Alligator mississippiensis taking care of young ones

9.6.5 Parental Care in Birds

All birds show parental care with 90% involved in bi-parental care, 9%
maternal and 1% paternal care. The parental care in birds can be divided into
226 three main stages.
Unit 9 Parental Care
Nest Building: Nests are the homes built by male birds that is used for egg
laying and incubation. It is one of the ways male attract females and also take
care of eggs and young ones. In nature there are many variety of bird nests.
Some birds use crevices among rocks or a cavity in the tree as nest (Fig. 9.25)
and lay their eggs there. But most birds build elaborate nests with variety of
material available to them. Many decorate it with pebbles and flowers to make
it attractive. Members of the same species build similar nests utilizing same
material.

Fig. 9.25: A male Blue-winged Parrotlet using tree cavity as nest

The Blackbirds make cup shaped nests of dried grasses (Fig. 9.26), where as
raptors make big platform nest (Fig. 9.27) high up on lofty trees that they use
for multiple breeding seasons.

Fig. 9.26: Common Blackbird cup nest Fig. 9.27: Osprey Platform nest

Weaverbirds and tailorbirds are named for their astonishing nest building
skills. Tailorbird sews the edges of a large single leaf to design beautiful home
out of it (Fig. 9.28). Weaverbird constructs a suspended pendant shaped nest
out grass and plant fiber (Fig. 9.29).

Fig. 9.28: Common

tailorbird’s nest

Fig. 9.29: A male village weaver (Ploceuscucullatus), building his nest 227
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Woodpeckers hollow out cavity in the tree (Fig. 9.30) and kingfisher excavates
riverbank to use it as nest.

Fig. 9.30: Black woodpecker with offspring

Some birds build colonial nest as in case of social weaverbird where multiple
birds build nests on the same tree or European bee-eater that lives in colonial
burrow nest (Fig. 9.31).

Fig. 9.31: European bee-eater colonial nest

Incubation of Eggs: Fertilized birds egg need to be kept at optimal

temperature constantly till they hatch. This is mostly done by sitting on eggs by
the parent birds. In most species both parents take turns to keep eggs warm,
in others its either male or female that incubate eggs (Fig. 9.32). In case of
emperor penguin, male incubates single egg in harsh winter for around two
months. He keeps egg on the top of his feet and keeps it warm till it hatches.
In case of Greylag goose and many species of ducks it's the mother who
incubates the eggs, but the father remains close by and guards them. In case
of doves, pigeons and sandpipers both female and male take turns to incubate
eggs. Period of incubation varies in different species of birds being 13 days in
a canary to 67 days in emperor penguins.

228 Fig. 9.32: Female Mallard Duck incubating her eggs

Unit 9 Parental Care
Taking care of young ones: In case of most birds, chicks require extensive
care post hatching. They need to be fed, cleaned and protected from the
predators. As described earlier precocial birds chick are born with feathers and
they can swim, run post hatching and are fairly independent. Still they require
parental presence to develop learned skills by observing adults. On the other
hand, altricial birds are born featherless, helpless and blind and remain inside
the nest for few weeks before they become independent.

In most cases, parental duties are shared between male and female in birds.
In emperor penguin once eggs hatch female takes over the chick and parental
duties, while male leaves for feeding (Fig. 9.33). Both parents after that take
turns, one brooding and other feeding and raise their chick together.

Crop milk is a
secretion from the
lining of the crop of
parent birds in some
species that is
regurgitated to young
Fig. 9.33: Emperor Penguin taking care of chick
birds. It is found
Feeding in birds involve collection of food by parent and placing in the mouth among all pigeons
and doves where it is
of chicks (Fig. 9.34). In pigeons, the epithelial lining of crop is shredded and
also referred to as
fed to the young one as pigeon milk. pigeon milk.

Imprinting is a type of
innate learning in
animals immediately
after birth or hatching
to recoginize their
caregiver for food and
protection. Birds after
hatching visually
imprint on their
parents during a
critical period of
development.
Fig. 9.34: Parent Swallow feeding its young ones

Many birds, especially raptors like osprey are very protective of their young
ones. They aggressively defend their nest from predators. Birds have long
term social bond with their hatchlings based on imprinting and offspring learn
lots of skills from parents till they leave the nest. 229
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9.6.6 Parental Care in Mammals
The term mammals is derived from presence of mammary glands in females
that produce milk to feed the young ones during their developmental stages.
Mammalian babies are mostly altricial or semi-precocial except seal where
seal pups are precocial. Mammals provide longest and most complex parental
care to their offspring compared to other animals. Mothers are the primary and
mostly the only caregiver (in 95% cases) in mammals (Fig. 9.35) with father
and other family members helping out in some species (5%).

Fig. 9.35: Harbor Seal pup feeding on Mother’s Milk

Parental Care in Prototheria

Duck-billed Platypus is oviparous and lays eggs with much of yolk. The mother
incubates the eggs till they hatch and feed the young ones milk till they
develop to take care of themselves.

Parental Care in Metatheria

Females are viviparous with short gestation period. Young metatherian are
usually born immature and finish their developmental stages in the mother’s
pouch (Fig. 9.36). Although mother feeds the baby milk, it is not highly bonded
with it and once fully developed baby mostly leaves the pouch.

Fig. 9.36: Marsupials: eastern grey kangaroo with baby, Virginia opossum, long-
230 nosed bandicoot, Monito del monte and Tasmanian devil
Unit 9 Parental Care
Parental care in Eutheria

They are called placental mammals as mother nurtures fetus during complete
gestation period via placental attachment. From oxygen, to nutrients, to
immunological protection, everything is provided to fetus and toxins are
removed utilizing placental connection. In eutheria, gestation period is directly
proportional to the weight of the animal i.e. larger the mammals longer will be
gestation. Hamster and mouse have shortest gestation period of 16-23 days
while African bush elephant have longest being around 665 days (Fig. 9.37).

Fig. 9.37: Family of African Bush Elephant

In mammals, post birth parental care involves feeding offspring, carrying it,
grooming (Fig. 9.38), guarding from predator and teaching social and survival
skills (Fig. 9.39).

Fig. 9.38: Japanese Snow Monkey: Mother grooming the young one

Many eutherians, like tiger (Fig. 9.39), bear and orangutan exhibit maternal
care only, wherein all the responsibilities are taken care of by the mother.
Few mammals like Indian striped hyenas show bi-parental care and in some
like meerkats, elephants, allo-parenting is seen.

Fig. 9.39: Mother Tigress with her cubs 231

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SAQ 4
a) Describe various types of nests across the phyla.

b) Why maternal care is most prevalent parental care type in mammals?

9.7 COST AND BENEFITS OF PARENTAL CARE

Most species do not show parental care. They produce large number of
gametes, hence large number of offspring and leave them to survive on their
own. In case of internal fertilization, fertilized eggs are abandoned post egg
laying.

The decision to provide some kind of parental care depends upon the how it
will increase the lifetime reproductive success rate of a caregiver. The
parental duties keep the caregiver away from mating opportunity. Also many
times parents risk their own life to safeguard offspring.

Parental investment: Any investment in the form of time, energy or resources

provided by the parent/s to the current offspring that improves the chances of
offspring’s survival and health at the cost of parent ability to invest in future
offspring.

9.7.1 Cost of Parenting

a) Reduced resources: Parents invest lots of resources on their progeny.

In most oviparous animals, eggs are loaded with yolk. The female parent
contributes nutrients in the yolk. Parents incubate eggs and spend their
body heat to maintain optimal temperature. Similarly, in many species
eggs are aerated, carried on back or brood pouches, which are all
energy consuming processes. In case of birds and mammals, post birth
offspring are fed multiple times. Mammalian females have mammary
glands just to feed the new born. This extracts valuable nutrients and
energy from parent/mother’s body to keep the child well fed.

b) Reduced mating opportunities: Parents spend lots of time and energy

on their progeny. It can range from few months to years in some cases.
During this time as they are busy with the child, they are unable to mate.
These lost mating opportunities reduce mating success of the parent/s.

c) Reduced survival rate: Parenting involves guarding the eggs or young

ones. To do so, many parents risk their life and face predators, which
under other circumstances they would have avoided. Most young
animals have weak immune system and they are highly susceptible to
diseases. Since they live in close proximity to parents, they can easily
transfer the pathogens to them. Also multiple tasking in parenting
makes it a stressful job for parents, making them physiologically weak
and immune suppressed. All these factors are capable of killing or
232 reducing the life span of parent.
Unit 9 Parental Care
Given the cost of parenting, then why do some species still provide parental
care to its offspring? It’s because the benefits of parenting in those species
outweigh the cost.

9.7.2 Benefits of Parenting

a) Parental care improves the chances of offspring survival into adulthood
and doing that improves the reproductive success of parent. Altricial
offspring won’t even survive without parental care.

b) Offspring growth is much better if provided with parental care v/s without
it. In birds it is seen that altricial species have better flight capacity and
forelimb function compared to precocial birds. Parental care also
increases offspring lifespan and fitness.

c) Good health of young ones improves their chances of finding mate in

adulthood and improves offspring reproductive success. This contributes
to the inclusive fitness of the species.

SAQ 5
What are cost and benefits of parental care?

9.8 EVOLUTIONARY STABLE STRATEGY

In 1973, J.M. Smith and G.R. Price explained Evolutionary Stable Strategy.
These are those strategies that can’t be supplanted with another strategy, as
they are important in adaptation of a population in a specific environment.
These are biologically heritable and encoded in the individuals of a population.
They are subjected to natural selection forces and get stabilized in response to
reproductive success. These strategies include individual response to various
conflicts that ultimately define reproductive success of a population. The
conflict can be sexual conflict, parent –offspring conflict, etc.

9.9 FAMILIAL CONFLICTS

9.9.1 Parents-Offspring Conflict
Parents invest huge resources in parental care and this is being done at the
cost of future mating success and even parental care to be provided to the
future offspring.

For an offspring, longer the parental care better the chances at survival and
reproduction. So what should be the length and the amount of care provided
by the parents?

According to Robert Trivers (1974), parents want to distribute their resources

almost equally among offspring both current and future, therefore in the later
stages of parental care, parent tries to reserve resources and energy for future
offspring rather than spending on the current one. This conflict decides
weaning age of current offspring. Till benefit vs cost ratio (B/C) of parenting is 233
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equal to or greater than 1, parents provide parental care. If B/C goes below 1
(cost of parenting is higher than benefit), parent especially mothers try to wean
off the offspring and offspring still tries to extract care from parents. This
conflict ends when both parent and offspring agree on the inclusive fitness of
the family and offspring decide to forego parental care in favor of its future
siblings with whom they are related 50% genetically. This happens when B/C
drops below ½.

Other factors such as life span of species and maternal age also contribute to
the investment in parental care. It has been observed that birds with longer life
span are less enthusiastic to protect their young ones from predators
compared to birds with shorter life span. This suggests, that species with
shorter life span and limited number of offspring invest more in parental care
verses species with long life span.

With age female reproductive fitness decreases, this leads to older females of
same species providing longer and better parental care compared to the
younger females. Thus as maternal age increases, so is parental investment.

9.9.2 Sibling-Sibling Conflict

Parental investment in a season is fixed and all the offspring compete to get
their share of resources. This created sibling-sibling conflict and older or in
some cases stronger sibling attacking the younger, weaker ones.

9.9.3 Parental Favoritism

Parents are related to their entire brood equally (50% genetically), but in some
species they might end up favoring one offspring over others. In many cases,
parents actively discriminate among children and favoring one and letting the
other starve or die by siblicide. The reason for this brood reduction is:

Insurance against failure: Many species do not want to invest in more than
one progeny per season. They keep other eggs as insurance. First healthy
offspring is taken care and rest of the offspring are left to die.

Trade off: Even if animal is capable of raising all the offspring, they choose to
raise the healthiest one and leave the rest to die. This saves their energy and
resources that can be used on future offspring. In Pandas, mother prefers to
raise one strong cub rather than two weak ones. So she trades off and lets
one of them to die (weaker one).

Environmental Uncertainty: Some species raise their complete clutch during

favorable conditions but resort to brood reduction during unfavorable times. In
case of limited resources, parents end up investing in healthiest offspring.
They have two types of clutch: core offspring (raised even in bad year) and
one marginal offspring (raised only during good year).

9.10 PARENTAL MANIPULATION

The reproductive success of parents is evaluated in term of life long
reproductive success. Sometimes parents manipulate current offspring to
increase their life long reproductive success. There are various strategies that
234 are employed to achieve this:
Unit 9 Parental Care
9.10.1 Maternal Manipulation in Eusocial Society
Eusocial structure in insect sociality stands on two strong pillars, inclusive
fitness (as stated by Hamilton) and maternal manipulation. Most insect
eusocial societies are made up of reproductive queen or pair (queen and king)
and non-reproductive workers, which are their offspring. In such structures,
queen (mother) manipulates its current offspring mostly daughters to be
workers and raise her offspring rather than have their own. Workers sacrifice
their reproductive fitness for the lifelong reproductive success of mother.

