On modeling cooperation: Linking the laboratory to the real world Richard Schuster Department of Psychology University of Haifa (Israel) University of Haifa

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On modeling cooperation Linking the laboratory
to the real world
Richard SchusterDepartment of Psychology
University of Haifa (Israel)University of
Haifas Forum on Research Methods 4 April, 2006
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On modeling cooperation Linking the laboratory
to the real world
Richard SchusterDepartment of Psychology
University of Haifa (Israel)University of
Haifas Forum on Research Methods 4 April, 2006
On explaining cooperation Linking Psychology
and Biology
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The focus of this series is on interdisciplinary
research
My talk is on cooperation. And how I use an
Interdisciplinary approach This can means
many things...
Today I mean explanation how do we answer the
question why cooperate? I will talk about the
advantages from distinguishing between two kinds
of explanation. Psychological and Evolutionary
.and then combining them within an integrated
psycho-biological framework
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A Behavioral/Psychological explanation looks at
the influences on the behavior of cooperating by
individuals at the time that the behavior is
performed.
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Behavioral / psychological explanation this
offers a proximate explanation in answer to the
question Why cooperate?
Aristotle called this an efficient cause whose
action produces a change of state the kinds of
causes that explain who behaves, when, how
often, in what form
Individual causes stimuli learning history
(memories) current states of motivation emotion
and physiology outcomes rewards, etc Social
causes presence of others which others, ranks,
past interactions
What kinds of things are explained? /who
cooperates and who does not (individual
differences)/how often a given individual will
engage in cooperation/how is cooperation
performed the behavior, coordinated,
cues/whether a given individual would choose
cooperation when non-cooperation is also
available as an option
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One Behavioral /Psychological factor I will
emphasize today is engaging in the behavior of
cooperating
By behaving cooperatively, I will refer to
dimensions of behavior associated with
cooperating that are not associated with
not-cooperating. For example, cooperation
includes non-random choosing of partners that
use each other to work together for shared
outcomes based on coordination, complementary
roles, social cues, etc etc.
I will ask whether behaving cooperatively is
associated with intrinsic states (emotions and
the like) that are also part of the answer to the
proximate question Why cooperate? Is
cooperating intrinsically pleasurable? rewarding?
motivating?
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• An Economic/Evolutionary explanation this
  looks mainly at outcomes that are linked to
  behaving cooperatively with the potential to
  benefit individuals (economic outcomes).
• Such benefits may not follow as direct outcomes
  from cooperating, they can occur at any time
  during an individuals lifetime and they have the
  potential to influence natural selection.

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Evolutionary / economic explanation this
provides what is often called an ultimate
explanation
The link to evolution offers an ultimate
explanation because it offers an answer to the
why question that is framed as why the
behavior might have come to exist as an
adaptation molded by evolution.
It asks not what influences individuals when they
cooperate but how much cooperation pays or is
beneficial/profitable to the individual at any
time during its lifetime ? how outcomes can
contribute to evolutionary fitness by increasing
individual success and ultimately reproductive
success.
In animals, outcome in this context
usually refers to cooperation that pays off
economically in things like more food from
hunting cooperatively, or a victory from fighting
cooperatively that leads to larger territory
and/or more mating partners.
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This economic evolutionary approach also uses
economic outcomes to distinguish between
cooperation and other social behaviors.

• e.g., Robert Trivers, The Evolution of Social
  Behavior (1985)
• social acts are classified by individual
  outcomes
• both parties gain (cooperative)..
• the actor confers a benefit but suffers a cost
  (altruistic),
• the actor gains while inflicting a cost (selfish)

