An

fMRI

Investigation

of

Emotional

Engagement

in

Moral

Judgment

Joshua

D.

Greene

et

al.

Science

-LRB-

-RRB-

DOI

science

The

following

resources

related

to

this

article

are

available

online

at

-LRB-

this

information

is

current

as

of

May

-RRB-

Updated

information

and

services

including

high-resolution

figures

can

be

found

in

the

online

version

of

this

article

at

Supporting

Online

Material

can

be

found

at

This

article

cites

articles

of

which

can

be

accessed

for

free

This

article

has

been

cited

by

article

-LRB-

s

-RRB-

on

the

ISI

Web

of

Science

This

article

has

been

cited

by

articles

hosted

by

HighWire

Press

see

This

article

appears

in

the

following

subject

collections

Psychology

Information

about

obtaining

reprints

of

this

article

or

about

obtaining

permission

to

reproduce

this

article

in

whole

or

in

part

can

be

found

at

Science

-LRB-

print

ISSN

online

ISSN

-RRB-

is

published

weekly

except

the

last

week

in

December

by

the

American

Association

for

the

Advancement

of

Science

New

York

Avenue

NW

Washington

DC

Copyright

by

the

American

Association

for

the

Advancement

of

Science

all

rights

reserved

The

title

Science

is

a

registered

trademark

of

AAAS

Downloaded

from

on

May

REPORTS

matic

assays

using

a

wide

range

of

in

vitro

conditions

Furthermore

once

the

proteins

are

prepared

proteome

screening

is

significantly

faster

and

cheaper

Using

similar

procedures

it

is

clearly

possible

to

prepare

protein

arrays

of

to

proteins

for

global

proteome

analysis

in

humans

and

other

eukaryotes

S.

Fields

Y.

Kohara

D.

J.

Lockhart

Proc

Natl.

Acad

Sci

U.S.A.

-LRB-

-RRB-

A.

Goffeau

et

al.

Science

-LRB-

-RRB-

P.

Ross-Macdonald

et

al.

Nature

-LRB-

-RRB-

J.

L.

DeRisi

V.

R.

Iyer

P.

O.

Brown

Science

-LRB-

-RRB-

E.

A.

Winzeler

et

al.

Science

-LRB-

-RRB-

P.

Uetz

et

al.

Nature

-LRB-

-RRB-

T.

Ito

et

al.

Proc

Natl.

Acad

Sci

U.S.A.

-LRB-

-RRB-

H.

Zhu

M.

Snyder

Curr

Opin

Chem

Biol

-LRB-

-RRB-

M.

R.

Martzen

et

al.

Science

-LRB-

-RRB-

H.

Zhu

et

al.

Nature

Genet

-LRB-

-RRB-

G.

MacBeath

S.

L.

Schreiber

Science

-LRB-

-RRB-

A.

Caveman

J.

Cell

Sci

-LRB-

-RRB-

P.

Arenkov

et

al.

Anal

Biochem

-LRB-

-RRB-

D.

A.

Mitchell

T.

K.

Marshall

R.

J.

Deschenes

Yeast

-LRB-

-RRB-

The

expression

vector

pEGH

was

created

by

inserting

an

epitope

tag

between

the

GST

gene

and

the

polycloning

site

of

pEG

-LRB-

KG

-RRB-

The

yeast

ORFs

were

cloned

using

the

strategy

described

previously

-LRB-

-RRB-

except

every

step

was

done

in

a

format

Plasmid

DNAs

confirmed

by

DNA

sequencing

were

reintroduced

into

both

yeast

-LRB-

-RRB-

and

E.

coli

-LRB-

-RRB-

The

library

contains

unique

ORFs

For

details

of

format

protein

purification

protocol

a

full

list

of

results

from

all

the

experiments

and

the

design

of

the

positive

identification

algorithms

please

visit

our

public

Web

site

-LRB-

http

bioinfo.mbb.yale.edu/proteinchip

-RRB-

and

supplementary

material

at

Science

Online

-LRB-

cgi/content/full

-RRB-

Biotinylated

calmodulin

-LRB-

CalBiochem

USA

-RRB-

was

added

to

the

proteome

chip

at

g

l

in

phosphatebuffered

saline

-LRB-

PBS

-RRB-

with

mM

calcium

and

incubated

in

a

humidity

chamber

for

hour

at

room

temperature

Calcium

-LRB-

mM

-RRB-

was

present

in

buffers

in

all

subsequent

steps

The

chip

was

washed

three

times

with

PBS

at

room

temperature

-LRB-

RT

C

-RRB-

streptavidin

-LRB-

Jackson

IR

USA

-RRB-

-LRB-

dilution

-RRB-

was

added

to

the

chip

and

incubated

for

min

at

RT.

After

extensive

washing

the

chip

was

spun

dry

and

scanned

using

a

microarray

scanner

the

data

was

subsequently

acquired

with

the

GenePix

array

densitometry

software

-LRB-

Axon

USA

-RRB-

S.

S.

Hook

A.

R.

Means

Annu

Rev.

Pharmacol

Toxicol

-LRB-

-RRB-

M.

S.

Cyert

R.

Kunisawa

D.

Kaim

J.

Thorner

Proc

Natl.

Acad

Sci

U.S.A.

-LRB-

-RRB-

D.

A.

Stirling

K.

A.

Welch

M.

J.

Stark

EMBO

J.

-LRB-

-RRB-

F.

Bohl

C.

Kruse

A.

Frank

D.

Ferring

R.

P.

Jansen

EMBO

J.

-LRB-

-RRB-

E.