In honeybees, it is achieved by pheromonal manipulation and differential

distribution of food. In honeybee colony, unfertilized egg will develop into male
and fertilized ones into female. The female larva that is fed with royal jelly with
some honey develops into queen and the ones fed only honey become
smaller worker bees. Queen bee is the largest and only reproductive female in
the colony. She secretes queen substance from her mandibular gland that
obstructs ovarian development in worker bees. It also prevents worker bees
from raising another queen by making them feed only honey to larvae.

In some species of ants, queen ant accumulates certain hormones in egg that
causes egg development into worker.

In social wasp and bees, queen maintains her reproductive status using
aggression on the worker and preventing them from breeding.

9.10.2 Brood Adjustment

In many birds, brood size is adjusted to raise either single offspring or only few
best among all the hatchling. In African black eagle, first hatched chick attacks
and kills its younger one while parents ignore eldest chick behaviour. In barn
swallow, the gape color decides which offspring will be fed more. The red
gape color of chick is indicator of better immunological health. So parents feed
chicks with brighter red color more food, compared to the duller colored one
and thus increasing their chances of survival. In coots, orange plume in chick
is indicator of health. They actively peck weaker babies and kill them to reduce
the brood size.

Hatching asynchrony: It is a subtype of brood adjustment . Birds need not

hatch their eggs asynchronously, but some of them do so that they have
option of brood adjustment . In black eagle, as soon first egg is laid, it starts
incubating it. This gives first laid egg advantage over others as it hatches early
and being stronger can kill off its younger siblings. In cattle egret, parents give
high dose of androgen to earlier laid egg and thus helps earliest hatched chick
to be aggressive so they kill off younger siblings.

SAQ 6
a) Describe evolutionary stable strategy.

b) What is parent-offspring conflict?

c) Explain maternal manipulation.

235
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9.11 BROOD PARASITISM
In some cases, one species deceives other member of same or different
species to provide parental care to its offspring.

9.11.1 Intra-species Brood Parasitism

When member of same species accept eggs of other members or raise
chick that is not their own (but same species) is called intra-specific
brood parasitism. This is usually a costly affair for the host parents as the
resources and energy meant for their offspring is shared with the adoptive
one. This is seen in species who live in colonies and sometimes chick wander
and join family that is not its own.

Example: Colonial nesting gull chicks if not fed properly leave its original nest
and join foster parent family. Ducks usually accept extra eggs or stray chick in
her brood (Fig. 9.40).

Why host tolerates parasite of offspring

1) To prevent accidental loss of own progeny: The method by which
parents recognize their own chicks is not so perfect in birds. If by
mistake, they kill or starve their own chicks due to confusion, the cost will
be too high for them. That is why, when in doubt most host parents still
feed adoptive chicks as they are not sure. The overall cost of feeding
adoptive chicks is still low as such incidences are very few in a colony as
compared to loss of own chicks by mistake.

2) Dilution of predator attack: Ducks are known to accept both unrelated

eggs and chicks. Ducklings are precocial and post imprinting follow
mother duck everywhere. This makes them highly susceptible to
predator attack. Acceptance of unrelated duckling dilutes the predator
attack on all ducklings as per probability now chance of being killed by
predator is reduced as number of duckling increases. And if it’s the
foster chick that dies in attack, the host parent will not have any genetic
loss.

Fig. 9.40: Golden eye duck usually lays eggs in nest of other females

9.11.2 Inter-species Brood Parasitism

The members of some species neither construct their own nest nor take care
of their young ones. They simply lay their eggs in the nests of unrelated birds
and deceive them to take care of their chicks. The new world cuckoo, old
236 world cuckoo and cowbirds (Fig. 9.41) are known brood parasites.
Unit 9 Parental Care

Fig. 9.41: Host rufous-collared sparrow feeding parasite shiny cowbird chick

Example: The famous example of inter-species brood parasitism is Reed

warbler raising common cuckoo chick (Fig. 9.42). Female cuckoo lays a single
egg in the nest of reed warbler. The chick as soon it is born, throws out all the
reed warbler eggs. In this case host parental investment gets completely lost
as it only raises the cuckoo chick. The cuckoo chick mimics and enhances
reed warbler chick signal that stimulates parental behaviour in Reed warbler.
Also, cuckoo lays its eggs in smaller birds nest, this helps its chick to hatch
before host’s own eggs. Also parent tendency to feed healthiest chick is
exploited as cuckoo chick grows very fast in very short time, even surpassing
host parent in size (Fig. 9.42).

Fig. 9.42: Reed warbler foster parent feeding oversized common cuckoo chick

Sometimes host species recognizes parasite eggs and if they have option they
abandon the old nest and re-nest in the same area. 237
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Why host tolerates parasite of offspring
1) Egg Mimicry: In this case parasite egg mimics the host egg. Even if
they are doubtful, they feed the parasite chick to prevent the loss of their
own chick by mistake. Mimetic egg is always accepted by the host
parents and raised as their own.

2) Nest site Hypothesis: Most brood parasites lay eggs in the nest of
species that are smaller in size. This makes it difficult for host to remove
the parasite egg from the nest. Nesting sites are limited and leaving a
nest sometime means loss of nest site even in subsequent seasons.
Instead host decides to raise the parasite chick. It is observed in case of
host warblers taking care of cowbirds chick, if they recognize parasite
egg/chick and if alternative nesting sites are available, they will abandon
the parasitized nest.

3) Mafia hypothesis: In this case the parasite bird monitors the host bird
and if the host bird recognizes and removes parasite egg they attack
host’s eggs. This is observed in spotted cuckoos. The host Magpie’s
eggs are attacked by parasite cuckoo if the host removes the parasite
egg. Magpie that removes parasite egg faces more predation i.e. 87%
compared to those who accepted it i.e. 12%.

SAQ 7
Differentiate between intra-species and inter-species brood parasitism.

9.12 SUMMARY
In this unit you have learnt that:

• Any behaviour that increases the chances of survival and future

reproductive success of an offspring comes under parental care.

• In animal world, parental care involves variety of practices like nest

building, incubation and protection of eggs, guarding young ones,
feeding offspring and teaching them valuable skills.

• There are two types of species depending upon how their offspring are
born : 1) altricial where an offspring is completely helpless and
dependent 2) precocial where offspring is born fairly mature and
independent

• Parental care is not shouldered equally by both sexes, in most species it

is the female who gives parental care, in some it is male and in others its
both parents who take care of progeny.

• Triver, Dawkins, Carlisle and Williams gave theory and hypothesis that
try to explain uneven investment by male and female in parental care.

• There are four types of parental care in nature: maternal (mother only),
paternal (father only), bi-parental (both parent) and allo-parenting (either
238 both or one parent along with other members of the family).
Unit 9 Parental Care
• Parental care can be observed across various phyla from insects, to fish
to mammals and there are multiple examples including honeybees,
seahorse, tiger, elephant, Penguin that exhibit it.

• Evolutionary stable strategy is a strategy that cannot be supplanted with

another strategy, as they are important in adaptation of a population in a
specific environment.

• Due to high investment nature of parental care, we see various types of

familial conflicts to reduce parental care cost. These conflicts include,
length of parental care that the parents are ready to provide to offspring,
parental favoritism and sibling-sibling competition.

• Parent manipulation is seen in the nature and it involves higher parental

reproductive success at the cost of progeny’s survival or reproductive
success.

• Many species cheat on parental care and manipulate other species to

take care of their young one. This is termed as brood parasitism and
most common example is common cuckoo as parasite on reed warbler
host.

9.13 TERMINAL QUESTIONS

1. What are various parental investment theories?
2. Explain different types of parental care seen in fishes?

3. How amphibians take care of their young ones? Explain.

4. What are the major types of parental care provided by birds? Discuss.
5. Why some species promote siblicide?

6. What is parental manipulation? Describe various ways in which parents

manipulate their progeny.

7. What is Brood parasitism? What are its subtypes? Why host species
tolerate parasite species eggs/young ones?

Activity 1

Locate any common bird nest with eggs/chicks in it and describe what kind of
parental care the bird provides.
Activity 2

Observe a female street dog with pups (from a far) and list all the parenting
duties performed by her.

9.14 ANSWERS
Self-Assessments Questions
1. a) Any behaviour on the part of either one or both parent that
enhances the chances of survival of offspring is termed as parental
care. Sometimes besides parents, other members of the family
also help in taking care of the offspring. 239
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Examples: Nest building by weaverbird, incubation of egg by
emperor penguin, guarding eggs by crocodile female and feeding
milk to young ones by mammalian mother are all examples of
parental care.
b)
Points of Altricial Species Precocial Species
Difference
Definition Species whose offspring are Species whose offspring
born completely helpless and are born relatively
require parents to feed them, mature and can move
take care of them during around from the moment
initial stages of life are called they are born or hatched
altricial species. are called precocial
species.
Examples Hawks, woodpecker, owls, Domestic chicken,
marsupials, domestic dogs, ducks, geese, guinea
cats, human pig, most ungulates

2. a) William’s hypothesis was given by George William in 1975 and this

correlates the parental association with embryo and parental care.
According to this in case of external fertilization, female lays eggs
in male territories that are fiercely guarded by males. Now males
will fertilize them and remain in close proximity and create an
association that causes males to take of them. In case of internal
fertilization, it is the female who is closely associated with embryo
and post fertilization takes care of the embryo.

b)
Points of Trivers’s Theory Dawkins and
Difference Carlisle’s Theory
Scientist Robert Trivers R Dawkins and T.R.
Carlisle
Year 1972 1976
Definition According to this the sex that It suggests that
invests less in parental care will parental care
invest more in the traits that help depends upon the
them getting chosen by a mate order of release of
(intra-sexual competition) and the gametes and the
sex that will invest more in chance of mate
parental care will be more choosy desertion.
and will be more selective of the
mate (inter-sexual selection).
Male vs. Since males invest in trait that In case of fishes, the
Female help them outcompete fellow female lays eggs
behavioural males of the same population, before fertilization by
response therefore they are less invested in male, giving them
240
Unit 9 Parental Care
parental care and since females chance to desert first
usually invest less in intra-sexual and leave the
competition and chooses mate fertilized eggs to be
very carefully, they are usually taken care by the
more involved in parental care male. On the other
giving. hand, internal
fertilization provides
males the chance to
desert first and leave
the offspring to be
taken care by
mother. This tries to
explain mostly
paternal care in
fishes and maternal
care in mammals.

c) Certainty of paternity hypothesis states that males invest in

parental care when they are sure of the paternity. In case of fishes
and amphibians, external fertilization takes place in aquatic
environment. In external fertilization, female lays eggs first as they
are larger in size and has less chances of being swept away by
water current. Male later fertilizes the eggs as sperms are smaller
and can get lost if released first. So in external fertilization males
are sure of paternity and paternal parental care is higher in fishes
and amphibians. On the other hand in internal fertilization males
are not sure of paternity and do not invest in parental care. In most
cases of internal fertilization, maternal care is seen. Male members
of the species contribute along with female in bi-parental care
mostly in monogamous relations as seen in the birds.

3. a) When other members of the family besides parents help in taking

care of the young ones, such a system is termed as allo-parental
care. This system is product of altruism and development of
eusociality. This improves reproductive success among the
members of species.

For mammalian mothers, allo-parenting helps in multiple ways.

With outside help, mammalian mother now will have more time to
feed young ones and more energy to produce rich milk. This
improves survival rate and fitness of offspring.

Example: Animals exhibiting allo-parenting are honeybees,

termites, ants, meerkats and elephants.

b)

Points of Maternal Care Paternal Care

Difference

Caregiver Female of the species Male of the species

241
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
Frequency Female only care in Male only parental care
mostly observed in is more common in
species with internal species with external
fertilization. Example of fertilization. Many
showing maternal only species of fishes and
care is seen in amphibian provide male-
crocodiles, most only parental care to
mammals and few birds their

Examples Tiger, leopard, and Various species of sea

Polar bears show horse is the most well
female only parental studied example of
care. paternal care.

c) When both parents work together to take care of offspring, such a

system is called bi-parental care. It is mostly seen in species,
where sex ratio in reproductively mature individual is well
balanced. This means species whose population has almost
similar number of sexually mature males and females. They are
most likely to engage in bi-parental care. This type of care involves
complete cooperation and coordination among the parents.