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Unlike a proximate or efficient cause, evolution
does not explain what triggers cooperation in any
individual during its lifetime.Instead, before
an individual is born, natural selection has
already weeded out genes that reduced the fitness
of ancestors, and the selected genes explain a
potential to cooperate that is only expressed
when the favorable conditions arise.
A proximate explanation is still needed to
explain what makes a given individual behave,
how, how often, with whom..
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Another way to look at the difference between
proximate and ultimate explanations is to look
only at outcomes and to distinguish between
short- and long-term outcomes following an act of
cooperation.
Assume a cooperative (or any) behavior is
performed The short-term outcomes are the
outcomes linked to the psychology of cooperation
by operating in the here-and now to determine
the likelihood of that an individual will engage
in the behavior via processes underlying
perceptions, motivations, incentives, emotional
states, reinforcements .at the time of
engaging in the behavior. The long term-outcomes
are the profitable, economic and
fitness-determining consequences at any time
during the life of an individual or its kin that
are linked to cooperation and determine its
evolution via natural selection. These can emerge
months or years later. The link between them
short-term outcomes can activate proximate
mechanisms that function to evoke behaviors that
are ultimately beneficial by leading eventually
to long-term profitable outcomes.
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There are four points to know about the
differences between short- and long-term outcomes
.
1) Short- and long-term outcomes can be different
in different currencies. Two examples play
and illness.
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Consider the widespread behavior of play in
animals and humans.
We can speak of a behavioral bias to play
because individuals usually young ones but not
only - expend time and energy when engaging in
play that is not immediately compensated by any
economic outcomes such as food.
Why do this? Many will say that play leads to
improving skills that will eventually increase
success over the life span ex. play hunting
and play fighting. This is the ultimate
evolutionary explanation.
But how do they know? The likely answer is that
they dont. So why do they do it? They are
having a good time. This then is the proximate /
behavioral explanation processes of motivation,
emotion and reward that evolved to make the
behavior happen in the first place and thereby
lead to the ultimate payoffs.
15
Another example of the difference between
proximate and ultimate illness
Proximate dimensions Disease symptoms such as
fever, pain, sweating, loss of appetite,
psychological states, misery are evoked by
reactions in the body to pathogens and their
effects on systems..
Ultimate dimensions Symptoms are signs of
evolved mechanisms that indicate the operation of
defenses against invading pathogens.the
symptoms are in fact linked to advantages that
increase survival.
As with play, the operation of proximate and
ultimate processes are both explanatory and
complementary for explaining disease ..
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2) Time lag There can be a substantial time lag
between the outcomes that influence behavior and
the outcomes that influence evolution. Long-term
outcomes can guide evolution, but they often
cannot be used as an explanation of behavior when
it occurs. The reason discounting..

• Discounting function rapid decrease in the value
  of delayed outcomes

Value of outcome
0....x Time
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3. Some kinds of outcomes can only influence
cooperation via proximate processes.
Events like intrinsic emotional states (pleasure,
empathy.) can be linked to engaging in
cooperation with others for shared outcomes
helping another costs me but it feels good
Such outcomes provide proximate explanations
when they function as incentives or goals that
motivate and reinforce cooperation when it is
performed.
But such outcomes do not in and of themselves
provide a contribution to fitness unless they
also lead later to beneficial economic outcomes
but this is not always guaranteed.
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4) The same beneficial economic outcomes can
sometimes operate as both proximate and ultimate
outcomes.
Some kinds of outcomes - like food, money, mating
partners etc are both economically valuable and
can be immediately gained following cooperation.
Such outcomes provide a proximate explanation
when they function as hedonic incentives or goals
that motivate and reinforce cooperate when it is
emitted the Law of Effect of learning theories.
Economists speak of utility or expected
utility as a psychological quantity that
represents the value of an outcome for an
individual an incentive to behaveand value as
we know can be a relative thing.
But the same outcomes also provide an ultimate
explanation if they are the kinds of beneficial
outcomes that influence natural selection.
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Which kind of explanation is preferable?It
would not matter if proximate outcomes mapped
onto long-term beneficial consequences that
elevate fitness. Immediate and long-term
benefits would then offer different measures of
the same cause. Either could be measured.
But the type of explanation does matter when
cooperation occurs at levels that exceed
predictions from the economic outcomes available
at the time of cooperating there is too much
cooperation which we call a bias to cooperate
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But first..I want to suggest that our
understanding of cooperation is incomplete and
even distorted when explanation and methodology
are dominated by an economic perspective.
I take advantage of a historical tilt or bias
towards the economic/evolutionary type of
explanation as capable of offering the more
powerful or fundamental way to answer the
question why cooperate? I will refer to three
variants of this perspective the
Evolutionary the Behaviorist and the
Game-theoretical
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The historical bias towards economic/evolutionary
explanations1. Explanation.
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Here is an example of the Behaviorist
perspective, anchored in the Law of Effect that
explains how the likelihood of cooperation is
determined by its reinforcing consequences
following the behavior of individuals
B.F. Skinner (Science and Human Behavior, 1953,
pp. 297-298) .... a 'social law' must be
generated by the behavior of individuals. It is
always an individual who behaves, .andhe
behaves with the same body and according to the
same processes as in a non-social situation."
Note that social behavior as a distinct
category of action has been defined into
non-existence
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Here is an example of the Evolutionary
perspective from behavioral ecologists who use
game theory models to focus on cooperation whose
evolution is determined by beneficial outcomes
that elevate fitness

• Since 1981, game theoretical models have been a
  large part of the literature of animal
  cooperation. .The game matrix focuses not on the
  nature of the behaviours involved but on their
  economic consequences. Game theory's significance
  as a tool for modeling cooperation hinges on the
  idea that one can determine which combinations of
  actions are cooperative by examining the game
  matrix. Hence, students of game theory prefer a
  view of cooperation that de-emphasizes the
  behavioural properties of an interaction, and
  focuses on the economic consequences of an
  interaction.
• We argue that the economic definition of
  cooperation is the best option because it can be
  objectively applied, and it offers us the
  powerful tools of game theory.
• Students of social behaviour should recognize as
  an economically defined interaction that may or
  may not involve coordination. we argue that
  cooperation is an outcome, not a mechanism.
• (and then go on to show cooperation without
  coordination or awareness in ants)
• D.W. Stephens J.P Anderson, Animal Behaviour,
  1997

