Michael E. Loik

Convergence across biomes to a common rain-use efficiency.

Water availability limits plant growth and production in almost all terrestrial ecosystems. However, biomes differ substantially in sensitivity of aboveground net primary production (ANPP) to between-year variation in precipitation. Average rain-use efficiency (RUE; ANPP/precipitation) also varies between biomes, supposedly because of differences in vegetation structure and/or biogeochemical constraints. Here we show that RUE decreases across biomes as mean annual precipitation increases. However, during the driest years at each site, there is convergence to a common maximum RUE (RUEmax) that is typical of arid ecosystems. RUEmax was also identified by experimentally altering the degree of limitation by water and other resources. Thus, in years when water is most limiting, deserts, grasslands and forests all exhibit the same rate of biomass production per unit rainfall, despite differences in physiognomy and site-level RUE. Global climate models predict increased between-year variability in precipitation, more frequent extreme drought events, and changes in temperature. Forecasts of future ecosystem behaviour should take into account this convergent feature of terrestrial biomes.

Assessing the Response of Terrestrial Ecosystems to Potential Changes in Precipitation

Abstract Changes in Earths surface temperatures caused by anthropogenic emissions of greenhouse gases are expected to affect global and regional precipitation regimes. Interactions between changing precipitation regimes and other aspects of global change are likely to affect natural and managed terrestrial ecosystems as well as human society. Although much recent research has focused on assessing the responses of terrestrial ecosystems to rising carbon dioxide or temperature, relatively little research has focused on understanding how ecosystems respond to changes in precipitation regimes. Here we review predicted changes in global and regional precipitation regimes, outline the consequences of precipitation change for natural ecosystems and human activities, and discuss approaches to improving understanding of ecosystem responses to changing precipitation. Further, we introduce the Precipitation and Ecosystem Change Research Network (PrecipNet), a new interdisciplinary research network assembled to encourage and foster communication and collaboration across research groups with common interests in the impacts of global change on precipitation regimes, ecosystem structure and function, and the human enterprise.

Thresholds, memory, and seasonality: understanding pulse dynamics in arid/semi-arid ecosystems

Changes in resource availability can alter the functioning of ecosystems, especially with regard to both population dynamics and the cycling of organic matter and nutrients. At perhaps one end of the spectrum, arid and semi-arid ecosystems represent an extreme in which essential resource availability (e.g., water) is discontinuously available and the availability of these resources impact the ecosystem as discreet pulse events interspersed among long periods of limited resource availability. The objective of the special section that follows is to contribute to our understanding of how pulsed events shape population dynamics, species interactions, and ecosystem processes in arid and semi-arid ecosystems. There has been a renewed interest in recent years in the relationships between precipitation and ecosystem processes in arid and semi-arid lands. For many years, studies focused on the effects of mean precipitation, averaged at annual, seasonal, or monthly time scales. These studies suggested broad-scale relationships between precipitation amounts and seasonal distribution and the productivity of ecosystems (Le Houerou 1984; Le Houerou et al. 1988), or the plant functional type composition of water-limited systems (e.g., Neilson 1995; Smith et al. 1997). Though some researchers asked questions about the effects of single rainfall events early on (e.g., Went and Westergaard 1949; Beatley 1974; Sala and Lauenroth 1982), this perspective received wider attention only in the last decade, in part through the development of stable isotope tracer methods in plant ecology (e.g., Caldwell et al. 1985; Ehleringer et al. 1991; Lin et al. 1996). More recently, studies also have begun to focus on intra-seasonal precipitation patterns, asking whether variability in rainfall event size, frequency, and timing alone affect the biological processes in water-limited ecosystems (e.g., Goldberg and Novoplansky 1997; Knapp et al. 2002; Schwinning et al. 2003). With a view to addressing these questions, an international workshop was held at the University of Arizona in Tucson, USA (http://ag.arizona.edu/research/schwinn/ workshop.html). This special issue contains both peerreviewed synthesis papers that were generated through the collaboration of workshop participants, as well as original research contributions related to pulse dynamics in arid and semi-arid ecosystems. A generation ago, two major paradigms for understanding rainfall effects in water-limited ecosystems were published: (1) the pulse-reserve paradigm of Noy-Meir (1974) and co-workers, and (2) Walter’s (1971) two-layer soil water-partitioning model. Our understanding has advanced significantly since then, moving from the largely conceptual paradigms to an invariably more complex body of data and theory. Reynolds et al. (2004) critically review both paradigms, concluding that the accurate description of soil moisture dynamics is pivotal for addressing the dynamics of primary production and soil water partitioning. While Walter (1971) envisioned water partitioning in vertical space to govern major patterns of plant functional type distribution, Ehleringer et al. (1991) and Lin et al. (1996) have provided strong evidence of this spatial partitioning. Evidence is mounting that partitioning over time may be at least as important (Reynolds et al. 2004; Schwinning et al. 2004a, b). However, the consequences of the hydraulic redistribution of water for species S. Schwinning (*) School of Natural Resources, University of Arizona, Biological Sciences East 325, Tucson, AZ, 85721 e-mail: schwinn@Ag.arizona.edu

