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Altruism as a Handicap: The Limitations of Kin Selection and Reciprocity

Journal of Avian Biology

March 1995
26(1)

DOI:10.2307/3677205

Authors:

Amotz Zahavi

Tel Aviv University

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Nordic Society Oikos

Altruism as a Handicap: The Limitations of Kin Selection and Reciprocity

Author(s): Amotz Zahavi

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Journal of Avian Biology,

Vol. 26, No. 1 (Mar., 1995), pp. 1-3

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Point-of-View

Altruism as a handicap

- the limitations

of kin selection and

reciprocity

Amotz

Zahavi,

Inst.

for Nature Conservation

Research,

Tel-Aviv

University,

Tel-Aviv

69978, Israel.

The phenomenon

of "helping

at the nest" has exposed

ornithologists

to the

problem

of the evolution of altruism,

i.e. why should a non-breeder invest in the fitness of

breeders rather than

trying

itself to breed. Until the 1960s

altruism was explained by many biologists through

mod-

els of "group

selection"

(GS), i.e. the investment of the

non-breeders

in the

breeding

of their

group

members was

justified by its contribution

to the group.

GS was later

rejected by most

evolutionary biologists (Maynard

Smith

1964, Lack 1966, Williams 1966), and another

model,

that of "kin-selection"

(KS) (Hamilton

1964) was sug-

gested

as an alternative solution

to the problem

of altru-

ism. In KS models

the investment

of the altruist is justi-

fied by the benefit to its relatives.

However,

the many

studies on group

living birds

which have resulted

from

the growing interest

in the dilemma

of altruism,

have

revealed that

the

helpers

are

in fact often

not related to the

breeders

(Ligon and Ligon 1990, Reyer 1990, Zahavi

1990). Trivers

(1971) suggested

an additional

model -

"reciprocal

altruism"

(RA)

- to interpret

altruistic

adapta-

tions

among

non-relatives.

This latter model

suggests

that

the investment of the altruist is compensated

by a gain

from a reciprocal

investment

by other

group

members.

However,

data from several

field studies have indicated

that

in many

cases the act of the non-related

altruist

was

not

reciprocated.

In these cases neither KS nor

RA could

explain helping

at the nest. It was evident,

however,

some of these studies (Ligon and Ligon 1990, Reyer

1990, Zahavi 1990) that many of the helpers

were en-

hancing

their own chances to breed. Helping by non-

birds, which is not reciprocated,

can therefore

only be interpreted

as a simple

selfish investment.

How-

ever,

these

findings

did

not

stimulate

further

research

into

the possibility

that "helpers"

in general gain direct

ad-

vantages

by their

apparently

altruistic

investment.

Weaknesses of previous models

In the following I note the inherent

instability

in the

models based on GS, KS or RA and propose what I

consider

to be a more

general

and

stable

model,

in which

helping

is considered

as a selfish

behaviour,

i.e. its initial

presentation

as an altruistic

adaptation

was misleading.

The model also has implications

for interpreting

other

altruistic

phenomena,

even among the eusocial insects,

but these will not be discussed

here.

Group

selection.

GS is not an illogical

model: it pays to

invest in a group,

if the benefit of having a successful

group

exceeds the investment of the individual members

in the general

welfare of the group.

However,

at present

GS is generally

not used

by most

evolutionary biologists,

because it has been shown to be vulnerable to social

parasitism

(Maynard

Smith

1964).

The reason is that,

in a

GS model,

a member

of the group

that does not invest

the group's

welfare

(a social parasite),

gains

as much

the individuals

that do invest in the group,

without

in-

curring

the cost of helping.

Kin selection.

According

to KS theory,

altruism is based

on a model

of individual

selection

in which the gene for

altruism is the selected unit (Dawkins

1989).

The theory

claims that the frequency

of the gene for altruism

in-

creases in the population

as a result of the altruistic

behaviour,

even though

it decreases the reproduction

the altruist

itself. Is this

really

so? The best

way to expose

the fallacy of this claim is to tell a variant

of a story

attributed

to J.B.S.

