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The Tibetan Plateau (TP) is experiencing high rates of climatic change. We present a novel combined mechanistic-bioclimatic modeling approach to determine how changes in precipitation and temperature on the TP may impact net primary production (NPP) in four major biomes (forest, shrub, grass, desert) and if there exists a maximum rain use efficiency (RUE(MAX)) that represents Huxman et al.'s "boundary that constrain[s] site-level productivity and efficiency." We used a daily mechanistic ecosystem model to generate 40-yr outputs using observed climatic data for scenarios of decreased precipitation (25-100%); increased air temperature (1 degrees - 6 degrees C); simultaneous changes in both precipitation (+/- 50%, +/- 25%) and air temperature (+1 to +6 degrees C) and increased interannual variability (IAV) of precipitation (+1 sigma to +3 sigma, with fixed means, where sigma is SD). We fitted model output from these scenarios to Huxman et al.'s RUE(MAX) bioclimatic model, NPP = alpha + RUE x PPT (where alpha is the intercept, RUE is rain use efficiency, and PPT is annual precipitation). Based on these analyses, we conclude that there is strong support (when not explicit, then trend-wise) for Huxman et al.'s assertion that biomes converge to a common RUE(MAX) during the driest years at a site, thus representing the boundary for highest rain use efficiency; the interactive effects of simultaneously decreasing precipitation and increasing temperature on NPP for the TP is smaller than might be expected from additive, single-factor changes in these drivers; and that increasing IAV of precipitation may ultimately have a larger impact on biomes of the Tibetan Plateau than changing amounts of rainfall and air temperature alone.
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Mudança Climática , Ecossistema , Monitoramento Ambiental , TibetRESUMO
Community and ecosystem responses to global environmental change are contingent on the magnitude of change and interacting global change factors. To reveal whether responses are also contingent on the magnitude of each interacting factor, multifactor, multilevel experiments are required, but are rarely conducted. We exposed model grassland ecosystems to six levels of atmospheric CO2 and six levels of nitrogen enrichment, applying the latter both chronically (simulating deposition) and acutely (simulating fertilization). The 66 treatments were maintained for 6 months under controlled growing conditions, with biomass harvested every 28 d and sorted to species. Aboveground plant productivity responses to CO2 were contingent on nitrogen amount, and the responses to nitrogen amount were dependent on whether applications were chronic or acute. Specifically, productivity responses to increasing CO2 concentrations were accentuated with higher nitrogen enrichments, and productivity was greater when higher nitrogen enrichments were applied acutely. Plant community composition was influenced only by nitrogen enrichment, where the co-dominant grass species with the greatest leaf trait plasticity increasingly dominated with higher nitrogen amounts. Community processes are considered to be unpredictable, but our data suggest that the prediction of the impacts of simultaneous global changes is more complex for ecosystem processes, given that their responses are contingent on the levels of interacting factors.
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Biota , Mudança Climática , Fenômenos Fisiológicos Vegetais , Biomassa , Dióxido de Carbono/farmacologia , Nitrogênio/farmacologia , Folhas de Planta/efeitos dos fármacos , Folhas de Planta/fisiologia , Fenômenos Fisiológicos Vegetais/efeitos dos fármacos , SoloRESUMO
Encroachment of woody plants into grasslands has generated considerable interest among ecologists. Syntheses of encroachment effects on ecosystem processes have been limited in extent and confined largely to pastoral land uses or particular geographical regions. We used univariate analyses, meta-analysis and structural equation modelling to test the propositions that (1) shrub encroachment does not necessarily lead to declines in ecosystem functions and (2) shrub traits influence the functional outcome of encroachment. Analyses of 43 ecosystem attributes from 244 case studies worldwide showed that some attributes consistently increased with encroachment (e.g. soil C, N), and others declined (e.g. grass cover, pH), but most exhibited variable responses. Traits of shrubs were associated with significant, though weak, structural and functional outcomes of encroachment. Our review revealed that encroachment had mixed effects on ecosystem structure and functioning at global scales, and that shrub traits influence the functional outcome of encroachment. Thus, a simple designation of encroachment as a process leading to functionally, structurally or contextually degraded ecosystems is not supported by a critical analysis of existing literature. Our results highlight that the commonly established link between shrub encroachment and degradation is not universal.