Example: In striped hyenas both parents take care of the young

ones. They are monogamous in nature and build den for the
family. Once offspring are born both male and female raise them
together.

4. a) Nests are shelter or safe space for the eggs or the offspring. Nest
building is used as a tool to attract the females and to take care of
fertilized eggs. There are variety of nests seen across various
phyla:

Nests in Fishes

Simple depression: simple depression at the floor of aquatic body

is used as nest in case of lungfishes.

Tunnel nest: Male stickleback fish makes his nest using rootlet
and other fragments from plants and it glues the material together
with adhesive secretion from his kidney. The nest in this case is
spherical in shape with a tunnel for the deposition of eggs.

Bubble Nest/Foam Nest: Male fish build these nests by producing

bubbles from mouth along with saliva or mouth secretions and they
float at the surface of aquatic body. These types of nests are
usually built in oxygen depleted stand still water bodies and fishes
that build them are called aphrophils. Examples of fishes that
build bubble nest are gouramis, eel and betta fish.

Nests in Amphibians

Foam Nest: Japanese tree frog (Rhacophorus schlegeli) covers

242 their fertilized eggs with mucus secretion. Both male and female
Unit 9 Parental Care
dig a tunnel together and leave their mucus-enveloped eggs in
there. The mucus keeps eggs hydrated and prevents their
desiccation.

Malabar gliding frog (Rhacophorus malabaricus) produces foam

nest on the trees, where female accumulates her eggs.

The female South American tree frog (Leptodactylus mystacinus)

produces foam nest in a hole near water body. In this case,
tadpoles directly enter the water as soon they hatch.

In case of Adelotus brevis males produce foam nest and also

defend the fertilized eggs actively till they hatch.

Mud Nest: Mud nest are prepared by male Brazilian tree frog
(Hyla fabre) by digging a hole or making a nursery in the mud of
shallow water. Female lays her eggs in the mud nest, where the
tadpole hatches and directly moves into the water body.

Leaf Nest: The tiger-striped leaf frog (Phyllomedusa tomopterna)

lays its eggs in gelatinous mass on the leaf above a pond. It then
folds the leaf and seals it to give the shape of a nest. When eggs
hatch, tadpole directly falls into the water below.

Tree Nest: Tree frog (Hyla resinfictrix) uses shallow tree cavity as
a nest. They line this cavity with beeswax obtained from the hives
of stingless bees. When these cavities are filled with rainwater,
female lays egg in it and their offspring develop in a protected
environment.

Nests in Birds

Cavity nest: Some birds use crevice among rocks or a cavity in

the tree as nest and lays their eggs there eg. parakeets.
Woodpeckers hollow out cavity in the trees and kingfisher
excavates riverbank to use it as nest.

Scrape nest: Some birds make a hollow in the ground to lay their
eggs. These hollows are called scrapes.

Platform nest: Some birds make very simple flat nests out of
twigs. These nests are called platform nestseg. large birds such as
Rooks, Magpies and Crows make these types of nests.

Cup shaped nest: These nests can be made from small twigs,
grass stalks and moss and it is cup shaped. It is most common
type of nest made by birds. Example: blackbirds, whitethroat,
Scottish crossbill, etc.

Woven nest: Weaverbird constructs a suspended pendant shaped

nest out grass and plant fiber. Tailorbird sews the edges of a large
single leaf to design beautiful home out of it.

b) The term mammals is derived from presence of mammary glands

in females that gives milk which is fed to the young ones during 243
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
their developmental stages. Mammals provide longest and most
complex parental care to their offspring compared to other
animals. Mothers are the primary and mostly the only caregiver (in
95% cases) in mammals with father and other family members
helping out in some species (5%). The reason for that is:

1) Due to internal fertilization females are closely associated

with the embryo and are more likely to take care of the
embryo and baby.

2) In mammalian female, due to gestation and lactation,

hormonal changes occur in the body of females, making
them physiologically predisposed to parental care.

5. Most species do no show parental care. The decision to provide some

kind of parental care depends upon how it will increase the lifetime
reproductive success rate of a caregiver. There are both cost and
benefit of parenting and based on its ratio, parental care is provided.

Cost of Parenting
• Reduced resources: Parents invest lots of resources on their
progeny. Most oviparous animal eggs are loaded with yolk. The
female parent contributes nutrients in the yolk. Parents incubate
eggs and spend their body heat to maintain optimal temperature.
Similarly in many species eggs are aerated, carried on back or
brood pouches, all of which are energy consuming processes. In
case of birds and mammals, post birth offspring are fed multiple
times. Mammalian females have mammary gland just to feed the
new born. This, extracts valuable nutrient and energy from
parent/mother’s body to keep the child well fed.

• Reduced mating opportunities: Parents spend lots of time and

energy on their progeny. It can range from few months to years in
some cases. During this time as they are busy with the child, they
are unable to mate. These lost mating opportunities reduce mating
success of the parent/s.

• Reduced survival rate: Parenting involves guarding the eggs or

young ones. To do so, many parents risk their life and face
predator, which under other circumstances they would have
avoided. Most young animals have weak immune system and they
are highly susceptible to diseases. Since they live in close
proximity to parents, they can easily transfer the pathogen to them.
Also multiple tasking of parenting makes it a stressful job for
parents, making them physiologically weak and
immunosuppressed. All these factors are capable of killing or
reducing life span of parent.

Benefits of Parenting

• Parental care improves the chances of offspring survival into

adulthood and doing that improves the reproductive success of
244 parent. Altricial offspring won’t even survive without parental care.
Unit 9 Parental Care
• Offspring growth is much better if provided with parental care v/s
without it. In birds it is seen that altricial species have better flight
capacity and forelimb function compared to precocial birds.
Parental care also increases offspring lifespan and fitness.

• Good health of young ones improves their chances of finding mate

in adulthood and improves offspring reproductive success. This
contributes to the inclusive fitness of the species.

6. a) Evolutionary Stable strategy is that strategy which cannot be

supplanted with another strategy, as they are important in
adaptation of a population in a specific environment. These are
biologically heritable and encoded in the individuals of a
population. They are subjected to natural selection forces and get
stabilized in response to reproductive success. These strategies
include individual response to various conflicts that ultimately
define reproductive success of a population.

b) Parents invest huge resource in parental care and this is being

done at the cost of future mating success and even parental care
to be provided to future offspring. For an offspring, longer parental
care means better chances at survival and reproduction.

Parents want to distribute their resources almost equally among

offspring both current and future, therefore in later stages of
parental care parent tries to reserve resources and energy for
future offspring rather than spending on current one. This conflict
decides weaning age of current offspring. Till benefit vs cost ratio
(B/C) of parenting is equal to or greater than 1, parents provide
parental care. If B/C goes below 1 (cost of parenting is higher than
benefit), parent especially mothers try to wean the offspring and
offspring still tries to extract care from parents. This conflict ends
when both parent and offspring agree on the inclusive fitness of
family and offspring decides to forego parental care in favor of its
future siblings with whom they are related 50% genetically. This
happens when B/C drops below ½.

c) Maternal manipulation is the manipulation of current offspring

mostly daughter to be workers and raise her offspring rather than
have their own. Workers sacrifice their reproductive fitness for the
lifelong reproductive success of mother.

Example: In honeybee, maternal manipulation is achieved by

pheromonal manipulation and differential distribution of food. In
honeybee colony unfertilized egg will develop into male and
fertilized one into female. The female larva that is fed with royal
jelly with some honey develops into queen and those are fed only
honey become smaller worker bees. Queen bee is the largest and
only reproductive female one colony. She secretes queen
substance from her mandibular gland that obstructs ovarian
development in worker bees. It also prevents worker bees from
raising another queen by making them feed only honey to larvae. 245
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
7.

Points of Intra-species Brood Inter-species Brood

Difference Parasitism Parasitism

Definition When members of the same The members of some

species accept eggs of other species neither
members or raise chick that construct their own nest
is not their own (but same nor take care of their
species) it’s called intra- young ones. They
specific brood parasitism. simply lay their eggs
into the nest of
unrelated birds and
deceive them to take
care of their chick.

Host and Belong to same species Belong to different

parasite species

Examples of Ducks, colonial nesting gulls Common cuckoo,

parasite cowbird

Terminal Questions
1. Refer to Section 9.4.

2. Refer to Subsection 9.6.2.

3. Refer to Subsection 9.6.3.

4. Refer to Subsection 9.6.5.

5. Refer to Subsection 9.9.3.

6. Refer to Section 9.10.

7. Refer to Section 9.11.

Acknowledgement of Figures
Fig. 9.1 By Wolfgang Wander, Papa Lima Whiskey (edit) - self-made /
[Link] CC BY-SA 3.0,
[Link]

Fig. 9.2 By Toby Hudson - Own work, CC BY-SA 3.0,

[Link]

Fig. 9.3 By Luis Miguel Bugallo Sánchez (Lmbuga

Commons)(LmbugaGalipedia)Publicadapor/Publish by: Luis
Miguel Bugallo Sánchez - Own work, CC BY-SA 3.0,
[Link]

Fig. 9.4 By Arian Zwegers - Bukit Lawang, orangutans, CC BY 2.0,

246 [Link]
Unit 9 Parental Care
Fig. 9.5 By photo taken by flickr user noisecollusion - flickr, CC BY 2.0,
[Link]

Fig. 9.6 By Francis C. Franklin - Own work, CC BY-SA 3.0,

[Link]

Fig. 9.7 By Diego Delso, CC BY-SA 4.0,

[Link]

Fig. 9.8 By Ian Alexander - Own work, CC BY-SA 4.0,

[Link]

Fig. 9.10 By Dunkelfalke - Own work (Original text: self-made), CC BY 2.0,

[Link]

Fig. 9.12 By The original uploader was MidgleyDJ at English Wikipedia. -

Transferred from [Link] to Commons., CC BY-SA 2.5,
[Link]

Fig. 9.14 By © Hans Hillewaert, CC BY-SA 4.0,

[Link]

Fig. 9.15 By Dinh D. Tran, FiMSeA -

[Link] CC BY 3.0,
[Link]

Fig. 9.17 Public Domain,

[Link]

Fig. 9.18 VikramGupchup, CC BY-SA 4.0

<[Link] via Wikimedia
Commons
Fig. 9.21 Gadow, Hans (1909) Amphibia and reptiles. MacMillan and Co.

Fig. 9.22 By Christian Fischer, CC BY-SA 3.0,

[Link]

Fig. 9.23 Edelcio Muscat and [Link]

releases/933847

Fig. 9.24 By Zygy - [Link] CC0,

[Link]
Fig. 9.25 By Dario Sanches - originally posted to Flickr as TUIM
(Forpusxanthopterygius ), CC BY-SA 2.0,
[Link]
Fig. 9.26 By JJ Harrison ([Link] - Own work, CC
BY-SA 3.0,
[Link]
Fig. 9.27 By 159766 - dr. TiborDuliskovich - [Link] -
Own work, CC BY-SA 3.0,
[Link]
Fig. 9.28 [Link] - Own work, CC BY-SA 3.0,
[Link] 247
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
Fig. 9.29 By Charles J. Sharp - Own work, from Sharp Photography,
sharpphotography, CC BY-SA 4.0,
[Link]

Fig. 9.30 By Alastair Rae from London, United Kingdom - Black

Woodpecker, CC BY-SA 2.0,
[Link]

Fig. 9.31 By User:Venan - Own work, CC BY-SA 2.5,

[Link]

Fig. 9.32 By Steve Brace from Herefordshire, UK - Duck Factory, CC BY

2.0, [Link]

Fig. 9.33 CC BY 2.5,

[Link]

Fig. 9.34 By Michael Gäbler, CC BY-SA 3.0,

[Link]

Fig. 9.35 By NevitDilmen (talk) - Own work, CC BY-SA 3.0,

[Link]

Fig. 9.36 By LittleJerry - Own work, CC BY-SA 4.0,

[Link]

Fig. 9.37 By Ikiwaner - Own work, GFDL 1.2,

[Link]

Fig. 9.38 By Rocktendo&amp; Adam Gman - Own work, CC BY-SA 4.0,

[Link]

Fig. 9.39 By Wikigringo -

[Link]
31&amp;pos=5&amp;pid=1246, CC BY-SA 3.0,
[Link]

Fig. 9.40 By Chad Teer from Coquitlam, Canada - [Link] - image

description page, CC BY 2.0,
[Link]

Fig. 9.41 By Dario Sanches from São Paulo, Brazil - VIRA-BOSTA Imaturo
(Molothrusbonariensis) e ZonotrichiaCapensis ), CC BY-SA 2.0,
[Link]

Fig. 9.42 By Per Harald Olsen - Own work, CC BY-SA 3.0,

[Link]

248
 UNIT 10
HUMAN BEHAVIOUR AND ITS
REGULATING
REGULATING FACTORS

Structure
10.1 Introduction 10.6 Linking Behaviours to Brain,
Genes, Hormones and
Objectives
Environment
10.2 Human Behaviour
Behaviour and Brain
10.3 Human Non-verbal
Behaviour and Genes
Communication
Behaviour and Hormones
Types of Non-verbal
Communication Behaviour and Environment

Comparison between Verbal 10.7 Summary

and Non-verbal
10.8 Terminal Questions
Communication
10.9 Answers
10.4 Mate Selection and Sexual
Strategy
Analysing Mate Selection

Sexual Strategy for Mate

Selection

10.5 Family Relationships and

Altruism
Family and Kinship

Altruism in the Context of

Family Relationships

10.1 INTRODUCTION
In the previous unit, we discussed the concept of parental care and got
familiarized with the idea that the level of parental care varies among different
species. The behaviour of our species has evolved such that we show a great
degree of parental care. The purpose of this unit is to discuss all the basic
aspects of our behaviour that make us distinctly human. It is fairly obvious that
human behaviour is a vast topic and therefore it is impossible for any piece of
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
literature to elaborate on all the aspects of human behaviour. In this Unit, we
will start with a generalized account of human behaviour and then focus on
some specific aspects, beginning with non-verbal communication. Later in
this Unit, we will discuss how humans go about the process of mate
selection. We will also briefly analyse the sexual strategy used by humans.
This will be followed by a close look at family relationships and how the
concept of family is related to the concept of altruism in the natural world.
Finally, we will try to understand how human behaviour gets influenced by
various intrinsic and extrinsic factors such as our brain, genes, hormones and
environment. In the next unit, which belongs to block 3 of the course, we will
learn to develop a scientific understanding of animal welfare.

