Being

Admired

or

Being

Liked

Classroom

Social

Status

and

Depressive

Problems

in

Early

Adolescent

Girls

and

Boys

This

study

investigates

associations

between

depressive

problems

and

classroom

social

status

in

a

large

population

cohort

of

Dutch

early

adolescents

-LRB-

N

age

girls

-RRB-

Depressive

problems

were

assessed

by

parent

and

self-reports

and

classroom

status

by

peer

nominations

We

assessed

peer

status

with

respect

to

both

achievement-related

-LRB-

being

a

good

learner

being

good

at

sports

being

good-looking

-RRB-

and

affection-related

-LRB-

being

liked

being

disliked

being

best

friend

-RRB-

areas

In

boys

depressive

problems

were

most

strongly

associated

with

not

being

good

at

sports

while

in

girls

the

association

was

strongest

for

not

being

liked

The

risk

of

a

low

status

in

one

area

could

largely

be

compensated

by

a

high

status

in

another

area

Depressive

problems

and

low

social

status

have

been

associated

since

ancient

times

In

fact

the

word

depression

is

derived

from

the

Latin

term

deprimere

which

means

being

brought

down

in

status

or

fortune

-LRB-

Jackson

-RRB-

The

association

between

depression

and

social

status

has

been

confirmed

empirically

in

numerous

studies

in

both

animals

and

humans

-LRB-

e.g.

Blanchard

Blanchard

Eaton

Harrison

Gilbert

Allan

Trent

Hecht

Inderbitzen

Bukowski

-RRB-

Social

status

can

be

defined

in

at

least

two

ways

in

terms

of

achievements

-LRB-

being

admired

-RRB-

and

in

terms

of

affection

-LRB-

being

liked

-RRB-

These

two

definitions

are

related

but

by

no

means

interchangeable

and

could

be

differentially

linked

to

depressive

problems

The

former

contains

an

element

of

competition

-LRB-

being

better

than

others

-RRB-

while

the

latter

relates

to

being

accepted

or

rejected

by

the

social

group

We

will

compare

the

relative

importance

of

both

of

them

with

respect

to

depressive

problems

in

early

adolescent

girls

and

boys

Adolescence

is

a

developmental

context

in

which

peer

status

is

particularly

salient

-LRB-

Connell

Dishion

-RRB-

Adolescents

spend

considerably

more

time

in

peer

interactions

than

younger

children

-LRB-

Larson

Richards

-RRB-

and

use

peers

as

primary

sources

for

social

comparison

and

self-appraisal

-LRB-

Prinstein

Aikins

-RRB-

Adolescence

is

also

a

highly

interesting

developmental

stage

for

studying

depressive

problems

because

the

prevalence

of

these

problems

increases

substantially

in

this

phase

of

life

especially

in

girls

-LRB-

e.g.

Hankin

et

al.

Oldehinkel

Wittchen

Schuster

-RRB-

Achievement

and

affection-related

social

status

Achievement-related

operationalizations

of

social

status

are

usually

based

on

Price

's

Social

Ranking

Theory

-LRB-

Gilbert

Allan

Price

-RRB-

Briefly

this

theory

postulates

that

in

pursuing

scarce

resources

-LRB-

e.g.

territories

sexual

opportunities

-RRB-

animals

will

have

to

fight

others

aiming

at

the

same

resources

To

prevent

harm

animals

who

are

likely

to

lose

these

conflicts

need

inhibitory

mechanisms

that

force

them

to

submit

-LRB-

withdraw

run

away

-RRB-

and

downgrade

their

aspirations

so

that

they

do

not

keep

engaging

in

fights

they

can

not

win

These

inhibitory

strategies

may

result

in

behavioral

and

physiological

phenomena

that

closely

resemble

depressive

symptoms

in

humans

-LRB-

e.g.

Blanchard

Blanchard

Sapolsky

-RRB-

Hence

according

to

this

theory

depression

is

an

involuntary

strategy

to

accept

defeat

in

-LRB-

ritual

-RRB-

agonistic

encounters

and

to

accommodate

what

would

otherwise

be

unacceptably

low

social

rank

-LRB-

Price

Sloman

Gardiner

Gilbert

Rohde

-RRB-

Human

social

hierarchy

does

not

need

actual

-LRB-

ritual

-RRB-

fighting

to

be

established

but

can

also

be

formed

by

imagining

what

would

happen

in

case

of

such

an

encounter

Just

knowing

the

strength

of

your

opponent

may

be

enough

to

experience

a

sense

of

inferiority

-LRB-

Rohde

-RRB-

which

in

turn

can

boost

depressive

problems

-LRB-

Brown

Harris

Hepworth

Gilbert

Allan

-RRB-

Internal

hierarchical

positions

are

also

fed

by

the

amount

of

social

attention

directed

by

others

-LRB-

Gilbert

-RRB-

As

opposed

to

most

other

species

humans

base

-LRB-

internal

-RRB-

hierarchies

not

just

on

physical

strength

but

on

other

capacities

related

to

the

availability

of

resources

such

as

intelligence

and

physical

attractiveness

-LRB-

e.g.

Anderson

John

Keltner

Kring

Nettle

-RRB-

as

well

Whereas

Price

's

Social

Ranking

Theory

is

achievement-oriented

and

implies

-LRB-

possibly

internal

-RRB-

social

comparison

and

grading

peer

status

can

also

be

defined

in

terms

of

affection

that

is

being

liked

or

disliked

-LRB-

rejected

-RRB-

While

achievement-related

status

mainly

refers

to

the

hierarchical

position

within

the

group

affection-related

status

may

also

have

implications

for

group

membership

in

itself

Throughout

human

history

being

banished

from

the

group

has

been

similar

to

a

death

sentence

so

it

is

not

surprising

that

being

accepted

by

other

people

represents

a

core

motive

-LRB-

e.g.

Baumeister

Leary

Fiske

-RRB-

Hence

social

rejection

is

a

significant

stressor

-LRB-

Coie

-RRB-

which

can

bring

about

feelings

of

loneliness

-LRB-

Asher

Wheeler

-RRB-

and

low

self-esteem

-LRB-

O'Brien

Bierman

-RRB-

and

may

contribute

to

the

development

of

depressive

problems

-LRB-

Boivin

Poulin

Vitaro

Monroe

Hadjiyannakis

-RRB-

Social

ranking

and

social

rejection

theories

offer

sound

theoretical

frameworks

to

predict

depressive

problems

in

those

with

a

low

status

but

are

less

explicit

regarding

consequences

of

having

a

high

status

Although

there

is

a

social

gradient

of

-LRB-

mental

-RRB-

ill-health

that

is

health

improves

with

each

step

one

moves

up

the

socioeconomic

ladder

-LRB-

e.g.

Siegrist

Marmot

-RRB-

this

may

not

be

translated

directly

to

the

association

between

peer

status

and

depressive

problems

in

adolescence

In

fact

animal

research

suggest

that

both

high

and

low-status

individuals

can

suffer

from

distress

among

other

things

depending

on

their

personality

and

the

stability

of

the

hierarchy

-LRB-

Sapolsky

-RRB-

Hence

high

and

low

peer

status

may

not

be

two

sides

of

the

same

coin

and

better

be

investigated

separately

Though

adolescents

social

status

is

multidimensional

-LRB-

Rohde

-RRB-

the

achievement-related

and

the

affection-related

approaches

to

social

status

have

evolved

independently

from

each

other

and

have

to

the

best

of

our

knowledge

never

been

studied

in

concert

Hence

it

is

not

known

how

the

two

are

interrelated

and

how

they

jointly

affect

depressive

problems

Multiple

hierarchies

exist

within

each

of

the

two

status

domains

as

well

achievement

includes

specific

areas

such

as

sports

and

school

performance

and

affection

may

relate

to

being

disliked

or

being

someone

's

best

friend

This

raises

the

question

if

and

how

a

low

status

in

one

of

the

areas

can

be

compensated

by

a

high

status

in

another

area

or

vice

versa

Being

caught

in

a

-LRB-

degrading

-RRB-

situation

where

no

escape

seems

possible

is

a

particularly

potent

trigger

of

depressive

problems

-LRB-

Brown

et

al.