Bertrand

et

al.

Mol

Cell

-LRB-

-RRB-

D.

C.

Winter

E.

Y.

Choe

R.

Li

Proc

Natl.

Acad

Sci

U.S.A.

-LRB-

-RRB-

C.

Schaerer-Brodbeck

H.

Riezman

Mol

Biol

Cell

-LRB-

-RRB-

K.

Homma

J.

Saito

R.

Ikebe

M.

Ikebe

J.

Biol

Chem

-LRB-

-RRB-

J.

Menendez

J.

Delgado

C.

Gancedo

Yeast

-LRB-

-RRB-

G.

Odorizzi

M.

Babst

S.

D.

Emr

Trends

Biochem

Sci

-LRB-

-RRB-

D.

A.

Fruman

et

al.

Annu

Rev.

Biochem

-LRB-

-RRB-

T.

F.

Martin

Annu

Rev.

Cell

Dev

Biol

-LRB-

-RRB-

S.

Wera

J.

C.

T.

Bergsma

FEMS

Yeast

Res

-LRB-

-RRB-

Liposomes

were

prepared

using

standard

methods

-LRB-

-RRB-

Briefly

appropriate

amounts

of

each

lipid

in

chloroform

were

mixed

and

dried

under

nitrogen

The

lipid

mixture

was

resuspended

in

TBS

buffer

by

vortexing

The

liposomes

were

created

by

sonication

To

probe

the

proteome

chips

l

of

the

different

liposomes

were

added

onto

different

chips

The

chips

were

incubated

in

a

humidity

chamber

for

hour

at

RT.

After

washing

with

TBS

buffer

for

three

times

streptavidin

-LRB-

dilution

-RRB-

was

added

to

the

chip

and

incubated

for

min

at

RT.

Positives

were

identified

using

a

combination

of

the

GenePix

software

which

computes

a

local

intensity

background

for

each

spot

and

a

series

of

algorithms

we

developed

Details

can

be

found

at

and

at

M.

C.

Costanzo

et

al.

Nucleic

Acids

Res

-LRB-

-RRB-

M.

Gerstein

Proteins

-LRB-

-RRB-

K.

Ansari

et

al.

J.

Biol

Chem

-LRB-

-RRB-

Y.

Barral

M.

Parra

S.

Bidlingmaier

M.

Snyder

Genes

Dev

-LRB-

-RRB-

Y.

Li

T.

Kane

C.

Tipper

P.

Spatrick

D.

D.

Jenness

Mol

Cell

Biol

-LRB-

-RRB-

I.

Arnold

et

al.

J.

Biol

Chem

-LRB-

-RRB-

S.

Chu

et

al.

Science

-LRB-

-RRB-

A.

Casamayor

et

al.

Curr

Biol

-LRB-

-RRB-

R.

Guerra

M.

L.

Bianconi

Biosci

Rep.

-LRB-

-RRB-

M.

Pardo

et

al.

Yeast

-LRB-

-RRB-

Single-letter

abbreviations

for

the

amino

acid

residues

are

as

follows

A

Ala

C

Cys

D

Asp

E

Glu

F

Phe

G

Gly

H

His

I

Ile

K

Lys

L

Leu

M

Met

N

Asn

P

Pro

Q

Gln

R

Arg

S

Ser

T

Thr

V

Val

W

Trp

and

Y

Tyr

X

indicates

any

residue

We

thank

K.

Nelson

and

S.

Dellaporta

for

providing

invaluable

help

We

also

thank

A.

Kumar

G.

Michaud

and

C.

Costigan

for

providing

comments

on

the

manuscript

This

research

is

supported

by

grants

from

NIH

H.Z.

A.C.

and

R.J.

were

supported

by

postdoctoral

fellowships

from

the

Damon

Runyon

Walter

Winchell

Foundation

the

Spanish

Ministerio

de

Ciencia

y

Tecnologia

and

by

an

IBM

Graduate

Research

Fellowship

respectively

May

accepted

July

Published

online

July

science

Include

this

information

when

citing

this

paper

References

and

Notes

An

fMRI

Investigation

of

Emotional

Engagement

in

Moral

Judgment

Joshua

D.

Greene

R.

Brian

Sommerville

Leigh

E.

Nystrom

John

M.

Darley

Jonathan

D.