Quantifying ecological memory in plant and ecosystem processes

The role of time in ecology has a long history of investigation, but ecologists have largely restricted their attention to the influence of concurrent abiotic conditions on rates and magnitudes of important ecological processes. Recently, however, ecologists have improved their understanding of ecological processes by explicitly considering the effects of antecedent conditions. To broadly help in studying the role of time, we evaluate the length, temporal pattern, and strength of memory with respect to the influence of antecedent conditions on current ecological dynamics. We developed the stochastic antecedent modelling (SAM) framework as a flexible analytic approach for evaluating exogenous and endogenous process components of memory in a system of interest. We designed SAM to be useful in revealing novel insights promoting further study, illustrated in four examples with different degrees of complexity and varying time scales: stomatal conductance, soil respiration, ecosystem productivity, and tree growth. Models with antecedent effects explained an additional 18-28% of response variation compared to models without antecedent effects. Moreover, SAM also enabled identification of potential mechanisms that underlie components of memory, thus revealing temporal properties that are not apparent from traditional treatments of ecological time-series data and facilitating new hypothesis generation and additional research.

Soil microbial and nutrient responses to 7 years of seasonally altered precipitation in a Chihuahuan Desert grassland

Soil microbial communities in Chihuahuan Desert grasslands generally experience highly variable spatiotemporal rainfall patterns. Changes in precipitation regimes can affect belowground ecosystem processes such as decomposition and nutrient cycling by altering soil microbial community structure and function. The objective of this study was to determine if increased seasonal precipitation frequency and magnitude over a 7-year period would generate a persistent shift in microbial community characteristics and soil nutrient availability. We supplemented natural rainfall with large events (one/winter and three/summer) to simulate increased precipitation based on climate model predictions for this region. We observed a 2-year delay in microbial responses to supplemental precipitation treatments. In years 3-5, higher microbial biomass, arbuscular mycorrhizae abundance, and soil enzyme C and P acquisition activities were observed in the supplemental water plots even during extended drought periods. In years 5-7, available soil P was consistently lower in the watered plots compared to control plots. Shifts in soil P corresponded to higher fungal abundances, microbial C utilization activity, and soil pH. This study demonstrated that 25% shifts in seasonal rainfall can significantly influence soil microbial and nutrient properties, which in turn may have long-term effects on nutrient cycling and plant P uptake in this desert grassland.

Differential daytime and night-time stomatal behavior in plants from North American deserts

Night-time stomatal conductance (g(night)) occurs in many ecosystems, but the g(night) response to environmental drivers is relatively unknown, especially in deserts. Here, we conducted a Bayesian analysis of stomatal conductance (g) (N=5013) from 16 species in the Sonoran, Chihuahuan, Mojave and Great Basin Deserts (North America). We partitioned daytime g (g(day)) and g(night) responses by describing g as a mixture of two extreme (dark vs high light) behaviors. Significant g(night) was observed across 15 species, and the g(night) and g(day) behavior differed according to species, functional type and desert. The transition between extreme behaviors was determined by light environment, with the transition behavior differing between functional types and deserts. Sonoran and Chihuahuan C(4) grasses were more sensitive to vapor pressure difference (D) at night and soil water potential (Ψ(soil)) during the day, Great Basin C(3) shrubs were highly sensitive to D and Ψ(soil) during the day, and Mojave C(3) shrubs were equally sensitive to D and Ψ(soil) during the day and night. Species were split between the exhibition of isohydric or anisohydric behavior during the day. Three species switched from anisohydric to isohydric behavior at night. Such behavior, combined with differential D, Ψ(soil) and light responses, suggests that different mechanisms underlie g(day) and g(night) regulation.