-RRB-

A

position

at

the

bottom

of

the

status

hierarchy

may

be

experienced

as

such

but

this

is

considerably

less

likely

in

adolescents

who

are

highly

regarded

in

other

respects

since

people

tend

to

value

most

the

hierarchy

in

which

they

rank

highest

-LRB-

Sapolsky

-RRB-

We

therefore

expected

that

the

effects

of

low

status

could

be

compensated

by

high

status

in

another

area

Gender

differences

Until

now

only

a

limited

number

of

studies

have

focused

on

gender

differences

in

associations

between

social

status

in

particular

domains

and

depressive

problems

With

respect

to

achievement-related

status

this

is

at

least

partly

due

to

the

fact

that

there

are

several

well-documented

strategies

to

induce

achievement-related

social

stress

in

males

-LRB-

e.g.

defeat

in

aggressive

encounters

-RRB-

but

hardly

any

for

females

among

other

things

because

it

is

hard

to

obtain

strong

dominance

relationships

in

female

animals

-LRB-

Haller

Fuchs

Halász

Makara

-RRB-

Rather

than

winning

or

losing

in

competitive

encounters

females

seem

to

be

sensitive

to

situations

which

disturb

their

relationships

with

other

animals

in

the

group

such

as

social

isolation

-LRB-

Holston

Scallet

Ali

Turner

-RRB-

and

instability

-LRB-

Haller

et

al.

-RRB-

There

is

some

evidence

for

gender

differences

in

humans

as

well

In

a

study

where

men

and

women

were

subjected

to

a

series

of

different

stress

conditions

men

appeared

physiologically

more

reactive

to

achievement

challenges

and

women

to

social

rejection

challenges

-LRB-

Stroud

Salovey

Epel

-RRB-

Consistent

with

this

work

problems

and

downward

social

mobility

-LRB-

i.e.

lower

achievement-related

status

-RRB-

have

been

found

to

be

associated

with

men

's

mental

health

in

particular

while

women

are

more

sensitive

to

problems

in

getting

along

with

dear

ones

and

peer

rejection

-LRB-

e.g.

Brendgen

Wanner

Morin

Vitaro

Kendler

Thornton

Prescott

Tiffin

Pearce

Parker

-RRB-

Taken

together

these

results

suggest

gender

differences

in

the

salience

of

specific

domains

of

social

status

We

propose

that

autonomy

and

dominance

are

more

important

sources

of

well-being

for

males

than

for

females

so

that

boys

will

be

especially

sensitive

to

achievement-related

life

events

and

status

loss

-LRB-

Price

et

al.

Rose

Rudolph

-RRB-

Girls

depressive

problems

on

the

other

hand

are

expected

to

be

more

strongly

associated

to

affection-related

peer

nominations

Compared

to

boys

adolescent

girls

exhibit

greater

affiliative

needs

increasing

their

sensitivity

to

conflict

and

rejection

within

interpersonal

relationships

-LRB-

Cyranowski

Frank

Young

Shear

Prinstein

Aikins

-RRB-

Present

study

We

have

studied

the

relationship

between

early

adolescents

depressive

problems

and

social

status

in

peers

comprising

an

everyday

social

network

We

chose

to

study

classroom

social

status

as

measured

by

peer

nominations

as

this

is

a

social

environment

in

which

adolescents

spend

a

large

part

of

the

day

and

are

engaged

in

comparable

activities

The

fact

that

school

attendance

is

compulsory

in

early

adolescence

hence

the

situation

is

inescapable

is

also

relevant

in

this

respect

because

entrapment

has

been

found

to

be

a

particularly

potent

elicitor

of

depressive

-LRB-

like

-RRB-

emotions

and

behaviors

-LRB-

Brown

et

al.

Gilbert

Allan

-RRB-

Peer

status

was

assessed

by

sociometric

nominations

which

are

generally

considered

the

most

reliable

and

valid

indices

of

peer

status

-LRB-

Coie

Dodge

-RRB-

Because

the

association

between

depression

and

peer

status

might

be

confounded

by

comorbid

conduct

problems

-LRB-

Connell

Dishion

Kiesner

-RRB-

we

adjusted

for

comorbidity

in

our

analyses

We

hypothesized

that

low

peer

status

in

the

achievement-related

domain

would

be

most

relevant

for

depressive

problems

in

boys

whereas

girls

depressive

problems

would

be

more

strongly

associated

with

low

affection-related

peer

status

A

further

hypothesis

was

that

high

peer

status

in

a

particular

area

although

not

necessarily

associated

with

the

probability

of

depressive

problems

in

itself

would

reduce

the

association

between

depressive

problems

and

low

status

in

another

status

area

Method

Sample

The

TRacking

Adolescents

Individual

Lives

Survey

-LRB-

TRAILS

-RRB-

is

a

prospective

cohort

study

of

Dutch

-LRB-

pre

-RRB-

adolescents

with

the

aim

to

chart

and

explain

the

development

of

mental

health

from

preadolescence

into

adulthood

which

started

in

Sample

selection

involved

two

steps

First

five

municipalities

in

the

North

of

the

Netherlands

including

both

urban

and

rural

areas

were

requested

to

give

names

and

addresses

of

all

inhabitants

born

between

and

-LRB-

first

two

municipalities

-RRB-

or

and

-LRB-

last

three

municipalities

-RRB-

yielding

names

Simultaneously

primary

schools

-LRB-

including

schools

for

special

education

-RRB-

within

these

municipalities

were

approached

with

the

request

to

participate

in

TRAILS

School

participation

was

a

prerequisite

for

eligible

children

and

their

parents

to

be

approached

by

the

TRAILS

staff

Of

the

primary

schools

within

the

municipalities

-LRB-

of

the

schools

accommodating

of

the

children

-RRB-

agreed

to

participate

in

the

study

If

schools

agreed

to

participate

parents

-LRB-

or

guardians

-RRB-

and

children

received

brochures

with

information

about

the

study

and

a

TRAILS

staff

member

visited

the

school

to

inform

eligible

children

about

the

study

Shortly

thereafter

a

TRAILS

interviewer

contacted

parents

by

telephone

to

ask

whether

they

and

their

child

were

willing

to

participate

in

the

study

Respondents

with

an

unlisted

telephone

number

were

requested

by

mail

to

pass

on

their

number

If

they

reacted

neither

to

that

letter

nor

to

a

reminder

letter

sent

a

few

weeks

later

staff

members

paid

personal

visits

to

their

house

Parents

who

refused

to

participate

were

asked

for

permission

to

call

back

in

about

months

If

both

parents

and

children

agreed

to

participate

parental

written

informed

consent

was

obtained

after

the

procedures

had

been

fully

explained

Children

were

excluded

from

the

study

if

they

were

incapable

of

participating

due

to

mental

retardation

or

a

serious

physical

illness

or

handicap

or

if

no

Dutch-speaking

parent

or

parent

surrogate

was

available

and

it

was

not

feasible

to

administer

any

of

the

measurements

in

the

parent

's

language

Of

all

children

approached

for

enrollment

in

the

study

-LRB-

N

-RRB-

were

excluded

because

of

mental

or

physical

incapability

or

language

problems

Of

the

remaining

children

-LRB-

N

mean

age

SD

girls

-RRB-

were

enrolled

in

the

study

-LRB-

i.e.

both

child

and

parent

agreed

to

participate

-RRB-

Responders

and

non-responders

did

not

differ

with

respect

to

the

prevalence

of

teacher-rated

problem

behavior

nor

regarding

associations

between

sociodemographic

variables

and

mental

health

outcomes

-LRB-

De

Winter

et

al.