Haldane,

who suggested

(1955)

that

one of two brothers

walking

beside a river,

were to fall

into

it and be in danger

of drowning,

it would

be reason-

able

for the other brother

to risk his life somewhat to save

the drowning

brother,

since by taking such a risk (i.e.

decreasing

his fitness), he may save his brother and

increase

the frequency

of genes similar

to his own in the

following generation.

The instability

of the model is clearly apparent

if the

same

story

is told

with three or more

brothers,

rather

than

two, walking along

the river. It is obvious

that

if one of

them

jumps

to the

rescue,

the other

sibling

(who does not

risk

himself),

gains

as much

as the

one who

risks

himself,

but

without

incurring any

cost. Thus,

in KS models,

as in

GS models,

the total

gain

of the selfish

brother

(the

social

parasite),

is higher than that of the altruist.

Eshel and

Motro

(1988), trying

to untangle

the problem

of multiple

1 JOURNAL OF AVIAN BIOLOGY 26:1 (1995) 1

potential rescuers, suggested an unrealistic

model, in

which there is no interaction or communication

among

the potential

rescuers,

and

consequently,

no way for the

brothers to predict

which of them, if any, is likely to

rescue the drowning

sibling.

Models of KS are in fact models of GS among

kin.

They are equally

unstable.

Although

the investment and

gain

in GS models are not

presented

by gene

frequencies,

any

investment is ultimately

turned into

gains

or losses in

fitness.

Thus,

because

of the

potential

advantage

to social

parasites

in models of KS and GS, both are equally

unstable over evolutionary

time.

The similarity

between GS and KS has recently

been

discussed

by Wilson and Sober

(1994). They suggested

that this similarity

justifies the use of GS, while I con-

sider both

GS and

KS to be equally

inadequate

to explain

social evolution. It is certainly

not

reasonable

to reject

and accept

KS.

Reciprocal

altruism. It is easy to show that reciprocal

altruism

(RA) is also inherently

unstable.

Trivers

(1971)

suggested

that

RA could

be stable

in cases in which there

are mechanisms that

ensure

reciprocation.

He suggested

that among higher

animals,

such as man and monkeys,

the social parasite,

i.e. the individual that

does not reci-

procate,

will be punished.

However,

like any other

char-

acter,

such a mechanism is costly to the individual

that

possesses

it (Zahavi 1981).

Sigmund

(1993)

discussed

the

difficulty

in enforcing

reciprocation

and

regarded

it as a

major

problem

for models of RA. Thus,

as in GS and KS

models,

selfish

individuals that

do not

invest

in punishing

a social parasite,

gain more than those that invest in

maintaining

reciprocation

within

their

group.

An alternative model

Our studies of the Arabian

Babbler

Turdoides

squam-

iceps, a group-breeding songbird,

suggest a completely

different interpretation

for the motivation of birds to

invest in helping

their

group:

an interpretation

that

may

provide

a general

solution to the problem

of altruism.

suggests

that the investment

in the welfare of the group,

or of its members,

functions to advertise the quality

and

motivation

of the helper.

The advertiser

gains from its

investment

by increasing

its "social prestige".

Helping

may thus be considered as a simple selfish character.

Earlier

(Zahavi

1976, 1990) I used the term

"social

sta-

tus"

instead of "social

prestige",

however,

"social

status"

is often

used as a synonym

for "social

rank".

To empha-

size the difference

- the rank order of babblers

does not

change as long as they remain

members

of the same

group,

whereas their

social

prestige

may

change

(without

a change

in rank),

i.e. other

individuals

may

compromise

with them

to a greater

or lesser

extent,

as a consequence

of their

performance.

Social

prestige

functions like

peacock's

tail

or the

song

of a songbird.

It attracts

collaborators

and deters

rivals.