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and here is the same game-theory bias from
economic theorists
Karl Sigmund, Ernst Fehr Martin A.
Nowak,Scientific American, January 2002It may
seem callous to reduce altruism to considerations
of costs and benefits, especially if these
originate in biological needs. Many of us prefer
to explain our generous actions simply by
invoking our good character. We feel better if we
help others and share with them. But where does
this inner glow come from? It has a biological
function. We eat and make love because we enjoy
it, but behind the pleasure stands the
evolutionary program commanding us to survive and
procreate. In a similar way, social emotions
such as friendship, shame, generosity and guilt
prod us toward achieving biological success in
complex social networks.
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• Game theory treats outcomes as both a proximate
  cause influencing individual motivation and
  behavior and as surrogates for fitness that
  determine evolution

Analogous economic processes are assumed to
operate, maximizing economically-important
outcomes that determine the emergence of
cooperation as a consequence of both proximate
psychological processes (learning, etc) and
ultimate processes of natural selection.
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The historical bias towards economic/evolutionary
explanations2. Methodology
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The issue of methodology arises because the
different kinds of explanations proximate vs.
ultimate - have been associated with different
ways of modeling and analyzing cooperation in the
laboratory.
The domination of evolutionary / economic
explanations is associated with 50 years of
experimental models that de-emphasize the
influence of behavior by exaggerating the role of
payoffs. The influence of behavior is dismissed
in both method and theory.
This was achieved by minimizing or totally
removing the social properties of behaving
cooperatively that are intrinsic to many kinds of
cooperation performed by humans and animals in
the natural world.
28
The Psychologist/ Behaviorist B.F. Skinner
cooperation is defined as an inter-dependent
contingency between the behavior of two or more
individuals and the outcomes achieved by each
whether or not there is social interaction.
B.F. Skinner (1953, p. 311) Cooperation when
the reinforcement of two or more individuals
depends on the behavior of both or all of them
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• 1) The explanation of cooperation resides in the
  contingency between the behaviors of 2 or more
  individuals and outcomes. This is a pure economic
  explanation.
• 2) The behaviors used when cooperating only
  address the secondary question of what
  individuals actually do when they cooperate but
  not why they do it. The why is selfish profit.
• 3) In the natural world, cooperating individuals
  might coordinate actions in complex ways or
  interact and communicate.
• 4) But from an economic perspective, cooperation
  can still be claimed in the complete absence of
  social interaction as long as an individual's
  outcomes also depend on the behaviors of others
  and benefits are obtained by all.
• 5) The neglect of cooperation as a social
  behavior is reflected in a long history of
  experimental models that eliminate the
  differences between the performance of
  cooperation and non-cooperation both are simple
  behaviors performed by anonymous and physically
  isolated subjects.

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Skinnerian laboratory models as simple as
individual reinforcements for two animals that
synchronize two simple, brief acts such as
pressing a bar..
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Non-social cooperation Models with no social
interaction whatsoever are still called
cooperative
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Note the close similarity with the ecological /
evolutionary perspective quoted before

• Hence, students of game theory prefer a view of
  cooperation that de-emphasizes the behavioural
  properties of an interaction, and focuses on the
  economic consequences of an interaction.
• Students of social behaviour should recognize
  cooperation as an economically defined
  interaction that may or may not involve
  coordination. we argue that cooperation is an
  outcome, not a mechanism.
• (D.W. Stephens J.P Anderson, Animal Behaviour,
  1997)

33
Game-theory models more complex inter-dependent
contingencies specified in game theory payoff
matrices that offer choice
Prisoners dilemma
Mutualism
34
But Game-theory models retain all the
impoverished social dimensions of the earlier
Skinnerian models
35
I suggest that this bias toward the
economic/evolutionary in method and theory
represents a historical error that has delayed
efforts to answer the question Why cooperate?
by confounding psychology and biology everything
has been linked to outcome and benefit
To me, this emphasis on economic decision-making
whether in Economics, Evolutionary Biology or
Psychology - represents an unnecessary
denigration of the psychological / proximate kind
of explanation as being somehow soft, imprecise
or impossible to confirm a soft science -
how it is done (psychology) but not why
(economics/evolution)
.whereas the economic approach offers the
security of counting and measuring explicit
events in a hard and mathematical science.This
is an updating of the old Behaviorist issue of
avoiding the kinds of events that allegedly
cannot be specified or confirmed.
..the hijacking of psychology by economics
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But economics alone cannot be used to explain
cooperation that is based on irreducible social
dimensions..
37
The study of cooperation in the natural
worldcooperation as a social behavior.
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Question Do models of isolation and anonymity
have relevance to cooperation in the natural
world?

