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Seasonal and annual changes in soil respiration in relation to soil temperature, water potential and trenching.

Soil respiration (rs), soil temperature (Ts) and volumetric soil water content were measured in a balsam fir (Abies balsamea (L.) Mill.) ecosystem from 1998 to 2001. Seasonal variation in root and microbial respiration, and covariation in abiotic factors confounded interpretation of the effects of Ts and soil water potential (Psis) on rs. To minimize the confounding effect of temperature, we analyzed the effect of Psis on rs during the summers of 1998-2000 when changes in Ts were slight. Soil respiration declined 25-50% in response to modest water stress (minimum Psis of -0.6 to -0.2 MPa), and between years, there was substantial variation in the relationship between rs and Psis. In the summer of 2000, 2-m2 plots were subjected to drought for 1 month and other plots were irrigated. The relationship between summertime rs and Psis in the experimental plots was similar to that estimated from the survey data obtained during the same summer. In late spring and early autumn of 2001, 2-m2 trenched and untrenched plots were subjected to drought or exposed to rainfall. It was dry in the early autumn and there was severe soil drying (Psis of -10 MPa in untrenched plots and -2 MPa in trenched plots). In spring, rs in untrenched plots responded more to modest water stress than rs in trenched plots, indicating that root respiration is more sensitive than microbial respiration to water stress at this time of year. The response to abiotic factors differed significantly between spring and autumn in untrenched plots but not in trenched plots, indicating that root activity was greater in early autumn than in late spring, and that roots acclimated to the sustained, severe water stress experienced before and during the autumn.

Abies↗

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine.

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.

Carbon Dioxide↗

Environmental controls on sap flow in a northern hardwood forest.

Our objective was to gain a detailed understanding of how photosynthetically active radiation (PAR), vapor pressure deficit (D) and soil water interact to control transpiration in the dominant canopy species of a mixed hardwood forest in northern Lower Michigan. An improved understanding of how these environmental factors affect whole-tree water use in unmanaged ecosystems is necessary in assessing the consequences of climate change on the terrestrial water cycle. We used continuously heated sap flow sensors to measure transpiration in mature trees of four species during two successive drought events. The measurements were scaled to the stand level for comparison with eddy covariance estimates of ecosystem water flux (Fw). Photosynthetically active radiation and D together explained 82% of the daytime hourly variation in plot-level transpiration, and low soil water content generally resulted in increased stomatal sensitivity to increasing D. There were also species-specific responses to drought. Quercus rubra L. showed low water use during both dry and wet conditions, and during periods of high D. Among the study species, Acer rubrum L. showed the greatest degree of stomatal closure in response to low soil water availability. Moderate increases in stomatal sensitivity to D during dry periods were observed in Populus grandidentata Michx. and Betula papyrifera Marsh. Sap flow scaled to the plot level and Fw demonstrated similar temporal patterns of water loss suggesting that the mechanisms controlling sap flow of an individual tree also control ecosystem evapotranspiration. However, the absolute magnitude of scaled sap flow estimates was consistently lower than Fw. We conclude that species-specific responses to PAR, D and soil water content are key elements to understanding current and future water fluxes in this ecosystem.

Acer↗

Influence of soil texture on hydraulic properties and water relations of a dominant warm-desert phreatophyte.

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.

Circadian Rhythm↗

Preformation in vegetative buds of Prunus persica: factors influencing number of leaf primordia in overwintering buds.

We investigated the influence of bud position, cultivar, tree age, tree carbohydrate status, sampling date, drought and light exposure on the number of leaf primordia formed in dormant vegetative peach buds (Prunus persica (L.) Batsch) relative to the number of primordia formed after bud break (neoformed). During winter dormancy, vegetative peach buds from California and Italy were dissected and the number of leaf primordia recorded. Between leaf drop and bud break, the number of leaf primordia doubled from about five to about 10. Parent shoot length, number of nodes on the parent shoot, cross-sectional area of the parent shoot, bud position along the parent shoot and bud cross-sectional area were correlated with the number of leaf primordia. Previous season light exposure, drought and tree carbohydrate status did not affect the number of leaf primordia present. The number of leaf primordia differed significantly among peach varieties and tree ages at leaf drop, but not at bud break. Our results indicate that neoformation accounted for all shoot growth beyond about 10 nodes. The predominance of neoformed shoot growth in peach allows this species great plasticity in its response to current-season conditions.

