Fundamental ecology is fundamental

Abstract

The primary reasons for conducting fundamental research are satisfying curiosity, acquiring knowledge, and achieving understanding. Here we develop why we believe it is essential to promote basic ecological research, despite increased impetus for ecologists to conduct and present their research in the light of potential applications. This includes the understanding of our environment, for intellectual, economical, social, and political reasons, and as a major source of innovation. We contend that we should focus less on short-term, objective-driven research and more on creativity and exploratory analyses, quantitatively estimate the benefits of fundamental research for society, and better explain the nature and importance of fundamental ecology to students, politicians, decision makers, and the general public. Our perspective and underlying arguments should also apply to evolutionary biology and to many of the other biological and physical sciences.

Full text

Fundamental
ecology
is
fundamental
Franck
Courchamp
1,2
,
Jennifer
A.
Dunne
3
,
Yvon
Le
Maho
4
,
Robert
M.
May
5
,
Christophe
The
´baud
6
,
and
Michael
E.
Hochberg
3,7,8,9
1
Laboratoire
d’Ecologie,Syste´
matique,
et
Evolution,
UMR
CNRS
8079,
Universite´
Paris-Sud,
Orsay,
France
2
Department
of
Ecology
and
Evolutionary
Biology
and
Center
for
Tropical
Research,
and
the
Institute
of
the
Environment
and
Sustainability,
University
of
California
Los
Angeles,
Los
Angeles,
CA
90095,
USA
3
Santa
Fe
Institute,
1399
Hyde
Park
Road,
Santa
Fe,
NM
87501,
USA
4
Institut
Pluridisciplinaire
Hubert
Curien,
UMR
CNRS
7178,
Universite´
de
Strasbourg,
Strasbourg,
France
5
Department
of
Zoology,
University
of
Oxford,
Oxford
OX1
3PS,
UK
6
Laboratoire
Ecologie
et
Diversite´
Biologique,
UMR
CNRS
5174,
Universite´
Paul
Sabatier,
Toulouse,
France
7
Institut
des
Sciences
de
l’Evolution,
UMR
5554,
Universite´
Montpellier
II,
Montpellier,
France
8
Kavli
Institute
for
Theoretical
Physics,
University
of
California,
Santa
Barbara,
CA
93106-4030,
USA
9
Wissenschaftskolleg
zu
Berlin,
14193
Berlin,
Germany
The
primary
reasons
for
conducting
fundamental
re-
search
are
satisfying
curiosity,
acquiring
knowledge,
and
achieving
understanding.
Here
we
develop
why
we
believe
it
is
essential
to
promote
basic
ecological
research,
despite
increased
impetus
for
ecologists
to
conduct
and
present
their
research
in
the
light
of
poten-
tial
applications.
This
includes
the
understanding
of
our
environment,
for
intellectual,
economical,
social,
and
political
reasons,
and
as
a
major
source
of
innovation.
We
contend
that
we
should
focus
less
on
short-term,
objective-driven
research
and
more
on
creativity
and
exploratory
analyses,
quantitatively
estimate
the
bene-
fits
of
fundamental
research
for
society,
and
better
ex-
plain
the
nature
and
importance
of
fundamental
ecology
to
students,
politicians,
decision
makers,
and
the
general
public.
Our
perspective
and
underlying
arguments
should
also
apply
to
evolutionary
biology
and
to
many
of
the
other
biological
and
physical
sciences.
What
is
fundamental
ecology?
Fundamental
ecology,
or
basic
ecology,
is
the
study
of
organismal
diversity
and
of
the
interactions
between
organisms
and
their
abiotic
and
biotic
environments
[1].
Its
main
goal
is
to
advance
knowledge
and
understand-
ing,
and
its
results,
even
if
sometimes
predictable,
are
not
known
with
certainty
in
advance.
By
contrast,
applied
ecology
is
usually
motivated
by
particular,
well-defined
objectives,
typically
to
solve
environmental
problems,
in-
cluding
the
management
of
natural
resources
such
as
land,
energy,
food,
or
biodiversity
[2].
Because
applied
ecology
often
involves
the
development
of
interventions
to
alter
events
(e.g.,
exotic
species
invasions,
endemic
species
de-
cline),
this
research
is
essentially
an
attempt
to
achieve
a
defined
objective.
Box
1
summarises
the
different
types
of
research
and
Figure
1
presents
a
conceptual
model
of
their
relationships.
One
of
the
central
objectives
and
achievements
of
fun-
damental
ecology
is
to
develop
and
test
general
theory
in
ecology
[3].
At
the
broadest
level,
a
general
theory
is
an
entire
domain
of
science
and
a
set
of
interwoven
funda-
mental
principles,
like
the
theory
of
evolution
by
natural
selection
[4].
Fundamental
research
is
sometimes
called
pure
science
or
‘blue-skies’
research.
The
term
blue-skies
research
has
its
roots
in
the
idea
of
curiosity-
and
inquiry-driven
studies
(Box
1).
It
is
said
that
under
Eisenhower’s
presidency
a
prominent
politician
unsympathetic
to
basic
research
claimed
‘I
don’t
care
what
makes
the
grass
green!’,
which
was
later
rephrased
as
‘what
makes
the
sky
blue’
[5].
In-
terestingly,
50
years
earlier,
using
vapours
and
light
beams
in
glass
tubes,
Tyndall
[6]
explained
the
basis
of
the
sky’s
colour
and
his
work
led
to
better
and
more
effective
processes
and
products
unforeseen
at
the
time
of
his
discovery.