• The validity of anonymous/isolation games seems
  relevant to those situations in which behaviors
  are performed individually and potentially have
  impact on other anonymous individuals or on
  society as a whole where to throw trash
  whether to make noise in a movie theater
  whether to drive a car with consideration for
  other drivers or pedestrians
• whether to wait in line or jump the queue
• whether to give money anonymously
• whether to conserve natural resources or rare
  wildlife
• whether to work for peace
• There is a social context but it is anonymous,
  lacks personal social interaction and reduces
  cooperation to an individual event.
• If there is a social relevance, it is limited in
  humans to things like reputation, impression on
  others

39
Another kind of cooperation that is also
widespread in humans and animals in the natural
world consists of actions by individuals that
work together by using each other to act
cohesively for jointly-obtained outcomes, as in
team sports, warfare or group hunting. .

• The irreducible social dimensions associated with
  this kind of natural cooperation include
• - cooperating individuals are familiar
• - they work together by using each others
  behaviors and locations to coordinate actions,
  sometimes with complementary roles
• - outcomes are jointly-obtained successes or
  failures
• - outcomes may lead to competition and dominance
  over allocation
• - adjunct social interactions
• (Schuster, Human Nature, 2002 Schuster
  Perelberg, Behavioral Processes, 2004 Schuster
  Berger, in press)

40
To understand this kind of cooperation, I will
try to show that
Levels of cooperation and preference for
cooperation can exceed predictions from
economic outcomes at the time of performance.
This bias is caused by proximate processes linked
to the behavior of cooperating, especially its
social dimensions. This can make cooperation
excessive and uneconomic in the short term.
The social dimensions of cooperation can be
incorporated into laboratory models that will
then evoke the same kinds of behaviors - and the
bias to cooperate - as in the natural world
Cooperation even when excessive or uneconomic
in the short-term can still be economic but
only in evolutionary terms when long-range
consequences can be traced to such behaviors that
influence the natural selection of cooperation
via ultimate processes.
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These issues will be addressed in four parts
1 - Behavior How do animals and humans cooperate
in their natural world?
2 Influence of Behavior Is there a bias to
cooperate that is influenced by the social
dimensions of cooperating?
3 - Models How can cooperation be represented by
laboratory models that incorporate the
behavioral dimensions of cooperation in the
natural world- and the bias?
4 - Explanation How can a proximate and ultimate
processes be incorporated within an integrated
explanatory framework?
42

• 1. Behavior How do animals and humans cooperate
  in their natural world?

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In the natural world, when cooperation is
expressed as joint actions for shared outcomes,
this kind of behavior is associated with
irreducible social dimensions that are absent
when not cooperating. This is widespread in
many species and in different contexts. hunting,
aggression, defense, reproduction...
Three animal examples dolphins, lions and
chimpanzees
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dolphins
Cooperative hunting of bottlenose
dolphins mms//vod4.haifa.ac.il/L/CRI/RM200604/dol
phins2.wmv
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Cooperative hunting of lions Zibalianja,
Botswana Intentional, planned and coordinated
hunt
water
mms//vod4.haifa.ac.il/L/CRI/RM200604/lions
linyanti short film.wmv
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Cooperative hunting by the common chimpanzee, Tai
Forest, Ivory CoastIntentional, planned and
coordinated hunt

• mms//vod4.haifa.ac.il/L/CRI/RM200604/chimp hunt
  Tai Oct 05.wmv

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Analogous examples of human behaviors.......
Teamwork with complementary roles

• Distributed cognition group performance
  divided among team members. While cooperating,
  each individual performs a role so specialized
  that it would be ineffectual if the individual
  was forced to perform the entire task alone. This
  kind of organization ..permits individuals to
  combine their efforts in ways that produce
  results that could not be produced by any
  individual... working alone. (E. Hutchins,
  Cognition in the Wild, 1995)

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1. In all these cases, cooperating becomes an
   extended action based on individuals acting
   cohesively

2. The only cues available during coordination
are social from locations and behaviors of
partners
3. While cooperating, there is unrestricted
social interaction
4. Outcomes are also a social event they are
jointly achieved and potentially shared, with
the possibility of competition
5. Strategies develop within groups for working
together? Pairs and groups therefore vary in
cues coordination roles
dominance allocation of outcomes
social behaviors .
49

• 2. Is there a bias to cooperate that is
  influenced by the social dimensions of
  cooperating?