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Ecossistema , Espécies Introduzidas , Modelos Biológicos , Plantas , Geografia , Dinâmica PopulacionalRESUMO
In the short-term heterotrophic soil respiration is strongly and positively related to temperature. In the long-term, its response to temperature is uncertain. One reason for this is because in field experiments increases in respiration due to warming are relatively short-lived. The explanations proposed for this ephemeral response include depletion of fast-cycling, soil carbon pools and thermal adaptation of microbial respiration. Using a > 15 year soil warming experiment in a mid-latitude forest, we show that the apparent 'acclimation' of soil respiration at the ecosystem scale results from combined effects of reductions in soil carbon pools and microbial biomass, and thermal adaptation of microbial respiration. Mass-specific respiration rates were lower when seasonal temperatures were higher, suggesting that rate reductions under experimental warming likely occurred through temperature-induced changes in the microbial community. Our results imply that stimulatory effects of global temperature rise on soil respiration rates may be lower than currently predicted.
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Adaptação Fisiológica/fisiologia , Temperatura Alta , Microbiologia do Solo , Biomassa , Análise de Regressão , Estações do Ano , Solo/análiseRESUMO
Patterns of resource availability and heterogeneity shape the composition, productivity, and dynamics of plant assemblages in a wide variety of terrestrial ecosystems. Despite this, the responses of plant assemblages to simultaneous changes in the availability and heterogeneity of more than a single resource are virtually unknown. To fill this gap, microcosms consisting of assemblages formed by Lolium perenne, Plantago lanceolata, Anthoxantum odoratum, Holcus lanatus, and Trifolium repens were grown in a factorial experiment with the following treatments: nutrient availability (NA), water availability (WA), spatial nutrient heterogeneity (NH), and temporal water heterogeneity (WH). Assemblages exhibited precise root foraging patterns in response to nutrient heterogeneity, which were modified by NA and WA. A series of two- and three-way interactions involving the four factors evaluated determined biomass production, the belowground: aboveground biomass ratio, the patterns of root biomass allocation with depth, and the relative contribution to aboveground biomass of Lolium and Anthoxanthum. In all cases, these interactions explained significant amounts of the variation found in the data. Our study demonstrates that considering the interactions between resource availability and heterogeneity allows for a refinement of predictions that can detectably reduce the error associated with extrapolating from single factor analyses.
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Ecossistema , Nitrogênio/fisiologia , Poaceae/fisiologia , Trifolium/fisiologia , Água/fisiologia , Biomassa , Fatores de TempoRESUMO
⢠We hypothesized that plant growth reduction under water stress is caused primarily by a reduction of leaf-area ratio (LAR, leaf area per unit of total plant dry mass). ⢠Two perennial Chihuahuan desert grass species (slow-growing Bouteloua eriopoda and fast-growing Eragrostis lehmanniana) were subjected over 6 wk to a combination of two water-supply regimes (control and drought) and two levels of atmospheric CO2 partial pressure (375 and 750 µmol mol-1 ). ⢠Drought reduced final biomass in Bouteloua by 60% regardless of CO2 concentration. Eragrostis experienced a similar biomass reduction at 375 µmol mol-1 , but large plants under elevated CO2 attained growth rates comparable to those of controls. Overall, for plants of similar size, drought reduced LAR in both species much more strongly than it affected net assimilation rate. This reduction in LAR was caused by reductions in both specific leaf area and leaf weight ratio. ⢠We conclude that reduced growth under drought can be considered as a byproduct of the same plastic, developmental responses that result in a reduced water loss.
