Plant strategies

A life history strategy is how an organism decides to use the energy it has
available for different life processes.
Plant strategies include the mechanisms and responses plants use to reproduce,
defend, survive, and compete in the landscape. Although the term "plant
strategy" has existed in the literature since at least 1965, multiple definitions
exist

Strategic response to competition, disturbance and stress

The concept that living organisms display ecological 'strategies' has advanced
rapidly in recent years. The r-K continuum of MacArthur and Wilson (1967) made an
important early contribution in contrasting the opportunistic r-species (with rapid
rates of population growth), which exploit temporary habitats, with the equilibrium K-
species of stable habitats in which competitive ability and survival of the individual is
more important than population growth. In the same way that differing species of
organism have gradually evolved over long periods of time (Darwin, 1859), such
strategies arose through the exertion of competitive natural selection upon varied
populations in which differences from the previous norm constantly arose. The CSR
model of Grime (1974,1979) -standing for Competitor-Stress tolerator-Ruderal -
made an important further advance in adding the stress-tolerators, organisms
capable of exploiting continuously unproductive environments or niches. The
competitors are equivalent to the K-species and the ruderals (living on disturbed
sites) approximate the r-species. This theory also recognizes that in plants there is a
separation of the established (adult) and regenerative (juvenile) strategies and they
may respond differently to their environment. This theory has been very thoroughly
applied over a long period of time in a number of ecosystems, and with the
publication of Plant Strategies, Vegetation Processes and Ecosystem Properties
(Grime, 2001), is becoming a major tool in the manipulation of vegetation and
ecological prediction.

The CSR triangle places plants into three categories based on their tolerance to stress and
disturbance, and they can be defined as competitors, stress-tolerators, and ruderals (CSR).
COMPETITORS are perennials that tend to take over where there is little stress or
disturbance.

Stress and disturbance are the two main external factors limiting the amount of living
and dead plant material in a habitat. Stress in this context consists of the external
constraints limiting the rate of growth (productivity, i.e. dry-matter production) of all or
part of the vegetation, including shortages of light, water and mineral nutrients and
the influence of suboptimal temperature.
Disturbance limits plant biomass by causing its partial or total destruction through
trampling, mowing, ploughing, the felling of trees, and the activities of pathogens and
other herbivores. It also results from wind damage, frost, drought, soil erosion and
fire.

Though no plants can long survive both high stress and high disturbance, there are
three combinations of these factors under which plants continue to exist. These have
led to the evolution of the three primary ecological strategies encapsulated in the
name CSR. Grime (2001) lists 18 ways in which plants possessing these strategies
differ from each other. These involve morphology, physiology, life history, palatability
to unspecialized herbivores (Section 5.7) and litter production. If both stress and
disturbance are absent then the composition of a plant community is determined
by competition between species. i.e. it is made up of competitors (C). Most trees are
competitors, exploiting conditions of low stress and low disturbance. Stress-
tolerators (S) endure high stress and low disturbance such as dog violet Viola
riviniana in the UK, growing in acidic, damp, shady woodlands, while ruderals (R)
such as annual meadow-grass Poa annua and Funaria hygrometrica (a moss
common in woods, often on recently burnt land) grow under conditions of high stress
and low disturbance. Competitors include herbs, shrubs and trees, whereas ruderals
are herbs or mosses. Stress-tolerators include various trees, shrubs, herbs and
particularly lichens (see Fig. 4.1).

The definition of competition adopted by Grime (1979) in the first edition of his well-
known book, is that it is the tendency of neighbouring plants to utilize the same
quantum of light, ion of mineral nutrient, molecule of water or volume of
space. Competition between plants occurs both above and below ground,
competition for light becoming particularly strong as the canopy begins to close so
that shoots of one plant are shaded by those of another (Section 1.4.2). Competitive
ability is a function of the area, the activity, and the distribution in space and time of
the surfaces through which resources are absorbed and as such it depends upon a
combination of plant characteristics. Very many plants possessing all the following
four features are extremely competitive:

(1) tall stature;

(2) a growth form which allows extensive and intensive exploitation of the
environment above and below ground, such as a densely branched rhizome as in
stinging nettle Urtica dioica and creeping soft-grass Holcus mollis, or an expanded
tussock structure as in tufted hair-grass Deschampsia cespitosa (of which a form is
frequent in the heavy soils in the oak woods of southern England);

(3) a high maximum potential relative growth rate (RGR); and

(4) a tendency to deposit a dense layer of litter on the ground surface.