These
included
a
test
for
optically
pure
air,
support
for
the
nonexistence
of
spontaneous
generation,
particle
filtering
of
lung
airways,
mould
destruction
by
Penicillium
bacteria,
and
even
the
precursors
of
the
flexi-
ble
gastroscope
and
bronchoscope
[6].
Financial
support
for
fundamental
research
in
ecology,
like
in
many
other
disciplines,
is
highly
competitive.
Many
perceive
stasis
or
a
continuous
decline
in
support,
but
accurate
numbers
are
difficult
to
obtain
to
substantiate
general
trends,
since
funding
categorization
can
be
open
to
alternative
interpretations
and
data
can
be
difficult
to
obtain.
This
perception
that
support,
be
it
financial
or
moral,
is
not
improving
is
worrisome
for
ecologists,
for
their
science,
and
ultimately
for
society.
Here
we
show
how
the
predominant
support
for
objective-driven,
applied
science
is
relatively
recent
and
why,
in
conjunction
with
this,
fundamental
research
has
seen
limited
investment
in
both
relative
and
absolute
terms.
We
then
advocate
in-
creased
and
less
constraining
support
for
basic
ecology,
by
presenting
the
principal
drivers
of
fundamental
research
and
some
of
its
main
benefits,
including
the
satisfaction
of
human
curiosity,
the
quest
to
explain
our
world,
the
crea-
tion
of
scientific
knowledge,
and
the
often
unintentional,
Opinion
0169-5347/
ß
2014
The
Authors.
Published
by
Elsevier
Ltd.
This
is
an
open
access
article
under
the
CC
BY-NC-ND
license
(http://creativeco mmons.org/license s/by-nc-nd/3.0/).
http://dx.doi.org/10.1016/j.tree.2014.11.005
Corresponding
authors:
Courchamp,
F.
(franck.courchamp@u-psud.fr);
Hochberg,
M.E.
(mhochber@univ-montp2.fr).
Keywords:
applied
ecology;
basic
ecology;
blue-skies
research;
fundamental
research;
research
priorities.
Trends
in
Ecology
&
Evolution,
January
2015,
Vol.
30,
No.
1
9

more
tangible,
economic,
social,
and
political
outcomes,
as
well
as
many
innovations.
Finally,
we
present
concrete
propositions
to
promote
further
support
for
fundamental
research
in
ecology.
Major
trends
in
research
funding
Basic
versus
applied
research
through
history
Historically,
fundamental
approaches
have
played
a
dom-
inant
role
in
scientific
research,
with
discoveries
of
all
kinds
and
importance
being
the
trademark
of
fundamental
scientific
research,
from
Antiquity
to
the
Age
of
Enlight-
enment.
The
recent
surge
in
the
growth
of
institutionalised
applied
research
stems,
in
part,
from
political
perspectives
of
the
role
of
scientific
research
in
society
[7].
This
has
created
a
new
market-oriented
and
objective-driven
ap-
proach
to
science
to
respond
to
economic
and
societal
expectations
[6].
Before
the
industrial
revolution,
funding
sources
were
very
different
from
what
they
are
today,
coming
from
personal
funds
(scientists
were
often
aristocrats,
with
education,
money,
and
time),
sponsorship
from
the
less-
educated
nobility
without
the
time
or
interest
to
do
research
themselves,
or
funding
sought
by
the
scientists
from
the
public
through
experimental
demonstrations
or
natural
history
displays
–
the
famed
‘cabinets
of
curiosi-
ties’.
After
the
Industrial
Revolution,
the
costs
of
scientific
research
changed
scale
and
funding
became
more
orga-
nised,
often
through
universities.
Over
time,
university
funding
has
become
more
tied
to
governmental
and
private
funding.
Now,
universities
and
research
institutions
often
seek
sources
through
tuition
fees,
patent
licensing,
endow-
ments,
private
sponsorship,
or
alumni
contributions
[8].
As
a
result,
and
not
unexpectedly,
the
emphasis
of
research
is
increasingly
being
based
on
the
expectations
of
these
funders
and
is
thus
more
often
expressed
in
terms
of
direct
and
immediate
benefits
to
society.
Some
funding
calls
prioritise
projects
in
which
scientists
associate
with
indus-
try
and
most
universities
now
adopt
business
models
[6]
and
develop
entrepreneurial
centres
to
encourage
related
technology
transfer
[9].
Current
sources
of
funding
The
private
sector
typically
seeks
short-term
(i.e.,
1–4
years),
low-risk
returns
on
investment,
which
is
incompat-
ible
with
the
unpredictability
and
long-term
(typically
decadal)
nature
of
returns
on
basic
research
[Dos
Reme-
dios,
C.
(2000)
The
value
of
fundamental
research.
Inter-
national
Union
for
Pure
and
Applied
Biophysics
(http://
iupab.org/publications/value-of-fundamental-research/)]
[6].
Because
private
investors
usually
dislike
uncertainty,
fundamental
research
is
still
mostly
supported
by
govern-
mental
institutions.
Basic
research
is
also
increasingly
funded
by
philanthropic
foundations
and
wealthy
person-
alities
[8,10].
The
approach
of
benefactors
has
changed
from
supporting
small
research
projects
to
large-scale
programmes
and
earmarked
research
networks
such
as,
in
our
field,
deep-ocean
exploration
in
the
search
of
giant
squid
or
paleontological
expeditions
to
find
remains
of
Tyrannosaurus
rex
[10].
In
parallel,
there
is
a
recent
trend
towards
smaller
project
budgets
being
sought
directly
from
the
public,
through
crowdfunding
[11].
In
both
cases,
there
is
thus
an
understandable
concern
that
funded
pro-
grammes
can
become
idiosyncratic,
at
the
expense
of
the
coverage
of
a
wider,
less
biased
range
of
basic
research
topics.