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Ever more frequent and closer involvement by clinical geneticists and counselors in the prenatal assessment of development mandates a better understanding of all stages of human ontogeny, but especially those of earliest development during which most of the lethal and all of the gross, multiple and complex defects of morphogenesis arise. Because of the phenomenon of universality, i.e., identical molecular inductive mechanisms involved in the process of embryonic pattern formation in all vertebrates, experimental animals indeed are a most valuable approach to an understanding of the causal and formal aspects of development and are beginning to forge essential, strong bonds between molecular biologists and clinicians in a mutually supportive discipline of developmental biology. However, to grieving parents of a stillborn fetus with, say, Pentalogy of Cantrell, sirenomelia or otocephaly, mouse data offer little comfort or reassurance about recurrence; thus, it is imperative to make ever more effective a science of human teratology (sensu lato) with participating reproductive geneticists, obstetricians, neonatologists, ultrasonographers, pediatric/fetal pathologists, cytogeneticists and pediatric geneticists to generate the diagnostic, pathogenetic and causal data necessary to counsel and to comfort the parents. Few molecular data exist on causes of blastogenetic defects in humans; however, the phenomenon of parsimony, whereby the same "morphogenetic" molecule, say, sonic hedgehog (SHH), is "deployed" simultaneously or sequentially during the morphogenesis (and even the histogenesis) of several/many embryonic primordia, makes it likely that a genetic/epigenetic disturbance of such an inductive system will have multiple effects on blastogenetic, organogenetic and perhaps also histogenetic events in the embryo. If causally defined, such a pattern of anomalies constitutes pleiotropy, and the embryo/fetus can be said to have a syndrome. If cause is unknown, the presumption of pleiotropy is less certain, and the fetus/infant may be said to have an "association" with low empiric recurrence risk.
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Blastocisto/patologia , Anormalidades Congênitas/classificação , Anormalidades Congênitas/embriologia , Desenvolvimento Embrionário e Fetal/genética , Anormalidades Congênitas/epidemiologia , Anormalidades Congênitas/genética , Embrião de Mamíferos/anormalidades , Feminino , HumanosRESUMO
It appears that polar regions of the Earth will bear the brunt of global temperature increases. Because of the ecological importance of the sedge Eriophorum vaginatum in the arctic and the large amount of data available on its growth and physiology, we chose this species as a test case to model the potential long-term response of arctic plants to global climate change. Our simulation model utilizes a mechanistic framework and includes the effects of light, temperature, season length, nitrogen availability, and CO2 concentration on E. vaginatum growth dynamics. The model was parameterized based on a series of published studies of the growth responses of E. vaginatum to nutrients and validated using (1) field studies on the growth responses of E. vaginatum to temperature and shading, and (2) the effects of elevated CO2 and temperature on E. vaginatum photosynthesis. The effect of a 50-yr period of climate change on peak biomass (overwintering biomass plus seasonal production) in E. vaginatum was explored. We use climate change here to refer to linear increases over a 50-yr period in temperature (from 8° to 13°C), season length (from 100 to 120 d), and atmospheric CO2 (from 340 to 680 @mL/L). Similarly, a wide range of nitrogen availabilities (from 9 to 18 g°m -2 °yr-1 ) was also examined because of its importance in productivity. The model predicts that a simultaneous increase in the direct effects of temperature, season length, and CO2 , with no change in nitrogen availability, will result in a slight decrease in peak biomass. A simulated long-term doubling of nitrogen availability results in an °70% increase in peak biomass, whereas with concurrent changes in climate and nitrogen availability, the model predicts a slight decline in peak biomass compared to increases in nitrogen alone. In essence, the model predicts that climate change will have substantial effects on E. vaginatum only indirectly through changes in nitrogen availability. Simulated peak biomass responds linearly up to a doubling of current nitrogen availabilities. Therefore, at low-to-moderate increases in nitrogen availability, the predicted response of E. vaginatum to climate change is linearly (and almost exclusively) dependent on our ability to predict the effects of climate change on nitrogen cycling. At nitrogen availabilities >2x current availabilities, the relationship flattens out very rapidly because the plant becomes limited by carbon uptake. Thus, if nitrogen availabilities more than double in the future, E. vaginatum may shift from being a nutrient-limited to a carbon-limited system and, consequently, increased season length and elevated CO2 concentrations may play an important role in controlling E. vaginatum productivity.
