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Fundamental ecology is fundamental

Article  in  Trends in Ecology & Evolution · November 2014

DOI: 10.1016/j.tree.2014.11.005

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 Opinion

Fundamental ecology is fundamental

Franck Courchamp1,2, Jennifer A. Dunne3, Yvon Le Maho4, Robert M. May5,
Christophe Thébaud6, and Michael E. Hochberg3,7,8,9
1
Laboratoire d’Ecologie,Systématique, et Evolution, UMR CNRS 8079, Université 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, Université de Strasbourg, Strasbourg, France
5
Department of Zoology, University of Oxford, Oxford OX1 3PS, UK
6
Laboratoire Ecologie et Diversité Biologique, UMR CNRS 5174, Université Paul Sabatier, Toulouse, France
7
Institut des Sciences de l’Evolution, UMR 5554, Université 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- research and Figure 1 presents a conceptual model of their
search are satisfying curiosity, acquiring knowledge, relationships.
and achieving understanding. Here we develop why One of the central objectives and achievements of fun-
we believe it is essential to promote basic ecological damental ecology is to develop and test general theory in
research, despite increased impetus for ecologists to ecology [3]. At the broadest level, a general theory is an
conduct and present their research in the light of poten- entire domain of science and a set of interwoven funda-
tial applications. This includes the understanding of our mental principles, like the theory of evolution by natural
environment, for intellectual, economical, social, and selection [4].
political reasons, and as a major source of innovation. Fundamental research is sometimes called pure science
We contend that we should focus less on short-term, or ‘blue-skies’ research. The term blue-skies research has
objective-driven research and more on creativity and its roots in the idea of curiosity- and inquiry-driven studies
exploratory analyses, quantitatively estimate the bene- (Box 1). It is said that under Eisenhower’s presidency a
fits of fundamental research for society, and better ex- prominent politician unsympathetic to basic research
plain the nature and importance of fundamental ecology claimed ‘I don’t care what makes the grass green!’, which
to students, politicians, decision makers, and the general was later rephrased as ‘what makes the sky blue’ [5]. In-
public. Our perspective and underlying arguments terestingly, 50 years earlier, using vapours and light
should also apply to evolutionary biology and to many beams in glass tubes, Tyndall [6] explained the basis of
of the other biological and physical sciences. the sky’s colour and his work led to better and more
effective processes and products unforeseen at the time
What is fundamental ecology? of his discovery. These included a test for optically pure air,
Fundamental ecology, or basic ecology, is the study of support for the nonexistence of spontaneous generation,
organismal diversity and of the interactions between particle filtering of lung airways, mould destruction by
organisms and their abiotic and biotic environments Penicillium bacteria, and even the precursors of the flexi-
[1]. Its main goal is to advance knowledge and understand- ble gastroscope and bronchoscope [6].
ing, and its results, even if sometimes predictable, are not Financial support for fundamental research in ecology,
known with certainty in advance. By contrast, applied like in many other disciplines, is highly competitive. Many
ecology is usually motivated by particular, well-defined perceive stasis or a continuous decline in support, but
objectives, typically to solve environmental problems, in- accurate numbers are difficult to obtain to substantiate
cluding the management of natural resources such as land, general trends, since funding categorization can be open to
energy, food, or biodiversity [2]. Because applied ecology alternative interpretations and data can be difficult to
often involves the development of interventions to alter obtain. This perception that support, be it financial or
events (e.g., exotic species invasions, endemic species de- moral, is not improving is worrisome for ecologists, for
cline), this research is essentially an attempt to achieve a their science, and ultimately for society. Here we show
defined objective. Box 1 summarises the different types of how the predominant support for objective-driven, applied
science is relatively recent and why, in conjunction with
Corresponding authors: Courchamp, F. ([email protected]); Hochberg,
this, fundamental research has seen limited investment in
M.E. ([email protected]). both relative and absolute terms. We then advocate in-
Keywords: applied ecology; basic ecology; blue-skies research; fundamental research; creased and less constraining support for basic ecology, by
research priorities.
presenting the principal drivers of fundamental research
0169-5347/ and some of its main benefits, including the satisfaction of
ß 2014 The Authors. Published by Elsevier Ltd. This is an open access article under
the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/3.0/). human curiosity, the quest to explain our world, the crea-
http://dx.doi.org/10.1016/j.tree.2014.11.005 tion of scientific knowledge, and the often unintentional,
Trends in Ecology & Evolution, January 2015, Vol. 30, No. 1 9
Opinion Trends in Ecology & Evolution January 2015, Vol. 30, No. 1

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

15.44%
n
drive
iosity
r
Cu
Fundamental
Discoveries

Innovaon

n
Applied

d ri ve
P ro d
Development

uc

on
u
27.03%

on
dr

ol
ive S
n

49.81%

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.