-RRB-

The

present

study

is

based

on

data

from

the

second

assessment

wave

of

TRAILS

-LRB-

-RRB-

which

was

held

to

years

after

baseline

assessment

-LRB-

mean

number

of

months

SD

range

-RRB-

During

this

wave

questionnaires

were

filled

out

by

the

adolescents

their

parents

and

their

teachers

The

adolescents

filled

out

their

questionnaires

at

school

in

the

classroom

under

the

supervision

of

one

or

more

TRAILS

assistants

In

addition

to

the

regular

questionnaires

which

were

filled

out

by

TRAILS

participants

only

the

assessment

wave

also

included

peer

nominations

which

were

collected

in

both

TRAILS

participants

and

their

classmates

at

a

separate

occasion

Of

the

baseline

participants

-LRB-

N

girls

-RRB-

participated

in

the

first

follow-up

assessment

-LRB-

-RRB-

Mean

age

at

was

SD

Subsample

with

peer

information

Peer

nominations

were

assessed

in

classes

with

at

least

three

regular

TRAILS

participants

In

total

school

classes

within

schools

met

this

criterion

and

agreed

to

participate

Two

schools

dropped

out

because

of

planning

difficulties

while

one

school

did

not

give

permission

to

assess

peer

nominations

within

the

class

At

this

school

we

collected

nominations

from

TRAILS

participants

only

providing

there

were

more

than

participants

per

class

Schools

provided

the

names

of

classmates

of

TRAILS

participants

Subsequently

all

eligible

students

received

an

information

letter

for

themselves

as

well

as

their

parents

in

which

they

were

asked

to

participate

in

this

part

of

the

data

collection

If

students

or

their

parents

refrained

from

participation

they

were

requested

to

send

a

reply

card

within

days

Passive

informed

consent

was

adequate

for

non-TRAILS

participants

because

they

were

used

as

informants

rather

than

being

study

subjects

themselves

A

total

number

of

students

of

whom

were

regular

TRAILS

participants

refused

to

participate

Peer

nominations

were

assessed

approximately

weeks

after

the

information

letter

had

been

sent

The

assessment

of

the

peer

nominations

lasted

about

min

and

took

place

during

the

regular

lessons

Two

participating

classes

were

excluded

due

to

too

many

recent

changes

in

the

composition

of

the

class

In

total

students

of

whom

were

regular

TRAILS

participants

filled

out

the

questionnaire

and

nominated

their

classmates

-LRB-

outdegree

-RRB-

whereas

students

of

whom

TRAILS

respondents

were

nominated

by

others

-LRB-

indegree

-RRB-

These

adolescents

did

not

differ

from

the

rest

of

the

TRAILS

participants

regarding

gender

χ2

-LRB-

N

-RRB-

p

or

depressive

problems

t

-LRB-

-RRB-

p

but

were

slightly

younger

-LRB-

mean

age

SD

versus

SD

-RRB-

t

-LRB-

-RRB-

p

Persons

with

missing

or

unreliable

data

on

depressive

problems

were

excluded

leaving

cases

for

analysis

Measures

Affection

and

achievement

related

status

Respondents

could

nominate

any

of

their

classmates

on

a

total

of

questions

covering

a

wide

range

of

issues

and

behaviors

-LRB-

e.g.

gossiping

helping

others

substance

abuse

-RRB-

For

the

purpose

of

this

study

we

selected

six

questions

three

of

which

referred

to

affection-related

status

-LRB-

Which

classmates

do

you

like

Which

classmates

do

you

totally

dislike

Which

classmates

are

your

best

friends

-RRB-

and

three

to

achievement-related

status

-LRB-

Who

are

good

at

sports

Who

are

good

learners

Who

are

good-looking

-RRB-

For

each

of

these

questions

we

used

the

proportion

of

nominations

-LRB-

indegree

-RRB-

as

a

measure

of

peer

status

Despite

the

fact

that

being

good-looking

is

usually

more

due

to

inherited

features

than

to

personal

accomplishments

it

was

classified

as

achievement-related

because

it

is

a

salient

social-ranking

dimension

in

adolescence

which

has

been

found

to

contribute

to

a

higher

social

ranking

and

influence

through

halo

effects

-LRB-

beautiful

is

good

-RRB-

-LRB-

Anderson

et

al.

Eagly

Ashmore

Makhijani

Longo

Feingold

-RRB-

Depressive

problems

Internalizing

and

externalizing

problem

behaviors

were

assessed

by

the

Child

Behavior

Checklist

-LRB-

CBCL

-RRB-

one

of

the

most

commonly

used

questionnaires

in

current

child

and

adolescent

psychiatric

research

-LRB-

Achenbach

Verhulst

Achenbach

-RRB-

It

contains

a

list

of

behavioral

and

emotional

problems

which

parents

can

rate

as

not

true

somewhat

or

sometimes

true

or

very

or

often

true

in

the

past

months

In

addition

to

the

CBCL

we

administered

the

self-report

version

of

this

questionnaire

the

Youth

Self-Report

-LRB-

YSR

Achenbach

-RRB-

The

original

empirically

derived

CBCL

and

YSR

scales

did

not

distinguish

between

anxiety

and

depressive

problems

In

order

to

improve

the

correspondence

with

clinical

diagnostic

categories

Achenbach

Dumenci

and

Rescorla

-LRB-

-RRB-

constructed

DSM-IV

scales

for

CBCL/YSR

problem

behaviors

based

on

international

experts

ratings

The

resulting

CBCL/YSR

Depressive

Problems

scale

consists

of

items

-LRB-

Cronbach

's

α

CBCL

YSR

-RRB-

covering

depressed

mood

anhedonia

loss

of

energy

feelings

of

worthlessness

and

guilt

suicidal

ideation

sleep

problems

and

eating

problems

The

scale

has

been

found

to

correspond

more

closely

to

DSM-IV

Major

Depressive

Disorder

than

the

original

CBCL/YSR

scales

-LRB-

Van

Lang

Ferdinand

Oldehinkel

Ormel

Verhulst

-RRB-

Test-retest

reliabilities

of

the

DSM-IV

scales

are

good

CBCL

r

YSR

r

-LRB-

Achenbach

et

al.

-RRB-

The

mean

item

score

of

the

CBCL-scale

was

-LRB-

SD

range

-RRB-

and

the

mean

item

score

of

the

YSR-scale

-LRB-

SD

range

-RRB-

while

-LRB-

CBCL

-RRB-

and

-LRB-

YSR

-RRB-

of

the

scores

were

greater

than

-LRB-

which

implies

that

more

than

half

of

the

symptoms

were

rated

as

at

least

somewhat

true

and/or

more

than

of

the

symptoms

as

very

true

-RRB-

The

agreement

between

parent-reported

and

adolescent-reported

depressive

problems

was

moderate

r

Because

depressive

problems

rated

as

present

by

both

parent

and

adolescent

are

assumed

to

be

more

severe

-LRB-

more

generalized

-RRB-

than

problems

rated

by

only

one

informant

we

used

the

mean

of

the

standardized

parent

and

adolescent

scores

as

a

measure

of

depressive

symptoms

in

this

study

When

data

of

one

informant

were

missing

or

unreliable

-LRB-

YSRs

n

CBCLs

n

-RRB-

the

composite

score

was

based

on

only

one

informant

Disruptive

behavior

Disruptive

behavior

was

included

to

adjust

for

comorbidity

Disruptive

behavior

was

assessed

with

the

CBCL

and

the

YSR

by

combining

the

items

for

Oppositional

Defiant

Disorder

and

Conduct

Disorder

which

resulted

in

a

scale

of

-LRB-

CBCL

Cronbach

's

α

-RRB-

or

-LRB-

YSR

Cronbach

's

α

-RRB-

items

Statistical

analysis

First

means

of

and

correlations

between

the

variables

used

in

the

study

were

calculated

and

gender

differences

tested

by

means

of

respectively

t-tests

and

z-tests

Subsequently

depressive

problems

were

predicted

by

peer-status

variables

in

a

linear

regression

model

Standard

errors

were

adjusted

for

possibly

dependent

observations

within

classes

using

the

cluster

option

available

in

the

statistical

package

of

STATA

-LRB-

StataCorp.