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Creosobebush (Larrea tridentata) fine litter was treated with either the general biocide HgCl2 and CuSO4 or water (controls) and buried 5 cm beneath the soil surface in the northern Chihuahuan Desert. The treated litter showed significantly less mass loss than controls during the three month summer-autumn field study; controls lost about 20% of the original mass while treated litter lost less than 2%. In addition, the total nitrogen content of the control litter increased from an initial concentration of about 14.08 g kg-1 to 17.62 g kg-1 dry weight by the end of the study, while treated litter nitrogen content decreased to 13.30 g kg-1. Results suggest abiotic processes other than leaching have little effect on the decomposition of buried litter in this environment.
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In previous papers we have described and verified a primary production model of the desert shrub Larrea tridentata. Here we address the validation phase of the evaluation of this model. Two versions of the model which differ in the priority scheme used for allocating carbon to reproductive or vegetative organs were compared on the basis of their usefulness and reliability over a range of soil-moisture conditions. Over an entire growing season when soil-moisture conditions were near "normal" both versions of the model were adequate predictors of total above-ground vegetative growth and one was an adequate predictor of reproductive growth as well. A more detailed analysis revealed that the versions varied in the range of soil-moisture conditions over which they were adequate and that neither was adequate when soil-moisture had remained high for extended periods. The validation process has revealed some likely areas for model improvement to increase adequacy.
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In arctic tundra soil, oxygen depletion associated with soil flooding may control plant growth either directly through anoxia or indirectly through effects on nutrient availability. This study was designed to evaluate whether plant growth and physiology of two arctic sedge species are more strongly controlled by the direct or indirect effects of decreased soil aeration. Eriophorum angustifolium and E. vaginatum, which originate from flooded and well-drained habitats, respectively, were grown in an in situ transplant garden at two levels of soil oxygen, nitrogen, and phosphorus availability over two growing seasons. In both species, N addition had a stronger effect on growth and biomass allocation than P addition or soil oxygen depletion. Net photosynthesis and carbohydrate concentrations were relatively insensitive to changes in these factors. Biomass reallocated from shoots to below-ground parts in response to limited N supply was equally divided between roots (nutrient acquisition) and perennating rhizomes (storage tissue formation) in E. angustifolium. E. Vaginatum only increased its allocation to rhizomes. In the flood-tolerant E. angustifolium, growth was improved by soil anoxia and biomass allocation among plant parts was not significantly affected. Contrary to our initial hypothesis, whole-plant growth in E. vaginatum improved in flooded soils; however, it only did so when N availability was high. Under low N availability growth in flooded soils was reduced by 20% compared to growth in the aerobic environment. Reduced biomass allocation to rhizomes and thus to storage potential under anaerobic conditions may reduce long-term survival of E. vaginatum in flooded habitats.