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

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

Box 2. Examples of basic research in ecology leading to environmental applications

In many areas of applied ecology, the urgency of problems such as to the questioning of the basic assumption that Allee effects are an
habitat destruction or species extinction often calls for a rapid intrinsic characteristic of populations and therefore cannot be created
response. Conservation science is unique in being both a field of by human activities. Thus, the standard paradigm was that humans
research and a field of action, but environmental challenges have could only drive populations down to sizes where a pre-existing,
naturally shifted emphasis towards action, sometimes at the expense unexpressed Allee effect would be activated. Questioning this
of building a strong fundamental framework. For example, a few premise led to the discovery of possible anthropogenic Allee effects,
decades ago biological invaders on islands were typically ignored through which human activities can exert inverse density-dependent
until their impact became a decisive factor for intervention, which was exploitation. According to this concept, the arbitrary value people
pursued without either pre-control survey or post-control monitoring attribute to rarity would confer an economic value on rare species that
[49]. Consequently, hysteresis and multiple stable states (including would maintain the incentive to exploit them, even at very high levels
extinction) were generally overlooked, resulting in a number of of rarity [63]. Rare species, being more valuable, would be more
‘surprise effects’, typically the failure of eradication, or the unex- exploited, thereby becoming even rarer, precipitating a vortex of
pected outbreak of other, hitherto neglected invasive species extinction. This process threatens, through many different wildlife-
[50]. Subsequently, incorporation of trophic-web theory allowed based markets, countless species of plants and animals.
assessments of mesopredator releases [51], hyperpredation pro- A final poignant illustration of the importance of basic science is the
cesses [52], competitor releases [53], and other ecological processes increasing need to promote ecosystem services, where possible
related to interspecific interactions and likely to interfere with or [64,65]. Only with an adequate understanding of the patterns and
facilitate eradication [54,55]. Similar observations apply to the underlying mechanisms of ecosystem functioning can one justifiably
management of fisheries, with long-lasting stock collapses [29,56], and reliably protect, restore, and value the services provided by that
algal blooms [57], eutrophication in freshwater and coastal marine ecosystem. Unfortunately, the current environmental crisis often elicits
[58,59], or regime shifts in terrestrial ecosystems [60]. rapid responses with little fundamental basis, in which avoidable (and
Another illustration comes from a fundamental programme aimed now irrecoverable) mistakes would have been averted. Recent
at better understanding the mechanisms underlying the Allee effect (a estimates of the global value of pollination for agriculture – US$153
positive relationship between the number of individuals in a billion annually – provides a good illustration of the importance of
population and their fitness [61,62]). Fundamental research has led better understanding the functioning and outputs of ecosystems [66].

on the stability and complexity of ecosystems shows future Promoting fundamental research
promise in the regulation of banks [42], while fundamental Ecologists can promote basic science in several important
studies in the behavioural ecology of social insects are ways.
being used in robotics [43]. First, we need to regularly assess the state of and
Nevertheless, that fundamental research often fuels progress made in fundamental ecology through the publi-
applications should not be the main argument for why cation of perspective, forum, review, and synthesis articles.
we need basic research. Unexpected applications of funda- In our view, one of the most influential ways to make both
mental discoveries are only providential byproducts of headway in specific areas and overall progress in the field
other objectives. We believe it is important that fundamen- is through discussing and recasting the most important
tal ecology is a major source of application, but the primary questions in fundamental ecology [44].
justifications for conducting fundamental research should Second, we must develop and draw on new approaches
remain firmly grounded in satisfying curiosity, acquiring to communicate results to politicians and granting agen-
knowledge, and achieving understanding. cies and to reassure them that giving liberty to scientists is

Box 3. Examples of basic research in ecology leading to applications in health sciences