For
example,
of
the
US$19
billion
of
private
funding
for
all
research
fields
in
2006,
nearly
a
quarter
was
spent
on
health-related
topics,
compared
with
less
than
3%
on
basic
biology,
a
considerable
decrease
compared
with
just
a
few
years
previously
[12].
Shift
of
governmental
support
from
basic
to
applied
Governments
are
the
primary
instrument
for
balancing
the
collective
benefits
of
a
research
strategy
and
of
personal
interests
and
funding
across
disciplines
so
that
research
embraces
the
full
spectrum
of
topics
[13].
However,
with
governmental
funding
showing
lower
trends
over
the
past
several
years
in
many
Organisation
for
Economic
Coopera-
tion
and
Development
(OECD)
countries
[14,15],
there
has
been
a
shift
in
strategy
towards
more
emphasis
on
the
short-
term
goals
of
applied
research,
the
creation
of
economic
value
even
becoming
a
legal
requirement
of
public
research
institutions
in
some
countries.
Thus,
many
institutions
dedicated
to
basic
research
are
expanding
their
associations
Box
1.
Different
types
of
research
We
define
fundamental
research
as
theoretical
or
experimental
work
undertaken
primarily
to
acquire
new
knowledge
of
the
underlying
foundations
of
phenomena
and
observable
facts
without
any
particular
application
in
view
[14].
Fundamental
ecology
is
therefore
often
exploratory
and
curiosity
driven.
By
contrast,
applied
research
focuses
on
finding
solutions
and
improving
them
and
is
thus
goal
driven.
The
distinction
in
objectives
is
important.
Conducting
research
with
a
specific
goal
in
mind
makes
achieving
it
more
likely
but
also
reduces
the
chances
of
obtaining
unexpected
results,
a
major
source
of
scientific
discovery.
Curiosity-driven
research
often
challenges
accepted
thinking
and
may
generate
new
fields
of
investigation.
Applied
research
feeds
to
some
extent
on
the
outcomes
of
fundamental
research
(see
Figure
1
in
main
text).
However,
basic
and
applied
research
are
not
entirely
discrete
alternatives
but
rather
can
be
viewed
as
a
continuum
[14].
This
is
especially
pertinent
to
ecological
research,
where
fields
such
as
conservation
biology,
fisheries
science,
or
global-change
biology
often
integrate
both
fundamental
and
applied
perspectives.
There
are
other,
less
discussed
but
nevertheless
important
research
approaches.
Development
research
aims
to
make
products
from
newly
discovered
technologies.
Strategic
research
is
primarily
directed
towards
understanding
the
fundamental
basis
of
an
applied,
ultimate
goal
[Dos
Remedios,
C.
(2000)
The
value
of
fundamental
research.
International
Union
for
Pure
and
Applied
Biophysics
(http://iupab.org/publications/value-of-fundamental-re-
search/)].
Finally,
translational
research
seeks
to
rapidly
transfer
findings
from
fundamental
research
into
practical
applications
of
direct
relevance
to
human
needs.
In
contrast
to
applied
research,
which
usually
represents
incremental
improvements
to
current
understanding,
translational
research
strives
to
deliver
break-
throughs,
notably
through
the
creative
and
multidisciplinary
exploration
of
results
from
fundamental
research.
Most
translational
research
is
aimed
at
innovative,
basic
biological
science
to
improve
medicine,
bypassing
the
typically
long
times
separating
basic
research
and
concrete
clinical
application
[39].
Ecology
provides
fertile
grounds
for
translational
research;
for
example,
in
conserva-
tion
science.
Unfortunately,
translational
research
often
competes
for
funding
and
attention
with
both
fundamental
and
applied
research,
although
its
success
obviously
relies
to
a
great
extent
on
progress
in
the
latter
[48].
In
principle,
the
boundaries
between
all
of
these
different
types
of
research
should
be
more
porous,
since
they
have
the
potential
to
interact
and
instruct
in
achieving
their
respective
aims.
Opinion Trends
in
Ecology
&
Evolution
January
2015,
Vol.
30,
No.
1
10

with
industry
and
are
becoming
increasingly
concerned
with
government-encouraged
transfer
to
application
[8].
In
OECD
countries,
funding
for
basic
research
over
the
past
few
decades
has
been
at
lower
levels
than
for
other
major
research
categories
[14]
[OECD
(2012)
Research
and
devel-
opment
statistics:
R&D
expenditure
by
sector
of
perfor-
mance
and
type
of
R&D.
OECD
Science,
Technology
and
R&D
Statistics
(http://www.oecd.org/statistics/)].
For
exam-
ple,
in
2011
funding
for
basic
research
was
at
14%
(US$85
billion),
whereas
applied
research
was
at
24%
(US$151
billion)
and
development
at
62%
(US$394
billion).
Sources
of
decisional
shifts:
politicians
and
the
public
The
increasing
influence
of
politicians
in
research
decision
making
[16]
has
lead
to
shorter
funding
timescales
corre-
sponding
to
political
mandates
and
priority
given
to
ques-
tions
of
direct
relevance
to
the
general
public
[Dos
Remedios,
C.
(2000)
The
value
of
fundamental
research.
International
Union
for
Pure
and
Applied
Biophysics
(http://iupab.org/
publications/value-of-fundamental-research/)].
There
is
a
growing
political
will
to
ensure
that
taxpayer-funded
re-
search
is
seen
to
benefit
the
public
[17].
Governments
face
numerous
competing
demands
for
public
funding,
including
some
of
more
immediate
and
obvious
benefits
to
society
such
as
controlling
emerging
diseases
or
increasing
agricultural
yields.