The

long-standing

rationalist

tradition

in

moral

psychology

emphasizes

the

role

of

reason

in

moral

judgment

A

more

recent

trend

places

increased

emphasis

on

emotion

Although

both

reason

and

emotion

are

likely

to

play

important

roles

in

moral

judgment

relatively

little

is

known

about

their

neural

correlates

the

nature

of

their

interaction

and

the

factors

that

modulate

their

respective

behavioral

influences

in

the

context

of

moral

judgment

In

two

functional

magnetic

resonance

imaging

-LRB-

fMRI

-RRB-

studies

using

moral

dilemmas

as

probes

we

apply

the

methods

of

cognitive

neuroscience

to

the

study

of

moral

judgment

We

argue

that

moral

dilemmas

vary

systematically

in

the

extent

to

which

they

engage

emotional

processing

and

that

these

variations

in

emotional

engagement

influence

moral

judgment

These

results

may

shed

light

on

some

puzzling

patterns

in

moral

judgment

observed

by

contemporary

philosophers

The

present

study

was

inspired

by

a

family

of

ethical

dilemmas

familiar

to

contemporary

moral

philosophers

-LRB-

-RRB-

One

such

dilemma

is

the

trolley

dilemma

A

runaway

trolley

is

headed

for

five

people

who

will

be

killed

if

it

proceeds

on

its

present

course

The

only

way

to

save

them

is

to

hit

a

switch

that

will

turn

the

trolley

onto

an

alternate

set

of

tracks

where

it

will

kill

one

person

instead

of

five

Ought

you

to

turn

the

trolley

in

order

to

save

five

people

at

the

expense

of

one

Most

people

say

yes

Now

consider

a

similar

problem

the

footbridge

dilemma

As

before

a

trolley

threatens

to

kill

five

people

You

are

Center

for

the

Study

of

Brain

Mind

and

Behavior

Department

of

Philosophy

Hall

of

Psychology

Green

Hall

Princeton

University

Princeton

NJ

USA

of

Psychiatry

University

of

Pittsburgh

Pittsburgh

PA

USA

To

whom

correspondence

should

be

addressed

Email

jdgreene@princeton.edu

standing

next

to

a

large

stranger

on

a

footbridge

that

spans

the

tracks

in

between

the

oncoming

trolley

and

the

five

people

In

this

scenario

the

only

way

to

save

the

five

people

is

to

push

this

stranger

off

the

bridge

onto

the

tracks

below

He

will

die

if

you

do

this

but

his

body

will

stop

the

trolley

from

reaching

the

others

Ought

you

to

save

the

five

others

by

pushing

this

stranger

to

his

death

Most

people

say

no

Taken

together

these

two

dilemmas

create

a

puzzle

for

moral

philosophers

What

makes

it

morally

acceptable

to

sacrifice

one

life

to

save

five

in

the

trolley

dilemma

but

not

in

the

footbridge

dilemma

Many

answers

have

been

proposed

For

example

one

might

suggest

in

a

Kantian

vein

that

the

difference

between

these

two

cases

lies

in

the

fact

that

in

the

footbridge

dilemma

one

literally

uses

a

fellow

human

being

as

a

means

to

some

independent

end

whereas

in

the

trolley

dilemma

the

unfortunate

person

just

happens

to

SCIENCE

VOL

SEPTEMBER

Downloaded

from

on

May

REPORTS

be

in

the

way

This

answer

however

runs

into

trouble

with

a

variant

of

the

trolley

dilemma

in

which

the

track

leading

to

the

one

person

loops

around

to

connect

with

the

track

leading

to

the

five

people

-LRB-

-RRB-

Here

we

will

suppose

that

without

a

body

on

the

alternate

track

the

trolley

would

if

turned

that

way

make

its

way

to

the

other

track

and

kill

the

five

people

as

well

In

this

variant

as

in

the

footbridge

dilemma

you

would

use

someone

's

body

to

stop

the

trolley

from

killing

the

five

Most

agree

nevertheless

that

it

is

still

appropriate

to

turn

the

trolley

in

this

case

in

spite

of

the

fact

that

here

too

we

have

a

case

of

using

These

are

just

one

proposed

solution

and

one

counterexample

but

together

they

illustrate

the

sort

of

dialectical

difficulties

that

all

proposed

solutions

to

this

problem

have

encountered

If

a

solution

to

this

problem

exists

it

is

not

obvious

That

is

there

is

no

set

of

consistent

readily

accessible

moral

principles

that

captures

people

's

intuitions

concerning

what

behavior

is

or

is

not

appropriate

in

these

and

similar

cases

This

leaves

psychologists

with

a

puzzle

of

their

own

How

is

it

that

nearly

everyone

manages

to

conclude

that

it

is

acceptable

to

sacrifice

one

life

for

five

in

the

trolley

dilemma

but

not

in

the

footbridge

dilemma

in

spite

of

the

fact

that

a

satisfying

justification

for

distinguishing

between

these

two

cases

is

remarkably

difficult

to

find

-LRB-

-RRB-

We

maintain

that

from

a

psychological

point

of

view

the

crucial

difference

between

the

trolley

dilemma

and

the

footbridge

dilemma

lies

in

the

latter

's

tendency

to

engage

people

's

emotions

in

a

way

that

the

former

does

not

The

thought

of

pushing

someone

to

his

death

is

we

propose

more

emotionally

salient

than

the

thought

of

hitting

a

switch

that

will

cause

a

trolley

to

produce

similar

consequences

and

it

is

this

emotional

response

that

accounts

for

people

's

tendency

to

treat

these

cases

differently

This

hypothesis

concerning

these

two

cases

suggests

a

more

general

hypothesis

concerning

moral

judgment

Some

moral

dilemmas

-LRB-

those

relevantly

similar

to

the

footbridge

dilemma

-RRB-

engage

emotional

processing

to

a

greater

extent

than

others

-LRB-

those

relevantly

similar

to

the

trolley

dilemma

-RRB-

and

these

differences

in

emotional

engagement

affect

people

's

judgments

The

present

investigation

is

an

attempt

to

test

this

more

general

hypothesis

Drawing

upon

recent

work

concerning

the

neural

correlates

of

emotion

-LRB-

-RRB-

we

predicted

that

brain

areas

associated

with

emotion

would

be

more

active

during

contemplation

of

dilemmas

such

as

the

footbridge

dilemma

as

compared

to

during

contemplation

of

dilemmas

such

as

the

trolley

dilemma

In

addition

we

predicted

a

pattern

of

behavioral

interference

similar

to

that

observed

in

cognitive

tasks

in

which

automatic

processes

can

influence

responses

such

as

the

Stroop

task

-LRB-

in

which

the

identity

of

a

color

word

can

interfere

with

participants

ability

to

name

the

color

in

which

it

is

displayed

e.g.