Photosynthetic responses of tree seedlings in grass and under shrubs in early-successional tropical old fields, Costa Rica

Only recently have studies addressed the effect of early-colonizing vegetation on tree seedling survival and growth during secondary succession in tropical old fields, and few studies have elucidated the physiological responses of tree seedlings to different vegetational communities. We compared growth and various photosynthetic parameters for seedlings of four rain-forest tree species, Cedrela tonduzii, Inga punctata, Ocotea whitei, and Tapirira mexicana, growing in areas of pasture grass and shrubs in early-successional abandoned pasture in Costa Rica; in addition, we made measurements for two species in forest gaps. We tested the general hypothesis that early-colonizing shrubs facilitate growth of forest tree seedlings. Specifically, we measured microclimate, growth, CO2 assimilation, stomatal conductance, photosystem II quantum yield (ΦPSII), and xanthophyll pigment pools for all seedlings. Photosynthetic flux density (PFD) was higher under grass than shrubs or forest gaps, but was highly variable in each growth environment. For three of the four species, height growth was greatest in the grass compared to the shrubs and forest gaps; growth was similar below grass and shrubs for O. whitei. Photosynthetic capacity, apparent quantum yield, and stomatal conductance did not vary across habitats, but light compensation point and PFD at light saturation tended to be higher in the grass compared to forest and shrub growth environments. Water use efficiency differed across growth environments for three of the species. For plants in ambient PFD and dark-adapted plants, the efficiency of excitation energy transfer through PSII was lowest for plants in the grass compared to shrubs and forest gaps and also differed across species. Measurement of steady-state responses of ΦPSII to increasing PFD indicated a significant effect of growth environment at low PFD for all species and significant effects at high PFD only for I. punctata. All species exhibited a high degree of midday xanthophyll de-epoxidation in the different growth environments. Xanthophyll pigment pool size on an area basis was highest in the grass compared to shrubs and forest gaps for all four species. The results suggest that shrubs do not provide a facilitative effect for growth or photosynthesis for ~1.5-year-old seedlings of these four species. We conclude that site differences in success of tree seedlings during succession are a result of complex interactions of facilitation and competition and are not simply based on physiological responses to PFD.

Utilizing intraspecific variation in phenotypic plasticity to bolster agricultural and forest productivity under climate change

Climate change threatens the ability of agriculture and forestry to meet growing global demands for food, fibre and wood products. Information gathered from genotype-by-environment interactions (G × E), which demonstrate intraspecific variation in phenotypic plasticity (the ability of a genotype to alter its phenotype in response to environmental change), may prove important for bolstering agricultural and forest productivity under climate change. Nonetheless, very few studies have explicitly quantified genotype plasticity-productivity relationships in agriculture or forestry. Here, we conceptualize the importance of intraspecific variation in agricultural and forest species plasticity, and discuss the physiological and genetic factors contributing to intraspecific variation in phenotypic plasticity. Our discussion highlights the need for an integrated understanding of the mechanisms of G × E, more extensive assessments of genotypic responses to climate change under field conditions, and explicit testing of genotype plasticity-productivity relationships. Ultimately, further investigation of intraspecific variation in phenotypic plasticity in agriculture and forestry may prove important for identifying genotypes capable of increasing or sustaining productivity under more extreme climatic conditions.

Genetic variation in circadian regulation of nocturnal stomatal conductance enhances carbon assimilation and growth

Circadian resonance, whereby a plants endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C3 plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole-plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian-regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.

Influence of summer marine fog and low cloud stratus on water relations of evergreen woody shrubs (Arctostaphylos: Ericaceae) in the chaparral of central California

Mediterranean-type climate (MTC) regions around the world are notable for cool, wet winters and hot, dry summers. A dominant vegetation type in all five MTC regions is evergreen, sclerophyllous shrubland, called chaparral in California. The extreme summer dry season in California is moderated by a persistent low-elevation layer of marine fog and cloud cover along the margin of the Pacific coast. We tested whether late dry season water potentials (Ψmin) of chaparral shrubs, such as Arctostaphylos species in central California, are influenced by this coast-to-interior climate gradient. Lowland coastal (maritime) shrubs were found to have significantly less negative Ψmin than upland interior shrubs (interior), and stable isotope (δ13C) values exhibited greater water use efficiency in the interior. Post-fire resprouter shrubs (resprouters) had significantly less negative Ψmin than co-occurring obligate seeder shrubs (seeders) in interior and transitional chaparral, possibly because resprouters have deeper root systems with better access to subsurface water than shallow-rooted seeders. Unexpectedly, maritime resprouters and seeders did not differ significantly in their Ψmin, possibly reflecting more favorable water availability for shrubs influenced by the summer marine layer. Microclimate and soil data also suggest that maritime habitats have more favorable water availability than the interior. While maritime seeders constitute the majority of local Arctostaphylos endemics, they exhibited significantly greater vulnerability to xylem cavitation than interior seeders. Because rare seeders in maritime chaparral are more vulnerable to xylem cavitation than interior seeders, the potential breakdown of the summer marine layer along the coast is of potential conservation concern.