Documentos de Académico
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INVITED PERSPECTIVE
Ch. Körner
present very fundamentally depend on these conditions, high mountains nor deserts are stressful for those natu-
while single individuals in any particular community rally living there, contrary to common belief (for in-
will almost always be operating a long way from their stance Callaway 2002). Stressed plants are commonly
growth-physiological maximum. The tundra vegetation, those which are unfit for a given situation, as may be
as we envisage it, is not limited by nutrients, though its humans who are visiting such places, hence the anthro-
biomass production is most likely nutrient-limited pocentric description of such conditions as 'hostile'. As
(Shaver & Chapin 1980). Alpine vegetation is not cold- with limitation, understanding the physiological mecha-
limited; it is low-temperature dependent, and warming nisms by which plants cope with stress is an important
can exert a threat to those plants, and most alpine species part of ecological science (e.g. Larcher 1987; Buchanan
die in a lowland climate. In fact alpine vegetation in 2000), but treating those conditions as adverse for the
temperate-zone mountains is not much less productive relevant community of species reflects a misconcep-
than a tropical rain forest, if the appropriate time reference tion. Fortunately the global change debate allows such
is chosen (e.g. per day of the growing season; Körner insight to slowly seep in. Warming the ‘cold-stressed’
1998, 1999). alpine is suddenly treated as a risk (a new stress?) rather
The ability to survive under exposure to specific than a relief for those ‘poor stressed creatures’.
environmental regimes can be achieved by: (1) evolu- Plants inhabiting such environmentally-demanding
tionary (phylogenetic) adaptation, (2) non-inherent, per- places may not thrive, but they profit from their exclusive
haps ontogenetic modifications, during the life of an capabilities – some more, others less – hence the
individual (or of its modules, such as leaves or tillers), or possibility of grouping plant species by such abilities
(3) reversible adjustment, often termed ‘acclimation’. (Grime 1989).
If, by any of these adaptive mechanisms, a plant ac-
quires the ability to grow and reproduce, it is fit. Natural “Stressful environments hardly exist”
selection usually sieves for fitness (Turesson 1925), but
not every trait is the result of selection for optimality of How can one claim that plants were limited, if the
a specific function (Grubb 1989). removal of the limitation terminates their existence?
Biomass production, irrespective of the species involved,
What is stress ? is almost always limited by one or several resources or
stressors, but the suite of taxa present in a given space
Stress is, in part, a synonym for resource limitation and time is not. This distinction, between the physio-
(e.g. water shortage), and in part it refers to physical logical- (process) or agronomic- (yield-) oriented
impacts (temperature, mechanical forces). It is not the meanings of limitation and stress, and the biodiversity-
diverging opinions about the term stress, as such, which oriented dimension of those terms, is crucial.
concern me, (see the refreshing debate in Grime 1989), Rather than teaching our students the “limited, stress-
but the commonly un-accounted change of its meaning ful nature of plant life’, it may be more fruitful to
with severeness. If any deviation of life conditions from explain these constraints of growth as vital elements for
those considered optimal for biomass increment is treated existence, coexistence and fitness. I feel increasingly
as stress, then we have just invented a new word for uncomfortable reading about ‘stressful environments’.
‘life’ and the term ‘stress’ becomes useless. Unfortu- They hardly exist for those webs of life which cope with
nately this is a very common definition of stress, even in them day by day, as stressful as these conditions may be
textbooks. to some isolated individuals involved. Deviation from
Deviations from physiological optimality are nor- physiological optimality is normal life at most places on
mal. They occur always, day by day. If we were able to earth. In our experiments we thus need to be alert for the
prevent these divergences from optimality for growth, presence of such deviations and the many genetic
we would remove all environmental conditioning and solutions that exist to cope with them. Hence, biolo-
adaptive ‘training’, and would rather rapidly ‘ruin’ the gically-relevant diversity, resource limitation and ‘stress’
affected organism. need to be made a part of experiments rather than being
So, the term, ‘stress’, is better restricted to extreme excluded from tests of responses to one specific envi-
situations. But what is an extreme? Once the ability to ronmental driver, be it climatic warming, nitrogen depo-
cope with environmental extremes has evolved, such sition, atmospheric CO2 enrichment, or the interaction
extremes become elements of ‘normal’ life (Körner with alien organisms, to name a few, if we want to make
1998). If we move plants to what, from our human these tests ecologically meaningful (Golley 1993;
perspective, might be a less stressful environment, Buchanan 2000; Körner 2001; Shaw et al. 2002).
most specialist plants would either die, or be sup-
pressed by species native to the new habitat. Neither
- Limitation and stress – always or never ? - 143