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

 explain the basic tenets of human behaviour,

 comprehend the concept of ‘human non-verbal communication’,

 differentiate between verbal and non-verbal communication,

 understand how humans perform ‘mate selection’,

 comprehend the basis of ‘sexual strategy’ used by humans,

 appreciate the concepts of ‘altruism’ and its relevance in ‘family

relationships’ of humans,

 comprehend how the physiology and anatomy of human brain affects

human behaviour,

 explain how genes and hormones control human behaviour, and

 appreciate how environment can alter human behaviour in short term

and long term.

10.2 HUMAN BEHAVIOUR

Human behaviour can be defined as the entire range of behaviours exhibited
by humans across our entire lifetimes. It is the most basic human action and is
regulated by a plethora of factors, including genetics, hormones,
psychology, culture, values, attitude, social status, etc. Since there are so
many variables that determine an individual’s behaviour, it shows a wide range
of variation among individuals. It is important to note here that human
behaviour consists of the behaviour directed towards self as well as towards
the environment, which includes other people also. In this context, the term
social behaviour is used specifically for the behaviour that is directed towards
other people.

An individual’s genetic makeup has a big impact on his/her behaviour. This is

true of other animals too; a fact that animal breeders have known for
centuries. Consider the domestication of dogs and horses for instance. Animal
250 breeders have been very successful in selecting and propagating favorable
Unit 10 Human Behaviour and Its Regulating Factors
behavioural patterns among animals. In humans, the effect of genetics on an
individual’s behaviour has been reported through studies on identical twins.
Studies involving comparison of behaviours of adopted children with those of
their biological and foster parents have also been performed. These studies
have generated a wealth of knowledge on how genetics and environment (in
the form of upbringing) impact human behaviour.

Humans are social beings, and therefore, our society affects the way we think
and act. Human societies have played a very important role in the evolution of
human civilization. Each society has its own culture, a system of beliefs and
practices, which in turn determines what is acceptable behaviour among
individuals of that society. Mostly, culture has a positive impact since a
society’s beliefs and practices are aimed at the welfare of the society at large.
However, sometimes it can lead to prejudices against other groups or
societies that have a different culture.

Incidentally, when compared to the brains of other species, human brain is

disproportionately larger. Even though it contributes only 2% to our body
weight, it uses 20% of our metabolic energy. Thus, we humans are paying a
heavy cost towards the upkeep of our brain. Obviously, the brain has
conferred a lot of benefits also to humans. In fact, human behaviour is
characterized by increased intelligence and consciousness; and much of that
can be attributed to our increased brain sizes. Another related aspect of
human behaviour is language. And even though many other animals have
been known to have their own language and communication systems through
which they convey a lot of information to each other, the linguistic capabilities
of human brain are unmatched. Many scientists, including Darwin, have been
of the opinion that language has had a tremendous impact on human
evolution. Verbal communication through language allows humans to
disseminate information faster and to a much larger audience. In the past, this
could have helped human ancestors to learn from each other’s experiences
and to form strategies to avoid predators. The importance of verbal
communication and language can be underlined with the fact that the human
brain has specific areas with dedicated machinery for these functions.

In any discussion about human behaviour, parental care deserves a special

mention (refer to Unit 9 for details). Humans provide extensive parental care
to their offspring. Parents invest a lot of time, energy and resources while
bringing up their children. They even help in the rearing of the offspring of their
offspring, i.e., their grandchildren.

Human behaviour shows a lot of diversity and complexity. It also poses a lot of
intriguing questions, especially in reference to the aspects of human behaviour
that seem to have no adaptive value. For example, some people develop anti-
social behaviour which causes harm to themselves and to the society in
general. The many complexities of human behaviour have always fascinated
and enthralled scholars from a variety of different fields such as sociologists,
psychiatrists, psychologists, anthropologists, social workers,
economists, etc. (Fig. 10.1). 251
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours

Sociology

Economics Psychology

Human
behaviour

Social work Anthropology

Fig. 10.1: The study of human behaviour provides useful insights into sociology,
psychiatry, psychology, anthropology, social work and economics.

10.3 HUMAN NON-VERBAL COMMUNICATION

Humans live in close-knit societies and have a very strong verbal
communication system in the form of speech. And while verbal communication
is our forte, we do have a considerable degree of non-verbal communication
too, even though many animals are arguably better at non-verbal
communication than us.

So, what is non-verbal communication? It refers to the transmission or

exchange of information or signals among individuals without using spoken or
written words but through a non verbal platform such as eye contact, gestures,
body language , facial expressions, etc.. Non-verbal communication can be
quite effective in conveying information regarding the emotions, intentions,
The scientific study of needs, attitudes and thoughts of individuals. However, it must be noted that
body language is non-verbal communication can be highly subjective and different people can
called kinesics. interpret the same signal in a variety of ways.

10.3.1 Types of Non-verbal Communication

Non-verbal communication mostly involves visual, auditory or tactile cues.
Most of the human non-verbal communication is through visual cues and
involves body language. The term body language includes a variety of
signals such as facial expressions, eye contact, gestures, postures, body
252 movements, etc.
Unit 10 Human Behaviour and Its Regulating Factors
As illustrated in Fig. 10.2, facial expressions are instrumental in conveying
information regarding emotions and feelings such as happiness, sadness,
fear, disgust, anger, surprise, etc. Particularly important facial expressions
include smile, frown, sneer, etc. Other visual cues, such as gestures; Oculesics is the
especially a nod, wink or thumbs-up, can also be efficient modes of non-verbal study of eye related
communication. Similarly, eye movements and eye contact (oculesics) can be non-verbal
very communicative too. Eye contact can also express warmth, interest and communication. It
concern, thereby facilitating and promoting more verbal communication. For includes gaze, eye
contact, eye
example, studies have revealed that people who make eye contact are better
movement, etc.
communicators since they find it easier to gain the confidence of their
listeners. On the other hand, those people who avoid eye contact are
perceived as deceitful, meek or unconfident.

Fig. 10.2: Some common facial expressions

Postures and body movements also communicate a wealth of information. An

upright posture and fluid body movements could convey confidence. On the
other hand, a rigid posture and stiff shoulders could signal stress and anxiety.
Similarly, even subtle movements of the body away from a person could
denote an aversion to that person. Fidgeting, i.e., making small repetitive
movements of hands and feet, indicates nervousness and restlessness. The
use of space is also an important non-verbal cue. Every individual has a sense
of personal space and any stranger that intrudes on that space is considered a
threat. For example, most students find it intimidating when a teacher stands
right next to them while they are studying or taking notes. This is because the
teacher is inadvertently encroaching on the student’s personal space. The
study of the space between individuals and its effect on their behaviour and
interpersonal communication is known as proxemics.

It is well known that spoken words constitute verbal communication. On the

other hand, paralinguistic cues include auditory cues such as the tone of
voice, pitch, speed, pace and volume. These provide non-verbal 253
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
communication. Thus, communication involves what we say (verbal
communication) and also, how we say it (non-verbal communication). For
instance, when something is said with a strong and upbeat tone, it conveys the
enthusiasm of the speaker. However, when something is said with a strong
and harsh tone, it conveys aggression of the speaker. Some recent studies
have compared the effectiveness of communication when the speakers use a
monotone versus when they modulate the pitch of their voice. Their findings
suggest that communication is more effective when the speakers practice
voice modulation.

The study of touch and the use of tactile stimuli for communication purposes is
known as haptics. Common tactile stimuli used in non-verbal communication
include handshakes, holding hands, high fives and patting someone on the
back. A gentle touch conveys feelings of affection, sympathy and familiarity.
On the other hand, a lingering touch may alarm the recipient and make them
uncomfortable. Recent studies suggest that humans learn to distinguish
between ‘good’ touch and ‘bad’ touch very early on in life. This proves that
tactile stimuli can be very powerful tools for communicating feelings and
intentions.

The use of time in communication is known as chronemics. This tells about

the personality traits such as punctuality, the speed of speech, etc of
sender/receiver.

The appearance of a person also indicates his personality viz the way he
dresses, his lifestyle, etc. This type of communication is known as artifactual
communication.

Sometimes, changes in non-verbal communication styles can be indicative of

mental health issues. For example, if someone’s facial expressions become
passive and less expressive, they avoid eye contact and give terse answers
with flat intonation, then they should be asked to consult a psychologist to
check if they could be suffering from clinical depression.

10.3.2 Comparison between Verbal and Non-verbal

Communication
Verbal communication is quite different from non-verbal communication. In
this, the words are used, whether in spoken or written form, to transmit
message from sender to receiver. As depicted in Table 10.1, these differ in a
variety of ways, such as precision, intentionality and of course, the mode of
signalling.

Table 10.1: Differences between Verbal and Non-verbal Communication.

Verbal Communication Non-verbal Communication

Direct and more precise. Indirect and less precise.

Relies only on a single type of signal Relies on a wide variety of signals

in the form of spoken or written that may be visual, auditory or tactile,
words. being transmitted simultaneously.
254
Unit 10 Human Behaviour and Its Regulating Factors
Non-verbal communication using
Involves interpretation of the literal auditory cues involves interpretation
meaning of spoken and written of the tone, pitch and manner of the
words. spoken words rather than their literal
meaning.

Based on a shared code in the form Based on abstract information,

of language with grammar, therefore, therefore, interpretation is usually
interpretation is usually same among subjective and can differ from person
all receivers. to person.

Mostly intentional Mostly unintentional

It is clear from Table 10.1 that even though verbal communication is more
direct and precise, non-verbal communication also conveys a lot of
information. Let us consider an example to illustrate how important non-verbal
communication is to the receiver of the information. Suppose someone claims
to be happy, but their facial expressions fail to show it; then as receivers of
that information, we are very likely to be suspicious of their claims of
happiness. The same point can be made for many other emotions. Thus, both
verbal and non-verbal communication are essential for proper transmission of
information.

SAQ 1
Define the following terms:

a) Oculesics ……………………………………………………………………….

b) Proxemics ………………………………………………………………………

c) Non-verbal Communication …...………………………………………………

10.4 MATE SELECTION AND SEXUAL STRATEGY

The fact that humans show a great degree of sexual dimorphism means that
both men and women choose their mates from among a pool of potential
mates, a phenomenon known as mate selection. Human mating behaviour is
quite complex and can vary a lot depending on the individual’s preferences,
and cultural and social background. Additionally, human mate selection is also
influenced by the emotion of love, a deep personal and romantic attachment
to another human being. However, since love is an abstract concept whose
effect on mate selection varies among different cultures and societies, there is
no empirical way to evaluate its impact on the mate selection process in
humans. Therefore, we need to apply a variety of different approaches in order
to gain a logical insight into the process.