50
The bias is known from observing human behavior
in the natural world of group decision-making
and joint action by groups Robert Frank,
Passion Within Reason, 1988 Sober Wilson,
Doing Unto Others, 1998In Anthropology
hunter/gatherers (pygmies, Bushmen, Ache) work
of Colin Turnbull.
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But Human subjects also cooperate more than
expected in the impoverished conditions of
game-theory experiments..
Fehr et al, 1999-2005 Haselhuhn and Mellers,
2005Reviews Dawes, 1986 Palameta Brown,
1999 Colman, 2003 and commentaries
(in Behavioral and Brain Sciences BBS)
52
Humans apparently play the games as if they have
social dimensions..

• If inform subjects about the participation of
  others? their behavior can be modified (e.g.,
  Baker Rachlin, 2002).
• Brain imaging Rilling et al 2002, 2004
• Social factors Suleiman et al, Ido Erev et al,
  Ilan Fischer.
• Emotions Haselhuhn and Mellers, 2005

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To explain the human bias towards cooperating, it
is suggested that humans are influenced by
mediating psychological states that are products
of culture/religion/moral systems social
orientation, group thinking, reputation, or
merely the desire to avoid sanctions or
embarrassment when caught acting selfishly.
Another proposal (by Fehr et al) is centered on a
strong motivation in cooperating humans to punish
selfish non-cooperators as a means to encourage
them to be more altruistic
But there are problems with this human cultural
learning model.
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One problem there is usually little direct
evidence for such states apart from the bias
towards cooperation. Such explanations risk the
danger of circularity.
A second problem the same bias can also
characterize animal cooperation when it includes
the kinds of social dimensions that characterize
cooperation in the natural world that is based on
joint actions for shared outcomes
55
Animals do not show the bias when cooperation
occurs in the absence of social interaction, as
in the isolated conditions of game theoretical
experiments ? the preference is now NOT to
cooperate e.g., In prisoners dilemma games,
animals prefer to defect towards non-cooperation
and the the larger potential outcome
Prisoners dilemma
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• Rats Flood, M, Lendenmann, K., and Rapoport, A.,
  1983. A 2 x 2 game played by rats different
  delays of reinforcement as payoffs. Behavioral
  Science, 28 65-78.
• Pigeons Baker, F. and Rachlin, H., 2002b.
  Self-control by pigeons in the prisoner's
  dilemma. Psychonomic Bulletin Review, 9
  482-488.
• Blue jays Clements, K.C. and Stephens, D.W.,
  1995. Testing models of non-kin cooperation
  mutualism and the Prisoners Dilemma. Animal
  Behavior, 50 527-535.

Animals are acting as if they are behaving alone
and either do not detect the inter-dependency of
the reinforcement contingency or are unaffected
by it..
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But animals reveal the bias when the social
dimensions of cooperation become
explicit..and this can be shown in their
natural world
Again, dolphins, lions and chimpanzees.
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Cooperation bias in dolphins

• Alliances among male adolescents? no obvious
  outcome at the time and for many years
• The Same animals as adults mate guard adult
  females for mating that is NOT always shared
  equally (Connor et al, 2000)

• The PhD work of Amir Perelberg on free-swimming
  dolphins that spontaneously approach trainers to
  receive petting at the Dolphin Reef tourist site
  in Eilat..
• The dolphins approach trainers singly or by
  coordinating their approach in pairs
• Petting can therefore be treated as a desired
  resource Is this what motivates the dolphins to
  approach?

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Is it costlier to be petted together?
Coordination level of each dolphin with petter

• 3.050.19 meanSE separations/min 2.540.14
  meanSE separations/min
• Wilcoxon Signed Ranks Test, N13, Z-2.27,
  P0.023

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Is there any preference for petting alone or
together?
Allocation of petting
Proportions of petting bouts count
Wilcoxon Signed Ranks
Dolphins prefer to be petted together even though
they pay a price in reduced physical contact
61
Cooperation bias in lions

• Learning Slow learning by cubs with almost NO
  material reinforcement from hunting together
  before about 2 years of age (Scheel Packer,
  1991), coordination is poor, the targets often
  wrong, and the probability of a kill minimal.
• ? if cooperation learning in lions is based on
  reinforcing outcomes, nutritional gain does not
  seem to be the exclusive or even the primary
  motive.
• Preference for cooperation in experienced
  huntersMore food can be obtained by adults that
  hunt alone even though the likelihood of making
  a kill increases, lions that hunt together must
  then share the prey (Packer, Scheel Pusey,
  1990) ?