Prestige may be gained by investing

in wasteful

charac-

ters (Zahavi 1977a, 1987) as well as by investing in

"altruistic"

activities. The investment

involved in the

altruistic activities

serves

as an honest

signal

of the

ability

of the babbler

to help its group and its quality as a

collaborator;

it also reflects

its quality

as a rivar

in intra-

group

conflicts.

It is interesting

to note

that,

contrary

to what

might

ex.ected from all three

models

in which the advantage

the

helper

is indirect

(GS,

KS & RA), babblers

are

highly

motivated

to invest

in their

group,

and the population

not infected

by social

parasites.

On the contrary,

individ-

uals compete

with each other

to invest

in the interests

the group,

often interfering

with the helping

of others.

Dominants

interfere

with the "altruistic"

activities of sub-

ordinates

in feeding the nestlings

(Carlisle

and Zahavi

1986),

in allofeeding

between

adults,

in sentinel

activities

(Zahavi,

unpubl.obs.),

in mobbing

(Carmeli

1988,

Anava

1992), and in the defence

of the common

territory

(Za-

havi, unpubl.

obs.). Dominants

often prevent

subordi-

nates from helping the group. They are most likely to

interfere

with older and experienced

individuals,

which

are one rank

lower than

themselves,

and are more

tolerant

towards

the investment

of young

and

inexperienced

birds

of lower ranks.

Competition

and interference

among

indi-

viduals acting as helpers

have been observed

in other

studies

of cooperatively

breeding

birds

(Reyer

1990),

but

these

phenomena

were

not

considered

by the observers

a challenge

to the theories

of indirect

selection.

These

kinds

of interference

cannot be explained by any

model of indirect

selection,

according

to which

individu-

als gain more when other

individuals,

rather than

them-

selves, invest in the altruistic

acts. However, if, as I

suggest,

helping

confers

a direct

advantage

on the

helper,

the interference

and

competition

for opportunities

to help

can easily

be explained by individual

selection,

and

there

is no need for alternative

models to explain

the phenom-

enon of helping.

Investment

in "altruistic"

activities

stabilises the

social

collaborations

by advertising

the motivation

of the altru-

ists to cooperate,

hence it may

erroneously

be considered

as a mechanism that

has been

selected

to ensure

reciproc-

ation. Reciprocation,

however, cannot explain the in-

terference

with the altruistic

acts of others,

or the

fact that

altruism

can also function

as threat.

The

evolution

of cooperation

in groups

of two, such

sexual

mates,

is no easier

to explain

than

that

of larger

cooperations.

Although

some sexual

partners

do abandon

their mates, or do not invest much in their offspring,

frequently

mates invest more than is required

of them.

Sexual

mates,

like individuals in a group

of babblers,

tend

to compete for the investment in the cooperation:

in-

cubating

birds

are

often

reluctant

to be replaced

by their

incoming

mate,

and

frequently

the incoming

bird has to

push

its mate

aside

in order to replace

it (Zahavi,

unpubl.

obs.). I suggest

that

even in collaborations

of two, a large

part

of the investment

can be explained

as an advertise-

ment of the

quality

of the

investor

and of its motivation to

2 JOURNAL OF AVIAN BIOLOGY 26:1 (1995)

continue collaborating, in order to decrease the partner's

tendency to cheat or desert.

It has been suggested that the solution to the "prison-

er's dilemma" could explain the problem of the evolution

of cooperation in general (Axelrod 1984). Hence it has

attracted

much attention among biologists. I suggest that

neither the dilemma, nor its solutions, has much to do

with the real biological world. In the real world, individu-

als assess the qualities and the motivations of their poten-

tial partners and invest in advertising their own qualities

and motivations - before they enter into any collab-

oration. They continue to test each other as long as they

continue to cooperate (Zahavi 1977b). In the prisoner's

dilemma the collaboration is neither preceded by commu-

nication, nor are the prisoners able to talk to one another

during the game. Hence, clever as the "dilemma" and its

solutions are, they have very little, if any, relation with

the world of biology and social behaviour.