California↗

Seasonal fluctuations and temperature dependence of leaf gas exchange parameters of co-occurring evergreen and deciduous trees in a temperate broad-leaved forest.

Seasonal fluctuations in leaf gas exchange parameters were investigated in three evergreen (Quercus glauca Thunb., Cinnamomum camphora Sieb. and Castanopsis cuspidata Schottky) and one deciduous (Quercus serrata Thunb.) co-occurring, dominant tree species in a temperate broad-leaved forest. Dark respiration rate (Rn), maximum carboxylation rate (Vcmax) and stomatal coefficient (m), the ratio of stomatal conductance to net assimilation rate after adjustment to the vapor pressure deficit and internal carbon dioxide (CO2) concentration, were derived inversely from instantaneous field gas exchange data (one-point method). The normalized values of Rn and Vcmax at the reference temperature of 25 degrees C (Rn25, Vcmax25) and their temperature dependencies (Delta Ha(Rn), Delta Ha(Vcmax)) were analyzed. Parameter Vcmax25 ranged from 24.0-40.3 micromol m(-2) s(-1) and Delta Ha(Vcmax) ranged from 29.1- 67.0 kJ mol(-1). Parameter Rn25 ranged from 0.6-1.4 micromol m(-2) s(-1) and Delta Ha(Rn) ranged from 47.4-95.4 kJ mol(-1). The stomatal coefficient ranged from 7.2-8.2. For the three evergreen trees, a single set of Vcmax25 and Rn25 parameters and temperature dependence curves produced satisfactory estimates of carbon uptake throughout the year, except during the period of simultaneous leaf fall and leaf expansion, which occurs in April and May. In the deciduous oak, declines in Vcmax25 were observed after summer, along with changes in Vcmax25 and Rn25 during the leaf expansion period. In all species, variation in m during periods of leaf expansion and drought should be considered in modeling studies. We conclude that the changes in normalized gas exchange parameters during periods of leaf expansion and drought need to be considered when modeling carbon uptake of evergreen broad-leaved species.

Climate↗

Variation in growth and osmotic regulation of roots of water-stressed maritime pine (Pinus pinaster Ait.) provenances.

Hydroponically cultivated Pinus pinaster Ait. seedlings of a drought-sensitive population from France (Landes) and of a more drought-adapted population from Morocco (Tamjoute) were subjected to a progressive increase in water stress by additions of an osmoticum (polyethylene glycol 600) to the nutrient solution. The final osmotic potential (Psi(ms)) of the nutrient solution was achieved over a period of up to 6 days, and ranged from -0.03 (control, no added osmoticum) to -0.8 MPa. In the 6 days during which water stress was imposed, roots elongated faster in the Moroccan provenance than in the French provenance, but the applied water deficits did not inhibit root elongation in either population. Among treatments, root dry weight per unit root length, total root dry weight and root/shoot dry weight ratio increased with decreasing Psi(ms) in both provenances. Both the water potential (Psi(w)) of the roots (apices) and the water potential difference between the roots and the nutrient solution decreased as Psi(ms) decreased. The reduction in Psi(w) was matched by a decrease of comparable magnitude in cell osmotic potential (Psi(pi)) so that root turgor was unaffected by the Psi(ms). Osmotic adjustment was greater, however, in the Moroccan provenance than in the French provenance. Consequently, under the osmotically imposed water stress, the water potential difference between root and nutrient solution was greater in the Moroccan provenance than in the French provenance. Similar changes in plant water relations were observed when seedlings were grown in drying sand.

Journal Article↗

The plant-growth-promoting rhizobacterium Paenibacillus polymyxa induces changes in Arabidopsis thaliana gene expression: a possible connection between biotic and abiotic stress responses.