10.4.1 Analysing Mate Selection

The first step towards analysing the process of mate selection in humans is to
understand the selection criteria or preferences of both the genders and then 255
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to compare those. In this regard, the most useful approach involves the study
of clinical correlation between variation in trait and variation in reproductive
success. However, there is a scarcity of data that unambiguously proves that a
particular variation of a trait is responsible for directly increasing or decreasing
the reproductive success of individuals. Consequently, we rely on indirect
approaches to generate the data that could help us analyse human mate
selection criteria. The most common of these approaches rely on the use of
questionnaires, personal advertisements, and computerized simulations of
faces and body shapes.

Out of all the strategies that can be employed to study human mate
preferences, a questionnaire-based survey is the easiest to perform. Such
studies have quite a few advantages, especially for the human subjects.
These are simpler to perform and need very little time investment from the
subjects. In this respect, the study performed by David Buss in 1989 is
particularly relevant. This survey included data from men and women from 37
different cultures and therefore represented a wide array of cultural, racial and
economic backgrounds. The key findings of this study were that women
choose mates on the basis of their earning potential, generosity, sociability
and reliability. Most women were found to value physical attractiveness less
than the aforementioned attributes. On the other hand, men prefer to partner
with women who are physically attractive and young. Clearly, the
questionnaire-based studies have provided deep insights into human mate
selection criteria. However, there are many pitfalls also of using such a
research methodology e.g., there have been instances where the subjects
have provided incorrect information or deliberately withheld information from
the researchers. The reasons for such a thing could be embarrassment or
deep resentment over one’s own actions and thoughts. Moreover, sometimes
the subjects tend to give the answers which they feel that the researcher
would prefer. Therefore, many scientists have questioned the authenticity of
the data collected through questionnaires.

Personal advertisements printed in newspapers can be very good sources of

information on human mate selection criteria. Broadly speaking, both personal
Body mass index advertisements as well as the questionnaire-based surveys generate similar
(BMI) is a rough kind of results for human mate selection criteria with very few exceptions.
estimate of the Personal advertisements have an advantage over questionnaire-based
amount of body fat in methods in that they are less susceptible to the phenomenon of the subjects
people with respect to
giving answers according to the expectations of the researchers who are
their heights. It is
calculated by dividing conducting the questionnaire-based study. Another advantage of using data
body weight (in from personal advertisements is that it is more reliable because it depicts real
kilograms) by the attempts of people to secure mates. However, the disadvantage is that the
square of height (in personal advertisement data corresponds to a very small proportion of the
meters). Adults with population and therefore cannot be a true reflection of the population as a
BMI between 18.5 to whole. In the past few years, online dating sites have also provided a wealth
24.9 kg/m2, are
of information on the mate selection criteria of humans.
considered to have a
normal weight.
Computerized simulations for analysing mate selection criteria entail the use
of visual stimuli such as line drawings, photos and animations. These
simulations are particularly suited to studies aiming to assess preferences for
256 particular facial features, body shapes and movement patterns. Modern
Unit 10 Human Behaviour and Its Regulating Factors
technology now allows us to perform manipulations in the pictures/videos so
that we can make subtle changes, e.g., altering the shape and size of a
particular physical attribute. A large number of human subjects are then shown
the visual stimuli and asked to rate the ones that they find more desirable in a
mate. Thus, we can study human mate preference in terms of facial features,
body type, body fat distribution and even physical movements such as gait.
The studies using computerized simulations indicate that both the sexes prefer
mates with symmetrical features. Further, it has been found that as far as
mate preference of human males is concerned, the distribution pattern of the
body fat is more important than the overall weight of the mate. An important
parameter in this respect is the body mass index (BMI). Men seem to prefer
women with average BMI which indicates normal weight, although historically,
this criterion has been shown to be highly variable. Another important
parameter for mate selection by men is the waist to hip ratio (WHR). The
WHR of healthy premenopausal women ranges from 0.67 to 0.8, while it
ranges from 0.85 to 0.95 for men. Studies indicate that men prefer a WHR of
0.7 while women seem to prefer a WHR of 0.9 in their mates.

10.4.2 Sexual Strategy for Mate Selection

Let us now try to understand how the mate selection criteria work as sexual
strategies in the evolutionary context. Both men and women invest a
considerable amount of time and resources towards child rearing. Therefore, it
is imperative that both the sexes be careful while selecting a mate. The
Darwinian explanation for the phenomenon of mate selection is that natural
selection has led to evolution of mechanisms that help humans show
preference for those traits that are indicators of high reproductive fitness in the
opposite sex.

The parental investment of mammalian females before the birth of the

offspring is very high. So, the main selection criterion used by females is
whether their prospective mate will be capable of providing for the children
after their birth. In strictly economic terms, this means that women tend to
prefer men with higher earning potential. Obviously, this type of behaviour has
an adaptive significance and therefore, women may have been evolutionarily
programmed to select mates based on their social and financial status. There
are other mate selection criteria also that are used by women such as
generosity, intellect, sense of humour, dominance and dependability. A simple
evolutionary explanation for all the criteria used by women for mate selection
would be that these personal qualities contribute to the success of a person
socially as well as economically. Thus, they could all be assessing whether the
chosen mate is capable of providing for the children. However, a different
approach to look at these criteria could be that they represent the cultural
conditioning of the woman choosing the mate.

Data using computerized audio/video simulations indicates that as far as

appearance is concerned, women seem to prefer men with well-developed
male secondary sexual characteristics such as deep voice, muscular upper
body and masculine facial features (e.g., strong cheekbones, prominent chin,
etc.) as mates. The existence of such a sexual selection strategy could be
explained on the basis of the immunocompetence hypothesis, proposed by 257
Block 2 Systematic Approach to Understand Important Animal and
Human Behaviours
Folstad and Karter in 1992. This hypothesis posits that the hormone
testosterone, which is responsible for the development of male sexual
characteristics, is also involved in compromising the immunocompetence of
males. In such a situation, only the males with strong immune system can
tolerate high testosterone levels. The variation in testosterone levels among
the men leads to differentially masculinized physical attributes. This, in turn, is
an indicator of their relative fitness as mates. Thus, it may be beneficial for the
women to choose the men with well-developed male secondary sexual
characteristics since their children would also inherit the father’s strong
immune system.

In order to understand the main selection criteria for males, we first need to
take a few basic facts into consideration. On an average, the fertility period of
women is from 13 to 45 years of age, while the fertility period of men is from
13 to 65 years of age. Therefore, youth and youth-related aspects like fertility
and physical attractiveness, are the main mate selection criteria for males. In
evolutionary terms, younger women have two advantages over older women:

i) They are more fertile, meaning that their chances of conceiving after a
mating are more.

ii) They have a higher residual reproductive value, meaning that they
have more reproductive years ahead of them and hence can have more
offspring in the future.

Broadly speaking, both these advantages would lead to more children and this
implies that mating with a younger woman would be more beneficial for a man,
as far as his reproductive success is concerned. Besides youth, men prefer
women whose features and secondary sexual characteristics are more
feminine. These criteria may be explained by the observation that all of these
traits are correlated with higher estrogen levels and hence could indicate
higher reproductive fitness of these women.

The question as to why men prefer women with low WHR can be answered by
the explanation that low WHR correlates with good physical health and high
fertility in women. Conversely, increased WHR is a risk factor for many health-
related problems like cardiovascular diseases, diabetes, hypertension and
some types of cancers. Further, recent studies have shown a direct link
between high WHR and fertility by demonstrating that women with higher
WHRs have more irregular menstrual cycles.

In 1995, a ground-breaking study by Wedekind [Link]. proved that body odour

plays a very important role in human mate selection. It has been postulated
that women prefer body odours of the men whose HLA (Human leucocyte
antigen) genes are dissimilar to their own. HLA genes are involved in antigen
presentation and hence are crucial for immune function. Many scientists have
proposed that mating between individuals whose HLA genes differ from each
other, leads to heterozygous offspring who have better immunity since their
immune system would be capable of neutralizing and destroying more types of
258 pathogens.
Unit 10 Human Behaviour and Its Regulating Factors
So far, we have only considered the sexual strategy with respect to
monogamous relationships. When compared to monogamy, polygamy is a
riskier sexual strategy. It involves multiple rounds of mate searching, an
activity that has proven to be very demanding in terms of time, energy and
Monogamy is a
other resources. However, many researchers claim that polygamy could
mating system
increase reproductive success by allowing men to sire more children with wherein an individual
multiple women. Another proposed advantage of polygamy is that it increases has a single mate for
the chances of the individuals of both sexes to mate with a partner with much a very long duration,
higher genetic quality than their own, who in a monogamous system, may or even throughout
have been unavailable to them as mates. The alleged advantages of life. On the other
polygamy could make one question why monogamy is so prevalent in most hand, polygamy is a
mating system
human cultures. This is because as a sexual strategy, monogamy is much
wherein an individual
safer. It assures a moderate level of reproductive success to every individual
has more than one
of both the sexes. Monogamy offers some other advantages also to men; it mates at the same
saves them from repeatedly spending energy (and resources) on mate search time.
and makes them more confident about their children’s paternity. For women,
monogamy offers the fathers’ protection for their children, thereby increasing
their chances of survival and eventually, reproduction.

A fascinating observation that emerges out of our studies of human mate

selection and sexual strategy is that both males and females seem to prefer
individuals whose facial features show symmetry over those that do not. Of
course, there are exceptions to this rule, but most humans seem to follow this
trend. So, what could be the basis behind such a phenomenon? The most
probable theory for this postulates that the development of perfectly
symmetrical features requires physiological precision which is only observed in
individuals with good metabolism. This implies that symmetry could be
advertising the genetic quality of the individual. Thus, it makes sense that both
the sexes prefer mates with symmetrical features.

Studies of men belonging to different cultures point out that men base their
perception of an ideal mate on the basis of the average of the women in their
immediate local surroundings. The same is true for faces and BMI too. Now,
there are two important things to consider here. The first is that the average of
a particular trait is preferred. The second is that it is the local average and not
the global average that seems to be more important. Let us start by explaining
the first observation. The preference for composite or average traits could be
explained using the idea that during evolution, each newly evolved trait
eventually undergoes a stabilizing selection and therefore the average would
represent the best adaptive strategy. In fact, for many complex traits, the
human population shows a bell-shaped distribution curve with most individuals
exhibiting the average value of the trait (Fig. 10.3). The second observation
that the local average is very crucial for mate section has particularly
interesting implications. This is because it indicates that cultural influences,
which are of course specific to a particular geographical region, play a very
important role in human sexual strategy.

In conclusion, there is no rational way as of now, to clearly discern how much

of the mate selection process in humans is adaptive in nature and how much
of it is based on cultural influences. Nonetheless, it is fascinating that many
cultural norms agree quite well with evolutionary predictions. This correlation
could imply that at least some social norms have adaptive value. 259
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Fig. 10.3: A bell-shaped distribution curve depicting data for men’s heights

SAQ 2
a) Which are the most common alternative approaches that are used to
gain an insight into human mate selection criteria?

b) What are the main postulates of the immunocompetence hypothesis?

10.5 FAMILY RELATIONSHIPS AND

ALTRUISM
You will recall from Units 6 and 7 of this course that the phenomenon of
altruism involves a naturally selected behaviour that benefits the recipient at
the cost of the individual who is performing the behaviour. You will also
remember that for a very long time, the concept of altruism baffled ethologists
as well as evolutionary biologists. This was because the idea of helping others
at the cost of self, appeared to be counterintuitive to the evolutionary theories
of the time. The problem was that there seemed to be no way that altruistic
behaviour could be seen to have adaptive value because the reproductive
fitness of the individual showing altruism was greatly reduced. Thus, it seemed
that this behavioural pattern should not have been passed from generation to
generation. And yet, there have been numerous instances of altruistic
behaviour in the animal kingdom (refer to Units 6 and 7 for details). In this
section, we will discern how altruism is prevalent in humans, especially in the
context of human families. We will also see how the theory of kin selection
can explain the existence of altruism directed towards an individual’s relatives.
Evolutionary scientists conjecture that altruism is a deep rooted human nature,
as helping and cooperating promotes the survival of its own species and is an
260 essential part of social instint.
Unit 10 Human Behaviour and Its Regulating Factors
10.5.1 Family and Kinship
There is enough scientific evidence to believe that right from the prehistoric
times, humans have been living in multilevel societies with more interaction,
cooperation and genetic relatedness among smaller groups. In the modern
times also, we witness a multilevel organizational structure wherein many
families collectively form residential societies, which then constitute a city, then
a state, followed by a nation and finally the entire human species spread
across the world.

Family is the smallest organizational structure in human populations.

Interestingly, the term ‘family’ is subjective and can mean different things to
different people and at different time points. Historically, the family units used
to be big aggregations, but nowadays, nuclear families have become the
norm in most human societies. These small associations are usually
characterized by genetic relatedness which is reinforced through emotional
ties and familial bonds. The most important exception to this need for genetic
relatedness within the family seems to be the spouses. While the spouses are
genetically unrelated to each other, they do share children, who in turn are
genetically related to both the parents. Such nuclear families can also be
described as conjugal families. The expansion of a nuclear family into an
extended family involves the inclusion of in-laws and cousins.