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Lions in the Serengeti
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Cooperation bias in chimpanzees
Even slower learning than in lions? minimal
success (little food reinforcement) for c. 10
years (Boesch, 2002 Nishida, film)
No evidence for food as primary motive ? not
related to hunger or absence of other food
(Mitani Watts, 2001)
Sharing outcomes with non-hunters for political
purposes
Unequal allocation of outcomes ?
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Unequal allocation of outcomes? dominants get
more meat is shared with females and some
cooperators get little or nothing (Boesch
Boesch, 1989)
Size of prey Keep all (respect) Theft Transfer Division
Small (infant or juvenile) 22 3 1 2
Large (adult) 3 1 4 25
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Question Is this still cooperation?

• Not from an economic rational perspective
  limited to outcomes
• Immediate beneficial outcomes may be minimal or
  absent
• Outcomes may not be allocated equally
• Behavior is effectively altruistic
• But from a behavioral perspective, the behavior
  is phenomenologically cooperative because it is a
  social and coordinated joint actionand because
  it captures the altruistic dimension of
  cooperating (doing for others) animals are
  using each others behaviors and locations for
  obtaining shared outcomes when more could be
  obtained by operating alone, or when the
  immediate economic outcome is not the primary
  goal
• If we assume that there is immediate
  reinforcement, I will suggest later that this
  behavior can be explained by additional immediate
  reinforcement that is not economic in the
  accepted sense because it arises from motives and
  emotions associated with the behavior of
  cooperating itself.

66
This also changes the way we characterize the
process of decision-making about whether or not
to cooperate
From an economic perspectivethe choice is
framed as a decision governed by outcomes and
their expected utility the form of the behavior
is irrelevant

• From a behavioral perspectiveThe choice is
  framed not only as a choice between different
  material outcomes such as food, money, etc.
• but also as a choice between different kinds of
  behaviors
• cooperation associated with irreducible social
  dimensions
• non-cooperation associated with the absence
  of those social dimensions.

67
This is consistent with a link to more basic
behavioral/ psychological processes in animals
linked to behaving cooperatively..and the
possibility that these are shared with humans.
68

• 3. Models How can cooperation be modeled in the
  laboratory to incorporate the dimensions of
  cooperation in the natural world and the
  cooperation bias?

69
Our goal was to incorporate more of the social
dimensions associated with cooperation into our
models in order to study the influences of
behaving cooperatively on performance and choice.
70
Cooperation pairs of laboratory rats that are
reinforced for working together, using each other
to coordinate behaviors
The behavior to coordinate movements within a
shared space with unlimited social interaction
Incorporates features of cooperative behaviors
such as group hunting and aggression as they
occur in the natural world
The economic reinforcement water sweetened with
saccharine
71
Zugia (Heb"pair) a model of social
cooperation
94 cm
mms//vod4.haifa.ac.il/L/CRI/RM200604/rats.wmv
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Contingency and control..

• Unlike isolation models with greater prediction
  and control over individual behaviors
• The behavior?outcome contingency in this kind of
  model is defined at the level of a dyad
• Within the limits set by the contingency, pairs
  are free to develop strategies with differences
  in dominance, control, roles, etc. each pair
  develops its own way of coordinating
• There are also differences across pairs in levels
  of coordination and social interaction

73
Non-cooperation single reinforcement for
individual performance of the same back-and-forth
shuttling, even if others are present
1 cup
Floor D
Floor N
Floor M
94 cm
74
1. Cooperating pairs learn to work together
Stage 1
75
Then every subject given a new, naïve partner for
cooperating
Stage 1
Stage 2
Group 1
Group 2
76
former cooperators learned faster with new naïve
partners than former non-cooperators
Stage 2
77
2. Pairs only learn to coordinate when reinforced
specifically for cooperating
Stage 1 Cooperative contingency
Stage 1 Individual independent
contingencies withmatched reinforcement
78
Sharing the same space is not enough to
coordinatecooperation is controlled by the
requirement to cooperate
But pairs coordinated only when reinforced for
cooperating.
Pairs Cooperative
Pairs Independent
79
What happens when choosing between these
behaviors andoutcomes are matched?
???
Economic perspective.?
Behavioral perspective.?
80
Stage 1First learned the two taskseach task
in a separate chamber 10 sessions
81
Stage 2 Then learned the locations of the
chambers 6 sessions/12 trials
Different floors
Insert guillotine doors
82
Stage 3Free choice 4 sessions/ 8 trials
83
Two experiments cooperation was not more
profitable in terms of reinforcement (28 subjects)
84
The 2nd experiment (n22 subjects)
85
Yet cooperation in Exp. 1 was strongly preferred
(n28)
86
And cooperation was again strongly preferred in
Exp. 2
87
Choice was not linked to relative
reinforcementRelationship between choices for
cooperation (Stage 3) and relative reinforcement
during Learning Stage 1 (sessions 6-10)
88
Choice was linked to a pairs level of
coordination.
High coordination -gt Low coordination
89
The bias to cooperate was also shown by
allocation of outcomes that did not strongly
influence learning and performance.
90
Experiment on competition over outcomes

