José M. Grünzweig

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2

Atmospheric CO2 enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO2 enrichment, with an emphasis on water relations. Increasing CO2 led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO2 enrichment were relatively greater in dry years. In contrast, CO2-induced aboveground biomass increases in the Texas C3/C4 grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO2 in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO2-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO2 were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO2 on plant and soil water relations may contribute substantially to experimentally induced CO2-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO2 effects on photosynthesis and those indirect CO2 effects via soil moisture as documented here.

Physiology-phenology interactions in a productive semi-arid pine forest

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D(L)) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g(s)) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g(s )sensitivity to D(L) below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis-phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.

Non-structural carbohydrates in woody plants compared among laboratories

Non-structural carbohydrates (NSC) in plant tissue are frequently quantified to make inferences about plant responses to environmental conditions. Laboratories publishing estimates of NSC of woody plants use many different methods to evaluate NSC. We asked whether NSC estimates in the recent literature could be quantitatively compared among studies. We also asked whether any differences among laboratories were related to the extraction and quantification methods used to determine starch and sugar concentrations. These questions were addressed by sending sub-samples collected from five woody plant tissues, which varied in NSC content and chemical composition, to 29 laboratories. Each laboratory analyzed the samples with their laboratory-specific protocols, based on recent publications, to determine concentrations of soluble sugars, starch and their sum, total NSC. Laboratory estimates differed substantially for all samples. For example, estimates for Eucalyptus globulus leaves (EGL) varied from 23 to 116 (mean = 56) mg g(-1) for soluble sugars, 6-533 (mean = 94) mg g(-1) for starch and 53-649 (mean = 153) mg g(-1) for total NSC. Mixed model analysis of variance showed that much of the variability among laboratories was unrelated to the categories we used for extraction and quantification methods (method category R(2) = 0.05-0.12 for soluble sugars, 0.10-0.33 for starch and 0.01-0.09 for total NSC). For EGL, the difference between the highest and lowest least squares means for categories in the mixed model analysis was 33 mg g(-1) for total NSC, compared with the range of laboratory estimates of 596 mg g(-1). Laboratories were reasonably consistent in their ranks of estimates among tissues for starch (r = 0.41-0.91), but less so for total NSC (r = 0.45-0.84) and soluble sugars (r = 0.11-0.83). Our results show that NSC estimates for woody plant tissues cannot be compared among laboratories. The relative changes in NSC between treatments measured within a laboratory may be comparable within and between laboratories, especially for starch. To obtain comparable NSC estimates, we suggest that users can either adopt the reference method given in this publication, or report estimates for a portion of samples using the reference method, and report estimates for a standard reference material. Researchers interested in NSC estimates should work to identify and adopt standard methods.

Towards an advanced assessment of the hydrological vulnerability of forests to climate change-induced drought

Preserving the integrity of the hydraulic system is crucial for survival of trees under dry conditions (Br eda et al., 2006). Maintaining a hydraulic safety margin (HSM) between the water potential (WP) associated with stomatal closure and the WP associated with cavitation can therefore be an important drought resistance trait in many tree species. Based on a broad survey of HSMs across tree species, it was recently concluded that all forest biomes are similarly and strongly threatened by climate changeinduced drought (Choat et al., 2012). We submit that while the prevalence of narrow HSMs across tree species and climates is remarkable, additional factors must be considered to assess vulnerability of trees and forests to drought, that is, the degree to which they are susceptible to, and unable to cope with, adverse impacts of drought (IPCC, 2007). In fact, such prevalence of narrowHSM is likely a basis, and a trigger, for the development of a range of strategies to cope with this situation in light of the everimminent drought stress. Furthermore, settling for an apparently small HSM alone may slow down research into critical aspects of water-use strategies in trees. For example, the accumulating evidence in recent years for rapid recovery from loss of hydraulic conductivity (Brodersen & McElrone, 2013, and references cited therein) indicates a lesser importance of the HSM and any temporary loss of conductivity than suggested. Here, we briefly review key issues in tree strategies to cope with drought that are critical to consider in addition to HSM, if we are to improve predictions of drought related vulnerability of forests.

Impact of forest conversion to agriculture on carbon and nitrogen mineralization in subarctic Alaska

Land-use change is likely to be a major component of global change at high latitudes, potentially causing significant alterations in soil C and N cycling. We addressed the biogeochemical impacts of land-use change in fully replicated black spruce forests and agricultural fields of different ages (following deforestation) and under different management regimes in interior Alaska. Change from forests to cultivated fields increased summer temperatures in surface soil layers by 4–5 °C, and lengthened the season of biological activity by two to three weeks. Decomposition of a common substrate (oat stubble) was enhanced by 25% in fields compared to forests after litter bags were buried for one year. In-situ net N mineralization rates in site-specific soil were similar in forests and fields during summer, but during winter, forests were the only sites where net N immobilization occurred. Field age and management had a significant impact on C and N mineralization. Rates of annual decomposition, soil respiration and summer net N mineralization tended to be lower in young than in old fields and higher in fallow than in planted young fields. To identify the major environmental factors controlling C and N mineralization, soil temperature, moisture and N availability were studied. Decomposition and net N mineralization seemed to be mainly driven by availability of inorganic N. Soil temperature played a role only when comparing forests and fields, but not in field-to-field differences. Results from soil respiration measurements in fields confirmed low sensitivity of heterotrophic respiration, and thus decomposition to temperature. In addition, both soil respiration and net N mineralization were limited by low soil water contents. Our study showed that (1) C and N mineralization are enhanced by forest clearing in subarctic soils, and (2) N availability is more important than soil temperature in controling C and N mineralization following forest clearing. Projecting the biogeochemical impacts of land-use change at high latitudes requires an improved understanding of its interactions with other factors of global change, such as changing climate and N deposition.