Family units are of particular interest for the study of altruism. This is because
family relationships necessitate a lot of self-sacrifice and concerted effort on
the part of all the family members. In many cases, there is a division of labour
based on the age and sex of the individuals. Thus, each family member
performs specific tasks and the benefits thus accrued (e.g., food, shelter and
other resources) are then shared within the family. Child-rearing is also a very
important aspect of family life and in many cases, the older children help the
parents raise their younger siblings, besides doing small chores for the family.

Besides humans, a few other primates, like chimpanzees and some baboons
also form societies consisting of multiple males and multiple females.
However, organization similar to that of human families is rarely, if ever, found
in any other primate group. The next obvious question is why human
populations would have started getting organized into families? This could be
explained by the fact that owing to the ecology and brain physiology of
humans, the day-to-day life of ancestral humans was much more complicated
than that of the other primates. For example, each individual was supposed to
perform multiple tasks such as acquiring and processing food, searching for
water, finding and guarding shelter, making protective clothes, preparing tools
for self-defence, maintaining social connections and information networks,
caring for the children, etc. Performing all of these tasks on a daily basis would
have been an almost impossible endeavour for any single individual. Also,
humans take a long time to mature into an adult and independent social entity.
Therefore, the help of spouse and older children, with the rearing of the
younger children, in the context of a family, could have been particularly
helpful for increasing the reproductive potential of the parents. Thus,
organization into family groups appears to have offered an effective solution
since this organizational system allowed a division of labour among the family
members. Many families may include the grandparents also, who make
significant contributions to the rearing and training of the grandchildren. They
may also perform quite a few tasks associated with daily sustenance. 261
Block 2 Systematic Approach to Understand Important Animal and
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Compared to all the other primates, the ability of humans to recognize kin is
remarkably well-developed. One of the most direct of the kin detection cues is
of course, facial resemblance. However, direct cues are very few and may be
unreliable at times. Therefore, in the practical world, the indirect cues (i.e.,
cues based on information obtained from a third party or from the
environment) become more important. Towards this end, several
neurobiologists have proposed that the human brain has kinship recognition
circuits (Fig. 10.4) which are very effective at recognizing kin and estimating
the degrees of kinship. These circuits have been hypothesized to operate
using the following mechanisms:

i) Maternal Perinatal Association (MPA): This mechanism is based on

observing the close association between a mother and her new-born
child after childbirth. MPA evolved as a useful cue for the observers to
associate the newborns with their mothers. Such a mechanism is
particularly useful for the recognition of younger siblings by the older
ones. If, for example, the older siblings see their mother taking care of
and breastfeeding a baby, then they identify the baby as their sibling.

ii) Co-residence duration monitoring: As early as 1891, Westermarck

suggested that close association and co-residence during childhood can
serve as powerful cues for kin detection. Thus, human brain appears to
monitor co-residence duration quite effectively and based on that,
determines siblingship. This mechanism is particularly useful for
identification of the older siblings by the younger ones.

iii) Additional indirect cue monitoring: These include mechanisms that

detect other indirect cues such as odours and HLA genes. It must be
noted here that this system has many elements in common with the
system that promotes sexual attraction towards individuals with
dissimilar HLA genes (see the previous section for details).

262 Fig. 10.4: Kinship detection using indirect cues

Unit 10 Human Behaviour and Its Regulating Factors
The inputs from all of these mechanisms are then integrated by a kinship
estimator in the brain, which in turn stimulates altruistic behaviour towards
one’s kin, while at the same time inhibiting sexual attraction towards them.
Thus, besides identifying genetic relatedness, kinship recognition circuits are
also useful for avoiding inbreeding among humans.

10.5.2 Altruism in the Context of Family Relationships

The level of cooperation seen in human families is truly unique and makes
human families excellent target groups for altruism-based studies. We have
already discussed how early evolutionary biologists and ethologists found it
particularly difficult to reconcile themselves to the idea of altruism which entails
an individual helping someone else, sometimes at a great personal cost. In
this context, the theory of kin selection is particularly significant as it offers an
evolutionary explanation for the existence of altruism. The theory was
formulated around 1963-64. The most significant contribution to the theory of
kin selection was that of William Hamilton. However, the term kin selection
was introduced by Maynard Smith in 1964. The theory posits that by helping
genetically related individuals (or kin) increase their evolutionary fitness, an
individual assures the transmission of shared genes to the next generation,
thereby increasing his/her own inclusive fitness.

He explained this phenomenon by arguing that in the evolutionary context, an

organism’s overall reproductive fitness or inclusive fitness is much more
pertinent as compared to its direct fitness which is the direct transfer of its
genes to its own offspring. Mathematically, the term inclusive fitness is
composed of two components. The first component is the direct fitness of the
animal in terms of the transmission of its genes to its offspring. Interestingly,
the second component is the indirect fitness of the individual, which is a
measure of the increase of the reproductive fitness of the recipient due to the
altruistic act. Hamilton argued that any altruistic behaviour that increases the
indirect fitness of the individual significantly, would have adaptive value
because this will ensure the transmission of the shared genes to the next
generation through the genetically related recipient. And any altruistic
behaviour that has adaptive value will get stabilized in the species through
natural selection. Thus, according to Hamilton, the transmission of an
individual’s genes to the next generation’s gene pool is more important than
the survival or reproduction of the individual per se. He then went on to define
the condition that would be conducible for the evolution of altruism, which later
became known as Hamilton’s rule and can mathematically be represented
as:

rB > C

where, r = Genetic relatedness or the proportion of shared genes between the

altruistic individual and the recipient.

B = Benefit provided to the recipient through the altruistic act in terms of the
gain of the recipient’s direct fitness.

C = Cost incurred by the altruistic individual in terms of the loss of its direct
fitness. 263
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In the equation, the product of r and B is the indirect fitness gain for the
altruistic individual. So, in simple words, an altruistic behaviour would get
naturally selected if the indirect fitness gain outweighs the cost (i.e., loss of
direct fitness) incurred while performing it. The higher the genetic relatedness
between the altruistic individual and the recipient, the higher would be the gain
in the indirect fitness and therefore, the more willing an altruistic individual
would be to help the recipient. Hamilton suggested that this was the reason
why individuals were more likely to help their own kin compared to others.
Strictly speaking, this kind of altruism is said to be genetic altruism because it
is directed towards individuals that are genetically related to the altruistic
individual. It has been observed that even outside of family groups, there is a
common tendency among humans to be altruistic. Such altruism, between
unrelated individuals, is called reciprocal altruism (refer to Unit 6 and 7 for
more details).

The advantages of altruism are not limited to increasing the individual’s

inclusive fitness; it has other advantages too. Neurobiologists have discovered
that performing altruistic acts activates the pleasure centres of human brain.
This feel-good factor motivates the individual to perform further altruistic
behaviour. Another advantage of altruism is that it may predispose others to
perceive the altruistic individual as a ‘good person’ and therefore, if such a
person is in distress, then they are more likely to get help from their kin.
Obviously, all the benefits of altruism are indirect because true altruism is, by
definition, a selfless act and is not performed for any direct personal gains.

SAQ 3
a) Name any two mechanisms that are important for the efficient
functioning of kinship recognition circuits.

b) Write down the mathematical representation of Hamilton’s rule.

10.6 LINKING BEHAVIOURS TO BRAIN, GENES,

HORMONES AND ENVIRONMENT
The ‘nature versus nurture’ debate has always confounded scientists as well
as philosophers. In fact, it is one of the longest debates in the history of
mankind. And despite all the scientific advancements of the modern era, we
are still not in a position to settle this debate and decide if it is our genes or our
external environment (which includes our circumstances) that determine who
we are and how we behave. However, what we do know is that quite a lot of
diverse factors, intrinsic as well as extrinsic, regulate human behaviour. The
most important ones include the brain, genes, hormones and environment.
Their impact on behaviour is interdependent on each other. For example, the
development of the brain is controlled by genes. In turn, the genes, along with
hormones and environment affect behaviour indirectly via the brain. First, the
brain receives the external and internal stimuli through the sensory receptors.
Then it processes all the inputs, determines an appropriate response and
finally relays this information to the various effector organs through
264 neurotransmitters and hormones.
Unit 10 Human Behaviour and Its Regulating Factors
10.6.1 Behaviour and Brain
Since time immemorial, mankind has known that there is a close association
between the human brain and behaviour; so much so, that when we behave
badly in a situation, we curse our brains for misleading us. Incidentally, there is
a lot of scientific evidence behind such assertions! The biggest evidence is the
fact that human brain is sexually dimorphic. In other words, the brains of males
and females differ in many structural and functional aspects. This explains
how males and females show sex-specific differences in behaviour.

In order to understand the regulation of behaviour by brain, let us first take a

look at the overall organization of the human brain (Fig. 10.5) and see which
parts are particularly relevant for regulating behaviour. The human brain is
composed of three main parts; forebrain, midbrain and hindbrain.

Fig. 10.5: Illustration of the overall organization of the human brain

i) Forebrain (prosencephalon): It is the forward-most portion of the brain

and is so big that it covers the midbrain and parts of the hindbrain.
Forebrain is responsible for carrying out higher cognitive functions,
regulating emotions and the final processing of the sensory information.
Therefore, it is not surprising that the human forebrain is proportionately
much larger compared to that of the other animals and is responsible for
generating the human-specific behaviours. The forebrain has many
specialized regions, out of which five major parts are cerebral cortex,
thalamus, hypothalamus, amygdala and hippocampus. Parts from
many different structures such as hypothalamus, hippocampus,
amygdala, thalamus and many smaller regions of the forebrain form the
limbic system. It is not a particular structure but rather a collection of
diverse parts of the forebrain that are coordinated together to regulate
the homeostatic machinery of the body. The limbic system thus
regulates a lot of processes including behaviour, emotions, long-term
memory, decision-making, appetite and olfaction. Besides being a part of
the limbic system, the hypothalamus also plays a major role in 265
Block 2 Systematic Approach to Understand Important Animal and
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generating motivation for behaviours like eating, drinking, aggression,
etc. Also, hypothalamus controls the pituitary gland, which in turn
controls the entire endocrine system. Let us now discuss the largest part
of the forebrain, the cerebral cortex, which is primarily composed of four
lobes (Fig. 10.6):

• Frontal lobe: It is in the foremost part of the cerebral cortex. The

sensory input from all the sense organs is finally relayed to the
frontal cortex. It then integrates the sensory input and uses this
information to produce higher cognitive function and behaviour
such as reasoning, abstract thought, planning and judgement. The
hind portion of the frontal cortex contains the primary motor cortex
which is responsible for controlling and coordinating body
movements. Frontal lobe also contains the Broca’s area which
plays a pivotal role in speech production.

• Parietal lobe: It is behind the frontal lobe. It has a limited role in

regulating behaviour and is generally involved in sensing inputs
from the skin, e.g., touch, pain and hot/cold sensation.

• Occipital lobe: It is behind the parietal lobe and just like the
parietal lobe, it too has a very limited role in regulating behaviour.
Its main function is the integration of visual inputs.

• Temporal lobe: It is situated laterally in the cerebral cortex. It

receives and processes auditory signals. It is also involved in
speech production and long-term memory formation.

Fig. 10.6: Diagrammatic representation of the four lobes of the cerebral cortex

i) Midbrain (mesencephalon): It is the middle portion of the brain and

plays a crucial role in processing sensory information and controlling
movement.

ii) Hindbrain (rhombencephalon): It is the part of the brain that connects

the rest of the brain to the spinal cord. Hindbrain is mostly responsible
for basic survival functions such as regulation of heart rate, blood
266 pressure, breathing rate, posture and movement. It is divided into three
Unit 10 Human Behaviour and Its Regulating Factors
subparts: medulla, pons and cerebellum. Out of all these subparts, the
cerebellum allegedly has the biggest influence on behaviour. Recent
studies claim that cerebellum might be involved in higher cognitive
functions as well as in processing of fear, pleasure and pain.

The effect of brain on behaviour is not fixed and can be modulated depending
upon external and internal stimuli. The brain has a tremendous ability to ‘learn’
from our experiences and adjust the behaviour accordingly in the future. For
example, when we perform a particular behaviour, its outcome could make the
brain perceive it as either pleasant or unpleasant. Having learnt from this
experience, in the future the brain would provide motivation to perform the
pleasant behaviour while it would inhibit the unpleasant behaviour.