• Completed coordination

Completed coordination
91
Measuring competition and dominance when a
single cup is presented, is it accessed by the
owner or invader
Cup presentations
Invader?
92
Dominance within pairs was expressed as the
degree of invasion of the partners cup by the
non-owner when presented alone
Invader displaces owner
Owner
5
Invader displaces owner
Invader
6
93
Majority of pairs eventually competed with cup
dominance level 5 (Scale 0?7) over Sessions 7-12
94
Effect of outcome dominance on cooperation? None
(n 48 pairs in 4 sub-groups Sessions 7-12)
Increasing dominance ?
95
Summary levels of cooperation and the decision
about whether or not to cooperate are also
influenced by the behavior of cooperating that
can lead to a bias..

• In animals and humans, this is present in the
  field
• In laboratory models with animals, this can be
  evoked when cooperation incorporates social
  dimensions analogous to those in the field(the
  issue of external validity)

96

• Question 4 - Explanation How can proximate
  behavioral processes and ultimate evolutionary
  processes be incorporated within an integrated
  explanatory framework?

97
Proximate Explanation if you are a
psychologistWhat proximate processes operating
at the time of behaving could explain the bias
towards cooperating?
98

• 1) Cooperation is reinforced by two kinds of
  immediate outcomes
• A) Material economic gains (if any) (food,
  mates, money)
• B) Intrinsic reinforcement from the act of
  cooperating with others
• from coordinating actions?
• from affiliative social behaviors?
• from relationships that develop while
  cooperating?

99

• Immediate outcomesfrom cooperating

Economic outcomes surrogates for fitness (food,
money, etc)
Intrinsic outcomes from states associated
with cooperating(emotions, etc.)
100
One behavioral function of intrinsic
reinforcement would be to provide an immediate
incentive when economic outcomes are absent,
insufficient or delayed at the time of cooperating
Act of cooperation
Immediate economic outcomes
Immediate intrinsic outcomes
Immediate economic outcomes

• when learning to cooperate is difficult
• when non-cooperation is more profitable
• when there is dominance and some cooperators
  receive less

101
Consistent with the rat, lion, chimpanzee and
dolphin data.
102
Emotions linked to coordinated and ritualized
behaviors ceremonial behaviors in groups
evoking excitement, affiliation, and
power (McNeill, 1995)Praying
mms//vod4.haifa.ac.il/L/CRI/RM200604/choir.wmv
Sports mms//vod4.haifa.ac.il/L/CRI/RM200604/nfl.
wmv
And also in humans.
103

• 2) There are two kinds of decision-making
• A) Explicit rational thinking based on analysis
  of economic data and estimations of likelihood
• B) Implicit (unconscious?) decision making based
  on influence of cues and contexts associated with
  cooperation
• from explicit social interaction
• from information about the involvement of others
  (Rilling et al, Baker Rachlin)

Analogous to Tversky and Kahnemans System I
(intuitive) and System II (deliberate, conscious,
rational) decision-making based on explicit
outcomes
104
Ex of implicit decision-making Humans in
Dictator game- Effects of eye-spots (being
observed)
10 keep or dividesingle trial Silent
(ear-covers) or no Eye spots or no
Mean allocationNo eyes non-silent mean
2.45 55 allocated Eyes non-silent
mean 3.79 88 allocated Eyes only mean
3.14 79 allocated (Haley Fessler, 2005,
Evolution and Human Behavior)
105
Can we call this a behavioral economics? the
influence of psychological dimensions associated
with cooperating on the likelihood of
cooperation? biases, etc? From work of Frans de
Waal, Schuster and others on animals Do
individuals prefer cooperation when
non-cooperation provides the same or less in
economic outcomes?Yes in rats, lions,
dolphins. Do individuals tend to share more
if another has helped in gaining the outcome?
Yes in capuchin monkeys, chimpanzees. Do
individuals cooperate more with other individuals
who have previously provided a benefit
(score-keeping)?Yes in a variety of animals
from vampire bats to chimpanzees Do individuals
react to unfairness by monopolizing outcomes
without sharing?Yes again in capuchin monkeys
and chimpanzees
106

• Ultimate explanation if you are a Biologist

• But bias invites the questionHow can
  cooperation evolve if it leads to uneconomic,
  seemingly irrational behavior that is costly in
  time and energy?
• Needed is an evolutionary economics that is
  adaptive and rational for a mindless ultimate
  process of natural selection even though the
  behaving individual may be incapable of knowing
  or intending the ultimate purpose of his action
  that influenced evolution

107
A short-acting immediate incentive can be
adaptive if it also leads to long-term adaptive
outcomes that elevate fitness
Act of cooperation
Immediate economic outcomes
Immediate intrinsic outcomes
Immediate economic outcomes
108
This glue or bridge between the short- and
long-term outcomes can be the development of
social bonds between individuals that engage in
joint actions for shared outcomes and these
bonds are what lead eventually to the beneficial
consequences that guide the evolution
Two filmed examples..lion and chimpanzee
109