Experiments to confront the environmental extremes of climate change

Extreme climate conditions can dramatically alter ecosystems and are expected to become more common in the future; however, our understanding of species and ecosystem responses to extreme conditions is limited. We must meet this challenge by designing experiments that cover broad ranges of environmental stress, extending to levels well beyond those observed currently. Such experiments are important because they can identify physiological, community, and biogeochemical thresholds, and improve our understanding of mechanistic ecological responses to climate extremes. Although natural environmental gradients can be used to observe a range of ecological responses, manipulation experiments – including those that impose drought and heat gradients – are necessary to induce variation beyond common limits. Importantly, manipulation experiments allow for determination of the cause and effect of species and ecosystem threshold responses. We present a rationale and recommendations for conducting extreme experiments that extend beyond the historical and even the predicted ranges of environmental conditions.

The role of mineral nutrients in the increased growth response of tomato plants in solarized soil

Soil solarization is a non-chemical disinfestation technique that frequently promotes plant growth in the absence of known major pathogens, a phenomenon termed increased growth response (IGR). The effect of solarization on plant nutrients and their role in the IGR was studied with tomato plants grown in solarized or non-solarized (control) sandy soil, under controlled conditions. Solarization considerably increased the soil concentrations of water extractable N, K, Ca, Mg and Na at most sites, whereas Cl and DTPA extractable Mn, Zn, Fe and Cu were decreased by the treatment. Plant growth and specific leaf area were enhanced in solarized as well as in N-supplemented control soil. In tomato plants grown in solarized soil, concentrations of most nutrients in the xylem sap, including N, were increased compared to the control, whereas Cl and SO4 levels decreased. The most significant increase in leaf nutrient concentration caused by soil solarization was recorded for N. Furthermore, leaf N concentration was highly and positively correlated with shoot growth. The concentration of Cu increased in leaves from the solarization vs. the control treatment, whereas that of SO4 and Cl decreased, the latter presumably below the critical toxicity level. The correlation between shoot growth and leaf concentration was positive for Cu and inverse for Cl and SO4. In conclusion, we found that soil solarization significantly affects nutrient composition in tomato plants, and provided strong evidence that N, and eventually also Cl, play a major role in IGR.

Biotic degradation at night, abiotic degradation at day: positive feedbacks on litter decomposition in drylands.

The arid and semi-arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO2 ) uptake, which depends largely on plant litter decomposition and the subsequent release of CO2 back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO2 efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO2 efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long-term trend of global carbon sequestration.

Impacts of climate change on biodiversity in Israel: an expert assessment approach

The Mediterranean region is both a global biodiversity hot spot and one of the biomes most strongly affected by human activities. Ecologists and land managers are increasingly required to advise on threats to biodiversity under foreseeable climate change. We used expert surveys to evaluate current understanding and uncertainties regarding climate change impacts on biodiversity in terrestrial, inland freshwater, and marine ecosystems of Israel. Finally, we propose a response strategy toward minimizing these changes. The surveys and the published literature indicated that the main climate change impacts in Israel include ongoing deterioration of freshwater habitats, decline of shrubland and woodland areas, and increased frequency and severity of forest fires. For the Mediterranean Sea, the surveys predict further introduction and establishment of invasive species from the Red Sea, accelerated erosion of coastal rocky habitat, and collapse of coastal rocky platforms. In the Gulf of Aqaba, Red Sea, corals may be resilient to foreseen climate change due to their high tolerance for rising water temperatures. Despite these predictions, science-based knowledge regarding the contribution of management toward minimizing climate change impacts on biodiversity is still lacking. Habitat loss, degradation, and fragmentation are presently the primary and immediate threats to natural ecosystems in Israel. Protection of natural ecosystems, including local refugia, must be intensified to maintain existing biodiversity under pressure from mounting urban development and climate change. This protection policy should include ecological corridors to minimize the consequences of fragmentation of existing natural habitats for species survival. A longer-term strategy should mandate connectivity across environmental and climatic gradients to maintain natural resilience by allowing reorganization of natural ecosystems facing climate change.

Involvement of endogenous gibberellins in the regulation of increased tomato shoot growth in solarized soil

Environmental factors often affect plant growth bymodifying the levels of endogenous gibberellins (GAs).In this study, the involvement of GAs in theregulation of enhanced shoot growth in tomato (Lycopersicon esculentum Mill.) plants grown in soiltreated by solarization (a soil disinfestation method)was investigated. Seedlings at the cotyledonary stagewere transplanted into either solarized or untreatedcontrol soil. Plants in both soils grew free of anydisease symptoms. As soon as four days after planting,seedlings in solarized soil had a higher dry weightcompared to the control. Throughout most of theexperimental period of 18 days, leaf weight ratio washigher in the solarized vs. the control soil. Treatingshoot tips of control plants with 0.1 mg.L-1GA3 resulted in enhanced leaf and stem growth,thus reaching values similar to those of plants grownin solarized soil. The opposite effect was obtained bytreating plants grown in solarized soil with1 mg.L-1 uniconazole, a GA biosynthesisinhibitor. Quantitative GC-MS analyses revealed thatGA1 content in one and two-weeks old transplantsgrown in various solarized soils was up to 1.8 fold,and that GA3 content in two-weeks old plants wasup to five fold the values in the control. Theseincreases were linearly correlated with the increasein leaf dry weight. It was concluded that theincreased quantities of GA1, and eventuallyGA3, play a role in the increased growth oftomato shoots in solarized soil as early as seven daysafter transplanting.