-RRB-

We

adjusted

for

disruptive

behavior

by

including

it

as

an

independent

variable

in

all

regression

analyses

Continuous

variables

were

standardized

to

mean

and

standard

deviation

to

ease

interpretation

of

the

regression

coefficients

In

order

to

be

able

to

distinguish

between

associations

with

low

status

and

associations

with

high

status

each

of

the

peer-status

variables

were

categorized

into

low

status

-LRB-

below

the

percentile

-RRB-

high

status

-LRB-

above

the

percentile

-RRB-

and

intermediate

status

-LRB-

between

the

and

percentile

-RRB-

For

being

disliked

high

status

referred

to

the

lowest

-LRB-

few

nominations

-RRB-

and

low

status

to

the

highest

-LRB-

many

nominations

-RRB-

Status

was

included

in

the

analyses

as

dummy

variables

with

intermediate

status

as

the

reference

category

We

analyzed

both

the

effects

of

each

status

area

individually

and

the

effects

of

all

areas

adjusted

for

each

other

The

analyses

were

stratified

by

gender

To

establish

if

gender

differences

were

statistically

significant

we

performed

a

two-step

regression

analysis

in

the

total

sample

In

the

first

step

main

effects

of

all

peer-status

variables

were

entered

in

the

model

as

well

as

gender

and

disruptive

problems

Interactions

between

gender

and

each

of

the

peer-status

variables

if

any

were

entered

in

a

second

step

by

forward

stepwise

selection

with

a

criterion

for

entry

of

p

Finally

we

examined

whether

low

status

in

-LRB-

at

least

-RRB-

one

area

could

be

compensated

by

high

status

in

one

or

more

other

areas

Peer-status

variables

that

were

significantly

related

to

depressive

problems

-LRB-

as

established

by

correlations

with

the

effects

of

gender

and

disruptive

behavior

partialled

out

-RRB-

were

used

to

create

four

combined

status

groups

low

high

referring

to

adolescents

with

a

low

status

-LRB-

i.e.