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We examined the extent to which carbon investment into secondary compounds in loblolly pine (Pinus taeda L.) is changed by the interactive effect of elevated CO2 and N availability and whether differences among treatments are the result of size-dependent changes. Seedlings were grown for 138 days at two CO2 partial pressures (35 and 70 Pa CO2) and four N solution concentrations (0.5, 1.5, 3.5, and 6.5 mmol l-1 NO3NH4) and concentrations of total phenolics and condensed tannins were determined four times during plant development in primary and fascicular needles, stems and lateral and tap roots. Concentrations of total phenolics in lateral roots and condensed tannins in tap roots were relatively high regardless of treatment. In the smallest seedlings secondary compound concentrations were relatively high and decreased in the initial growth phase. Thereafter condensed tannins accumulated strongly during plant maturation in all plant parts except in lateral roots, where concentrations did not change. Concentrations of total phenolics continued to decrease in lateral roots while they remained constant in all other plant parts. At the final harvest plants grown at elevated CO2 or low N availability showed increased concentrations of condensed tannins in aboveground parts. The CO2 effect, however, disappeared when size differences were adjusted for, indicating that CO2 only indirectly affected concentrations of condensed tannins through accelerating growth. Concentrations of total phenolics increased directly in response to low N availability and elevated CO2 in primary and fascicular needles and in lateral roots, which is consistent with predictions of the carbon-nutrient balance (CNB) hypothesis. The CNB hypothesis is also supported by the strong positive correlations between soluble sugar and total phenolics and between starch and condensed tannins. The results suggest that predictions of the CNB hypothesis could be improved if developmentally induced changes of secondary compounds were included.
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It has long been observed that leaf nitrogen concentrations decline with depth in closed canopies in a number of plant communities. This phenomenon is generally believed to be related to a changing radiation environment and it has been suggested by some researchers that plants allocate nitrogen in order to optimize total whole canopy photosynthesis. Although optimization theory has been successfully utilized to describe a variety of physiological and ecological phenomena, it has some shortcomings that are subject to criticism (e.g., time constraints, oversimplifications, lack of insights, etc.). In this paper we present an alternative to the optimization theory of plant canopy nitrogen distribution, which we term coordination theory. We hypothesize that plants allocate nitrogen to maintain a balance between two processes, each of which is dependent on leaf nitrogen content and each of which potentially limits photosynthesis. These two processes are defined as Wc, the Rubiscolimited rate of carboxylation, and Wj, the electron transport-limited rate of carboxylation. We suggest that plants allocate nitrogen differentially to, leaves in different canopy layers in such a way that Wc and Wj remain roughly balanced. In this scheme, the driving force for the allocation of nitrogen within a canopy is the difference between the leaf nitrogen content that is required to bring Wc and Wj into balance and the current nitrogen content. We show that the daily carbon assimilation of a canopy with a nitrogen distribution resulting from this internal coordination of Wc and Wj is very similar to that obtained using optimization theory.
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While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO(2), soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO(2). Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO(2) or nutrient heterogeneity. Under elevated CO(2) conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity-productivity relationships.
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Dióxido de Carbono/análise , Ecossistema , Raízes de Plantas/fisiologia , Poaceae/crescimento & desenvolvimento , Solo/análise , Análise de Variância , Atmosfera/química , Biomassa , Carbono , Nitrogênio , Especificidade da EspécieRESUMO
In this millennium, global drylands face a myriad of problems that present tough research, management, and policy challenges. Recent advances in dryland development, however, together with the integrative approaches of global change and sustainability science, suggest that concerns about land degradation, poverty, safeguarding biodiversity, and protecting the culture of 2.5 billion people can be confronted with renewed optimism. We review recent lessons about the functioning of dryland ecosystems and the livelihood systems of their human residents and introduce a new synthetic framework, the Drylands Development Paradigm (DDP). The DDP, supported by a growing and well-documented set of tools for policy and management action, helps navigate the inherent complexity of desertification and dryland development, identifying and synthesizing those factors important to research, management, and policy communities.