the

ability

to

say

green

in

response

to

the

word

red

written

in

green

ink

-RRB-

-LRB-

-RRB-

In

light

of

our

proposal

that

people

tend

to

have

a

salient

automatic

emotional

response

to

the

footbridge

dilemma

that

leads

them

to

judge

the

action

it

proposes

to

be

inappropriate

we

would

expect

those

-LRB-

relatively

rare

-RRB-

individuals

who

nevertheless

judge

this

action

to

be

appropriate

to

do

so

against

a

countervailing

emotional

response

and

to

exhibit

longer

reaction

times

as

a

result

of

this

emotional

interference

More

generally

we

predicted

longer

reaction

times

for

trials

in

which

the

participant

's

response

is

incongruent

with

the

emotional

response

-LRB-

e.g.

saying

appropriate

to

a

dilemma

such

as

the

footbridge

dilemma

-RRB-

We

predicted

the

absence

of

such

effects

for

dilemmas

such

as

the

trolley

dilemma

which

according

to

our

theory

are

less

likely

to

elicit

a

strong

emotional

response

In

each

of

two

studies

Experiments

and

we

used

a

battery

of

practical

dilemmas

-LRB-

-RRB-

These

dilemmas

were

divided

into

moral

and

non-moral

categories

on

the

basis

of

the

responses

of

pilot

participants

-LRB-

-RRB-

-LRB-

Typical

examples

of

non-moral

dilemmas

posed

questions

about

whether

to

travel

by

bus

or

by

train

given

certain

time

constraints

and

about

which

of

two

coupons

to

use

at

a

store

-RRB-

Two

independent

coders

evaluated

each

moral

dilemma

using

three

criteria

designed

to

capture

the

difference

between

the

intuitively

up

close

and

personal

-LRB-

and

putatively

more

emotional

-RRB-

sort

of

violation

exhibited

by

the

footbridge

dilemma

and

the

more

intuitively

impersonal

-LRB-

and

putatively

less

emotional

-RRB-

violation

exhibited

by

the

trolley

dilemma

-LRB-

-RRB-

Moral

dilemmas

meeting

these

criteria

were

assigned

to

the

moralpersonal

condition

the

others

to

the

moralimpersonal

condition

Typical

moral-personal

dilemmas

included

a

version

of

the

footbridge

dilemma

a

case

of

stealing

one

person

's

organs

in

order

to

distribute

them

to

five

others

and

a

case

of

throwing

people

off

a

sinking

lifeboat

Typical

moral-impersonal

dilemmas

included

a

version

of

the

trolley

dilemma

a

case

of

keeping

money

found

in

a

lost

wallet

and

a

case

of

voting

for

a

policy

expected

to

cause

more

deaths

than

its

alternatives

Participants

responded

to

each

dilemma

by

indicating

whether

they

judged

the

action

it

proposes

to

be

appropriate

or

inappropriate

In

each

experiment

nine

participants

-LRB-

-RRB-

responded

to

each

of

dilemmas

-LRB-

-RRB-

while

undergoing

brain

scanning

using

f

MRI

-LRB-

-RRB-

Figures

and

describe

brain

areas

identified

in

Experiment

by

a

thresholded

omnibus

analysis

of

variance

-LRB-

ANOVA

-RRB-

performed

on

the

functional

images

-LRB-

-RRB-

In

each

case

the

Fig.

Effect

of

condition

on

activity

in

brain

areas

identified

in

Experiment

R

right

L

left

B

bilateral

Results

for

the

middle

frontal

gyrus

were

not

replicated

in

Experiment

The

moral-personal

condition

was

significantly

different

from

the

other

two

conditions

in

all

other

areas

in

both

Experiments

and

In

Experiment

the

medial

frontal

and

posterior

cingulate

gyri

showed

significant

differences

between

the

moral-impersonal

and

non-moral

conditions

In

Experiment

only

the

posterior

cingulate

gyrus

was

significantly

different

in

this

comparison

Brodmann

's

Areas

and

Talairach

-LRB-

-RRB-

coordinates

-LRB-

x

y

z

-RRB-

for

each

area

are

as

follows

-LRB-

left

to

right

in

graph

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Fig.