below

the

percentile

-RRB-

in

at

least

one

area

and

a

high

status

-LRB-

above

the

percentile

-RRB-

in

at

least

one

area

only

low

designating

adolescents

with

a

low

status

in

at

least

one

area

and

no

high

status

only

high

representing

adolescents

with

a

high

status

in

at

least

one

area

and

no

low

status

and

intermediate

that

is

adolescents

with

an

intermediate

status

in

all

areas

Depressive

problems

in

the

only-low

only-high

and

intermediate

group

were

compared

to

those

in

the

low

high

group

-LRB-

the

reference

category

-RRB-

by

linear

regression

analysis

adjusting

for

disruptive

behaviors

and

for

dependent

observations

within

classes

Gender

differences

in

each

of

the

associations

were

tested

by

interaction

effects

A

lower

depression

score

in

the

low

high

group

than

in

the

only-low

group

suggests

that

compensation

of

a

low

status

by

a

high

status

in

another

area

is

possible

a

lower

depression

score

in

the

only-high

group

than

in

the

low

high

group

suggests

that

a

low

status

counteracts

the

benefits

of

a

high

status

in

another

area

Comparison

of

the

intermediate

group

with

the

low

high

group

provides

insight

in

the

differences

between

being

extraordinary

-LRB-

in

any

direction

-RRB-

and

being

average

To

take

into

account

that

the

specific

areas

may

represent

the

two

broad

domains

of

achievement-related

and

affection-related

status

disproportionately

this

procedure

was

repeated

for

the

strongest

predictor

of

each

For

all

analyses

a

p-value

smaller

than

was

considered

statistically

significant

Because

we

performed

multiple

statistical

tests

the

results

may

suffer

from

capitalization

on

chance

one

would

expect

some

of

the

associations

examined

to

be

significant

merely

on

the

basis

of

chance

and

the

probability

to

find

at

least

one

significant

effect

increases

with

every

additional

statistical

test

We

therefore

recommend

to

consider

the

general

pattern

of

associations

rather

than

interpreting

each

individual

effect

as

if

it

were

the

only

one

tested

Results

Descriptive

statistics

Table

presents

descriptive

statistics

of

the

variables

used

in

this

study

Girls

had

more

nominations

for

being

good

learners

and

good-looking

while

boys

were

nominated

more

often

as

being

good

at

sports

Affection-related

status

domains

showed

no

significant

gender

differences

Girls

had

more

depressive

problems

and

boys

more

disruptive

behavior

problems.Table

-LRB-

standard

deviation

-RRB-

of

peer

status

and

mental

health

by

genderMean

-LRB-

SD

-RRB-

Girls

-LRB-

N

-RRB-

Boys

-LRB-

N

-RRB-

Difference

t-testbEffect

sizecProportion

nominations

for

-LRB-

-RRB-

-LRB-

-RRB-

t

p

-LRB-

-RRB-

-LRB-

-RRB-

t

p

Best

-LRB-

-RRB-

-LRB-

-RRB-

t

p

Good

at

-LRB-

-RRB-

-LRB-

-RRB-

t

p

A

good

-LRB-

-RRB-

-LRB-

-RRB-

t

p

-LRB-

-RRB-

-LRB-

-RRB-

t

p

Mental

healthDepressive

-LRB-

-RRB-

-LRB-

-RRB-

t

p

Disruptive

-LRB-

-RRB-

-LRB-

-RRB-

t

p

aFor

descriptive

purposes

the

means

are

based

on

mean

unstandardized

scores

of

parent

and

adolescent

reports

The

tests

are

based

on

mean

standardized

scores.bEffects

are

adjusted

for

possible

dependence

of

observations

within

classes.cCohen

's

d

-LRB-

-RRB-

σpooled

where

is

the

mean

of

the

first

group

is

the

mean

of

the

second

group

and

σpooled

is

the

square

root

of

the

mean

variance

of

the

two

groups

-LRB-

-RRB-

Bivariate

associations

Correlations

between

the

peer

nominations

were

generally

moderate

to

high

-LRB-

Table

-RRB-

Being

a

good

learner

was

not

significantly

related

to

being

good-looking

and

in

girls

neither

to

being

liked

and

being

someone

's

best

friend

Strongest

associations

were

found

among

the

affection-related

variables

-LRB-

being

liked

being

disliked

and

being

someone

's

best

friend

-RRB-

In

girls

depressive

problems

were

most

strongly

associated

with

being

disliked

in

boys

the

correlation

was

highest

for

being

good

at

sports.Table

between

peer

status

and

mental

health

in

girls

-LRB-

above

the

diagonal

-RRB-

and

boys

-LRB-

below

the

diagonal

-RRB-

DepressiveDisruptiveLikedDislikedBest

friendGood

at

sportsGood

learnerGood-lookingproblemsbehaviorLiked

Disliked

Best

Friend

Good

at

sports

A

good

learner

Good-looking

Depressive

Problemsa

Disruptive

Behaviora

Note

Bold

p

italics

significant

gender

difference.aStandardized

scores

based

on

mean

standardized

scores

of

parent

and

adolescent

reports

Hierarchy

of

status

areas

To

distinguish

between

associations

with

low

and

high

status

each

of

the

peer-status

variables

was

categorized

into

low

status

-LRB-

below

the

percentile

-RRB-

high

status

-LRB-

above

the

percentile

-RRB-

and

intermediate

status

-LRB-

between

the

and

percentile

-RRB-

For

being

disliked

high

status

referred

to

the

lowest

-LRB-

few

nominations

-RRB-

and

low

status

to

the

highest

-LRB-

many

nominations

-RRB-

The

distribution

of

the

peer

nominations

precluded

that

the

high-status

group

of

being

disliked

contained

approximately

of

the

cases

of

the

adolescents

were

nominated

by

no

one

and

hence

classified

as

having

high

disliked

status

Table

shows

gender-specific

regression

coefficients

for

each

of

the

low

and

high-status

groups

vis-à-vis

the

intermediate

group

both

without

and

with

adjustment

for

the

other

peer-status

dimensions

All

effects

were

adjusted

for

disruptive

behavior

Together

the

peer-status

variables

explained

of

the

variance

in

boys

and

in

girls.Table

of

depressive

problems

by

low

and

high

peer-status

in

several

areas

adjusted

for

disruptive

behavior

stratified

by

genderB

-LRB-

p

-RRB-

Unadjusted

for

other

areasAdjusted

for

other

areasStatus

areaStatusGirlsBoysGirlsBoysAffectionLikedLow

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

High

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

DislikedLowa

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Higha

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Best

friendLow

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

High

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

AchievementGood

at

sportsLow

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

High

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Good

learnerLow

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

High

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Good-lookingLow

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

High

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

-LRB-

-RRB-

Note

Effects

of

low

and

high

status

are

with

regard

to

intermediate

status

Depressive

problems

were

standardized

to

mean

and

standard

deviation

Effects

are

adjusted

for

possible

dependence

of

observations

within

classes.aAs

opposed

to

other

status

areas

a

low

status

at

being

disliked

reflects

many

nominations

and

a

high

status

few.Bold

p

It

is

interesting

to

note

that

in

girls

only

low

status

appeared

associated

with

depressive

problems

while

in

boys

high

status

tended

to

be

relevant

as

well

especially

in

the

affection

domain

Both

without

and

with

adjustment

for

other

status

areas

girls

regression

coefficients

were

highest

for

a

low

status

in

being

liked

and

boys

coefficients

for

a

low

status

in

sports

indicating

that

girls

and

boys

with

few

nominations

regarding

these

areas

had

the

highest

number

of

estimated

depressive

problems

When

all

status

areas

were

analyzed

jointly

these

two

associations

were

the

only

significant

ones

Without

adjustment

for

disruptive

problems

the

effects

remained

largely

the

same

except

that

the

effect

of

a

high

status

for

being

disliked

-LRB-

reflecting

no

nominations

-RRB-

became

significant

in

boys

B

p

Gender

differences

in

the

associations

between

peer

status

and

depressive

problems

were

tested

by

interaction

effects

of

gender

and

each

of

the

peer-status

variables

in

the

total

sample

using

a

forward

stepwise

selection

procedure

-LRB-

with

criterion

for

entry

p

-RRB-

after

all

main

effects

had

been

entered

in

the

model

Gender

interactions

with

a

low

status

on

being

liked

B

p

and

a

low

status

on

being

good

at

sports

B

p

were

both

selected

by

this

procedure

but

the

latter

only

showed

a

trend

and

did

not

reach

the

significance

criterion

of

p

Other

status

variables

did

not

the

meet

the

entry

criterion

Across-area

compensation

With

the

effects

of

gender

and

disruptive

behavior

partialled

out

all

peer-status

variables

were

significantly

associated

with

depressive

problems

except

being

a

good

learner

r

p

This

variable

was

excluded

from

further

analyses

hence

the

index

describing

the

combined

statuses

was

based

on

five

areas

Of

the

total

sample

fell

in

the

low

high

group

in

the

only-low

group

in

the

only-high

group

and

in

the

intermediate

group

This

distribution

was

approximately

equal

for

girls

and

boys

χ2

-LRB-

N

-RRB-

p

The

regression

coefficients

for

the

only-low

only-high

and

intermediate

group

in

relation

to

the

high

low

group

are

presented

in

Table

Table

of

depressive

problems

by

cross-area

peer

status

groups

adjusted

for

disruptive

behaviorStatus

groupnB

-LRB-

p

-RRB-

Low

high

-LRB-

reference

category

-RRB-

Only

-LRB-

-RRB-

Only

-LRB-

-RRB-

-LRB-

-RRB-

Note

Low

High

low

status

in

at

least

one

area

and

high

status

in

at

least

one

area

Only

Low

low

status

in

at

least

one

area

and

no

high

status

Only

High

high

status

in

at

least

one

area

and

no

low

status

Intermediate

intermediate

status

in

all

areas

Effects

are

adjusted

for

possible

dependence

of

observations

within

classes.Bold

p

The

coefficients

indicate

that

the

only-low

group

had

more

depressive

problems

than

the

low

high

group

while

the

only-high

and

intermediate

group

did

not

differ

significantly

from

the

low

high

group

Thus

these

results

suggest

that

the

risk

associated

with

low

status

in

one

area

can

be

compensated

by

a

high

status

in

another

area

while

the

opposite

-LRB-

i.e.

counteraction

of

the

beneficial

effects

of

a

high

status

by

a

low

status

in

another

area

-RRB-

is

not

true

at

least

not

to

the

same

degree

of

the

associations

showed

gender

differences

-LRB-

all

p-values

-RRB-

To

test

the

robustness

of

these

findings

we

also

constructed

a

combined

status

index

-LRB-

with

the

categories

only

low

only

high

low

high

and

intermediate

as

defined

in

the

method

section

-RRB-

based

on

only

the

strongest

predictors

from

each

of

the

two

broad

domains

of

achievement-related

and

affection-related

status

that

is

being

liked

and

being

good

at

sports

respectively

The

use

of

this

alternative

status

measure

yielded

similar

results

more

depressive

problems

in

the

only-low

group

than

in

the

low

high

group

B

p

no

significant

differences

between

the

low

high

group

and

respectively

the

only-high

group

B

p

and

the

intermediate

group

B

p

and

no

gender

differences

-LRB-

all

p-values

-RRB-

Discussion

Striving

for

status

has

been

proposed

as

a

universal

human

motive

and

a

wealth

of

evidence

has

indicated

that

persons

status

within

their

group

influences

their

psychological

and

physiological

well-being

-LRB-

e.g.

Anderson

et

al.

-RRB-

For

adolescents

one

of

the

most

salient

social

groups

is

made

up

by

their

classmates

In

this

study

on

associations

between

classroom

peer

status

and

depressive

problems

in

young

adolescents

from

the

general

population

the

peer

status

variables

explained

about

of

the

variance

in

depressive

problems

-LRB-

unadjusted

for

measurement

error

hence

underestimating

the

true

variance

-RRB-

which

is

a

modest

yet

noteworthy

amount

We

hypothesized

that

peer

status

in

the

achievement-related

domain

would

be

most

relevant

for

depressive

problems

in

boys

whereas

girls

depressive

problems

would

be

more

strongly

associated

to

their

affection-related

peer

nominations

A

further

hypothesis

was

that

high

alternative

hierarchical

positions

would

-LRB-

wholly

or

partly

-RRB-

counteract

the

association

between

depressive

problems

and

low

status

in

a

particular

status

area

These

hypotheses

were

largely

supported

by

the

data

although

it

should

be

noted

that

the

expected

patterns

were

not

found

in

all

status

areas

investigated

Compared

with

other

status

areas

not

being

good

at

sports

hence

a

low

achievement-related

status

was

most

strongly

associated

with

depressive

problems

in

boys

while

not

being

liked

hence

a

low

status

on

an

affection-related

area

had

the

strongest

association

with

depressive

problems

in

girls

However

our

data

suggest

that

not

being

good

at

sports

may

affect

girls

well-being

as

well

and

the

gender

difference

regarding

this

status

domain

although

showing

a

trend

was

not

statistically

significant

Not

being

liked

on

the

other

hand

appeared

to

be

related

to

depressive

problems

in

girls

only

Hence

girls

seem

to

be

more

sensitive

than

boys

to

affection-related

status

particularly

to

a

lack

of

peers

that

like

them

Low

status

versus

high

status

In

general

the

risk

associated

with

a

low

group

status

was

stronger

than

the

resilience

associated

with

a

high

status

especially

in

girls

and

when

the

effects

of

all

status

areas

were

adjusted

for

each

other

the

only

variables

that

remained

significantly

associated

with

depressive

problems

reflected

a

low

status

Low

peer

status

may

lead

to

depressive

problems

through

a

variety

of

pathways

such

as

a

reduced

opportunity

to

make

friends

controlling

or

dominating

behavior

by

others

and

expressions

of

disapproval

or

contempt

-LRB-

Deater-Deckard

-RRB-

All

of

these

may

undermine

self-esteem

and

invoke

feelings

of

loneliness

-LRB-

Asher

Wheeler

O'Brien

Bierman

-RRB-

which

are

known

risk

factors

for

depressive

problems

-LRB-

e.g.