Limited
and
sometimes
erroneous
information
obtained
through
education
and
the
media
combine
to
ex-
plain
why
some
taxpayers
are
opposed
to
research
that
has
no
immediately
obvious
benefits
for
society.
Stimulating
themes
in
fundamental
ecology
such
as
understanding
bio-
logical
diversity
–
even
with
the
applied
aim
of
species
conservation
–
can
easily
be
made
to
sound
frivolous
and
counterproductive
in
the
context
of
economic
growth
and
societal
challenges
[18].
Some
scientists
have
echoed
this
trend
by
calling
for
‘more
projects
focusing
on
applied
challenges,
arguing
that
public
funding
should
focus
on
public
problems
rather
than
fundamental
curiosities’
[19],
while
some
have
claimed
that
‘blue
sky
research
should
be
brought
back
to
Earth’
[20].
With
current
environmental
challenges,
ecologists,
especially
younger
scientists,
are
increasingly
drawn
to-
wards
applied
ecology.
In
parallel,
the
current
obsession
in
academia
for
quantity
[21]
reduces
creativity,
reflection,
and
risk
taking
and
therefore,
arguably,
opportunities
for
fundamental
discoveries.
What
drives
fundamental
research?
Breakthroughs
in
research
often
result
from
a
combination
of
curiosity,
creativity,
intelligence,
passion,
perseverance,
and
even
chance
[22],
with
curiosity
being
arguably
the
main
driver
of
fundamental
science.
Exploring
the
wonders
of
life
and
the
nature
of
things
for
the
sake
of
knowledge
alone
is
possibly
one
of
the
most
ancient
and
noble
of
human
aspirations.
Another
driver
of
fundamental
research
is
the
innate
desire
to
understand
inherently
complex
systems.
Most
if
not
all
ecologists
marvel
at
understanding
the
intricate
beauty
of
systems
involving
many
interacting
components,
be
they
molecules,
individuals,
or
populations.
That
many
ecologists
are
now
trained
in
the
more
quantitative
sciences,
such
as
physics,
computer
science
and
mathemat-
ics,
is
indicative
of
the
general
affinity
across
disciplines
for
understanding
complex
concepts
and
systems.
Promoting
the
importance
of
fundamental
ecology
today
seems
a
key
Fundamental
Applied
Development
27.03%
49.81%
15.44%
Innovaon
Discoveries
C
u
r
i
o
s
i
t
y
d
r
i
v
e
n
S
o
l
u

o
n
d
r
i
v
e
n
P
r
o
d
u
c

o
n
d
r
i
v
e
n
TRENDS in Ecology & Evolution
Figure
1.
A
conceptual
model
showing
the
links
between
different
types
of
scientific
research
(Box
1).
The
width
of
the
curved
arrows
is
an
indication
of
the
importance
of
the
transfers
between
research
types.
For
example,
fundamental
research
is
the
main
basis
of
applied
research,
but
outcomes
of
applied
research
may
fuel,
in
turn,
new
studies
in
fundamental
research.
Similarly,
transfers
from
fundamental
research
to
development
research
are
often
called
‘translational
research’;
the
reverse
results
in
innovative
scientific
equipment
and
technologies
that
can,
in
turn,
open
new
lines
of
fundamental
research.
The
green
arrows
indicate
gains
for
society;
yellow
arrows
represent
relative
funding
[OECD
(2012)
Research
and
development
statistics:
R&D
expenditure
by
sector
of
performance
and
type
of
R&D.
OECD
Science,
Technology
and
R&D
Statistics
(http://www.oecd.org/statistics/)]
based
on
average
funding
for
the
past
10
years
in
Organisation
for
Economic
Cooperation
and
Development
(OECD)
countries.
Opinion Trends
in
Ecology
&
Evolution
January
2015,
Vol.
30,
No.
1
11

component
in
continuing
to
attract
skilled,
creative,
and
curious
students
to
ecology
who
can
help
to
build
and
test
a
coherent
body
of
fundamental
ecological
knowledge.
Why
fundamental
ecology
is
important
Understanding
Ecology
is
still
a
young
discipline
and
we
are
only
starting
to
reach
an
understanding
of
how
species
function
and
interact
and
of
the
processes
underlying
patterns
in
biodi-
versity.
Nevertheless,
there
are
numerous
examples
of
progress
in
understanding
basic
ecology
and
in
the
funda-
mental
ecological
frameworks
influencing
applications.
For
instance,
theory
has
contributed
immensely
to
identi-
fying
relationships
involving
different
temporal,
spatial,
and
biological
scales
[3].
Using
systems
as
diverse
as
hare–
lynx
interactions
(e.g.,
[23,24]),
host–parasite
relation-
ships
(e.g.,
[25,26]),
and
insect
pest
outbreaks
(e.g.,
[27])
ecologists
have
shown
that
mechanistic
models
can
out-
perform
many
data-fitting
statistical
models
in
under-
standing
how
these
complex
systems
function
and
in
predicting
future
trends.
Fundamental
studies
of
trophic
networks
have
shed
light
on
the
role
of
trophic
cascades
in
ecosystem
functioning
and
ecosystem
services,
in
both
oceanic
[28,29]
and
terrestrial
communities
[30].
Funda-
mental
studies
on
population
dynamic
modelling
have
highlighted
the
need
to
understand
the
interaction
of
demographic
and
genetic
factors
in
extinction
[31,32],
which
led
to
the
International
Union
for
the
Conservation
of
Nature
(IUCN)
Red
List
of
threatened
species
[33].