Brain

areas

exhibiting

differences

in

activity

between

conditions

shown

in

three

axial

slices

of

a

standard

brain

-LRB-

-RRB-

Slice

location

is

indicated

by

Talairach

-LRB-

-RRB-

z

coordinate

Data

are

for

the

main

effect

of

condition

in

Experiment

Colored

areas

reflect

the

thresholded

F

scores

Images

are

reversed

left

to

right

to

follow

radiologic

convention

SEPTEMBER

VOL

SCIENCE

Downloaded

from

on

May

REPORTS

ANOVA

identified

all

brain

areas

differing

in

activity

among

the

moral-personal

moral-impersonal

and

non-moral

conditions

Planned

comparisons

on

these

areas

revealed

that

medial

portions

of

Brodmann

's

Areas

-LRB-

BA

-RRB-

and

-LRB-

medial

frontal

gyrus

-RRB-

BA

-LRB-

posterior

cingulate

gyrus

-RRB-

and

BA

-LRB-

angular

gyrus

bilateral

-RRB-

were

significantly

more

active

in

the

moralpersonal

condition

than

in

the

moral-impersonal

and

the

non-moral

conditions

Recent

functional

imaging

studies

have

associated

each

of

these

areas

with

emotion

-LRB-

-RRB-

Areas

associated

with

working

memory

have

been

found

to

become

less

active

during

emotional

processing

as

compared

to

periods

of

cognitive

processing

-LRB-

-RRB-

BA

-LRB-

middle

frontal

gyrus

right

-RRB-

and

BA

-LRB-

parietal

lobe

bilateral

-RRB-

both

associated

with

working

memory

-LRB-

-RRB-

were

significantly

less

active

in

the

moral-personal

condition

than

in

the

other

two

conditions

In

BA

-LRB-

bilateral

-RRB-

BA

and

BA

-LRB-

bilateral

-RRB-

there

was

no

significant

difference

between

the

moral-impersonal

and

the

non-moral

condition

-LRB-

-RRB-

Experiment

served

to

replicate

the

results

of

Experiment

-LRB-

-RRB-

and

to

provide

behavioral

data

concerning

participants

judgments

and

reaction

times

Planned

comparisons

on

the

seven

brain

areas

identified

in

Experiment

yielded

results

nearly

identical

to

those

of

Experiment

with

the

following

differences

In

Experiment

there

was

no

difference

in

BA

between

the

moral-impersonal

and

non-moral

conditions

and

no

differences

were

found

for

BA

-LRB-

-RRB-

Reaction

time

data

from

Experiment

are

described

by

Fig.

Our

theory

concerning

emotional

interference

predicted

longer

reaction

times

for

emotionally

incongruent

responses

which

occur

when

a

participant

responds

appropriate

in

the

moral-personal

condition

-LRB-

e.g.

