TREE vol. 13, no. 4 April 1998 133
known about the pollinators, seed disper-
sal, seed banks or gene flow for the plants.
However, she reported that most plans in-
clude the gathering of more information
about pollinators, breeding system and
seed dispersal. The number of plans listing
knowledge of pollinators as a high-priority
task has increased significantly. While this
is encouraging, no plan included manage-
ment of pollinators in the recovery of the
endangered species.
All participants agreed that plant–pol-
linator interactions will often be important
in the restoration or re-establishment of
damaged ecosystems. Direct effects of de-
graded pollination systems are often easily
recognized and are receiving more atten-
tion from restoration and conservation bi-
ologists. It is the less obvious effects of de-
graded pollination systems on population
and genetic structure that are in need of
more attention. Perhaps most importantly,
we need to shift the emphasis in resto-
ration biology from individual species to
community interactions and ecosystem
function.
Paul R. Neal
Dept of Biology, Cox Science Building,
University of Miami, Coral Gables,
FL 33124, USA
(pneal@fig.cox.miami.edu)
NEWS & COMMENT
Copyright © 1998, Elsevier Science Ltd. All rights reserved. 0169-5347/98/$19.00 PII: S0169-5347(97)01293-7
O
ver the past several years, ecologists
have begun to recognize that the role
played by positive plant–plant interac-
tions in communities is underappreciated.
In a recent paper in Oikos
1
, Brooker and
Callaghan offer an interesting perspective
and new insights to this ongoing cathar-
sis. They argue that plant interactions are
best characterized as a balance between
positive and negative components and that
this balance is tipped in favor of net posi-
tive or negative effect by environmental
factors. This argument has also recently
been made by Callaway and Walker
2
and
Holmgren, Scheffer and Huston
3
. Clearly,
this is an idea whose time has arrived.
Brooker and Callaghan’s perspective on
this balance, however, is unique in that
they see the issue through the eyes of arc-
tic plant ecologists, and envision physical
disturbance rather than environmental
stress as the trigger that shifts the balance
between positive and negative effects.
Brooker and Callaghan begin by noting
the lack of attention paid to positive plant–
plant interactions by ecologists over the
past three decades, and argue that a num-
ber of recent studies suggest that positive
interactions among plant neighbors are
commonly important. They then observe
that recent examples of positive interac-
tions playing an important role in commu-
nity structure have two things in common.
First, most examples are from relatively
harsh physical environments. Second, the
environmental factor constraining plant
growth or survival in these examples can
be alleviated by the physical presence of
other plants. They then argue that posi-
tive components in plant neighbor inter-
actions are probably common in all habi-
tats, and that, in general, plant interactions
are best characterized as a balance be-
tween positive and negative components.
Fine so far.
The paper then takes a bit of a twist.
They define environmental severity as a
combination of both physical stress and
disturbance (following Grime
4
) and con-
cede that since the relationship between
the roles played by competition and physi-
cal stress in structuring communities is
contentious, they will focus instead on the
role played by disturbance in tipping the
balance between positive and negative
plant neighbor effects. This is perhaps
surprising, since they had just made such
a good case for a strong relationship be-
tween positive interactions and physical
stress. Indeed, few of their examples were
unambiguously associated with disturb-
ance or independent of physical stress.
Brooker and Callaghan suggest that
while it is widely accepted that increased
disturbance reduces the intensity and im-
portance of competition in plant commu-
nities, as a by-product of reducing the level
of competition, disturbances must increase
the importance and intensity of positive
neighbor interactions. They argue that
since disturbance decreases the intensity
of competition and that plant–plant inter-
actions are a direct balance between nega-
tive and positive effects, disturbance must
increase the relative importance and in-
tensity of positive interactions. (But surely
any balance between negative, competitive
and positive, ameliorating components of
interactions must have a mechanistic ba-
sis? Disturbances remove competitors, but
only remove cooperators if there is some-
thing to cooperate about.) They then ar-
gue that recent examples of positive inter-
actions in plant communities are driven
by environmental severity which includes
disturbance. In their discussion of these
examples, however, physical stress al-
leviation implicitly remains prominent.
They conclude that ‘community manipu-
lation experiments are more likely to reveal
positive plant–plant interactions in highly
disturbed communities … and as disturb-
ance is reduced either through space or
time, competition becomes more impor-
tant and in parallel, positive interactions
become less important, so that competition
becomes the dominant selective force in
plant–plant interactions’.
The paper closes strongly with an in-
teresting and thoughtful discussion of the
selective impact of positive plant interac-
tions drawn primarily from their work on
arctic plant communities. Decreased seed
dispersal distances, compact plant mor-
phologies, limited responses to improved
conditions, increased frequency of mycor-
rhizal associations and increased domi-
nance of clonal plants are all suggested to
be traits selected for in extreme environ-
ments where plants can ameliorate the
harsh conditions.
This is an interesting, thought-
provoking, if sometimes perplexing paper.
Brooker and Callaghan challenge ecolo-
gists to get a better grasp on the role
played by positive interactions in commu-
nities, and correctly, I think, characterize
plant–plant interactions as a balance be-
tween negative and positive components.
Their examples, and discussion, however,
are more consistent with physical stresses
that are ameliorated by plant cover trig-
gering the dominance of positive interac-
tions in communities than disturbances
triggering a shift to net positive effects
5
.
I suspect that their views are strongly in-
fluenced by the arctic plant communities
they study where plants can protect one
another from disturbance, but that disturb-
ances also typically create stressful physi-
cal conditions that plant cover can amelio-
rate. In physically benign habitats, where
disturbances lead to safe sites and intense
competition among recruits, their disturb-
ance-based model should break down. In
focusing on disturbance and simplicity,
their discussion also entirely ignores con-
sumers and the prominent positive roles
played by plant neighbors and associ-
ational defenses
6,7
. Any realistic model of
the role played by direct positive interac-
tions in the structure of plant communities
Searching for the role of positive
interactions in plant communities