These
are
only
a
few
among
many
examples
demonstrating
methodological
or
applied
advances
that
arose
from
fun-
damental,
entirely
curiosity-driven
work
in
population
ecology.
A
general
illustration
comes
from
the
recent
com-
pilation
by
the
British
Ecological
Society
of
‘100
influential
papers’
from
the
past
100
years
[34],
most
of
which
involve
fundamental
ecology.
We
cannot
anticipate
all
the
biological
and
environmen-
tal
challenges
that
humanity
will
face
in
the
future.
A
fundamental
understanding
of
the
problems
underlying
the
current
environmental
and
biodiversity
crises
is
the
most
reasonable
path
to
solving
them.
It
is
also
probably
a
safer,
less
expensive,
and
ultimately
time-saving
option.
This
is
yet
another
reason
to
foster
the
acquisition
of
knowledge
with
as
few
preconceived
routes
as
possible.
Economic,
social,
and
political
perspectives
Economic
theory
has
demonstrated
the
importance
of
fun-
damental
research.
Since
the
output
of
basic
research
is
inherently
intangible,
unpredictable,
and
difficult
for
researchers
to
appropriate,
it
provides
some
of
the
largest
spill-over
benefits
to
society
[35–38].
Although
there
are
strong
conceptual
and
methodological
difficulties
in
asses-
sing
the
many
economic
benefits
of
publicly
funded
basic
research,
several
studies
have
demonstrated
its
impor-
tance,
arguing
for
high
levels
of
continuous
investment,
particularly
by
governments
(reviewed
in
[13]).
There
are
numerous
ways
through
which
benefits
from
research
flow
into
the
economy
and
society,
including:
(i)
increase
in
the
stock
of
useful
knowledge;
(ii)
supply
of
skilled
graduates
and
researchers;
(iii)
creation
of
new
scientific
instrumen-
tation
and
methodologies;
(iv)
development
of
networks
and
stimulation
of
social
interactions;
(v)
enhancement
of
problem-solving
capacities;
(vi)
creation
of
new
firms;
and
(vii)
provision
of
social
knowledge
[13].
Because
most
attempts
to
assess
the
socioeconomic
benefits
of
basic
research
focus
on
one
or
only
a
few
of
these
(usually
the
first),
the
total
benefits
are
often
underestimated
[13].
The
current
trend
for
seeking
an
applied
component
to
ecological
research,
independent
of
the
potential
economic
benefits
of
its
fundamental
components,
may
also
reflect
how
society
and,
ultimately,
policy
makers
often
view
ecology.
This
is
exacerbated
by
the
common
confusion
between
ecology
and
environmentalism
and
suggests
that
the
perception
of
ecology
should
be
corrected
through
education
to
explain
what
exactly
is
the
science
of
ecology
and
why
it
is
important
to
understand
its
underlying
processes
[4].
In
this
regard,
an
emphasis
is
needed
on
various
outreach
activities,
including
formal
programs
early
in
education,
and
more
systematic
popularisation
of
fundamental
ecology,
for
example
through
scientifically
sound
nature
documentaries.
Fundamental
ecology
is
also
important
for
political
credibility.
With
the
unprecedented
biodiversity
and
envi-
ronmental
crises,
ecologists
have
a
responsibility
to
pro-
vide
insights
into
the
functioning
of
highly
complex
systems.
Politicians
and
decision
makers
need
global
pre-
dictions
of
ecosystem
and
biodiversity
trajectories
and
ways
of
assessing
the
quality
and
uncertainty
associated
with
these
predictions,
but
they
also
require
an
accurate
understanding
of
the
underlying
processes
governing
pat-
terns
and
predictions.
Finally,
fundamental
research
in
ecology
–
and
in
other
fields
–
is
crucial
for
the
development
of
societies.
It
is
a
great
accomplishment
of
humanity
that
some
individuals
are
encouraged
to
advance
and
spread
knowledge
and
understanding
for
the
potential
benefit
of
all.
Worldwide,
societies
have
long
been
providing
resources
for
research
regardless
of
practical,
short-term
returns
[22].
Indeed,
it
has
been
argued
that
fundamental
research
is
not
a
luxury,
but
rather
a
cultural
achievement
and
even
the
foundation
of
many
types
of
benefit
for
the
entire
society
[22].
A
source
of
innovation
Probably
the
most
common
argument
defending
basic
research
is
that
it
potentially
leads
to
new
discoveries.
Novel
applications
without
prior
development
are
notori-
ously
uncommon,
ostensibly
because
major
discoveries
seldom
emerge
from
strategically
planned
research.
Inno-
vations,
when
they
do
arise,
often
stem
from
surprising
translations
or
recombinations
of
existing
knowledge
[19,39].
There
are
countless
examples
of
unexpected
appli-
cations
from
basic
research
(see
Boxes
2
and
3
for
examples
in
ecology).
A
typical
illustration
is
the
increased
focus
on
organismal
and
system
oddities
with
hopes
of
direct
appli-
cations
to
industry.
For
example,
The
Biomimicry
Institute
(http://biomimicry.net/about/biomimicry38/institute/)
compiled
over
2000
examples
of
technologies
inspired
by
basic
research
in
the
fields
of
ecology
and
evolution.
Appli-
cations
can
also
extend
to
fields
remote
from
basic
ecology.
For
example,
models
of
prey–predator
dynamics
developed
last
century
are
now
being
used
in
both
industrial
econom-
ics
[40]
and
political
economics
[41],
insights
from
research
Opinion Trends
in
Ecology
&
Evolution
January
2015,
Vol.
30,
No.
1
12

on
the
stability
and
complexity
of
ecosystems
shows
future
promise
in
the
regulation
of
banks
[42],
while
fundamental
studies
in
the
behavioural
ecology
of
social
insects
are
being
used
in
robotics
[43].