This paper addresses changes in plant gene expression induced by inoculation with plant-growth-promoting rhizobacteria (PGPR). A gnotobiotic system was established with Arabidopsis thaliana as model plant, and isolates of Paenibacillus polymyxa as PGPR. Subsequent challenge by either the pathogen Erwinia carotovora (biotic stress) or induction of drought (abiotic stress) indicated that inoculated plants were more resistant than control plants. With RNA differential display on parallel RNA preparations from P. polymyxa-treated or untreated plants, changes in gene expression were investigated. From a small number of candidate sequences obtained by this approach, one mRNA segment showed a strong inoculation-dependent increase in abundance. The corresponding gene was identified as ERD15, previously identified to be drought stress responsive. Quantification of mRNA levels of several stress-responsive genes indicated that P. polymyxa induced mild biotic stress. This suggests that genes and/or gene classes associated with plant defenses against abiotic and biotic stress may be co-regulated. Implications of the effects of PGPR on the induction of plant defense pathways are discussed.

Arabidopsis↗

Water availability affects reproduction in deer mice.

Water restriction impaired sperm production in deer mice, a seasonally breeding mammal that encounters aperiodic droughts in its natural habitat throughout North America. Water-induced spermatogenic responses were sorted into three categories based upon epididymal sperm numbers: aspermic, oligospermic, and euspermic. Average gonadal mass was reduced after 10 wk of limited water consumption. Inter-individual variation in gonadal response to a simulated drought was similar to phenotypic differences in reproductive function in response to other environmental cues that direct annual reproductive cycles. Our findings suggest that water availability may act as a cue to suppress gametogenesis in deer mice independently from food, temperature, and day length.

Animals↗

Coalitions of relatives and reproductive skew in cooperatively breeding white-winged choughs.

We used DNA fingerprinting to examine reproductive skew in cooperatively breeding white-winged choughs, Corcorax melanorhamphos, which live in groups of up to 20 individuals. Before a severe drought, groups that had been stable for multiple years were characterized by long-term monogamy involving a single breeding pair (high skew). After the drought, new groups formed from the amalgamation of multiple individuals and coalitions of relatives. At most one member of each faction succeeded in breeding, such that skew was dependent on the number of unrelated factions, and not group size. In the new groups, dominant males and females with supporting relatives were always successful. Whereas most females without support also gained breeding positions, many males without family support failed to breed. Thus subordinates gain indirect fitness by first helping related males to secure a breeding position, and then helping to raise their young. Our study demonstrates the advantage of operating in coalitions, and suggests that the acquisition of future allies may be a major benefit of helping behaviour in this species.

Animals↗

Determinants of the abundance of invasive annual weeds: community structure and non-equilibrium dynamics.

The dynamics of an annual pasture community are described from a five-year experimental and monitoring study. The community was dominated by two grasses (Lolium rigidum and Vulpia bromoides) and a legume (Trifolium subterraneum). Fits of population dynamic models to per capita rates of population change indicate that interactions between the grasses were generally strong, while interactions between the grasses and legumes were weaker. Most, but not all, of the net effects of competition on population growth could be attributed to interactions occurring during plant growth. Phase-plane analysis indicated that, for a constant environment, a joint equilibrium of the two grasses is unstable since interspecific competition between Lolium and Vulpia is stronger than intraspecific competition. Consequently, the community will tend to a mixture of only one or other of the grass species and T. subterraneum, depending on the founding composition of the pasture. Analysis of data taken from a year in which a drought occurred (1993-1994) demonstrated profound effects on all three species. Modelling of the long-term impacts of the effects of repeated droughts showed that disturbance of this form overrides the founder effect observed under constant conditions. Consequently, Vulpia is ultimately able to invade any mixture of the other species in environments where stochastic disturbances occur.

Ecosystem↗

Soil moisture mediates association between the winter North Atlantic Oscillation and summer growth in the Park Grass Experiment.

Several aspects of terrestrial ecosystems are known to be associated with the North Atlantic Oscillation (NAO) through effects of the NAO on winter climate, but recently the winter NAO has also been shown to be correlated with the following summer climate, including drought. Since drought is a major factor determining grassland primary productivity, the hypothesis was tested that the winter NAO is associated with summer herbage growth through soil moisture availability, using data from the Park Grass Experiment at Rothamsted, UK between 1960 and 1999. The herbage growth rate, mean daily rainfall, mean daily potential evapotranspiration (PE) and the mean and maximum potential soil moisture deficit (PSMD) were calculated between the two annual cuts in early summer and autumn for the unlimed, unfertilized plots. Mean and maximum PSMD were more highly correlated than rainfall or PE with herbage growth rate. Regression analysis showed that the natural logarithm of the herbage growth rate approximately halved for a 250 mm increase in maximum PSMD over the range 50-485 mm. The maximum PSMD was moderately correlated with the preceding winter NAO, with a positive winter NAO index associated with greater maximum PSMD. A positive winter NAO index was also associated with low herbage growth rate, accounting for 22% of the interannual variation in the growth rate. It was concluded that the association between the winter NAO and summer herbage growth rate is mediated by the PSMD in summer.