judging

it

appropriate

to

push

the

man

off

the

footbridge

in

the

footbridge

dilemma

-RRB-

but

which

do

not

occur

in

the

moral-impersonal

and

non-moral

conditions

As

predicted

responses

of

appropriate

-LRB-

emotionally

incongruent

-RRB-

were

significantly

slower

than

responses

of

inappropriate

-LRB-

emotionally

congruent

-RRB-

within

the

moral-personal

condition

and

there

was

no

significant

difference

in

reaction

time

between

responses

of

appropriate

and

inappropriate

in

the

other

two

conditions

In

fact

the

data

exhibit

a

trend

in

the

opposite

direction

for

the

other

two

conditions

-LRB-

-RRB-

with

responses

of

inappropriate

taking

slightly

longer

than

responses

of

appropriate

In

each

of

the

brain

areas

identified

in

both

Experiments

and

the

moral-personal

condition

had

an

effect

significantly

different

from

both

the

moral-impersonal

and

the

non-moral

conditions

All

three

areas

showing

increased

relative

activation

in

the

moral-personal

condition

have

been

implicated

in

emotional

processing

The

behavioral

data

provide

further

evidence

for

the

increased

emotional

engagement

in

moral-personal

condition

by

revealing

a

reaction

time

pattern

that

is

unique

to

that

condition

and

that

was

predicted

by

our

hypothesis

concerning

emotional

interference

Moreover

the

presence

of

this

interference

effect

in

the

behavioral

data

strongly

suggests

that

the

increased

emotional

responses

generated

by

the

moral-personal

dilemmas

have

an

influence

on

and

are

not

merely

incidental

to

moral

judgment

-LRB-

-RRB-

These

data

also

suggest

that

in

terms

of

the

psychological

processes

associated

with

their

production

judgments

concerning

impersonal

moral

dilemmas

more

closely

resemble

judgments

concerning

non-moral

dilemmas

than

they

do

judgments

concerning

personal

moral

dilemmas

The

trolley

and

footbridge

dilemmas

emerged

as

pieces

of

a

puzzle

for

moral

philosophers

Why

is

it

acceptable

to

sacrifice

one

person

to

save

five

others

in

the

trolley

dilemma

but

not

in

the

footbridge

dilemma

Here

we

consider

these

dilemmas

as

pieces

of

a

psychological

puzzle

How

do

people

manage

to

conclude

that

it

is

acceptable

to

sacrifice

one

for

the

sake

of

five

in

one

case

but

not

in

the

other

We

maintain

that

emotional

response

is

likely

to

be

the

crucial

difference

between

these

two

cases

But

this

is

an

answer

to

the

psychological

puzzle

not

the

philosophical

one

Our

conclusion

therefore

is

descriptive

rather

than

prescriptive

We

do

not

claim

to

have

shown

any

actions

or

judgments

to

be

morally

right

or

wrong

Nor

have

we

argued

that

emotional

response

is

the

sole

determinant

of

judgments

concerning

moral

dilemmas

of

the

kind

discussed

in

this

study

On

the

contrary

the

behavioral

influence

of

these

emotional

responses

is

most

strongly

suggested

in

the

performance

of

those

participants

who

judge

in

spite

of

their

emotions

What

has

been

demonstrated

is

that

there

are

systematic

variations

in

the

engagement

of

emotion

in

moral

judgment

The

systematic

nature

of

these

variations

is

manifest

in

an

observed

correlation

between

-LRB-

i

-RRB-

certain

features

that

differ

between

the

trolley

dilemma

and

the

footbridge

dilemma

and

-LRB-

ii

-RRB-

patterns

of

neural

activity

in

emotion-related

brain

areas

as

well

as

patterns

in

reaction

time

Methodological

constraints

led

us

to

characterize

these

certain

features

by

means

of

a

highly

regimented

distinction

between

actions

that

are

personal

and

impersonal

-LRB-

-RRB-

This

personal-impersonal

distinction

has

proven

useful

in

generating

the

present

results

but

it

is

by

no

means

definitive

We

view

this

distinction

as

a

useful

first

cut

an

important

but

preliminary

step

toward

identifying

the

psychologically

essential

features

of

circumstances

that

engage

-LRB-

or

fail

to

engage

-RRB-

our

emotions

and

that

ultimately

shape

our

moral

judgments

judgments

concerning

hypothetical

examples

such

as

the

trolley

and

footbridge

dilemmas

but

also

concerning

the

more

complicated

moral

dilemmas

we

face

in

our

public

and

private

lives

A

distinction

such

as

this

may

allow

us

to

steer

a

middle

course

between

the

traditional

rationalism

and

more

recent

emotivism

that

have

dominated

moral

psychology

-LRB-

-RRB-

The

present

results

raise

but

do

not

answer

a

more

general

question

concerning

the

relation

between

the

aforementioned

philosophical

and

psychological

puzzles

How

will

a

better

understanding

of

the

mechanisms

that

give

rise

to

our

moral

judgments

alter

our

attitudes

toward

the

moral

judgments

we

make

J.

J.

Thomson

Rights

Restitution

and

Risk

-LRB-

Harvard

Univ.

Press

Cambridge

-RRB-

pp.

A

loose

but

potentially

illuminating

analogy

can

be

made

between

this

and

the

Chomskyan

question

How

is

that

most

people

can

speak

grammatically

without

being

able

to

exhaustively

cite

the

rules

of

grammar

A.

R.

Damasio

Descartes

Error

-LRB-

Putnam

New

York

-RRB-

R.

J.

Davidson

W.

Irwin

Trends

Cogn

Sci

-LRB-

-RRB-

E.

M.

Reiman

J.

Clin

Psychiatry

-LRB-

suppl

-RRB-

-LRB-

-RRB-

J.

R.

Stroop

J.

Exp

Psychol

-LRB-

-RRB-

C.

M.

MacLeod

Psychol

Bull

-LRB-

-RRB-

Testing

materials

-LRB-

dilemmas

-RRB-

are

available

from

Science

Online

at

The

three

criteria

are

as

follows

First

coders

indicated

for

each

dilemma

whether

the

action

in

question

could

reasonably

be

expected

to

lead

to

serious

bodily

harm

Second

they

were

asked

to

indicate

whether

this

harm

would

be

the

result

of

deflecting

an

existing

threat

onto

a

different

party

Our

use

of

this

criterion

which

parallels

a

distinction

made

by

Thomson

-LRB-

-RRB-

is

a

an

attempt

to

operationalize

an

intuitive

notion

of

agency

Intuitively

when

a

harm

is

produced

by

means

of

deflecting

an

existing

threat

the

agent

has

merely

edited

and

not

authored

the

resulting

harm

and

thus

its

contemplation

is

less

emotionally

engaging

Lastly

coders

were

asked

to

indicate

whether

the

resulting

harm

would

befall

a

particular

person

or

a

member

or

members

of

a

particular

group

of

people

Here

the

question

in

intuitive

terms

is

whether

the

victim

is

on

stage

in

the

dilemma

The

moral

dilemmas

of

which

the

coders

said

that

the

action

in

question

-LRB-

a

-RRB-

could

reasonably

be

expected

to

lead

to

serious

bodily

harm

-LRB-

b

-RRB-

to

a

particular

person

or

a

member

or

members

of

a

particular

group

of

people

-LRB-

c

-RRB-

where

this

harm

is

not

the

result

of

deflecting

an

existing

threat

onto

a

different

party

were

assigned

to

the

moral

References

and

Notes

Fig.

Mean

reaction

time

by

condition

and

response

type

in

Experiment

A

mixed-effects

ANOVA

revealed

a

significant

interaction

between

condition

and

response

type

-LSB-

F

-LRB-

-RRB-

P

-RRB-

Reaction

times

differed

significantly

between

responses

of

appropriate

and

inappropriate

in

the

moral-personal

condition

-LSB-

t

-LRB-

-RRB-

P

-RSB-

but

not

in

the

other

conditions

-LRB-

P

-RRB-

Error

bars

indicate

two

standard

errors

of

the

mean

SCIENCE

VOL

SEPTEMBER

Downloaded

from

on

May

REPORTS

personal

condition

the

others

were

assigned

to

the

moral-impersonal

condition

Participants

were

five

male

and

four

female

undergraduates

in

Experiment

four

male

and

five

female

in

Experiment

All

participants

provided

written

informed

consent

Dilemmas

were

presented

in

random

order

in

a

series

of

six

blocks

of

ten

trials

each

in

Experiment

twelve

blocks

of

five

trials

each

in

Experiment

Participants

responses

to

versions

of

the

trolley

and

footbridge

dilemmas

were

consistent

with

the

intuitions

described

above

-LRB-

-RRB-

Stimuli

-LRB-

dilemmas

-RRB-

were

presented

on

a

visual

display

projected

into

the

scanner

Each

dilemma

was

presented

as

text

through

a

series

of

three

screens

the

first

two

describing

a

scenario

and

the

last

posing

a

question

about

the

appropriateness

of

an

action

one

might

perform

in

that

scenario

-LRB-

e.g.