Nevertheless,
that
fundamental
research
often
fuels
applications
should
not
be
the
main
argument
for
why
we
need
basic
research.
Unexpected
applications
of
funda-
mental
discoveries
are
only
providential
byproducts
of
other
objectives.
We
believe
it
is
important
that
fundamen-
tal
ecology
is
a
major
source
of
application,
but
the
primary
justifications
for
conducting
fundamental
research
should
remain
firmly
grounded
in
satisfying
curiosity,
acquiring
knowledge,
and
achieving
understanding.
Promoting
fundamental
research
Ecologists
can
promote
basic
science
in
several
important
ways.
First,
we
need
to
regularly
assess
the
state
of
and
progress
made
in
fundamental
ecology
through
the
publi-
cation
of
perspective,
forum,
review,
and
synthesis
articles.
In
our
view,
one
of
the
most
influential
ways
to
make
both
headway
in
specific
areas
and
overall
progress
in
the
field
is
through
discussing
and
recasting
the
most
important
questions
in
fundamental
ecology
[44].
Second,
we
must
develop
and
draw
on
new
approaches
to
communicate
results
to
politicians
and
granting
agen-
cies
and
to
reassure
them
that
giving
liberty
to
scientists
is
Box
2.
Examples
of
basic
research
in
ecology
leading
to
environmental
applications
In
many
areas
of
applied
ecology,
the
urgency
of
problems
such
as
habitat
destruction
or
species
extinction
often
calls
for
a
rapid
response.
Conservation
science
is
unique
in
being
both
a
field
of
research
and
a
field
of
action,
but
environmental
challenges
have
naturally
shifted
emphasis
towards
action,
sometimes
at
the
expense
of
building
a
strong
fundamental
framework.
For
example,
a
few
decades
ago
biological
invaders
on
islands
were
typically
ignored
until
their
impact
became
a
decisive
factor
for
intervention,
which
was
pursued
without
either
pre-control
survey
or
post-control
monitoring
[49].
Consequently,
hysteresis
and
multiple
stable
states
(including
extinction)
were
generally
overlooked,
resulting
in
a
number
of
‘surprise
effects’,
typically
the
failure
of
eradication,
or
the
unex-
pected
outbreak
of
other,
hitherto
neglected
invasive
species
[50].
Subsequently,
incorporation
of
trophic-web
theory
allowed
assessments
of
mesopredator
releases
[51],
hyperpredation
pro-
cesses
[52],
competitor
releases
[53],
and
other
ecological
processes
related
to
interspecific
interactions
and
likely
to
interfere
with
or
facilitate
eradication
[54,55].
Similar
observations
apply
to
the
management
of
fisheries,
with
long-lasting
stock
collapses
[29,56],
algal
blooms
[57],
eutrophication
in
freshwater
and
coastal
marine
[58,59],
or
regime
shifts
in
terrestrial
ecosystems
[60].
Another
illustration
comes
from
a
fundamental
programme
aimed
at
better
understanding
the
mechanisms
underlying
the
Allee
effect
(a
positive
relationship
between
the
number
of
individuals
in
a
population
and
their
fitness
[61,62]).
Fundamental
research
has
led
to
the
questioning
of
the
basic
assumption
that
Allee
effects
are
an
intrinsic
characteristic
of
populations
and
therefore
cannot
be
created
by
human
activities.
Thus,
the
standard
paradigm
was
that
humans
could
only
drive
populations
down
to
sizes
where
a
pre-existing,
unexpressed
Allee
effect
would
be
activated.
Questioning
this
premise
led
to
the
discovery
of
possible
anthropogenic
Allee
effects,
through
which
human
activities
can
exert
inverse
density-dependent
exploitation.
According
to
this
concept,
the
arbitrary
value
people
attribute
to
rarity
would
confer
an
economic
value
on
rare
species
that
would
maintain
the
incentive
to
exploit
them,
even
at
very
high
levels
of
rarity
[63].
Rare
species,
being
more
valuable,
would
be
more
exploited,
thereby
becoming
even
rarer,
precipitating
a
vortex
of
extinction.
This
process
threatens,
through
many
different
wildlife-
based
markets,
countless
species
of
plants
and
animals.
A
final
poignant
illustration
of
the
importance
of
basic
science
is
the
increasing
need
to
promote
ecosystem
services,
where
possible
[64,65].
Only
with
an
adequate
understanding
of
the
patterns
and
underlying
mechanisms
of
ecosystem
functioning
can
one
justifiably
and
reliably
protect,
restore,
and
value
the
services
provided
by
that
ecosystem.
Unfortunately,
the
current
environmental
crisis
often
elicits
rapid
responses
with
little
fundamental
basis,
in
which
avoidable
(and
now
irrecoverable)
mistakes
would
have
been
averted.
Recent
estimates
of
the
global
value
of
pollination
for
agriculture
–
US$153
billion
annually
–
provides
a
good
illustration
of
the
importance
of
better
understanding
the
functioning
and
outputs
of
ecosystems
[66].
Box
3.
Examples
of
basic
research
in
ecology
leading
to
applications
in
health
sciences
Since
each
species
potentially
has
unique
biological
properties,
it
is
unsurprising
that
an
estimated
60%
of
antitumour
and
antimicrobial
drugs
are
of
natural
origin
[67].
Basic
research
is
at
the
origin,
for
example,
of
the
discovery
in
the
naked
mole
rat
(Heterocephalus
glaber)
of
anticancer
mechanisms
[68].