It has often been claimed that the observations that

individuals of many species tend to cooperate with their

kin, support

the theory of KS. However, this is not neces-

sarily so. When a new group is formed, members invest

time, energy and risk in advertising their qualities and

assessing the qualities and motivations of their potential

collaborators before they decide with whom to collab-

orate, and how much to invest in the collaboration with a

particular partner. Collaboration with kin may save a

large amount of this investment, because in a group of kin

the individuals know each other, and their rank order and

social prestige have been established long before the

establishment of the new collaboration. In babblers, new

groups composed of non-related individuals, take much

longer to settle down to breed than groups composed of

kin (Zahavi, unpubl. obs.). The fact that it is easier to

collaborate with kin may have an effect on the gene

frequency of the following generation. I term this 'kin

effect'. However, if this effect would cause individuals to

compromise their own direct fitness, in order to increase

their "inclusive fitness", the path would be open for the

success of social parasites and for the disintegration of

the social system.

To conclude, I suggest that all three models, GS, KS

and RA, sophisticated as they seem to be, are not useful

for interpretating

social interactions, because natural se-

lection has evolved a much more stable solution to the

problem: individuals invest in their collaboration in order

to increase their social prestige. They do so by advertis-

ing their qualities and their motivation to collaborate. The

benefit to the group is a consequence of, rather than the

factor that selects for the investment. It is now generally

accepted that a handicap involving waste, such as the

peacock's tail, is a logical and stable strategy to advertise

quality. Altruistic handicaps may operate equally well.

Thus, the problem of altruism is solved when the in-

vestment in the altruistic phenomenon is considered as a

handicap.

Acknowledgements

- The author wishes to acknowledge

the

Program

for Alternative

Thinking

for its financial support.

Thanks are also due to Yoram Yom-Tov and Arnon Lotem

for

their constructive

suggestions,

and to Avishag

Zahavi for her

major

contribution

in discussing

the arguments

and their

pre-

sentation.

I thank Nomi Paz for her editorial

assistance.

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1"* JOURNAL OF AVIAN BIOLOGY 26:1 (1995) 3

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Cooperative breeding occurs when helpers provide alloparental care to the offspring of a breeding pair. One hypothesis of why helping occurs is that helpers gain valuable skills that may increase their own future reproductive success. However, research typically focuses on the effect of helping on short-term measures of reproductive success. Fewer studies have considered how helping affects long-term fitness measures. Here, we analyse how helping experience affects key breeding and fitness-related parameters in the Seychelles warbler (Acrocephalus sechellensis). Importantly, we control for females that have co-bred (reproduced as a subordinate by laying an egg within a territory in which they are not a dominant breeder), as they already have experience with direct reproduction. Helping experience had no significant association with any of the metrics considered, except that helpers had an older age at first dominance. Accounting for helping experience, females that had co-bred produced more adult offspring (≥1 year) after acquiring dominance and had a higher lifetime reproductive success than females that had never co-bred. Our results suggest that, in the Seychelles warbler, helping experience alone does not increase the fitness of helpers in any of the metrics considered, and highlights the importance of separating the effects of helping from co-breeding. Our findings also emphasise the importance of analysing the effect of helping at various life-history stages, as higher short-term fitness may not translate to an overall increase in lifetime reproductive success.

Better to Bend Than to Break? Effects of Rule Behavior on Dominance, Prestige, and Leadership Granting

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Full-text available

Dec 2023

How people handle rules can influence their social standing in the eyes of others, including their appeal as leaders. It stands to reason that people prefer to grant leadership to individuals who follow rather than break the rules. However, preferences for rule abiders are less evident than one might expect. To enhance understanding of people’s responses to (counter)normative behavior, we (a) introduce the concept of rule bending—behavior that infringes a rule without technically breaking it—and (b) draw on the dominance/prestige framework of social rank to illuminate the underlying processes that drive responses to such behavior. In two experiments (Study 1: N = 149; Study 2: N = 480, preregistered), we show that rule breaking (compared to rule abiding) signals relatively high dominance and low prestige, which undermine leadership granting to rule breakers. We further found that rule benders are seen as relatively high on both prestige and dominance, which renders them more attractive as leaders than rule breakers. Finally, we show that the attractiveness of nonabiders as leaders increases under competition when their apparent dominance becomes an asset. We discuss how rule bending relates to rule abiding and rule breaking and consider implications for understanding and managing rule-bending behavior.