Poaceae↗

Environmental refugees: a growing phenomenon of the 21st century.

There is a new phenomenon in the global arena: environmental refugees. These are people who can no longer gain a secure livelihood in their homelands because of drought, soil erosion, desertification, deforestation and other environmental problems, together with the associated problems of population pressures and profound poverty. In their desperation, these people feel they have no alternative but to seek sanctuary elsewhere, however hazardous the attempt. Not all of them have fled their countries, many being internally displaced. But all have abandoned their homelands on a semi-permanent if not permanent basis, with little hope of a foreseeable return. In 1995, environmental refugees totalled at least 25 million people, compared with 27 million traditional refugees (people fleeing political oppression, religious persecution and ethnic troubles). The total number of environmental refugees could well double by the year 2010, and increase steadily for a good while thereafter as growing numbers of impoverished people press ever harder on overloaded environments. When global warming takes hold, there could be as many as 200 million people overtaken by sea-level rise and coastal flooding, by disruptions of monsoon systems and other rainfall regimes, and by droughts of unprecedented severity and duration.

Disasters↗

Sources or sinks? The responses of tropical forests to current and future climate and atmospheric composition.

How tropical rainforests are responding to the ongoing global changes in atmospheric composition and climate is little studied and poorly understood. Although rising atmospheric carbon dioxide (CO2) could enhance forest productivity, increased temperatures and drought are likely to diminish it. The limited field data have produced conflicting views of the net impacts of these changes so far. One set of studies has seemed to point to enhanced carbon uptake; however, questions have arisen about these findings, and recent experiments with tropical forest trees indicate carbon saturation of canopy leaves and no biomass increase under enhanced CO2. Other field observations indicate decreased forest productivity and increased tree mortality in recent years of peak temperatures and drought (strong El Niño episodes). To determine current climatic responses of forests around the world tropics will require careful annual monitoring of ecosystem performance in representative forests. To develop the necessary process-level understanding of these responses will require intensified experimentation at the whole-tree and stand levels. Finally, a more complete understanding of tropical rainforest carbon cycling is needed for determining whether these ecosystems are carbon sinks or sources now, and how this status might change during the next century.

Atmosphere↗

Inactivation of AtRac1 by abscisic acid is essential for stomatal closure.

Plant water homeostasis is maintained by the phytohormone abscisic acid (ABA), which triggers stomatal pore closure in response to drought stress. We identified the Arabidopsis small guanosine triphosphatase (GTPase) protein AtRac1 as a central component in the ABA-mediated stomatal closure process. ABA treatment induced inactivation of AtRac GTPases and disruption of the guard cell actin cytoskeleton. In contrast, in the ABA-insensitive mutant abi1-1, which is impaired in stomatal closure, neither AtRac inactivation nor actin cytoskeleton disruption was observed on ABA treatment. These observations indicate that AtRac1 inactivation is a limiting step in the ABA-signaling cascade leading to stomatal closure. Consistent with these findings, expression of a dominant-positive mutant of AtRac1 blocked the ABA-mediated effects on actin cytoskeleton and stomatal closure in wild-type plants, whereas expression of a dominant-negative AtRac1 mutant recapitulated the ABA effects in the absence of the hormone. Moreover, the dominant-negative form of AtRac1 could also restore stomatal closure in abi1-1. These results define AtRac1 as a central element for plant adaptation to drought.

Abscisic Acid↗

Winter at the alpine timberline. Why does embolism occur in norway spruce but not in stone pine?

