Journal of Vegetation Science 14: 141-143, 2003
© IAVS; Opulus Press Uppsala.
- Limitation and stress – always or never ? -
INVITED PERSPECTIVE
Limitation and stress – always
or never ?
Ch. Körner
Institute of Botany, Schönbeinstrasse 6, University of
Basel, CH-4056 Basel, Switzerland
E-mail ch.koerner@unibas.ch
Few terms are as abundant and as confusing in the
ecological literature as 'limitation' and 'stress'. Limita-
tion is the more general term, and stress is a specific
(extreme) part of it. Limitation has its roots in economy
and in agriculture; stress has its roots in physics, and
was later adopted by psychologists, physiologists and in
medicine before it made its way to ecology.
What is limitation ?
‘Limitation’ is considered the alpha and omega of
ecology – the science of limiting resources and nature’s
house-keeping (Malthus 1798, cited in Grubb 1989).
Most often, ‘limitation’ is used as if there were general
agreement on what the target of its action is, and on how
the action arises. However, depending on what is
considered ‘limited’, plants may be either never, or
almost always limited, so the term is obscure.
Many people use ‘limitation’ while implicitly as-
suming limitation of certain metabolic processes or
growth. This is the classical meaning in plant physiology
and in agriculture (for example Hiler & Howell 1983),
where the term has, doubtless, its rightful place, referring
to sub-maximal rates of processes, or yields of biomass,
or certain biomass compounds (shortages can be in
mineral nutrients or water, but also in thermal energy or
light). A farmer can reduce such limitations for a specific
crop, and so enhance yield and income. The global
patterns of annual biomass production clearly reflect
such obvious growth limitations by resources (Field et
al. 1998).
As can easily be demonstrated, this same concept
loses its meaning when applied to plants in multispecies
communities in which species occur, driven by their
own preferences, abilities and biotic linkages, rather
than being sown or planted. As Kerner (1897, 1898)
141
Christian Körner has been Professor of Botany at the University
of Basel, Switzerland since 1989. He got his PhD in 1977 at
the University of Innsbruck, Austria. His research fields include
plant water and carbon relations, forest ecology, alpine ecology
and global change biology (atmospheric CO2-enrichment, links
between biodiversity and ecosystem functioning). Christian
Körner is a member of the steering committee of IGBP, chairs
the Global Mountain Biodiversity Assessment (GMBA) of
DIVERSITAS and acts as convening lead author of the moun-
tain chapter for the Millennium Assessment. His editorial
duties include the chief-editorship of Oecologia and member-
ship of the board of referees of the journal Science. Consult
http://www.unibas.ch./botschoen/koerner/
wrote of plants, “the reason for their living together lies
in the nature of the habitat”. If one changes the nature of
the habitat by adding a previously growth-limiting re-
source, the so-claimed ‘limited’ plants will commonly
disappear sooner or later, as a result of a cascade of
induced community processes. Watering the desert, fer-
tilizing the tundra, warming the alpine zone, cooling a
hot spring, or removing shade from an understorey
community, are commonly fatal for the plants living
there, creating a completely new assemblage of species,
with the former ‘limited’ ones being eliminated. Even
subtle changes in supply of relevant resources, such as
atmospheric CO2-enrichment, will exert such bio-
diversity effects (Körner 2000).
All this may sound trivial, not requiring any re-
emphasising, but it does not seem to be widely
appreciated, hence the many unspecific statements on
the limitations of biota to be found in the literature. A
given assemblage of organisms is nature’s answer to a
certain combination of growth limitations. The species
142 Körner, Ch.
present very fundamentally depend on these conditions,
while single individuals in any particular community
will almost always be operating a long way from their
growth-physiological maximum. The tundra vegetation,
as we envisage it, is not limited by nutrients, though its
biomass production is most likely nutrient-limited
(Shaver & Chapin 1980). Alpine vegetation is not cold-
limited; it is low-temperature dependent, and warming
can exert a threat to those plants, and most alpine species
die in a lowland climate. In fact alpine vegetation in
temperate-zone mountains is not much less productive
than a tropical rain forest, if the appropriate time reference
is chosen (e.g. per day of the growing season; Körner
1998, 1999).