Monroe

Hadjiyannakis

Pelkonen

Marttunen

Aro

-RRB-

Although

a

high

peer

status

does

not

seem

to

be

very

effective

in

preventing

depressive

problems

in

itself

this

may

be

different

for

those

with

a

low

status

in

another

area

a

high

status

appeared

to

have

the

potential

to

compensate

the

risk

associated

with

a

low

status

Adolescents

with

a

low

status

in

at

least

one

area

and

no

high

status

to

counterbalance

this

had

most

depressive

problems

which

supports

the

idea

that

depression

is

particularly

likely

when

no

escape

from

the

degrading

situation

seems

possible

-LRB-

Brown

et

al.

-RRB-

and

underscores

once

more

the

importance

of

emphasizing

and

stimulating

adolescents

strengths

and

talents

particularly

for

adolescents

with

weaknesses

in

other

areas

Gender

differences

The

gender

differences

found

in

this

study

notably

the

fact

that

a

direct

association

between

affection-related

status

and

depressive

problems

was

found

only

in

girls

add

evidence

to

the

notion

that

interpersonal

stressors

are

more

depressogenic

for

adolescent

girls

than

for

boys

-LRB-

e.g.

Brendgen

et

al.

Shih

Eberhart

Hammen

Brennan

-RRB-

Associations

with

depressive

problems

were

adjusted

for

disruptive

behaviors

to

take

into

account

comorbidity

of

the

two

Without

adjustment

for

disruptive

behaviors

being

disliked

was

significantly

related

to

boys

depressive

problems

which

may

indicate

that

disliked

boys

tend

to

show

disruptive

behavior

rather

than

-LRB-

solely

-RRB-

depressive

problems

Achievement-related

status

was

most

strongly

related

to

depressive

problems

in

boys

Compared

to

girls

boys

focus

more

on

agentic

goals

including

their

own

dominance

on

the

peer

group

-LRB-

e.g.

Rose

Rudolph

-RRB-

which

apparently

makes

them

more

vulnerable

to

low

status

in

relevant

areas

However

we

found

some

indications

that

sports-related

achievements

may

be

relevant

for

girls

too

This

finding

could

hint

at

a

dwindling

of

traditional

socio-cultural

gender

differences

but

it

could

as

well

be

due

to

the

fact

that

not

being

good

at

sports

reflects

not

only

a

low

probability

of

winning

in

ritual

fights

but

also

lack

of

physical

energy

and

activity

which

is

likely

to

occur

in

both

depressed

boys

and

depressed

girls

The

issue

of

reverse

causality

is

discussed

more

extensively

hereafter

Differences

between

specific

status

areas

Within

the

achievement-related

domain

sporting

performance

was

more

strongly

associated

with

depressive

problems

than

being

a

good

learner

and

being

good-looking

Sports

more

than

theoretical

subjects

and

outward

appearances

reflect

ritual

agonistic

behavior

with

the

end

being

signified

by

the

loser

yielding

and

signals

of

superiority

by

the

winner

-LRB-

Rohde

-RRB-

In

other

words

sports

are

much

more

closely

related

to

social

defeat

than

the

other

two

achievement

domains

But

other

factors

may

account

for

the

relatively

exceptional

position

of

sports

as

well

To

start

with

adolescents

who

are

good

at

sports

may

do

relatively

more

physical

exercise

which

can

help

to

reduce

depressive

problems

-LRB-

e.g.

Nabkasorn

et

al.

-RRB-

so

there

may

be

a

direct

effect

of

sports

on

mental

health

on

top

of

the

effect

through

peer

status

Furthermore

being

good

at

sports

increases

the

likelihood

of

team

membership

which

in

turn

may

result

into

admiration

and

new

friendships

The

latter

could

be

especially

relevant

for

boys

who

tend

to

have

multiple

activity-related

friendships

rather

than

a

few

close

friends

-LRB-

e.g.

Moller

Hymel

Rubin

-RRB-

The

association

between

being

a

good

learner

and

having

a

high

peer

status

is

probably

more

equivocal

than

that

between

being

good

at

sports

and

peer

popularity

Although

some

studies

found

positive

correlations

between

grade

point

average

and

popularity

among

peers

-LRB-

e.g.

Cauce

Green

Forehand

Beck

Vosk

-RRB-

others

suggested

that

intelligence

is

not

associated

with

peer

status

-LRB-

Luthar

McMahon

-RRB-

and

that

overachievers

do

not

have

a

secure

position

in

their

peer

group

-LRB-

Hartup

-RRB-

In

fact

Bukowski

Sippola

and

Newcomb

-LRB-

-RRB-

showed

that

good

classroom

behavior

starts

to

loose

its

attraction

for

peers

during

early

adolescence

in

favor

of

more

aggressive

behaviors

As

concerning

the

role

of

being

considered

good-looking

our

results

seem

to

contradict

earlier

findings

that

physically

unattractive

adolescents

are

likely

to

receive

more

negative

feedback

-LRB-

Burns

Farina

Cash

-RRB-

and

that

body

dissatisfaction

is

an

independent

risk

factor

for

-LRB-

girls

-RRB-

depressive

problems

in

adolescence

-LRB-

e.g.

Stice

Hayward

Cameron

Killen

Taylor

-RRB-

That

peer

nominations

for

being

good-looking

did

not

independently

contribute

to

adolescents

-LRB-

lack

of

-RRB-

depressive

problems

in

our

study

could

be

due

to

the

fact

that

peer

judgments

have

little

to

do

with

one

's

own

body

image

-LRB-

Rosenblum

Lewis

-RRB-

Furthermore

not

being

good-looking

is

not

necessarily

equivalent

to

being

ugly

in

other

words

the

question

Who

is

unattractive

might

have

been

a

stronger

correlate

of

depressive

problems

than

the

question

Who

is

good-looking

Within

the

affection-related

domain

which

appeared

to

be

relevant

to

girls

in

particular

not

being

liked

was

the

most

salient

factor

in

relation

to

depressive

problems

In

other

words

it

does

not

seem

to

matter

much

whether

or

not

some

people

dislike

you

as

long

as

there

are

still

others

that

like

you

A

lack

of

sympathizers

is

a

threat

to

mental

health

From

an

evolutionary

point

of

view

this

makes

good

sense

because

being

liked

by

others

can

offer

social

support

and

protection

against

being

expelled

from

the

group

and

the

need

to

belong

is

a

fundamental

motivation

-LRB-

Baumeister

Leary

Fiske

-RRB-

Limitations

strengths

and

recommendations

for

future

research

Our

study

has

some

limitations

To

start

with

the

CBCL/YSR

Depressive

Problems

scale

was

not

developed

to

assess

depressive

problems

according

to

DSM-IV

criteria

but

constructed

on

the

basis

of

expert

ratings

of

the

original

empirically

derived

CBCL

and

YSR

scale

items

Consequently

the

items

do

not

represent

one-to-one

counterparts

of

all

DSM-IV

criteria

and

inferences

about

associations

between

peer

status

and

DSM-IV

Major

Depression

are

putative

Furthermore

we

performed

multiple

statistical

tests

which

increases

the

probability

of

chance

findings

and

warrants

replications

in

other

samples

A

final

limitation

of

the

study

is

the

cross-sectional

and

non-experimental

nature

of

our

data

which

does

not

allow

causal

interpretation

of

the

associations

found

-LRB-

Kraemer

et

al.

-RRB-

Most

likely

however

the

association

between

peer

status

and

depressive

problems

is

bidirectional

not

only

does

low

peer

status

induce

depressive

problems

depressed

children

and

adolescents

tend

to

display

maladaptive

interpersonal

behavior

as

well

-LRB-

e.g.

Altman

Gotlib

Baker

Milich

Manolis

-RRB-

A

bidirectional

association

is

especially

plausible

with

respect

to

sporting

performance

Being

looked

down

on

because

you

are

bad

at

sports

may

reduce

self

confidence

and

through

this

incite

the

development

of

depressive

problems

but

it

is

just

as

likely

that

depressive

problems

which

often

include

lack

of

energy

and

motivation

have

a

negative

impact

on

adolescents

sporting

achievements

Similarly

one

could

argue

that

the

association

between

depressive

problems

and

not

being

liked

is

a

two-way

one

Having

no

allies

in

a

peer

group

can

be

devastating

for

your

self-worth

and

hence

trigger

depression

but

depressed

behavior

may

also

fend

off

others

-LRB-

e.g.