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Clima Desértico , Ecossistema , Meio Ambiente , Animais , Biodiversidade , Ecologia , Humanos , Política Pública , SoloRESUMO
BACKGROUND AND AIMS: Size-asymmetric competition occurs when larger plants have a disproportionate advantage in competition with smaller plants. It has been hypothesized that nutrient heterogeneity may promote it. Experiments testing this hypothesis are inconclusive, and in most cases have evaluated the effects of nutrient heterogeneity separately from other environmental factors. The aim of this study was to test, using populations of Lolium perenne, Plantago lanceolata and Holcus lanatus, two hypotheses: (a) nutrient heterogeneity promotes size-asymmetric competition; and (b) nutrient heterogeneity interacts with both atmospheric CO2 partial pressure (P(CO2)) and nutrient availability to determine the magnitude of this response. METHODS: Microcosms consisting of monocultures of the three species were grown for 90 d in a factorial experiment with the following treatments: P(CO2) (37.5 and 70 Pa) and nutrient availability (NA; 40 and 120 mg of N added as organic material) combined with different spatial distribution of the organic material (NH; homogeneous and heterogeneous). Differences in the size of individual plants within populations (size inequality) were quantified using the coefficient of variation of individual above-ground biomass and the combined biomass of the two largest individuals in each microcosm. Increases in size inequality were associated with size-asymmetric competition. KEY RESULTS: Size inequality increased when the nutrients were heterogeneously supplied in the three species. The effects of NH on this response were more pronounced under high nutrient supply in both Plantago and Holcus (significant NA x NH interactions) and under elevated P(CO2) in Plantago (significant P(CO2) x NA x NH interaction). No significant two- and three-way interactions were found for Lolium. CONCLUSIONS: Our first hypothesis was supported by our results, as nutrient heterogeneity promoted size-asymmetric competition in the three species evaluated. Nutrient supply and P(CO2) modified the magnitude of this effect in Plantago and Holcus, but not in Lolium. Thus, our second hypothesis was partially supported.
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Dióxido de Carbono/metabolismo , Ecossistema , Holcus/crescimento & desenvolvimento , Lolium/crescimento & desenvolvimento , Plantago/crescimento & desenvolvimento , Pressão Atmosférica , Biomassa , Holcus/anatomia & histologia , Holcus/metabolismo , Lolium/anatomia & histologia , Lolium/metabolismo , Nitrogênio/metabolismo , Pressão Parcial , Fósforo/metabolismo , Plantago/anatomia & histologia , Plantago/metabolismo , SoloRESUMO
Predicting future plant and ecosystem responses to elevated CO(2) also requires an understanding of the role of other factors, especially soil nitrogen. This is particularly challenging for global aridlands where total N and the relative amounts of nitrate and ammonia vary both spatially and seasonally. We measured gas exchange and primary and secondary C metabolites in seedlings of two dominant aridland shrub species (Prosopis flexuosa [S America] and P. glandulosa [N America]) grown at ambient (350 ppm) or elevated (650 ppm) CO(2) and nitrogen at two levels (low [0.8 mM] and high [8.0 mM]) and at either 1 : 1 or 3 : 1 nitrate to ammonia. Whereas elevated CO(2) increased assimilation rate, water use efficiency, and primary carbon metabolites in both species, these increases were strongly contingent upon nitrogen availability. Elevated CO(2) did not increase secondary metabolites (i.e., phenolics). For these important aridland species, the effects of elevated CO(2) are strongly influenced by nitrogen availability and to a lesser extent by the relative amounts of nitrate and ammonia supplied, which underscores the importance of both the amount and chemical composition of soil nitrogen in mediating the potential responses of seedling growth and establishment of aridland plants under future CO(2)-enriched atmospheres.
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Interactive effects of atmospheric CO(2) concentration ([CO(2)]), soil nutrient availability and soil nutrient spatial distribution on the structure and function of plant assemblages remain largely unexplored. Here we conducted a microcosm experiment to evaluate these interactions using a grassland assemblage formed by Lolium perenne, Plantago lanceolata, Trifolium repens, Anthoxanthum odoratum and Holcus lanatus. Assemblages exhibited precise root foraging patterns, had higher total and below-ground biomass, and captured more nitrogen when nutrients were supplied heterogeneously. Root foraging responses were modified by nutrient availability, and the patterns of N capture by interactions between nutrient distribution, availability and [CO(2)]. Greater above-ground biomass was observed under elevated CO(2) only under homogeneous conditions of nutrient supply and at the highest availability level. CO(2) interacted with nutrient distribution and availability to determine foliar percentage N and below : above-ground biomass ratios, respectively. Interactions between nutrient distribution and CO(2) determined the relative contribution to above-ground biomass of four of the species. The responses of dominant and subordinate species to [CO(2)] were dependent on the availability and distribution of nutrients. Our results suggest that soil nutrient distribution has the potential to influence the response of plant species and assemblages to changes in [CO(2)] and nutrient availability.