Biological Markets: The Ubiquitous Influence of Partner Choice on the Dynamics of Cleaner Fish–Client Reef Fish Interactions

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Full-text available

Oct 2003

The latest interdisciplinary research on the evolutionary models of cooperation. Current thinking in evolutionary biology holds that competition among individuals is the key to understanding natural selection. When competition exists, it is obvious that conflict arises; the emergence of cooperation, however, is less straightforward and calls for in-depth analysis. Much research is now focused on defining and expanding the evolutionary models of cooperation. Understanding the mechanisms of cooperation has relevance for fields other than biology. Anthropology, economics, mathematics, political science, primatology, and psychology are adopting the evolutionary approach and developing analogies based on it. Similarly, biologists use elements of economic game theory and analyze cooperation in "evolutionary games." Despite this, exchanges between researchers in these different disciplines have been limited. Seeking to fill this gap, the 90th Dahlem Workshop was convened. This book, which grew out of that meeting, addresses such topics as emotions in human cooperation, reciprocity, biological markets, cooperation and conflict in multicellularity, genomic and intercellular cooperation, the origins of human cooperation, and the cultural evolution of cooperation; the emphasis is on open questions and future research areas. The book makes a significant contribution to a growing process of interdisciplinary cross-fertilization on this issue.

Alloparental Care

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May 2022

Amotz Zahavi

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Apr 2021

Manpal Singh Bhogal

The Three Brothers' Problem: Kin Selection with More than One Potential Helper. 1. The Case of Immediate Help

Article

Full-text available

Jan 1988
AM NAT

The evolution of reciprocal altruism

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Full-text available

Jan 1971

Robert Trivers

Cooperative nesting in Eurasian birds

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Full-text available

Jan 1976

Amotz Zahavi

The Three Brothers' Problem: Kin Selection with More than One Potential Helper. 2. The Case of Delayed Help

Article

Full-text available

Oct 1988

In situations involving a single relative in distress and several potential helpers, the advantage of leaving the risky job of aiding the relative in need to any of the other potential helpers is quite obvious. Hence, even in Hamilton's condition for altruism in the one-to-one situation is satisfied, it might seem that altruistic behaviour would not evolve if there is more than one potential helper. The evolutionarily stable strategy (ESS) of helping behaviour in situations involving several potential helpers and a necessarily instantaneous response are examined here. If Hamilton's condition is met, the ESS is usually a mixed strategy of altruism and selfishness, with a decreasing (yet positive) probability of helping as the number of potential helpers increases. -from Authors

Pied Kingfishers: ecological causes and reproductive consequences of cooperative breeding

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Apr 1990

H.-U. Reyer

Cooperative breeding is an unusual kind of social behaviour, found in a few hundred species worldwide, in which individuals other than the parents help raise young. Understanding the apparently altruistic behaviour of helpers has provided numerous challenges to evolutionary biologists. This book includes detailed first-hand summaries of many of the major empirical studies of cooperatively breeding birds. It provides comparative information on the demography, social behaviour and behavioural ecology of these unusual species and explores the diversity of ideas and the controversies which have developed in this field. The studies are all long-term and consequently the book summarises some of the most extensive studies of the behaviour of marked individuals ever undertaken. Graduate students and research workers in ornithology, sociobiology, behavioural ecology and evolutionary biology will find much of value in this book.

Green Woodhoopoes: life history traits and sociality

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