Prinstein

Borelli

Cheah

Simon

Aikins

-RRB-

It

seems

plausible

however

that

if

depression

were

a

cause

of

not

being

liked

we

would

expect

that

to

happen

in

both

boys

and

girls

and

not

in

girls

only

We

thus

assume

that

not

being

liked

is

actually

a

risk

factor

for

depression

at

least

in

girls

Compared

to

the

vast

literature

on

peer

processes

predicting

conduct

problems

research

on

peer

processes

in

depression

has

been

relatively

rare

-LRB-

Connell

Dishion

Deater-Deckard

-RRB-

To

the

best

of

our

knowledge

our

study

is

one

of

the

first

to

examine

the

association

between

status-related

peer

nominations

and

depressive

problems

in

a

large

population

cohort

of

early

adolescents

In

addition

to

the

size

and

representativeness

of

our

sample

important

assets

of

this

study

relate

to

the

use

of

multiple

informants

multiple

aspects

of

peer

status

and

multiple

outcomes

which

allowed

adjusting

for

comorbid

disruptive

behavior

Hence

we

feel

that

it

is

an

excellent

starting

point

for

further

research

We

propose

three

lines

of

extension

Firstly

the

cross-sectional

associations

should

be

extended

to

-LRB-

bidirectional

-RRB-

patterns

of

associations

across

time

to

be

better

able

to

distinguish

between

concomitants

and

risk

factors

-LRB-

Kraemer

et

al.

-RRB-

Secondly

the

study

could

be

extended

to

include

other

outcomes

not

only

within

the

domain

of

psychopathology

but

also

regarding

somatic

health

Thirdly

it

is

highly

relevant

from

a

public-health

point

of

view

to

-LRB-

further

-RRB-

explore

to

what

extent

and

how

peer

processes

can

be

manipulated

in

order

to

prevent

aversive

outcomes

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 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 B I O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 I O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 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 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 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 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 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 B I O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 B 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 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 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 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 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 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 B 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 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 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 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 O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 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 B 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 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 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