Taking
full
advantage
of
such
a
potential
‘gold
mine’
would
not
have
been
possible
without
research
on
understanding
the
causes
of
mortality
in
this
species
through
necroscopies.
Another
example
is
the
incubating
male
king
penguin
(Aptenodytes
patagonicus),
which
preserves
fish
in
its
stomach
for
up
to
3
weeks
to
feed
to
its
young
at
hatching,
should
the
mother
be
absent
when
the
juvenile
begs
for
food
[69].
Searching
for
the
mechanism
permitting
fish
conservation
at
378C
led
to
the
discovery
of
spheniscin,
a
small
antimicrobial
and
antifungal
peptide.
This
molecule
has
since
been
shown
to
reduce
the
growth
of
the
two
main
agents
of
hospital-acquired
infections,
the
bacterium
Staphylo-
coccus
aureus
and
the
fungus
Aspergillus
fumigatus
[70].
Basic
research
on
plant–insect
interactions
has
also
led
to
important
applications,
such
as
alternative
strategies
for
the
protection
of
crops
like
maize
and
rice
against
herbivores.
Plants
possess
many
defence
mechanisms
based
on
secondary
metabolites
or
defensive
proteins
whose
toxic
or
deterring
properties
are
harmful
to
herbivores.
However,
it
was
not
known
until
recently
that
some
plants
release
volatile
signals
that
attract
herbivore
natural
enemies,
such
as
parasitic
wasps
above
ground,
and
entomopathogenic
nematodes
below
ground
[71].
These
defences
are
regulated
by
conserved
signalling
pathways
[72].
In
another
striking
example,
fundamental
studies
of
the
microbial
ecology
of
hot
springs
at
Yellowstone
National
Park
resulted
in
the
discovery
of
hyperthermophiles,
which
incidentally
led
to
the
identification
of
the
Taq
polymerase,
since
then
used
in
PCR
[73].
This
unexpected
finding
was
key
to
innovations
in
agriculture
and
medicine
and
helped
create
the
new
field
of
biotechnology,
which
includes
genomics,
recombinant
gene
technol-
ogies,
applied
immunology,
and
the
development
of
pharmaceutical
therapies
and
diagnostic
tests.
These
are
just
a
few
examples
of
how
basic
ecology
can
unexpectedly
provide
the
molecular
basis
of
important
applications.
Environmental
health
is
another
field
where
basic
ecological
science
has
repeatedly
proved
to
be
of
major
importance.
With
the
development
of
mathematical
models
of
host–parasite
relationships
[74,75],
ecological
epidemiology
has
resulted
in
a
better
under-
standing
of
the
spread
of
pathogens
and
parasites,
paving
the
way
for
applications
to
manage
the
spread
of
infectious
diseases.
Examples
include
host–parasite
modelling
of
epidemics
such
as
measles,
which
subsequently
informed
control
campaigns
against
the
2001
outbreak
of
foot
and
mouth
disease
in
the
UK
[76–78].
Unfortunately,
basic
science
was
not
sufficiently
developed
(or
not
identified
as
such)
to
realise
its
full
potential
in
contributing
to
the
control
of
the
spread
of
avian
(H5N1)
and
swine
(H1N1)
flu
and
the
current
Ebola
epidemic.
Opinion Trends
in
Ecology
&
Evolution
January
2015,
Vol.
30,
No.
1
13

not
necessarily
at
the
expense
of
accountability
[6].
It
is
essential
that
funders
realise
that
academic
research
pro-
grammes
once
thrived
on
intellectual
freedom
and
that
this
is
essential
for
discovery
[16].
Defining
clearly
what
ecology
is
and
what
ecologists
are
(i.e.,
not
environmen-
talists)
is
key
in
this
regard.
Third,
we
should
consider
creating
programmes
giving
large
and
unconstrained
blue-sky
research
grants
to
prom-
ising
or
experienced
researchers.
Such
initiatives
obviously
should
not
be
at
the
expense
of
other
research
programmes.
Even
a
modest
investment
in
supplementary,
long-term
grants,
without
externally
imposed
objectives,
should
con-
tribute
to
considerable
breakthroughs
[16].
Fourth,
we
need
to
establish
norms
for
funding
funda-
mental
research
that
will
facilitate
adherence
by
agencies
and
promote
the
long-term
future
of
fundamental
science.
One
possibility
is
to
establish
an
international
norm
of
a
minimum
level
and/or
percentage
of
research
funding
that
participating
agencies
pledge
to
maintain
for
fundamental
science.
Participating
national,
international,
and
private
funding
bodies
would
then
officially
declare
that
they
adhere
to
international
norms.
This
highlights
the
neces-
sity
that
politicians/decision
makers,
agencies,
and
scien-
tists
all
be
involved
in
institutional-,
national-,
and
international-level
discussion
to
establish
such
norms
and
to
determine
the
extent
of
their
application.
We
stress
that
promoting
basic
ecological
research
requires
contin-
ued
dialogue
and
need
not
result
in
reduced
budgets
for
other
research
approaches
(Box
1).
Fifth,
we
must
find
ways
to
quantitatively
value
funda-
mental
findings.
This
would
involve
estimating
the
pay
off
and
timescale
of
the
returns
of
basic
research,
which,
although
known
to
be
important
[36,37],
remain
largely
unquantified
[16].
This
could
be
one
way
of
demonstrating
unambiguously
the
importance
of
fundamental
research
in
ecology.
Sixth,
we
need
to
reduce
the
pressure
on
ecologists
to
systematically
provide
short-term
results
to
predefined
questions.
One
approach
would
be
to
modify
tenure
rules
to
account
for
the
importance
of
longer-term,
less
output-
oriented
research.