Since the brain has such a profound impact on behaviour, it is obvious that
damage to selective parts of the brain or their malformation can result in
specific behavioural problems. For example, a recent study has indicated that
in people who display lifelong antisocial behaviour, the cerebral cortex is
thinner at particular sites. Also, such people have reduced cortical surface
area in the regions of the brain which are involved in controlling emotions,
motivations and thoughts.

10.6.2 Behaviour and Genes

The field of behavioural genetics entails the analysis of how variations in a
particular gene can cause variations in behaviour. It is thanks to the modern
sequencing methods, that we can now assign certain behaviour patterns (such
as aggressiveness and response to stress) to the underlying gene mutations.
For example, the gene that codes for serotonin transporter regulates
Reflex action (or
neurotransmission throughout the brain. It has two allelic variants: long and
reflex behaviour) is a
short. The short allele is dominant and is associated with reduced expression fast and involuntary
and function of serotonin transporter. Consequently, individuals carrying this response to a
allele are very susceptible to anxiety and depression. stimulus. It does not
need conscious
Generally, behaviour is affected by genes as well as the external environment. thought.
However, you will recall from Unit 3 that innate (or instinctive) behaviour is
solely controlled by genes, with the environment hardly playing any role.
Therefore, all the members of a species will show the same innate behaviour
when exposed to a particular stimulus. Humans have very few such Epigenetic
behaviours. In fact, reflex actions are the only truly innate behaviours in modifications
humans. An example of an innate behaviour in humans is the sucking reflex in include methylation of
DNA sequences and
babies. Due to this reflex, newborns instinctively start sucking whenever a
post translational
nipple is placed in their mouth. Another example of innate behaviour in
modifications (e.g.,
humans is the knee-jerk reflex which is seen when the tendon below our methylation,
kneecap is tapped with a rubber hammer. Instantaneously, our lower leg gets acetylation,
lifted involuntarily into a kick. phosphorylation, etc.)
of the histone
Surprisingly, it is not just the sequence of genes but also their epigenetic proteins.
modifications that can determine the behavioural traits of individuals.
Eventually, these modifications can regulate gene expression so that they may
decide which genes get to express themselves and to what extent. Some
studies claim that behavioural traits such as boldness, shyness, parental care
and anxiety can be inherited from the parents through epigenetic modifications
of the relevant genes. 267
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Human Behaviours
10.6.3 Behaviour and Hormones
Hormones can be defined as organic chemical messengers that are
synthesized and released by specialized ductless glands called the endocrine
glands. These glands release hormones directly into the bloodstream, thereby
allowing the hormones to act on distant target organs. Hormones can impact
the behaviour and physiology of the animal in numerous ways. The scientific
study pertaining to the interaction between behaviours and hormones is known
as behavioural endocrinology. In Unit 5, we discussed the important
hormones that regulate animal behaviour.

Notably, the interaction between hormones and behaviour is bidirectional. This

means that not only can the hormones affect behaviour, but also the behaviour
can affect hormone levels via feedback mechanisms. For example, in humans,
it has been observed that winning a combat, or even a simulated battle like
chess, can increase testosterone concentrations. On the other hand, the
testosterone levels of the losing side are significantly reduced.
Let us now see how the endocrine system is regulated by the brain (Fig. 10.7).
The hypothalamus region of the brain receives information from the external
and internal stimuli. In accordance with the information received, it influences
the pituitary gland via releasing hormones. The pituitary gland, in turn,
secretes tropic hormones, which help it exert control over the functioning of
most of the other endocrine glands, which then release their own hormones.
The hormones thus produced stimulate their own target organs. The
information relayed from hypothalamus to pituitary to the other endocrine
glands via hormones is very specific. Hence, only specific target organs are
targeted, leading to a very specific response. At the same time, due to
increased plasma concentration of a particular hormone, a negative feedback
loop gets activated which signals to the hypothalamus and the pituitary gland
to decrease the production of that hormone.

Hypothalamus

Releasing hormone

Pituitary gland

Tropic hormone

Other endocrine glands

Hormone

Target organs

Fig. 10.7: Regulation of the endocrine system. The hypothalamus region of the
brain controls the entire endocrine system through the pituitary gland.
Conversely, the increase in the concentration of a particular hormone
provides negative feedback (shown with red arrows) to the pituitary
268 and hypothalamus
Unit 10 Human Behaviour and Its Regulating Factors
The hormones that have the most significant effect on behaviour include:

• cortisol (mediates stress responses)

• estrogen and progesterone (improve cognitive performance, elicit

maternal behaviour, and regulate sexual motivation and performance)

• testosterone (promotes male-specific sexual behaviour, territoriality and

aggression)

• oxytocin and vasopressin (regulate parental care, pair-bonding and

social bonding)

• prolactin (promotes many behaviours associated with parental care).

Besides the behaviours listed above, hormones are involved in mediating a lot
of other behaviours also such as feeding, drinking, learning, memory,
sleeping, wakefulness, etc. Therefore, it is hardly surprising that hormonal
imbalances can lead to many behaviour-related psychopathologies such as
eating disorders, anxiety, insomnia, depression, etc.

10.6.4 Behaviour and Environment

The environment provides the external or extrinsic stimuli that influence the
brain as well as the behaviour. Sometimes environmental stimuli affect
behaviour indirectly by eliciting an emotional response which subsequently
defines our behaviour. The term ‘environment’ is a heterogenous term, The term conspecific
encompassing a multitude of biotic and abiotic components of our is used to describe
surrounding, including but not limited to air, water, food, light, temperature, the individuals that
other individuals (conspecific as well as heterospecific), etc. And as we have belong to same
species while the
already discussed, genes are also important determinants of our behaviour.
term heterospecific
However, the environment and the genes affect behaviour in different ways. is used to describe
While our genes create a predisposition within us for a certain kind of the individuals that
behaviour, the environment provides the actual trigger for the manifestation of belong to different
that particular behaviour. species.

The environment can have a positive or negative impact on our behaviour. Let
us take a look at a few examples of positive influences of the environmental
factors:

• Usually, social and cultural values have been observed to have a

positive influence, especially during childhood and adolescence.

• Children growing up in a safe and organized family environment report

higher levels of satisfaction and happiness in life. In a long run, this
promotes cooperative behaviour in the children and ultimately allows
them to develop mutually beneficial social relationships.

• Children growing up in mentally stimulating family environments have

been observed to have greater cognitive abilities.

• A good family environment is instrumental in instilling good dietary

practices and providing proper nutrition, thereby increasing the physical
and psychological well-being of all the family members. 269
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Human Behaviours
• People with unhindered access to nature and greenery have often
reported an overall sense of well-being with reduced stress levels
besides better cognition and concentration.

• Patients living in well-lit and airy rooms report a more positive outlook
and tend to be more cooperative towards their caretakers. As an added
advantage, this speeds up their recovery process.

The examples mentioned above have very clearly established the fact that
certain environmental factors are essential for normal human behaviour. For
example, the impact of light on behaviour is demonstrated by the occurrence
of seasonal affective disorder (SAD) that affects people during periods with
reduced sunlight, e.g., the winter season. The symptoms include depression,
hopelessness, reduced concentration, decreased interest in activities, etc.

Feral children provide a much more striking example of the importance of

proper environment for normal human behaviour. Feral children are those
human children who have been isolated from all human contact from a very
young age. Thus, a feral child has no significant experience of human
behaviour, affection, care or language. Surprisingly, feral children find it
extremely difficult to acquire normal social skills even after being provided
human company and guidance. Such cases provide strong evidence that
environment plays a very important role in the development of normal
behavioural patterns, especially during childhood.

Humans are territorial beings by nature and therefore, have a heightened

sense of personal space. Invasion of our personal space invokes a stressful
state of mind. And while it is normal to want to avoid crowds, some people
develop an acute fear of crowds. Such a condition is known as
enochlophobia. People having this condition perceive crowds as excessively
dangerous and show irrational behaviour and thoughts whenever they find
themselves among a crowd. On the other hand, in certain occasions people
report that crowds create positive energy, especially at a concert, cricket
match or on the dance floor. On such occasions, humans have been known to
tolerate (and even enjoy!) loud noises, which is also something that we tend to
avoid under normal circumstances. These apparent aberrations can be
explained by the fact that besides being territorial, humans are social beings
too. Heightened emotions during these crowded events create a feeling of
cohesion among the people. Plus, our prior experience of having enjoyed
these events in the past, could also suppress our natural tendency to avoid
crowds and loud noises. Incidentally, much of our behaviour is based on
learning and conditioning. Our past experiences with particular circumstances
have a great bearing on how we would react to the similar circumstances in
the future.

Some environmental factors, such as stress and pathogens, have a negative

impact on behaviour. Stress can alter our brain chemistry, cause dysregulation
of hormonal levels, and affect our mood and behaviour adversely. Similarly,
pathogens in the environment can cause infections and illness, which in turn
270 can make the patients unhappy and asocial.
Unit 10 Human Behaviour and Its Regulating Factors
To conclude, the study of human behaviour is fascinatingly complex and
infinitely rewarding. It finds applications in our daily lives as well as a wide
variety of scientific disciplines. People’s genetic background, brain physiology,
past experiences, surroundings, hormones and many other factors play a role
in determining their behaviour.

SAQ 4
a) Give the scientific terms for forebrain, midbrain and hindbrain.

b) What role does the hormone oxytocin play in human behaviour?

10.7 SUMMARY
In this unit you have learnt that:

• Human behaviour is regulated by many factors such as genetics,

hormones, psychology, culture, values, attitude, social status, etc.

• Non-verbal communication mostly involves visual, auditory or tactile

cues.

• Most of the human non-verbal communication is through visual cues and

involves body language.

• The most common approaches being employed to analyse mate

selection rely on the use of questionnaires, personal advertisements,
and computerized simulations of faces and body shapes.

• The strategy of mate selection varies between the sexes.

• Many social norms for mate selection agree quite well with evolutionary
predictions of the optimal sexual strategy.

• Altruism involves a behaviour that benefits the recipient at the cost of the
individual who is performing the behaviour.

• The limbic system is composed of parts from many different structures

such as hypothalamus, hippocampus, amygdala, thalamus and many
smaller regions of the forebrain.

• Reflex behaviours are the only truly innate behaviours in humans.

• Feral children provide a striking example of the importance of proper

environment for normal human behaviour.

• The sequence of our genes as well as the epigenetic modifications

associated with them, play a substantial role in determining our
behavioural traits.

• The scientific study pertaining to the interaction between behaviours and

hormones is known as behavioural endocrinology. 271
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Human Behaviours

10.8 TERMINAL QUESTIONS

1. Indicate whether the following statements are true or false:

a) Paralinguistic cues are involved in verbal communication.

b) Fidgeting serves as a non-verbal cue.

c) When compared to younger women, older women have a higher

residual reproductive value.

d) Women with higher residual reproductive value have more

reproductive years ahead of them and hence can have more
offspring in the future.

e) When compared to monogamy, polygamy is a riskier sexual

strategy.

f) Forebrain is also known as rhombencephalon.

g) The occipital lobe has Broca’s area which plays a pivotal role in
speech production.

2. Fill in the blanks.

a) ……………… is the scientific term for study of facial expressions,

eye contact, gestures, postures and body movements.

b) The study of touch and the use of tactile stimuli for communication
purposes is known as ……………... .

c) Parts from many different structures such as hypothalamus,

hippocampus, amygdala, thalamus and many smaller regions of
the forebrain form the ………………. .

d) Studies suggest that women prefer body odours of the men whose
……………….. genes are dissimilar to their own.

e) ……………… theory can explain the existence of altruism directed

towards an individual’s relatives.

3. Give the full form of the following terms:

a) MPA

b) BMI

c) WHR

d) SAD

Activity 1

Notice the body language of the people around you. Record your observations
and see if you can pick up some unique patterns that are indicative of their
feelings or emotions.

272
Unit 10 Human Behaviour and Its Regulating Factors

10.9 ANSWERS
Self-Assessment Questions
1. a) Oculesics is defined as the study of eye related non-verbal
communication that involves visual cues such as gaze, eye
contact, eye movement, etc.

b) Proxemics is defined as the study of the space between individuals

and its effect on their behaviour and interpersonal communication.

c) Non-verbal communication is defined as the transmission or

exchange of information among individuals without using spoken
or written words.

2. a) Questionnaires, personal advertisements, and computerized

simulations of faces and body shapes.

b) Immunocompetence hypothesis posits that the hormone

testosterone, which is responsible for the development of male
sexual characteristics, is also involved compromising the
immunocompetence of males. In such a situation, only the males
with strong immune system can tolerate high testosterone levels.
The variation in testosterone levels among the men leads to
differentially masculinized physical attributes, and this in turn, is an
indicator of their relative fitness as mates.