Since each species potentially has unique biological properties, it is below ground [71]. These defences are regulated by conserved
unsurprising that an estimated 60% of antitumour and antimicrobial signalling pathways [72]. In another striking example, fundamental
drugs are of natural origin [67]. Basic research is at the origin, for studies of the microbial ecology of hot springs at Yellowstone
example, of the discovery in the naked mole rat (Heterocephalus National Park resulted in the discovery of hyperthermophiles, which
glaber) of anticancer mechanisms [68]. Taking full advantage of such incidentally led to the identification of the Taq polymerase, since then
a potential ‘gold mine’ would not have been possible without used in PCR [73]. This unexpected finding was key to innovations in
research on understanding the causes of mortality in this species agriculture and medicine and helped create the new field of
through necroscopies. Another example is the incubating male king biotechnology, which includes genomics, recombinant gene technol-
penguin (Aptenodytes patagonicus), which preserves fish in its ogies, applied immunology, and the development of pharmaceutical
stomach for up to 3 weeks to feed to its young at hatching, should therapies and diagnostic tests. These are just a few examples of how
the mother be absent when the juvenile begs for food [69]. Searching basic ecology can unexpectedly provide the molecular basis of
for the mechanism permitting fish conservation at 378C led to the important applications.
discovery of spheniscin, a small antimicrobial and antifungal peptide. Environmental health is another field where basic ecological
This molecule has since been shown to reduce the growth of the two science has repeatedly proved to be of major importance. With the
main agents of hospital-acquired infections, the bacterium Staphylo- development of mathematical models of host–parasite relationships
coccus aureus and the fungus Aspergillus fumigatus [70]. [74,75], ecological epidemiology has resulted in a better under-
Basic research on plant–insect interactions has also led to important standing of the spread of pathogens and parasites, paving the way for
applications, such as alternative strategies for the protection of crops applications to manage the spread of infectious diseases. Examples
like maize and rice against herbivores. Plants possess many defence include host–parasite modelling of epidemics such as measles, which
mechanisms based on secondary metabolites or defensive proteins subsequently informed control campaigns against the 2001 outbreak
whose toxic or deterring properties are harmful to herbivores. of foot and mouth disease in the UK [76–78]. Unfortunately, basic
However, it was not known until recently that some plants release science was not sufficiently developed (or not identified as such) to
volatile signals that attract herbivore natural enemies, such as realise its full potential in contributing to the control of the spread of
parasitic wasps above ground, and entomopathogenic nematodes avian (H5N1) and swine (H1N1) flu and the current Ebola epidemic.

