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Recent observational evidence suggests that nighttime temperatures are increasing faster than daytime temperatures, while in some regions precipitation events are becoming less frequent and more intense. The combined ecological impacts of these climatic changes on crassulacean acid metabolism (CAM) plants and their interactions with other functional groups (i.e., grass communities) remain poorly understood. Here we developed a growth chamber experiment to investigate how two CAM–grass communities in desert ecosystems of the southwestern United States and northern Mexico respond to asymmetric warming and increasing rainfall variability. Grasses generally showed competitive advantages over CAM plants with increasing rainfall variability under ambient temperature conditions. In contrast, asymmetric warming caused mortality of both grass species (Bouteloua eriopoda and Bouteloua curtipendula) in both rainfall treatments due to enhanced drought stress. Grass mortality indirectly favored CAM plants even though the biomass of both CAM species Cylindropuntia imbricata and Opuntia phaeacantha significantly decreased. The stem's volume-to-surface ratio of C. imbricata was significantly higher in mixture than in monoculture under ambient temperature (both P < 0.0014); however, the difference became insignificant under asymmetric warming (both P > 0.1625), suggesting that warming weakens the negative effects of interspecific competition on CAM plant growth. Our findings suggest that while the increase in intra-annual rainfall variability enhances grass productivity, asymmetric warming may lead to grass mortality, thereby indirectly favoring the expansion of co-existing CAM plants. This study provides novel experimental evidence showing how the ongoing changes in global warming and rainfall variability affect CAM–grass growth and interactions in dryland ecosystems. [ABSTRACT FROM AUTHOR]

Background and aims: The intensity of extreme rainfall events is increasing in many regions of the world, with important impacts on community dynamics and ecosystem functioning especially in water-limited ecosystems. The impact of the intensification of extreme precipitation on mixed communities of CAM plants and dryland C4 grasses remains poorly understood. Methods: Through a set of greenhouse experiments we investigated the effect of increasing intraseasonal rainfall variability on two separate pairs of CAM and C4 grass species (C. imbricata and B. eriopoda; O. phaeacantha and B. curtipendula) that coexist in arid grasslands across the southwest USA. Results: The increased rainfall variability did not significantly change the biomass of C. imbricata while increasing the biomass of B. eriopoda when these two species coexisted. More extreme rainfall regimes caused a 24.8% decrease in B. curtipendula (grass) biomass, compared with a 71.3% decline in the O. phaeacantha CAM plant. Significantly lower nocturnal carbon assimilation rates and higher stem volume-to-surface area ratios were found in CAM plants in mixture than in monoculture. Conclusions: Our study suggests that C4 grasses may outcompete CAM plants through greater access to deep soil water under increased rainfall variability and highlights the important role of hydrologic conditions as determinants of the competitive relations between CAM plants and grasses, their community composition, and ecological resilience in dryland ecosystems. [ABSTRACT FROM AUTHOR]

Background and aims: We investigated the response of a model facultative CAM plant (Mesembryanthemum crystallinum) and its competition with a C3 grass (Bromus mollis) to changes in rainfall regime.Methods: Seedlings of M. crystallinum and B. mollis in both monoculture and mixtures growing in shallow and deep pots were subjected to three levels of intra-seasonal rainfall variability and rainfall seasonality in both high water and low water conditions. Response of plants were evaluated by nocturnal carboxylation and biomass.Results: A high rate of water drainage beneath root zones in coarse soil led to a negative response of M. crystallinum and B. mollis in monoculture under increased intra-seasonal rainfall variability. Seasonal rainfall shifts to later dates during the growing season generally favored the growth of M. crystallinum and B. mollis in monoculture, with the exception of high water stress conditions whereby drought-intolerant species B. mollis was disfavored. Rainfall seasonality but not intra-seasonal rainfall variability affected nocturnal carboxylation by M. crystallinum in monoculture.Conclusions: We suggest that soil texture, root depth, and rainfall gradient are important mediators of plant growth under increased intra-seasonal rainfall variability. Drought severity and the ability of a plant to tolerate drought and can greatly affect its response to the seasonal timing of rainfall. Nocturnal carboxylation by M. crystallinum in response to rainfall variability depends on the timescale. [ABSTRACT FROM AUTHOR]