3. a) Maternal Perinatal Association/ Co-residence duration monitoring/

Odour and HLA monitoring

b) rB > C

where, r = Genetic relatedness or the proportion of shared genes

between the altruistic individual and the recipient; B = Benefit
provided to the recipient through the altruistic act in terms of the
gain of the recipient’s direct fitness; C = Cost incurred by the
altruistic individual in terms of the loss of its direct fitness.

4. a) Prosencephalon, mesencephalon and rhombencephalon,

respectively.

b) Regulates parental care, pair-bonding and social bonding.

Terminal Questions
1. a) True/ False, b) False, c) True, d) False, e) True, f) True,
g) False, h) False

2. a) Kinesics

b) Haptics

c) Limbic system

d) HLA (Human leucocyte antigen)

e) Kin selection 273

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Human Behaviours
3. a) Maternal Perinatal Association

b) Body Mass Index

c) Waist to Hip Ratio

d) Seasonal Affective Disorder

Acknowledgement of Figures
Fig. 10.2: [Link]

Fig. 10.3: [Link]

new/srme/modules/mod1/8/[Link]

Fig. 10.5: [Link]

Fig. 10.6: [Link]

274
Volume 1 Animal Welfare-I
GLOSSARY
Action potential : The neural signal; a self-regenerating change in
membrane electrical charge that travels the
length of a nerve cell and may elicit changes in
the activity of a neighboring nerve cell or muscle.

Adaptation : A characteristic that confers higher fitness on

individuals than any other existing alternative
exhibited by other individuals within the
population; a trait that has spread, or will spread,
or is being maintained in a population as a result
of natural selection.

Adaptive value : The contribution that a trait or gene makes to

fitness or individual genetic success.

Allele : A form of a gene; different alleles typically code

for distinctive variants of the same protein.

Alturism : Helpful behavior that raises the recipient’s direct

fitness while lowering the donor’s direct fitness.

Arbitrary culture : The view that human behavior is the arbitrary

theory product of whatever cultural traditions people are
exposed to within a society; thus, our actions are
not expected to be explicable in evolutionary
terms.

Artificial selection : A process that is identical to natural selection,

except that humans control the reproductive
success of alternative phenotypes within the
selected population.

Associated : A seasonal change in reproductive behavior that

reproductive pattern is associated with changes in the gonads and
hormones in contrast to a disassociated
reproductive pattern in which the onset of
reproductive activity is not triggered by a sharp
change in certain hormone concentrations.

Batesian mimic : An edible animal that deceptively resembles

another inedible species and gains protection
against predators as a result.

Causal question : In the scientific method, a question about the

cause of a natural phenomenon.

Central pattern : A group of cells in the central nervous system

generator that can produce a particular pattern of signals
that riggers a functional behavioral response.

Chase-away selection : A form of sexual selection that occurs when

males differ in their ability to exploit the sensory
systems of females in ways that raise male and 275
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reduce female reproductive success, leading to
counter-selection on females favoring those that
are increasingly resistant to male courtship
signals.

Circadian rhythm : A roughly 24-hour cycle of behavior that

expresses itself independent of environmental
changes.

Circannual rhythm : A annual cycle of behavior that expresses itself

independent of environmental changes.

Cladogram : A diagram of the evolutionary relationships

among species or groups of species; a
phylogenetic tree.

Coefficient of : The probability that an allele present in one

relatedness individual will be present in a close relative; the
proportion of the total genotype of one individual
present in another as a result of shared ancestry.

Communication : The cooperative transfer of information from a

signaler to a receiver.

Convergent evolution : The independent acquisition over time of similar

characteristics in unrelated species subject to
similar selection pressures.

Cooperation : A mutually helpful interaction between two

individuals.

Critical period : A phases in an animal’s life when certain

experiences are particularly likely to have a
potent developmental effect.

Cryptic female choice : The ability of a female that has received sperm
from more than one male to choose which male’s
sperm will fertilize her eggs.

Cumulative selection : The effect of repeated bouts of natural selections,

resulting in the accumulation of many small
adaptive changes in an evolving population,
which can add up to a large evolutionary change
over time.

Darwinian puzzle : A trait that appears to reduce the fitness of

individuals that possess it.

Developmental : The capacity of developmental mechanisms

homeostasis within individuals to produce adaptive traits
despite the potentially disruptive effects of mutant
genes and suboptimal environmental conditions.

Dilution effects : Safety in numbers that comes from swamping the

276 consumption capacity of local predators.
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Display : A stereotyped action used as a communication
signal by individuals.
Divergent evolution : The evolution of differences among closely
related species because of differing selection
pressures in their different environments.
Dominance hierarchy : A social ranking within a group in which some
individuals give way to others; often conceding
useful resources to others without a fight.
Ethology : The study of the proximate mechanisms and
adaptive value of animal behavior.
Eusocial : Of or relating to societies that contain specialized
nonreproducing castes that work on behalf of the
reproductive members of the society.
Fertility insurance : An increased likelihood of egg fertilization,
which females can achieve by mating with
several males.
Fitness : A measure of the genes contributed to the next
generation by an individual, often stated in terms
of the number of offspring that survive to
reproduce.
Fixed action pattern : An innate, highly stereotyped response that is
triggered by a well-defined, simple stimulus; once
the pattern is activated, the response is
performed in its entirety.
Free-running cycle : The cycle of activity of an individual that is
expressed in a constant environment.
Frequency-dependent : A form of natural selection in which the fitness of
selection a phenotype depends upon its relative frequency
in the population.
Game theory : An optimality approach to the study of behavioral
adaptation in which the fitness payoffs to an
individual from a particular behavioral tactic are
dependent on what the other members of the
group are doing.
Genetic compatibility : The ability of the genes present in some sperm to
complement the genes present in some eggs,
resulting in an increased likelihood that the
zygote will develop into an especially competent
individual.
Genomic imprinting : The phenomenon in which the phenotypic effect
of a gene depends upon the parent that supplied
it.

Good parent theory : An explanation for mate choice in which choosy

individuals select partners on the basis of how
well they will care for their offspring. 277
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Group selection : Selection that occurs when groups differ in their
collective attributes and those differences affect
the survival chances of those groups.
Hamilton’s rule : The theoretical point made by W. D. Hamilton
that altruism can spread when rb>c (with r being
the coefficient of relatedness between the altruist
and the individual helped, b being the fitness
benefit received by the helped individual, and c
being the cost of altruism in terms of the altruist’s
lost direct fitness.)
Home range : An area that an animal occupies but does not
defend, in contrast to a territory, which is
defended.
Honest signal : A signal that conveys accurate information to
receivers about the signaler’s phenotypic quality
which may be relevant to its fighting ability or
quality as a potential mate.
Hotshot : A male whose attributes are especially appealing
to sexually receptive females.
Hotspot : A location whose properties attract sexually
receptive females to the males able to hold the
size.
Imprinting : A form of learning in which individuals exposed to
certain key stimuli, usually early in life, form an
association with the stimulus and may later
exhibit sexual behavior towards individuals with
similar stimuli.
Inbreeding depression : The tendency of the descendants of individuals
that mate with close relatives to have lower
fitness than non-inbred members of their species.

Inclusive fitness : The sum of an individual’s direct and indirect

fitness.

Indirect fitness : The genes contributed to the next generation by

an individual indirectly by helping nondescendant
kin, in effect creating relatives that would not
have existed without the help of the individual.

Instinct : A behavior pattern that reliably develops in most

individuals and that promotes a functional
response to a releaser or sign stimulus the first
time the stimulus elicits the response.

Interneuron : A neuron that relays messages either from

receptor neurons to the central nervous system
(a sensory interneuron) or from the central
nervous system to neurons commanding muscle
278 cells (a motor interneuron).
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Kin select ion : A form of natural selection that occurs when
individuals differ in ways that affect their parental
care or helping behavior, and thus the survival of
their own offspring or the survival of non
descendant kin.

Learning : A durable and usually adaptive change in an

individual’s behavior traceable to a specific
experience in that individual’s life.

Lek : A traditional display site that females visit to

select a mate from among the several to many
males displaying on small, resource-free
territories.

Mate guarding : Actions taken by males (usually) to prevent a

sexual partner from acquiring sperm from other
males.

Natural selection : The process that produces evolutionary change

when individuals differ in heritable traits that are
correlated with differences in their individual
reproductive success; also known as direct
selection.

Nuptial gift : A food item or other useful present transferred by

a male to a female just prior to or during
copulation.

Operant conditioning : A kind of learning based on trial and error, in

which an action, or operant, becomes more
frequently performed if it is rewarded.

Operational sex ratio : The ratio of receptive males to receptive females

during a given period.

Optimal foraging : An explanation for the feeding decisions made by

theory animals based on the assumption that individuals
are attempting to maximize their net fitness gains
by, for example, maximizing caloric intake or
some other dietary gain that correlates with
reproductive success.

Parental investment : Costly parental activities that increase the

likelihood of survival for some existing off-spring,
but that reduce the parent’s chances of producing
offspring in the future.

Parent-offspring : The clash of interests that occurs when parents

conflict can gain fitness by withholding parental care or
resources from some offspring in order to invest
in another round of reproduction at a time when
existing offspring would benefit from receipt of the
investment. 279
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Phenotype matching : A proximate mechanism of kin discrimination in
which an individual’s behavior toward another is
based on how similar they are in some way, such
as odor or appearance.

Pheromone : A volatile chemical released by an individual as a

scent signal for another.

Phylogenetic tree : A diagram of the evolutionary relationships

among species or clusters of species.

Phylogeny : An evolutionary genealogy of the relationships

among a number of species or clusters of
species.

Polyandry : A mating system in which a female has several

partners in a breeding season.

Polygyny : A mating system in which a male fertilizes the

eggs of several females in a breeding season.
This mating system takes many forms including
female defense polygyny (in which males guard
groups of mates), lek polygyny (in which males
with unusually attractive traits or display
territories attract several mates), resource
defense polygyny (in which males with
unusually large amounts of resources attract
several mates), and scramble competition
polygyny (in which males that are unusually
good at finding mates become polygynous).

Polygyny threshold : An explanation for why females might enter into a

model polygynous mating relationship when the
resources gained by joining an already mated
male on his territory may exceed those to be
gained by forming a partnership with an unmated
male.

Polymorphism : The coexistence of two or more distinctive forms

or traits within a species.

Proximate cause : An immediate underlying cause based on the

operation of internal mechanisms possessed by
an individual.

Reciprocity : Reciprocal altruism in which a helpful action is

repaid at a later date by the recipient of
assistance. (Indirect reciprocity involves
reciprocal altruism in which a helpful action is
repaid at a later date by observers of the helpful
individual.)

Releaser : A sign stimulus given by an individual as a social

280 signal to another.
Volume 1 Animal Welfare-I
Reproductive success : The number of surviving offspring produced by an
individual; direct fitness.
Runaway selection : A form of sexual selection that occurs when
female mating preferences for certain male
attributes create a positive feedback loop favoring
both males with these attributes and females that
prefer them.
Satellite male : A male that intercepts females drawn to a calling
or resources-defending rival of the satellite.

Search image : A perceptual screening mechanism used by

predators to search visually for cryptic, edible
prey.
Selection : The effect of differences among individuals in
their ability to transmit copies of their genes to the
next generation.
Selfish herd : A group of individuals whose members use
others as living shields against predators.
Sensory exploitation : A situation in which a signaler is able to tap into a
preexisting sensitivity or bias in the perceptual
system of a receiver, thereby gaining an
advantage in transmitting a message to that
receiver.
Sex role reversal : The occurrence of competition among females for
access to males, which may choose selectively
among potential mates.
Sexual dimorphism : A difference between males and females of the
same species.
Sexual selection : A form of natural selection that occurs when
individuals differ in their ability to compete with
others for mates or to attract members of the
opposite sex.
Sign stimulus : The effective component of an action or object
that triggers a fixed action pattern.
Sociobiology : A discipline that uses evolutionary theory as a
foundation for the study of social behavior; often
used to refer to adaptationist studies of human
reproductive and social behavior.
Stimulus filtering : The capacity of neurons and neural networks to
ignore stimuli that could potentially elicit a
response from them.
Strategy : A set of rules for behavior controlled by an
inherited proximate mechanism such that the
differences between individuals in their strategies
are linked to differences in their genes. 281
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Synapse : The point of near contact between once nerve
cell and another.

Tactic : A behavior pattern that is specified by an evolved

strategy.

Territory : An area that an animal defends against intruders.

Test : In the scientific method, actual results that permit

one to evaluate a hypothesis by comparing the
evidence against the predicted results.

Ultimate cause : The evolutionary, historical reason why

something is the way it is.

282