• Lions Lion cooperative hunting by males and/or
  females may not pay in food gained per
  individual..
• But cooperative hunting can eventually pay off
  when ?- adult females cooperatively defend
  their cubs against -infanticidal males (Packer,
  Scheel Pusey, 1990), -members of other prides
  (McComb, Packer Pusey, 1994) -hyenas (films).
• -adult males cooperatively fight together when
  taking over a pride of females
• mms//vod4.haifa.ac.il/L/CRI/RM200604/lions2.wmv

110

• Chimpanzees Food and sex do not seem to be the
  primary motivations for cooperative hunting
  (Mitani Watts, 2001) Male-male associations
  and social behaviors were the best predictors
• But cooperative hunting eventually can pay
  offwhen adult males later engage in inter-group
  warfare with direct consequences for mating
  success and fitness (Watts Mitani,
  2001).mms//vod4.haifa.ac.il/L/CRI/RM200604/chim
  p warfare Gombe.wmv

111

• Summary The links between behaviors and
  outcomes and between proximate and ultimate
  explanations - are complex
• (1) the immediate consequencess for cooperating
  include intrinsic social emotions and social
  relationships that are problematical for
  explanations anchored in rationality
• (2) fitness is linked to material consequences
  that may be realized only in the long-term and
  they are also far from certain
• (3) the fitness-enhancing consequences can also
  be indirect, occurring in a different context
  and in a different currency from the original
  cooperation.
• (Schuster, 2002 Schuster Perelberg, 2004
  Perelberg Schuster, in prep)

Do we want the term rational to apply when only
long-term adaptive outcomes are maximized?
112
Implications for game theory matrices combining
economic gains (E) with intrinsic
(psychological) reinforcement (P)e.g., a
Prisoners dilemma game
113
Take-home messages

• In nature
• Cooperation is a class of behaviors incorporating
  irreducible social dimensions that are absent
  when engaging in non-cooperation

Laboratory Models Models should have external
validity, incorporating tasks influenced by the
irreducible social dimensions of cooperation.
Immediate Outcomes Cooperation is not always
economic at the time of cooperating -explicit
material reinforcement -intrinsic reinforcement
from emotions linked to behaving cooperatively
Implications for evolutionary explanations Fitnes
s is determined by beneficial (economic)
outcomes that may only be realized long after the
cooperative behavior is performed. The bridge
between the outcomes is the development of social
relationships among cooperating individuals that
experience the intrinsic reinforcements
associated with engaging in joint actions for
shared outcomes.
Explanation Only by incorporating proximate
psychological processes with ultimate
evolutionary processes can we explain both the
behavior and its link to fitness
114

• Anatol Rapoport (BBS, 2003) quoting David Hume
  (1739)
• Reason is, and ought to be, the slave of
  passions, and can never pretend to any other
  office than to serve and obey them.
• Rapoport adds
• In other words, effective means to reach
  specific goals can be prescribed, but not the
  goals.
• Herbert Simon Rationality in Human Affairs The
  neocortex is the hired gun of the paleocortex.

115
Recent summaries

• R. Schuster B.D. Berger, An animal model for
  studying the behavior of cooperating. In M.
  Anderson (ed.), Tasks and Techniques A Sampling
  of Methodologies for the Investigation of Animal
  Learning, Behavior and Cognition. In press.
• R. Schuster A. PerelbergBehavioural Processes,
  2004, 66, 261-277
• R. Schuster
• Human Nature, 2002, 13, 47-83
• R. Schuster
• Proceedings of 6th Annual Symposium on the
  Science of Behavior Social Behavior. University
  of Guadalajara, Mexico
• Revista mexicana de análisis de la conducta
  (Mexican Journal of Behavior Analysis), 2001,
  27, 165-200)

116
Collaborators
Barry D. Berger, University of Haifa Peter R.
Killeen, Arizona State University Howard
Rachlin, State University of NY at Stony
Brook Heidi Swanson, Netherlands Institute for
Brain Research Shlomo Hareli, University of Haifa
Moussa B.M. Youdim, Technion- Israel
Institute of Technology, Faculty of
Medicine, Director of the Eve Topf and USA
National Parkinson Foundation Centers of
Excellence for Neuro- degenerative
Diseases Gadi Katzir, University of Haifa,
Biology Dietmar Tödt, Free University of Berlin,
Zoology Shai Shoham
Students
Sonia del Canho Tamar Borovitch Keren
Gavish Yona Rubin
Michael Tsoory Steve Arnautof Corina Dollingher
Amir Perlberg Edna Cohen Shai Sela