Another
possibility
is
longer
grant
per-
iods,
to
promote
the
tackling
of
challenging
and
risky
questions
with
possible
unexpected
outcomes.
Despite
risk
being
increasingly
presented
as
a
positive
criterion,
the
current
mindset
on
productivity
discourages
risky
projects.
Typically,
reviewers
are
required
explicitly
to
express
their
opinions
on
the
expected
impact
of
grant
proposals,
often
in
terms
of
output
quality
and
certainty
[45,46].
Precluding
risk
can
constrain
creativity
and,
therefore,
discovery
in
the
longer
term
[47].
Seventh,
supervisors
and
mentors
need
to
encourage
their
students
to
conduct
side
projects
and
pilot
projects
to
foster
the
spark
of
curiosity
and
creativity.
Supervisors
and
mentors
should
insist
on
the
benefits
of
devoting
time
to
thinking,
to
exploratory
research,
and
to
the
importance
of
‘toying’
with
concepts
and
data
(and
thinking
‘outside
the
box’)
rather
than
emphasising
high
scientific
productivity
or
being
drawn
into
applied
research
on
the
sole
basis
of
better
funding
opportunities.
They
should
also
insist
on
the
importance
of
asking
difficult
(but
tractable)
questions
–
that
is,
steeping
out
of
one’s
comfort
zone.
Finally,
researchers
themselves
need
to
prioritise
how
to
effectively
defend
basic
research.
Politicians
have
long
recognised
that
research
free
of
practical
constraints
is
at
the
heart
of
technology
and
industry.
For
example,
Van-
nevar
Bush,
scientific
advisor
to
Franklin
D.
Roosevelt,
stated
in
1945
that
scientific
progress
resulted
‘from
the
free
interplay
of
free
intellectuals,
working
on
subjects
of
their
own
choice,
in
the
manner
dictated
by
their
curiosity
for
exploration
of
the
unknown’
[22].
Yet
many
politicians
still
view
basic
science
as
an
unaffordable
luxury,
especial-
ly
in
times
of
financial
stress
[9].
Young
and
more
senior
ecologists
must
engage
debate
with
policymakers
to
correct
the
common
misconception
that
fundamental
research
is
a
luxury
[Dos
Remedios,
C.
(2000)
The
value
of
fundamental
research.
International
Union
for
Pure
and
Applied
Bio-
physics
(http://iupab.org/publications/value-of-fundamen-
tal-research/)]
[9].
Concluding
remarks
We
have
developed
several
lines
of
reasoning
supporting
the
promotion
of
fundamental
research
in
ecology,
al-
though
the
same
or
similar
arguments
could
apply
to
evolutionary
biology
[18]
and
to
other
sciences.
We
empha-
sise
that
we
do
not
claim
that
applied
research
has
less
merit
or
intrinsic
value
than
basic
research
or
that
pro-
moting
fundamental
science
should
negatively
impact
bud-
gets
for
applied
research.
We
argue,
however,
that
basic
science
is
at
the
foundation
of
ecology
and
requires
active
support
if
it
is
to
function
at
the
highest
level
and
continue
to
create
intellectual
capital
in
the
future.
This
support
encompasses
project
funding,
but
also
endorsement
by
academics,
governments,
and
society
as
a
whole.
Support
includes
fostering
the
building
blocks
of
the
ecological
sciences
such
as
taxonomy,
as
exemplified
by
the
Global
Taxonomy
Initiative
(http://www.cbd.int/gti/default.sht).
It
is
easy
to
depict
a
caricatured
vision
of
the
world
a
century
from
now,
should
fundamental
ecology
not
gain
greater
support.
Acquired
knowledge
and
understanding
would
probably
be
put
to
good
use,
through
high-level
engineering,
but
this
recycling
of
science
would
likely
lead
to
fewer
breakthroughs
and
fewer
challenges
to
existing
paradigms.
Intellectual
capital
would
be
‘consumed’
faster
than
it
is
replaced
[16].
It
might
be
a
world
where
ecological
tinkerers
would,
perhaps
brilliantly,
build
on
the
current
foundations
of
science,
but
those
foundations
would
cease
to
be
developed
or
would
develop
more
slowly,
with
associ-
ated
risks
of
failure
to
face
future,
novel
challenges.
To
explain
the
living
world
around
us,
we
need
to
meet
the
intellectual
challenge
of
understanding
life
in
complex,
changing
environments;
to
this
end,
fundamental
ecology
is
our
most
fundamental
instrument.
Acknowledgements
This
paper
stemmed
from
talks
presented
at
the
INTECOL
Symposium
‘Emphasizing
the
importance
of
basic
science
in
ecology’.
The
authors
thank
John
Harte,
Hal
Mooney,
Stephen
Pacala,
Jens
Rolff,
Sam
Scheiner,
and
Alan
Hastings
for
their
comments
and
discussions,
the
Socie
´te
´Franc¸aise
d’Ecologie
for
funding
this
symposium,
and
grants
from
the
Biodiversa
EraNet
(FFII)
to
F.C.,
the
National
Science
Foundation
(CNH-1313830)
to
J.A.D.,
Eranet
Netbiome
(ISLAND-BIODIV),
the
Fondation
pour
la
Recherche
sur
la
Biodiversite
´(CESAB-ISLANDS),
and
the
Laboratoire
d’Excellence
TULIP
(ANR-10-LABX-41)
to
C.T.,
and
Opinion Trends
in
Ecology
&
Evolution
January
2015,
Vol.
30,
No.
1
14

the
Wissenschaftskolleg
zu
Berlin,
the
James
S.
McDonnell
Foundation
(220020294),
and
the
CNRS
(PlCS05313)
to
M.E.H.
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