turning

the

trolley

-RRB-

Participants

were

allowed

to

read

at

their

own

pace

pressing

a

button

to

advance

from

the

first

to

the

second

screen

and

from

the

second

to

the

third

screen

After

reading

the

third

screen

participants

responded

by

pressing

one

of

two

buttons

-LRB-

appropriate

or

inappropriate

-RRB-

Participants

were

given

a

maximum

of

s

to

read

all

three

screens

and

respond

The

intertrial

interval

-LRB-

ITI

-RRB-

lasted

for

a

minimum

of

s

-LRB-

seven

images

-RRB-

in

each

trial

allowing

the

hemodynamic

response

to

return

to

baseline

after

each

trial

Baseline

activity

was

defined

as

the

mean

signal

across

the

last

four

images

of

the

ITI

Task-related

activity

was

measured

using

a

floating

window

of

eight

images

surrounding

-LRB-

four

before

one

during

and

three

after

-RRB-

the

point

of

response

-LRB-

This

window

includes

three

post-response

images

in

order

to

allow

for

the

to

delay

in

hemodynamic

response

to

neural

activation

-RRB-

This

floating

window

technique

combined

the

benefits

of

an

eventrelated

design

with

the

flexibility

required

to

image

a

complex

and

temporally

extended

psychological

process

that

inevitably

proceeds

at

its

own

pace

In

Experiment

functional

images

were

acquired

in

axial

slices

parallel

to

the

AC-PC

-LRB-

anterior

commisure

posterior

commisure

-RRB-

line

-LSB-

spiral

pulse

sequence

repetition

time

-LRB-

TR

-RRB-

ms

echo

time

-LRB-

TE

-RRB-

ms

flip

angle

field

of

view

-LRB-

FOV

-RRB-

mm

isotropic

voxels

-RSB-

using

a

GE

Signa

whole-body

scanner

In

Experiment

functional

images

were

acquired

in

axial

slices

parallel

to

the

AC-PC

line

-LRB-

echoplanar

pulse

sequence

TR

ms

TE

ms

flip

angle

FOV

mm

isotropic

voxels

interslice

spacing

-RRB-

using

a

Siemens

Allegra

head-dedicated

scanner

Before

statistical

analysis

images

for

all

participants

were

coregistered

using

a

automatic

algorithm

Images

were

smoothed

with

an

fullwidth

at

half

maximum

-LRB-

FWHM

-RRB-

Gaussian

filter

In

Experiment

the

images

contained

in

each

response

window

were

analyzed

with

the

use

of

a

voxelwise

mixed-effects

ANOVA

with

participant

as

a

random

effect

and

dilemma-type

block

and

response-relative

image

as

fixed

effects

Statistical

maps

of

voxelwise

F-ratios

were

thresholded

for

significance

-LRB-

P

-RRB-

and

cluster

size

-LRB-

voxels

-RRB-

In

Experiments

and

planned

comparisons

for

significant

differences

between

conditions

-LRB-

P

cluster

size

voxels

-RRB-

were

made

for

each

area

identified

by

the

thresholded

ANOVA

in

Experiment

R.

J.

Maddock

Trends

Neurosci

-LRB-

-RRB-

S.

M.

Kosslyn

et

al.

Neuroreport

-LRB-

-RRB-

E.

M.

Reiman

et

al.

Am

J.

Psychiatry

-LRB-

-RRB-

W.

C.

Drevets

M.

E.

Raichle

Cognition

Emotion

-LRB-

-RRB-

E.

E.

Smith

J.

Jonides

Cognit

Psychol

-LRB-

-RRB-

J.

D.

Cohen

et

al.

Nature

-LRB-

-RRB-

In

BA

-LRB-

right

-RRB-

a

small

minority

of

voxels

-LRB-

of

-RRB-

showed

a

significant

difference

between

the

moral-impersonal

and

non-moral

conditions

Due

to

magnetic

susceptibility

artifact

we

were

unable

to

image

the

orbitofrontal

cortex

an

area

thought

by

some

to

play

an

important

role

in

moral

judgment

-LRB-

-RRB-

Experiments

and

were

not

identical

-LRB-

-RRB-

Experiment

employed

some

modified

versions

of

dilemmas

from

Experiment

as

well

as

some

new

dilemmas

in

order

to

avoid

a

confound

present

in

the

design

of

the

behavioral

aspect

of

Experiment

-LRB-

-RRB-

The

replicated

results

for

BAs

and

bilateral

were

achieved

at

a

higher

significance

threshold

in

Experiment

-LRB-

P

-RRB-

than

in

Experiment

A

potential

confound

in

the

design

of

the

behavioral

aspect

of

the

present

study

deserves

attention

One

might

suppose

that

participants

respond

more

slowly

when

giving

an

unconventional

response

i.e.