Abstract Plants with crassulacean acid metabolism ( CAM) are increasing their abundance in drylands worldwide. The drivers and mechanisms underlying the increased dominance of CAM plants and CAM expression (i.e., nocturnal carboxylation) in facultative CAM plants, however, remain poorly understood. We investigated how nutrient and water availability affected competition between Mesembryanthemum crystallinum (a model facultative CAM species) and the invasive C3 grass Bromus mollis that co-occur in California's coastal grasslands. Specifically we investigated the extent to which water stress, nutrients, and competition affect nocturnal carboxylation in M. crystallinum. High nutrient and low water conditions favored M. crystallinum over B. mollis, in contrast to high water conditions. While low water conditions induced nocturnal carboxylation in 9-week-old individuals of M. crystallinum, in these low water treatments, a 66% reduction in nutrient applied over the entire experiment did not further enhance nocturnal carboxylation. In high water conditions M. crystallinum both alone and in association with B. mollis did not perform nocturnal carboxylation, regardless of the nutrient levels. Thus, nocturnal carboxylation in M. crystallinum was restricted by strong competition with B. mollis in high water conditions. This study provides empirical evidence of the competitive advantage of facultative CAM plants over grasses in drought conditions and of the restricted ability of M. crystallinum to use their photosynthetic plasticity (i.e., ability to switch to CAM behavior) to compete with grasses in well-watered conditions. We suggest that a high drought tolerance could explain the increased dominance of facultative CAM plants in a future environment with increased drought and nitrogen deposition, while the potential of facultative CAM plants such as M. crystallinum to expand to wet environments is expected to be limited. [ABSTRACT FROM AUTHOR]

Abiotic drivers of environmental stress have been found to induce CAM expression (nocturnal carboxylation) in facultative CAM species such as Mesembryanthemum crystallinum. The role played by biotic factors such as competition with non-CAM species in affecting CAM expression, however, remains largely understudied. This research investigated the effects of salt and water conditions on the competition between M. crystallinum and the C grass Bromus mollis with which it is found to coexist in California's coastal grasslands. We also investigated the extent to which CAM expression in M. crystallinum was affected by the intensity of the competition with B. mollis. We found that M. crystallinum had a competitive advantage over B. mollis in drought and saline conditions, while B. mollis exerted strong competitive effects on M. crystallinum in access to light and soil nutrients in high water conditions. This strong competitive effect even outweighed the favorable effects of salt or water additions in increasing the biomass and productivity of M. crystallinum in mixture. Regardless of salt conditions, M. crystallinum did not switch to CAM photosynthesis in response to this strong competitive effect from B. mollis. Disturbance (i.e., grass cutting) reduced the competitive pressure by B. mollis and allowed for CAM expression in M. crystallinum when it was grown mixed with B. mollis. We suggest that moderate competition with other functional groups can enhance CAM expression in M. crystallinum, thereby affecting its plasticity and ability to cope with biological stress. [ABSTRACT FROM AUTHOR]

Past studies have largely focused on hydraulic redistribution (HR) in trees, shrubs, and grasses, and recognized its role in interspecies interactions. HR in plants that conduct crassulacean acid metabolism (CAM), however, remains poorly investigated, as does the effect of HR on transpiration in different vegetation associations (i.e., tree-grass, CAM-grass, and tree-CAM associations). We have developed a mechanistic model to investigate the net direction and magnitude of HR at the patch scale for tree-grass, CAM-grass, and tree-CAM associations at the growing season to yearly timescale. The modeling results show that deep-rooted CAM plants in CAM-grass associations could perform hydraulic lift at a higher rate than trees in tree-grass associations in a relatively wet environment, as explained by a significant increase in grass transpiration rate in the shallow soil layer, balancing a lower transpiration rate by CAM plants. By comparison, trees in tree-CAM associations may perform hydraulic descent at a higher rate than those in tree-grass associations in a dry environment. Model simulations also show that hydraulic lift increases the transpiration of shallow-rooted plants, while hydraulic descent increases that of deep-rooted plants. CAM plants transpire during the night and thus perform HR during the day. Based on these model simulations, we suggest that the ability of CAM plants to perform HR at a higher rate may have different effects on the surrounding plant community than those of plants with C or C photosynthetic pathways (i.e., diurnal transpiration). [ABSTRACT FROM AUTHOR]

Plants with crassulacean acid metabolism (CAM) are increasing their cover in many dryland regions around the world. Their increased dominance has been related to climate warming and atmospheric CO fertilization, while the effects of interspecies interactions and the role of CAM plant facilitation by trees and grasses remain poorly understood. Woody plants are known for their ability to directly facilitate CAM plants through amelioration of the abiotic environment. Mechanisms of indirect facilitation of trees on CAM plants in tree-grass-CAM associations, however, have received less attention. It is also unclear whether grasses might facilitate CAM plants in mixed tree-grass-CAM communities. For instance, the inclusion of grasses in tree-CAM associations could enhance hydraulic lift and facilitate CAM plants in their access to shallow soil moisture at the expenses of deep-rooted trees. If this effect outweighs the competitive effects of grasses on CAM plants, grasses could overall facilitate CAM plants through hydraulic lift. Here we develop a process-based ecohydrological model to investigate the direct and indirect facilitation in tree-CAM-grass associations; the model quantifies transpiration of CAM plants when isolated as well as in associations with trees and/or grasses. It is found that woody plants having a high root overlap with CAM plants indirectly facilitate CAM plants by significantly reducing grass transpiration in shaded conditions. For situations of a low-to-moderate root overlap, facilitation may occur both directly and indirectly. Conversely, grasses are unable to indirectly facilitate CAM plants through the mechanism of hydraulic lift because the competitive effects of grasses on CAM plants outweigh the facilitation induced by hydraulic lift. [ABSTRACT FROM AUTHOR]