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Dióxido de Carbono/metabolismo , Lolium/crescimento & desenvolvimento , Plantago/crescimento & desenvolvimento , Poaceae/crescimento & desenvolvimento , Trifolium/crescimento & desenvolvimento , Biomassa , Dióxido de Carbono/farmacologia , Modelos Biológicos , Nitrogênio/metabolismo , Especificidade da EspécieRESUMO
The 'two-layer' and 'pulse-reserve' hypotheses were developed 30 years ago and continue to serve as the standard for many experiments and modeling studies that examine relationships between primary productivity and rainfall variability in aridlands. The two-layer hypothesis considers two important plant functional types (FTs) and predicts that woody and herbaceous plants are able to co-exist in savannas because they utilize water from different soil layers (or depths). The pulse-reserve model addresses the response of individual plants to precipitation and predicts that there are 'biologically important' rain events that stimulate plant growth and reproduction. These pulses of precipitation may play a key role in long-term plant function and survival (as compared to seasonal or annual rainfall totals as per the two-layer model). In this paper, we re-evaluate these paradigms in terms of their generality, strengths, and limitations. We suggest that while seasonality and resource partitioning (key to the two-layer model) and biologically important precipitation events (key to the pulse-reserve model) are critical to understanding plant responses to precipitation in aridlands, both paradigms have significant limitations. Neither account for plasticity in rooting habits of woody plants, potential delayed responses of plants to rainfall, explicit precipitation thresholds, or vagaries in plant phenology. To address these limitations, we integrate the ideas of precipitation thresholds and plant delays, resource partitioning, and plant FT strategies into a simple 'threshold-delay' model. The model contains six basic parameters that capture the nonlinear nature of plant responses to pulse precipitation. We review the literature within the context of our threshold-delay model to: (i) develop testable hypotheses about how different plant FTs respond to pulses; (ii) identify weaknesses in the current state-of-knowledge; and (iii) suggest future research directions that will provide insight into how the timing, frequency, and magnitude of rainfall in deserts affect plants, plant communities, and ecosystems.
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Clima Desértico , Ecossistema , Modelos Biológicos , Fenômenos Fisiológicos Vegetais , Chuva , Fatores Etários , Simulação por Computador , Estações do Ano , Solo/análise , Fatores de TempoRESUMO
The conceptual framework for a simulation model of primary productivity, decomposition and nitrogen cycling in a shrub-dominated desert ecosystem in southern New Mexico is presented. This model is based on our previous attempt to simulate carbon allocation patterns in the desert shrub Larrea tridentata Cov., which demonstrated that moisture patterns alone are insufficient to predict desert productivity. These results, as well as others, suggest that mineral nutrients, especially nitrogen, may also be an important determinant of productivity in arid environments. Our current research in the Chihuahuan desert is directed towards elucidating the numerous biotic and abiotic interactions that determine the rates and directions of carbon, nitrogen and water fluxes in this ecosystem. The development of this working model will serve as a tool to accomplish three major objectives: (1) to synthesize the large amount of existing data on decomposition and nitrogen cycling in deserts, (2) to quantify our present state of knowledge about the structure and function of ecosystem components important in carbon and nitrogen dynamics in deserts, and (3) to address hypotheses concerning the complex mechanisms of interactions and feedbacks among the organisms involved in carbon and nitrogen exchanges in deserts.