Figure 4.1 Model describing the various equilibria between competition, stress and
disturbance in vegetation and the location of primary and secondary strategies. C,
Competitor; S, stress-tolerator; R, ruderal; C-R, competitive-ruderal; S-R, stress-
tolerant ruderal; C-S, stress-tolerant competitor; C-S-R,

C-S-R strategist. Ic, relative importance of competition (-); Is, relative importance of
stress ( ); Id, relative importance of disturbance

(-----).The strategic range of three life forms is also shown (a) trees and shrubs (b)
bryophytes and (c) lichens. (Redrawn from Grime et al., 1988. Comparative Plant
Ecology. Unwin Hyman.)

To represent these strategies, Grime et al. (1988) have developed a triangular

diagram whose points are the maxima of competition, stress and disturbance (Fig.
4.1). It is here that the three primary strategies are situated. The position of a
particular species is determined in relation to the indices of competition (Ic), stress

(Is) and disturbance (7d). The four main types of secondary strategy (C-R, C-S, S-R
and the central C-S-R) occur in intermediate positions. Grime et al. (1988) derive the
position of a particular species by using a dichotomous key involving various
morphological, behavioural and reproductive characteristics. Each strategy (also
called a functional type) can be represented within the triangular diagram by a set of
C, S and R co-ordinates. These co-ordinates relate to a whole set of attributes that
contribute to a species' ability to survive under given conditions of productivity and
disturbance.

Figure 4.2 illustrates the strategies employed by some common species of the UK
woodland field layer. Woodlands are usually stable; indeed the only ruderal present
in the main figure is common chickweed Stellaria media which is occasionally
encountered at disturbed woodland margins. Like other vegetation dominants, the
stinging nettle Urtica dioica can generate high shoot thrust, pushing aside foliage of
other species and resisting physical displacement. The leaves of such perennial
competitors, capable of rapid growth, also have high nitrogen contents; this
coincides with high concentrations of the enzyme 1,5-biphosphate
carboxylase/oxygenase (Rubisco) which appear to facilitate rapid rates of
photosynthesis in shady conditions. This latter feature is also aided by foliar
phosphorus concentrations that are much higher than in slow-growing species.
Shoots of stinging nettle die back completely in winter so associates such as the
annual competitive-ruderal goosegrass Galium aparine, low-growing rough meadow-
grass Poa trivialis (Fig. 4.2) and the moss Brachythecium rutabulum, which climbs
stems of other plants, are able to exploit relatively high light intensities in spring and
autumn.

Though the main focus of attention in earlier studies of plant strategies, and indeed
of Grime (1979), was that of life-history traits (i.e. the strategies of life-history events
such as germination, early survival, seed production and longevity), it is now
apparent that the primary strategies outlined above also concern other attributes
(resource capture and utilization, tissue chemistry and life-span, anti-herbivore
defence, rates of decomposition). As Grime (2001) remarks these 'have obvious and
direct connections to the functioning of ecosystems'; much of his second edition is
designed to establish the nature and usefulness of these connections.

4.1.2 Influence of forest clearance in Prince Edward Island, Canada

The extensive survey made by Sobey (1995a, b) of the vegetation of Prince Edward
Island in eastern Canada is of particular interest in its demonstration of the very long
period required for a succession to culminate in climax hardwood forest here. Some
reproductive strategies are exceedingly long term! The
Figure 4.2 C-S-R ordination diagram of woodland field layer species in the UK. This
describes the various equilibria between competition, stress and disturbance in
vegetation. The point shown for each species indicates the position of its maximum
percentage occurrence in a matrix of vegetation types classified according to the
strategies of the component species in Grime et al. (1988). Where points are not
provided the data for the species concerned are insufficiently clear. The small
diagram shows the strategic range of herbs. (From Packham and Cohn, 1990.
Arboricultural Journal 14.)

Figure 4.2 C-S-R ordination diagram of woodland field layer species in the UK. This
describes the various equilibria between competition, stress and disturbance in
vegetation. The point shown for each species indicates the position of its maximum
percentage occurrence in a matrix of vegetation types classified according to the
strategies of the component species in Grime et al. (1988). Where points are not
provided the data for the species concerned are insufficiently clear. The small
diagram shows the strategic range of herbs