Arid environments are characterized by limited and variable rainfall that supplies resources in pulses. Resource pulsing is a special form of environmental variation, and the general theory of coexistence in variable environments suggests specific mechanisms by which rainfall variability might contribute to the maintenance of high species diversity in arid ecosystems. In this review, we discuss physiological, morphological, and life-history traits that facilitate plant survival and growth in strongly water-limited variable environments, outlining how species differences in these traits may promote diversity. Our analysis emphasizes that the variability of pulsed environments does not reduce the importance of species interactions in structuring communities, but instead provides axes of ecological differentiation between species that facilitate their coexistence. Pulses of rainfall also influence higher trophic levels and entire food webs. Better understanding of how rainfall affects the diversity, species composition, and dynamics of arid environments can contribute to solving environmental problems stemming from land use and global climate change.

I Whether or not competition intensity increases or is similar along productivity gradients has been highly controversial for a number of years, but empirical results bearing on this question are quite variable and no consistent answer has yet emerged. We have developed a more general hypothesis that includes these contradictory predictions as special cases that apply under different types of resource dynamics and different types of interactions between the growth and survival components of fitness. 2 The two-phase resource dynamics hypothesis of plant interactions along productivity gradients is based on the fact that soil resources are usually supplied in pulses rather than continuously, as assumed by most formal theory for community dynamics of terrestrial plants. When soil resource supply is temporally variable, individual plants will experience two distinct phases of resource availability: pulse periods when resources are high and most growth and resource accumulation occurs, and interpulse periods when resources are too low for most plants to take up and most mortality due to resource deficits takes place. 3 Competitive effects on growth should occur during pulses at both high and low productivity. In productive environments, interpulse intervals should be relatively mild and infrequent and therefore competitive effects during pulses will usually be important for individual and population persistence. However, as productivity decreases, the frequency of pulses (as well as or in place of their magnitude) often decreases and the duration of interpulse periods increases. Therefore, we suggest that processes occurring during interpulse intervals become increasingly important for individual and population persistence as interpulse intervals become longer. Whether or not competition occurs under low productivity will then depend on (i) the extent to which the asymptotically low resource availability during interpulse periods is determined by plant uptake or by abiotic factors such as leaching, drainage, evaporation and volatization, and (ii) the extent to which decreased growth due to competition during pulses results in decreased survival during interpulse periods. 4 According to the two-phase resource dynamics hypothesis, Grime's hypothesis that competition is unimportant at low productivity will hold when soil resource availability between pulses in unproductive environments is controlled by abiotic factors and when survival during interpulse intervals is independent of or even negatively correlated with growth during pulse periods. In contrast, Newman's and Tilman's hypothesis that competition is equally important along productivity gradients will apply when either of these conditions is not true. We predict that the conditions for Grime's hypothesis to apply are more likely for productivity gradients driven by water than by mineral nutrients and when response to competition is measured for community structure or individual survival rather than for individual growth. 5 We tested the predictions about water vs...

Plants are limited in their ability to choose their neighbours, but they are able to orchestrate a wide spectrum of rational competitive behaviours that increase their prospects to prevail under various ecological settings. Through the perception of neighbours, plants are able to anticipate probable competitive interactions and modify their competitive behaviours to maximize their long-term gains. Specifically, plants can minimize competitive encounters by avoiding their neighbours; maximize their competitive effects by aggressively confronting their neighbours; or tolerate the competitive effects of their neighbours. However, the adaptive values of these non-mutually exclusive options are expected to depend strongly on the plants' evolutionary background and to change dynamically according to their past development, and relative sizes and vigour. Additionally, the magnitude of competitive responsiveness is expected to be positively correlated with the reliability of the environmental information regarding the expected competitive interactions and the expected time left for further plastic modifications. Concurrent competition over external and internal resources and morphogenetic signals may enable some plants to increase their efficiency and external competitive performance by discriminately allocating limited resources to their more promising organs at the expense of failing or less successful organs.

Summary 1Competition usually involves the allocation of limiting resources to non-reproductive functions. Natural selection is expected to favour mechanisms that increase competition with non-self neighbours and limit wasteful competition with self. 2 We used Pisum sativum plants that had two roots and 'double plants' with two shoots and two roots that could be either longitudinally separated into two genetically identical but physiologically distinct individuals or left intact. 3 Root development was significantly greater in split-root plants whose neighbours belonged to different plants. Furthermore, root development was relatively greater in the presence of roots of a different plant, regardless of its identity. This discrimination had a vectorial component whereby plants developed more and longer lateral roots towards neighbouring roots of different plants than towards other roots of the same plant.4 The results thus demonstrate a mechanism of avoiding self-competition that is based on physiological co-ordination among different organs of the same plant rather than on allochemical recognition that depends on genetic differences. 5 The ability to discriminate between self and non-self could be expected to increase resource use efficiency and ecological performance in plants. It could also be expected that tight physiological co-ordination will be selected for between organs of the same plant that have greater probability of being engaged in direct competitive interactions with each other.

Recent evidence suggests that self͞non-self discrimination exists among roots; its mechanisms, however, are still unclear. We compared the growth of Buchloe dactyloides cuttings that were grown in the presence of neighbors that belonged to the same physiological individual, were separated from each other for variable periods, or originated from adjacent or remote tillers on the same clone. The results demonstrate that B. dactyloides plants are able to differentiate between self and non-self neighbors and develop fewer and shorter roots in the presence of other roots of the same individual. Furthermore, once cuttings that originate from the very same node are separated, they become progressively alienated from each other and eventually relate to each other as genetically alien plants. The results suggest that the observed self͞non-self discrimination is mediated by physiological coordination among roots that developed on the same plant rather than allogenetic recognition. The observed physiological coordination is based on an as yet unknown mechanism and has important ecological implications, because it allows the avoidance of competition with self and the allocation of greater resources to alternative functions.Buchloe dactyloides ͉ competition ͉ development ͉ phenotypic plasticity ͉ physiological coordination V irtually all multicellular organisms possess recognition systems that allow them to distinguish self from non-self with precision (1-4). For example, self͞non-self recognition systems enable the prevention of inbreeding by self-pollination in plants (4-5) as well as the cooperation between kin in various clonal marine invertebrates (6-8). Because competition entails allocation of limiting resources to nonreproductive functions (9-11), natural selection is expected to favor mechanisms that minimize wasteful competition among parts of the same individual (12), clonemates, and kin (13).Recent evidence suggests that roots are able to alter their growth according to the presence or absence of specific neighbors (14) and to segregate spatially in ''territories'' (15). The evidence also suggests the existence of two different types of self͞non-self discrimination among roots. (24,31), it is usually not studied in the context of self͞non-self interactions. However, the results of previous studies suggest the involvement of physiological coordination in self͞non-self root discrimination (18,20,21). The prevention of contact inhibition between roots of the same A. dumosa plant (18) as well as the avoidance of self competition between roots of F. chiloensis (20) was found to be at least partially based on physiological coordination between roots that develop on the same plant. In an earlier study, P. sativum plants were grown so that they had two roots and two shoots that could be either longitudinally separated into two genetically identical but physiologically distinct individuals or left intact. Root growth was significantly greater in the presence and the direction of roots that belonged to different plants,...