a

response

that

differs

from

that

of

the

majority

One

might

suppose

further

that

the

moralpersonal

condition

makes

greater

use

of

dilemmas

for

which

the

emotionally

incongruent

response

is

also

the

unconventional

response

-LRB-

as

in

judging

that

one

may

push

the

man

off

the

footbridge

in

the

footbridge

dilemma

-RRB-

thus

confounding

emotional

incongruity

with

unconventionality

in

participants

responses

Therefore

an

effect

that

we

attribute

to

emotional

engagement

may

simply

be

an

effect

of

the

conventionality

of

participants

responses

To

deconfound

these

factors

in

Experiment

we

included

additional

moral-personal

dilemmas

for

which

the

conventional

response

was

emotionally

incongruent

rather

than

congruent

For

example

one

dilemma

asked

whether

it

is

appropriate

to

smother

one

's

crying

baby

to

death

in

order

to

prevent

its

crying

from

summoning

enemy

soldiers

who

will

kill

oneself

the

baby

and

a

number

of

others

if

summoned

Most

participants

judged

this

action

to

be

appropriate

in

spite

of

their

putative

emotional

tendencies

to

the

contrary

As

predicted

by

our

hypothesis

reaction

times

in

such

cases

were

significantly

longer

-LSB-

t

-LRB-

-RRB-

P

-RSB-

than

the

reaction

times

for

conventional

and

emotionally

congruent

responses

as

were

typically

made

in

response

to

the

footbridge

dilemma

Thus

after

controlling

for

conventionality

reaction

times

in

the

moral-personal

condition

are

longer

for

trials

which

according

to

our

theory

reflect

a

judgment

that

is

emotionally

incongruent

rather

than

congruent

Although

our

conclusion

concerning

the

behavioral

influence

of

the

observed

emotional

responses

does

not

require

that

the

emotion-related

areas

identified

in

Experiments

and

be

different

from

areas

that

show

increased

activity

in

response

to

more

basic

kinds

of

emotional

stimuli

one

might

wonder

to

what

extent

they

do

differ

from

such

areas

We

made

a

preliminary

attempt

to

answer

this

question

in

the

form

of

an

addendum

study

to

Experiment

Five

participants

responded

to

moral-personal

and

moral-impersonal

dilemmas

as

in

Experiments

and

Participants

also

performed

a

task

in

which

they

named

the

colors

of

visually

presented

emotional

and

neutral

words

a

task

similar

to

the

one

used

by

Isenberg

et

al.

-LRB-

-RRB-

The

emotional

word

stimuli

were

extracted

from

the

text

of

the

moral

dilemmas

by

three

independent

coders

Neutral

words

and

additional

emotional

words

were

drawn

from

materials

used

by

Isenberg

et

al.

-LRB-

-RRB-

A

comparison

of

the

emotional

and

neutral

word

conditions

-LRB-

t

test

P

cluster

size

voxels

-RRB-

revealed

no

significant

activation

in

the

emotion-related

areas

identified

in

Experiment

and

only

a

marginal

activation

-LRB-

out

of

voxels

-RRB-

in

one

of

the

working

memory

areas

-LRB-

left

BA

-RRB-

This

comparison

did

however

reveal

activations

in

numerous

other

areas

A

comparison

of

the

moral-personal

and

moral-impersonal

conditions

from

the

same

five

sessions

replicated

the

activations

observed

in

Experiments

and

in

BA

-LRB-

of

voxels

at

P

-RRB-

and

left

BA

-LRB-

of

voxels

at

P

-RRB-

These

results

demonstrate

at

the

very

least

that

the

effects

observed

in

Experiments

and

in

the

medial

frontal

gyrus

-LRB-

BA

-RRB-

can

not

be

attributed

to

the

mere

reading

of

emotional

words

This

area

more

than

any

of

the

others

we

have

identified

is

likely

to

play

a

role

in

the

integration

of

emotion

and

cognition

in

complex

decision-making

-LRB-

-RRB-

J.

D.

Haidt

Psych

Rev.

in

press

N.

Isenberg

et

al.

Proc

Natl.

Acad

Sci

U.S.A.

-LRB-

-RRB-

J.

Talairach

P.

Tournoux

A

Co-Planar

Stereotaxic

Atlas

of

the

Human

Brain

-LRB-

Thieme

New

York

-RRB-

We

gratefully

acknowledge

M.

Gilzenrat

N.

Isenberg

P.

Jablonka

J.

Kroger

and

T.-Q

Li

for

their

contributions

to

this

project

Supported

in

part

by

grants

from

the

Pew

Charitable

Trusts

-LRB-

no.

-RRB-

and

the

National

Science

Foundation

-LRB-

no.

-RRB-

May

accepted

July

BAFF-R

a

Newly

Identified

TNF

Receptor

That

Specifically

Interacts

with

BAFF

Jeffrey

S.

Thompson

Sarah

A.

Bixler

Fang

Qian

Kalpit

Vora

Martin

L.

Scott

Teresa

G.

Cachero

Catherine

Hession

Pascal

Schneider

Irene

D.

Sizing

Colleen

Mullen

Kathy

Strauch

Mohammad

Zafari

Christopher

D.

Benjamin

Jurg

Tschopp

Jeffrey

L.

Browning

Christine

B

cell

homeostasis

has

been

shown

to

critically

depend

on

BAFF

the

B

cell

activation

factor

from

the

tumor

necrosis

factor

-LRB-

TNF

-RRB-

family

Although

BAFF

is

already

known

to

bind

two

receptors

BCMA

and

TACI

we

have

identified

a

third

receptor

for

BAFF

that

we

have

termed

BAFF-R

BAFF-R

binding

appears

to

be

highly

specific

for

BAFF

suggesting

a

unique

role

for

this

ligand-receptor

interaction

Consistent

with

this

the

BAFF-R

locus

is

disrupted

in

A/WySnJ

mice

which

display

a

B

cell

phenotype

qualitatively

similar

to

that

of

the

BAFF-deficient

mice

Thus

BAFF-R

appears

to

be

the

principal

receptor

for

BAFF-mediated

mature

B

cell

survival

The

TNF

family

ligand

BAFF

also

known

as

THANK

BLyS

and

-LRB-

-RRB-

enhances

B

cell

survival

in

vitro

-LRB-

-RRB-

and

has

recently

emerged

as

a

key

regulator

of

peripheral

B

cell

populations

in

vivo

Mice

overexpressing

BAFF

display

mature

B

cell

hyperplasia

and

symptoms

of

systemic

lupus

erythaematosus

-LRB-

SLE

-RRB-

-LRB-

-RRB-

Likewise

some

SLE

patients

have

significantly

increased

levels

of

BAFF

in

their

SEPTEMBER

VOL

SCIENCE

Downloaded

from

on

May

O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O B O O O O O O O O O O B O O O O O B O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O B O O O O O O O O O O B O O O O B O O O O O B O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O B O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O