Conifers growing at the alpine timberline are exposed to frost drought and freeze-thaw cycles during winter-stress factors known to induce embolism in tree xylem. The two dominant species of the European Central Alps timberline were studied: Norway spruce (Picea abies [L.] Karst) and stone pine (Pinus cembra), which usually reaches higher altitudes. We hypothesized to find embolism only at the timberline and to observe less embolism in stone pine than in Norway spruce due to avoidance mechanisms. Seasonal courses of embolism and water potential were studied at 1,700 and 2,100 m during two winter seasons and correlated to vulnerability (to drought-induced embolism), leaf conductance, and micrometeorological data. Embolism was observed only at the timberline and only in Norway spruce (up to 49.2% loss of conductivity). Conductivity losses corresponded to low water potentials (down to -3.5 MPa) but also to the number of freeze-thaw events indicating both stress factors to contribute to embolism induction. Decreasing embolism rates-probably due to refilling-were observed already in winter. Stone pine did not exhibit an adapted vulnerability (50% loss of conductivity at -3.5 MPa) but avoided critical potentials (minimum -2.3 MPa): Cuticulare conductance was 3.5-fold lower than in Norway spruce, and angles between needles and axes were found to decrease in dehydrating branches. The extent of conductivity losses in Norway spruce and the spectrum of avoidance and recovery mechanisms in both species indicates winter embolism to be relevant for tree line formation.

Acclimatization↗

1-Aminocyclopropane-1-Carboxylic Acid Transported from Roots to Shoots Promotes Leaf Abscission in Cleopatra Mandarin (Citrus reshni Hort. ex Tan.) Seedlings Rehydrated after Water Stress.

The effect of water stress and subsequent rehydration on 1-aminocyclopropane-1-carboxylic acid (ACC) content, ACC synthase activity, ethylene production, and leaf abscission was studied in Cleopatra mandarin (Citrus reshni Hort. ex Tan.) seedlings. Leaf abscission occurred when drought-stressed plants were allowed to rehydrate, whereas no abscission was observed in plants under water stress conditions. In roots of water-stressed plants, a high ACC accumulation and an increase in ACC synthase activity were observed. Neither increase in ACC content nor significant ethylene production were detected in leaves of water-stressed plants. After rehydration, a sharp rise in ACC content and ethylene production was observed in leaves of water-stressed plants. Content of ACC in xylem fluid was 10-fold higher in plants rehydrated for 2 h after water stress than in nonstressed plants. Leaf abscission induced by rehydration after drought stress was inhibited when roots or shoots were treated before water stress with aminooxyacetic acid (AOA, inhibitor of ACC synthase) or cobalt ion (inhibitor of ethylene-forming enzyme), respectively. However, AOA treatments to shoots did not suppress leaf abscission. The data indicate that water stress promotes ACC synthesis in roots of Cleopatra mandarin seedlings. Rehydration of plants results in ACC transport to the shoots, where it is oxidized to ethylene. Subsequently, this ethylene induces leaf abscission.

Journal Article↗

Genotypic variability in vulnerability of leaf xylem to cavitation in water-stressed and well-irrigated sugarcane.

Genotypic variability in vulnerability of leaf xylem to water-stress-induced cavitation was determined in four sugarcane (Saccharum sp.) clones using detached leaf segments in a hydraulic conductivity apparatus. Vulnerability curves were constructed by plotting the percentage of maximum conductivity versus leaf water potential (psi(I)) and fitting curves using a Weibull function. The psi(I) at which each clone lost 10, 50, and 80% of maximum conductivity was determined. Maximum conductivity per unit of leaf width was positively associated with metaxylem vessel diameter. The commercial clone H65-7052 exhibited the highest and the nondomesticated S. spontaneum exhibited the lowest conductivity. All four clones lost substantial conductivity at values of psi(I) less negative than -1.4 MPa, but H65-7052 was able to maintain 50% conductivity to lower psi(I) than the other clones. S. spontaneum sustained the most negative psi(I) (-1.99 MPa) before reaching the 80% conductivity loss point. Clone H69-8235 was consistently the most vulnerable to initial loss of conductivity. These vulnerability functions were used in conjunction with field measurements of psi(I) to estimate diurnal losses in leaf hydraulic conductivity under irrigated and droughted conditions. H69-8235 lost up to 50% of its conductivity during the day, even when well irrigated, and more than 80% when subjected to drought. The other clones exhibited lower conductivity losses. These losses are apparently reversed overnight by root pressure. Despite their close genetic relationships, these clones exhibited large differences in conductivity, in the vulnerability of their xylem to cavitation, and in gas exchange behavior. The potential for altering water relations by selecting for particular hydraulic characteristics is discussed.

Journal Article↗