depressive symptoms
social hierarchy
adolescent
gender


Defense-Inducing Volatiles : In Search of the Active Motif Herbivore-induced volatile organic compounds -LRB- VOCs -RRB- are widely appreciated as an indirect defense mechanism since carnivorous arthropods use VOCs as cues for host localization and then attack herbivores . Another function of VOCs is plant -- plant signaling . That VOCs elicit defensive responses in neighboring plants has been reported from various species , and different compounds have been found to be active . In order to search for a structural motif that characterizes active VOCs , we used lima bean -LRB- Phaseolus lunatus -RRB- , which responds to VOCs released from damaged plants with an increased secretion of extrafloral nectar -LRB- EFN -RRB- . We exposed lima bean to -LRB- Z -RRB- -3-hexenyl acetate , a substance naturally released from damaged lima bean and known to induce EFN secretion , and to several structurally related compounds . -LRB- E -RRB- -3-hexenyl acetate , -LRB- E -RRB- -2-hexenyl acetate , 5-hexenyl acetate , -LRB- Z -RRB- -3-hexenylisovalerate , and -LRB- Z -RRB- -3-hexenylbutyrate all elicited significant increases in EFN secretion , demonstrating that neither the -LRB- Z -RRB- - configuration nor the position of the double-bond nor the size of the acid moiety are critical for the EFN-inducing effect . Our result is not consistent with previous concepts that postulate reactive electrophile species -LRB- Michael-acceptor-systems -RRB- for defense-induction in Arabidopsis . Instead , we postulate that physicochemical processes , including interactions with odorant binding proteins and resulting in changes in transmembrane potentials , can underlie VOCs-mediated signaling processes . Introduction Plants respond to herbivore damage with the release of volatile organic compounds -LRB- VOCs -RRB- that signal the presence of herbivore prey to predators and parasites and thereby serve as an indirect defense mechanism -LRB- e.g. , Heil 2008 -RRB- . Research has demonstrated that these VOCs can also be perceived by neighboring plants or intact , systemic parts of the damaged plant -LRB- Baldwin et al. 2006 ; Heil 2008 -RRB- . Particularly , green-leaf volatiles -LRB- GLVs -RRB- have been associated with induced resistance in intact plants -LRB- Arimura et al. 2000 ; Engelberth et al. 2004 ; Farag et al. 2005 ; Ruther and Kleier 2005 ; Kost and Heil 2006 ;-RRB- . However , little is known about the identity of VOCs that are active in this context or about a structural motif that active VOCs might have in common . In an attempt to explain the inducing activity of such VOCs on gene expression , it has been suggested that GLVs with an α,β-unsaturated carbonyl group such as -LRB- E -RRB- -2-hexenal can trigger defense responses in Arabidopsis through their activity as reactive electrophile species -LRB- Almeras et al. 2003 -RRB- . In the present study , we used lima bean -LRB- Phaseolus lunatus L. -RRB- to search for traits that characterize defense-inducing VOCs . Lima bean responds to herbivore damage with the jasmonate-mediated production of VOCs and extrafloral nectar -LRB- EFN -RRB- , an aqueous , sugar-containing secretion on nonreproductive plant organs that attracts predatory arthropods -LRB- mainly ants -RRB- . The natural blend of VOCs that is released from a herbivore-damaged lima bean induces EFN secretion in intact neighboring plants -LRB- Kost and Heil 2006 -RRB- and serves as a within-plant signal -LRB- Heil and Silva Bueno 2007 -RRB- . Among the quantitatively dominant VOCs that are released from induced lima bean , however , only -LRB- Z -RRB- -3-hexenyl acetate significantly induced EFN secretion when used as pure compound -LRB- Kost and Heil 2006 -RRB- . In the present study , we applied different structurally related esters to lima bean plants and monitored their EFN secretion . Materials and Methods Plants were cultivated from seeds collected in the coastal area of Puerto Escondido , Oaxaca , México -LRB- 15 ° 55.596 N and 097 ° 09.118 W , elevation 15 m -RRB- . Seedlings were cultivated under ambient conditions in 250 ml pots filled with soil from the original growing site . The plants were watered daily and fertilized 6 weeks after germination with commercial fertilizer : 10 ml per plant of a solution of 3 mg l − 1 of `` Fertilisante foliar de alta concentración '' -LRB- Grupo Bioquímico Mexicano , Aaltillo , Coah. , Mexico -RRB- . Experiments were conducted with plants of an age of 8 -- 10 weeks . The following compounds were dissolved in Lanolin paste -LRB- all at 0.1 μg μl − 1 -RRB- : -LRB- A -RRB- -LRB- Z -RRB- -3-hexenyl acetate , -LRB- B -RRB- -LRB- E -RRB- -3-hexenyl acetate , -LRB- C -RRB- -LRB- E -RRB- -2-hexenyl acetate , -LRB- D -RRB- 5-hexenyl acetate , -LRB- E -RRB- -LRB- Z -RRB- -3-hexenyl isovalerate , -LRB- F -RRB- -LRB- Z -RRB- -3-hexenyl butyrate , -LRB- G -RRB- 2-ethylhexanol , -LRB- H -RRB- -LRB- E -RRB- -2-hexanal , -LRB- I -RRB- decanal , and -LRB- K -RRB- nonanal -LRB- Lanolin paste and all compounds were purchased from Sigma Aldrich -RRB- . Each 0.25 g paste per plant was applied on green plastic stripes -LRB- MAX Bindeband -RRB- attached to the plant in order to avoid direct contact of the paste with the plant . The plants were then packed in perforated PET foil bags -LRB- Bratenschlauch , Toppits , Minden , Germany -RRB- and in nets to protect them from EFN consumers . Amounts of VOCs released from the lanolin paste into the plants ' headspace under these conditions were monitored in a parallel experiment that used a closed-loop stripping system as described previously -LRB- Kost and Heil 2006 -RRB- . EFN secretion was quantified from the five youngest leaves 24 h later . In short , EFN concentration was measured with a portable refractometer , and nectar volume was measured with glass capillaries -LRB- Kost and Heil 2006 -RRB- to calculate the total amount of soluble solids secreted , which in the case of lima bean EFN are mainly glucose , fructose , and sucrose . Leaves were then collected and dried to calculate EFN secretion as soluble solids secreted per gram leaf dry mass and per 24 h. Plants to which Lanolin paste without any compound added had been applied served as controls -LRB- Lanolin control , LC -RRB- . Per day , two groups each comprising all 11 treatments were investigated , and in total eight plants per treatment were used . Results and Discussion Amounts of VOCs released from the lanolin paste into the plant 's headspace resembled those released from an induced plant , i.e. , all individual VOCs were present at 80 -- 130 % of the amount of -LRB- Z -RRB- -3-hexenyl acetate that is released under comparable conditions from five induced leaves of lima bean -LRB- data not shown , for amounts of VOCs released from an induced plant see Kost and Heil 2006 -RRB- . The different VOCs to which lima bean plants had been exposed affected their EFN secretion significantly -LRB- General linear model : F77 ,10 = 10.05 , P < 0.001 -RRB- . Compounds A -- F consisted of acyl hexenols in which the configuration and position of the double bond was systematically shifted from the polar head to the aliphatic terminus of the molecule . Moreover , the importance of the size of the acyl moiety was evaluated by using acetates , butyrate , and isovalerate of -LRB- Z -RRB- -3-hexenol . Although only one of the tested VOCs is naturally released from lima bean , exposure to the six compounds -LRB- A -- F -RRB- elicited EFN secretions that were significantly higher than those of the controls -LRB- LSD posthoc analysis : P < 0.05 , see Fig. 1 -RRB- . These compounds were -LRB- Z -RRB- -3-hexenyl acetate , -LRB- E -RRB- -3-hexenyl acetate , -LRB- E -RRB- -2-hexenyl acetate , 5-hexenyl acetate , -LRB- Z -RRB- -3-hexenyl isovalerate , and -LRB- Z -RRB- -3-hexenyl butyrate . Hence , EFN secretion by lima bean can be induced by VOCs that are not released naturally from this species , while several VOCs that are released from induced plants did not significantly change EFN secretion in previous experiments -LRB- Kost and Heil 2006 -RRB- . Fig. 1EFN secretion in μg soluble solids secreted per gram leaf dry mass and per 24 h is depicted . Plants were exposed for 24 h to -LRB- A -RRB- -LRB- Z -RRB- -3-hexenyl acetate , -LRB- B -RRB- -LRB- E -RRB- -3-hexenyl acetate , -LRB- C -RRB- -LRB- E -RRB- -2-hexenyl acetate , -LRB- D -RRB- 5-hexenyl acetate , -LRB- E -RRB- -LRB- Z -RRB- -3-hexenyl isovalerate , -LRB- F -RRB- -LRB- Z -RRB- -3-hexenyl butyrate , -LRB- G -RRB- 2-ethylhexanol , -LRB- H -RRB- -LRB- E -RRB- -2-hexanal , -LRB- I -RRB- decanal , and -LRB- K -RRB- nonanal dissolved in Lanolin paste . Plants to which pure Lanolin paste had been applied served as controls -LRB- LC -RRB- . Lower and upper whiskers represent the 5 % and the 95 % percentile , lower and upper margins of boxes the 25 % and the 75 % percentile , the lines within boxes indicate medians . Different letters appearing above boxes mark treatment effects that differ significantly from each other -LRB- P < 0.05 according to LSD posthoc analysis -RRB- , and structures of compounds leading to a significant induction of EFN secretion as compared to the control are graphically presented . LogP values of all compounds were calculated with ChemDraw Ultra 6.0 and are given under the structures -LRB- compounds A -- F -RRB- and in the insert -LRB- G -- K -RRB- How are VOCs perceived by plants , and via which mechanisms do they affect defense expression patterns ? It has been suggested that molecules with an α,β-unsaturated carbonyl group can trigger defenses in Arabidopsis through their activity as reactive electrophile species -LRB- Almeras et al. 2003 -RRB- . In principle , VOCs could also be perceived by binding to specific receptor-proteins or to odorant binding proteins with a preference for a class of compounds , similar to animal olfactory systems . Apparently , neither the configuration nor the position of the double bound is a critical factor . Compounds reported to prime or induce gene activity or phenotypic defenses in intact corn plants comprise -LRB- Z -RRB- -3-hexen-1-ol , -LRB- Z -RRB- -3-hexenal , and -LRB- Z -RRB- -3-hexenyl acetate -LRB- Engelberth et al. 2004 ; Farag et al. 2005 ; Ruther and Kleier 2005 -RRB- . We found that -LRB- Z -RRB- -3-hexenyl acetate , -LRB- E -RRB- -3-hexenyl acetate , -LRB- E -RRB- -2-hexenyl acetate , and 5-hexenyl acetate all elicited particularly high EFN secretion rates in lima bean . The majority of these substances lack an α,β-unsaturated carbonyl group , which thus can not be a generally required motif . In addition , a compound such as 5-hexenyl acetate can not yield an electrophile such as -LRB- E -RRB- -2-hexenal via the sequence of ester hydrolysis , oxidation , and isomerization , as it is principally possible for -LRB- E or Z -RRB- -3-hexenyl acetate or -LRB- E or Z -RRB- -2-hexenyl acetate . Moreover , unlike -LRB- Z -RRB- -3-hexenyl acetate the potential end product of the transformation sequence , -LRB- E -RRB- -2-hexenal , had only a weak effect on the nectar flow . Similarly , the observation that -LRB- Z -RRB- -3-hexenyl acetate and -LRB- E -RRB- -3-hexenyl acetate elicited almost identical EFN secretion rates contradicts the involvement of classical receptor proteins , since these usually require a specific stereochemistry of the interacting molecule . We further calculated LogP values to check for putative importance of the compound 's lipophilicity but did not find any clear relation between the EFN-inducing activity of a compound and its octanol-water partition coefficient -LRB- Fig. 1 -RRB- . More studies are required to understand which VOCs induce plant defenses via which mechanism . However , our results point to a new mechanism . Changes in transmembrane potentials -- occurring through modulations of ion fluxes -- are involved in early signaling events in the cellular response to stress -LRB- Maffei et al. 2007 -RRB- , and exposition to VOCs indeed changes membrane potentials in intact lima bean leaves -LRB- M. Maffei , personal communication -RRB- . It is thus tempting to speculate that the dissolving of VOCs in the membranes coupled to interactions with membrane proteins , similar to the odorant binding proteins of insects -LRB- Campanacci et al. 2001 -RRB- , leads to changes in transmembrane potentials and thereby induces gene activity . This hypothesis gains support from the observation that the induction of EFN secretion by VOCs appears to be a gradual one rather than a clear ` yes-or-know ' response . Several of the compounds that are released naturally from lima bean showed a trend towards an increase in EFN secretion , although the difference was not significant : EFN secretion in response to Linalool was on average higher by 40 % than in control plants , and DMNT -LRB- -LRB- 3E -RRB- -4,8-dimethylnona-1,3,7-triene -RRB- increased EFN secretion by 50 % -LRB- see Table 1 in Kost and Heil 2006 -RRB- . Similarly , plants exposed to ethylhexanol or -LRB- E -RRB- -2-hexanal in the present study showed a trend towards higher EFN secretions than controls -LRB- Fig. 1 -RRB- . EFN responds to a comparably wide variety of structures , and slight changes in the molecular structure gradually alter the induction effect . This observation is best explained by a comparably simple physicochemical process , whose detailed nature remains to be elucidated . How can signals evolve that apparently lack chemical specificity ? The answer might be simply that plants seldom are exposed to GLVs or chemically related compounds that are not released from attacked plants . In evolutionary terms , the probability of this situation was probably too low to cause any selection towards a more specific signal perception .

O O O O O O O O O O O O O O O O O O O O O B I O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 B I O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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 O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O 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

herbivore-induced volatiles
hexenyl acetate
induced defense
plant-plant communication