The ability to survive under exposure to specific
environmental regimes can be achieved by: (1) evolu-
tionary (phylogenetic) adaptation, (2) non-inherent, per-
haps ontogenetic modifications, during the life of an
individual (or of its modules, such as leaves or tillers), or
(3) reversible adjustment, often termed ‘acclimation’.
If, by any of these adaptive mechanisms, a plant ac-
quires the ability to grow and reproduce, it is fit. Natural
selection usually sieves for fitness (Turesson 1925), but
not every trait is the result of selection for optimality of
a specific function (Grubb 1989).
What is stress ?
Stress is, in part, a synonym for resource limitation
(e.g. water shortage), and in part it refers to physical
impacts (temperature, mechanical forces). It is not the
diverging opinions about the term stress, as such, which
concern me, (see the refreshing debate in Grime 1989),
but the commonly un-accounted change of its meaning
with severeness. If any deviation of life conditions from
those considered optimal for biomass increment is treated
as stress, then we have just invented a new word for
‘life’ and the term ‘stress’ becomes useless. Unfortu-
nately this is a very common definition of stress, even in
textbooks.
Deviations from physiological optimality are nor-
mal. They occur always, day by day. If we were able to
prevent these divergences from optimality for growth,
we would remove all environmental conditioning and
adaptive ‘training’, and would rather rapidly ‘ruin’ the
affected organism.
So, the term, ‘stress’, is better restricted to extreme
situations. But what is an extreme? Once the ability to
cope with environmental extremes has evolved, such
extremes become elements of ‘normal’ life (Körner
1998). If we move plants to what, from our human
perspective, might be a less stressful environment,
most specialist plants would either die, or be sup-
pressed by species native to the new habitat. Neither
high mountains nor deserts are stressful for those natu-
rally living there, contrary to common belief (for in-
stance Callaway 2002). Stressed plants are commonly
those which are unfit for a given situation, as may be
humans who are visiting such places, hence the anthro-
pocentric description of such conditions as 'hostile'. As
with limitation, understanding the physiological mecha-
nisms by which plants cope with stress is an important
part of ecological science (e.g. Larcher 1987; Buchanan
2000), but treating those conditions as adverse for the
relevant community of species reflects a misconcep-
tion. Fortunately the global change debate allows such
insight to slowly seep in. Warming the ‘cold-stressed’
alpine is suddenly treated as a risk (a new stress?) rather
than a relief for those ‘poor stressed creatures’.
Plants inhabiting such environmentally-demanding
places may not thrive, but they profit from their exclusive
capabilities – some more, others less – hence the
possibility of grouping plant species by such abilities
(Grime 1989).
“Stressful environments hardly exist”
How can one claim that plants were limited, if the
removal of the limitation terminates their existence?
Biomass production, irrespective of the species involved,
is almost always limited by one or several resources or
stressors, but the suite of taxa present in a given space
and time is not. This distinction, between the physio-
logical- (process) or agronomic- (yield-) oriented
meanings of limitation and stress, and the biodiversity-
oriented dimension of those terms, is crucial.
Rather than teaching our students the “limited, stress-
ful nature of plant life’, it may be more fruitful to
explain these constraints of growth as vital elements for
existence, coexistence and fitness. I feel increasingly
uncomfortable reading about ‘stressful environments’.
They hardly exist for those webs of life which cope with
them day by day, as stressful as these conditions may be
to some isolated individuals involved. Deviation from
physiological optimality is normal life at most places on
earth. In our experiments we thus need to be alert for the
presence of such deviations and the many genetic
solutions that exist to cope with them. Hence, biolo-
gically-relevant diversity, resource limitation and ‘stress’
need to be made a part of experiments rather than being
excluded from tests of responses to one specific envi-
ronmental driver, be it climatic warming, nitrogen depo-
sition, atmospheric CO2 enrichment, or the interaction
with alien organisms, to name a few, if we want to make
these tests ecologically meaningful (Golley 1993;
Buchanan 2000; Körner 2001; Shaw et al. 2002).
 - Limitation and stress – always or never ? -
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