13
Opinion Trends in Ecology & Evolution January 2015, Vol. 30, No. 1

not necessarily at the expense of accountability [6]. It is Finally, researchers themselves need to prioritise how
essential that funders realise that academic research pro- to effectively defend basic research. Politicians have long
grammes once thrived on intellectual freedom and that recognised that research free of practical constraints is at
this is essential for discovery [16]. Defining clearly what the heart of technology and industry. For example, Van-
ecology is and what ecologists are (i.e., not environmen- nevar Bush, scientific advisor to Franklin D. Roosevelt,
talists) is key in this regard. stated in 1945 that scientific progress resulted ‘from the
Third, we should consider creating programmes giving free interplay of free intellectuals, working on subjects of
large and unconstrained blue-sky research grants to prom- their own choice, in the manner dictated by their curiosity
ising or experienced researchers. Such initiatives obviously for exploration of the unknown’ [22]. Yet many politicians
should not be at the expense of other research programmes. still view basic science as an unaffordable luxury, especial-
Even a modest investment in supplementary, long-term ly in times of financial stress [9]. Young and more senior
grants, without externally imposed objectives, should con- ecologists must engage debate with policymakers to correct
tribute to considerable breakthroughs [16]. the common misconception that fundamental research is a
Fourth, we need to establish norms for funding funda- luxury [Dos Remedios, C. (2000) The value of fundamental
mental research that will facilitate adherence by agencies research. International Union for Pure and Applied Bio-
and promote the long-term future of fundamental science. physics (http://iupab.org/publications/value-of-fundamen-
One possibility is to establish an international norm of a tal-research/)] [9].
minimum level and/or percentage of research funding that
participating agencies pledge to maintain for fundamental Concluding remarks
science. Participating national, international, and private We have developed several lines of reasoning supporting
funding bodies would then officially declare that they the promotion of fundamental research in ecology, al-
adhere to international norms. This highlights the neces- though the same or similar arguments could apply to
sity that politicians/decision makers, agencies, and scien- evolutionary biology [18] and to other sciences. We empha-
tists all be involved in institutional-, national-, and sise that we do not claim that applied research has less
international-level discussion to establish such norms merit or intrinsic value than basic research or that pro-
and to determine the extent of their application. We stress moting fundamental science should negatively impact bud-
that promoting basic ecological research requires contin- gets for applied research. We argue, however, that basic
ued dialogue and need not result in reduced budgets for science is at the foundation of ecology and requires active
other research approaches (Box 1). support if it is to function at the highest level and continue
Fifth, we must find ways to quantitatively value funda- to create intellectual capital in the future. This support
mental findings. This would involve estimating the pay off encompasses project funding, but also endorsement by
and timescale of the returns of basic research, which, academics, governments, and society as a whole. Support
although known to be important [36,37], remain largely includes fostering the building blocks of the ecological
unquantified [16]. This could be one way of demonstrating sciences such as taxonomy, as exemplified by the Global
unambiguously the importance of fundamental research in Taxonomy Initiative (http://www.cbd.int/gti/default.sht).
ecology. It is easy to depict a caricatured vision of the world a
Sixth, we need to reduce the pressure on ecologists to century from now, should fundamental ecology not gain
systematically provide short-term results to predefined greater support. Acquired knowledge and understanding
questions. One approach would be to modify tenure rules would probably be put to good use, through high-level
to account for the importance of longer-term, less output- engineering, but this recycling of science would likely lead
oriented research. Another possibility is longer grant per- to fewer breakthroughs and fewer challenges to existing
iods, to promote the tackling of challenging and risky paradigms. Intellectual capital would be ‘consumed’ faster
questions with possible unexpected outcomes. Despite risk than it is replaced [16]. It might be a world where ecological
being increasingly presented as a positive criterion, the tinkerers would, perhaps brilliantly, build on the current
current mindset on productivity discourages risky projects. foundations of science, but those foundations would cease
Typically, reviewers are required explicitly to express their to be developed or would develop more slowly, with associ-
opinions on the expected impact of grant proposals, often in ated risks of failure to face future, novel challenges. To
terms of output quality and certainty [45,46]. Precluding explain the living world around us, we need to meet the
risk can constrain creativity and, therefore, discovery in intellectual challenge of understanding life in complex,
the longer term [47]. changing environments; to this end, fundamental ecology
Seventh, supervisors and mentors need to encourage is our most fundamental instrument.
their students to conduct side projects and pilot projects to
foster the spark of curiosity and creativity. Supervisors and
Acknowledgements
mentors should insist on the benefits of devoting time to
This paper stemmed from talks presented at the INTECOL Symposium
thinking, to exploratory research, and to the importance of ‘Emphasizing the importance of basic science in ecology’. The authors
‘toying’ with concepts and data (and thinking ‘outside the thank John Harte, Hal Mooney, Stephen Pacala, Jens Rolff, Sam
box’) rather than emphasising high scientific productivity Scheiner, and Alan Hastings for their comments and discussions, the
or being drawn into applied research on the sole basis of Société Française d’Ecologie for funding this symposium, and grants from
the Biodiversa EraNet (FFII) to F.C., the National Science Foundation
better funding opportunities. They should also insist on the
(CNH-1313830) to J.A.D., Eranet Netbiome (ISLAND-BIODIV), the
importance of asking difficult (but tractable) questions – Fondation pour la Recherche sur la Biodiversité (CESAB-ISLANDS),
that is, steeping out of one’s comfort zone. and the Laboratoire d’Excellence TULIP (ANR-10-LABX-41) to C.T., and

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Opinion Trends in Ecology & Evolution January 2015, Vol. 30, No. 1

the Wissenschaftskolleg zu Berlin, the James S. McDonnell Foundation 32 Lande, R. (1993) Risks of population extinction from demographic and
(220020294), and the CNRS (PlCS05313) to M.E.H. environmental stochasticity and random catastrophes. Am. Nat. 142,
911–927
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