A 10-year demographic study of rabbitbrush (Chrysothamnus nauseosus): growth,survival and water limitation

A population of 126 adult rabbitbrush (Chrysothamnus nauseosus var. consimilis) was followed for 10 consecutive years from 1983 through 1992 in a sanddune ecosytem on the north shore of Mono Lake, California. The study area received an average (1981–1992) of 160 mm total annual precipitation of which 55 mm fell from April through October, making this site drier than other known sites in the Great Basin desert where C. nauseosus occurs. Cover of all shrubs was 18.0%, and that of C. nauseosus 7.5%, in the 3077 m² study plot. Water was limiting for this population of C. nauseosus, exerting large effects on new canopy growth and reproductive effort in a given year, on survival, and on establishment of juveniles into the adult population. Growth and survivorship of the adult shrubs were significantly correlated with the depth to ground water, which varied from 3.6 to 6.0 m below the surface during the study, and not with any measure of precipitation in a given year or adjacent years. Shrubs in this population exhibited a net decrease in size between 1983 and 1992. The region experienced a significant drought from 1987 onwards, with some increase in precipitation after 1990. Mortality of shrubs began in 1987 and reached 22% by 1992. Individuals that died were smaller on average initially than individuals surviving throughout the study: 50% or more of the individuals having 4000 cm² of canopy or less at the start of the study died by 1992. Elevation (m msl) of each shrub in the study area, presumably correlated with depth to ground water, had no effect on growth but a weak effect on survival during the study period. Specific microsite had significant effects, with shrubs on steep slopes being smaller, having more negative growth slopes, and being more likely to die during the drought compared to individuals occurring on flatter gradients. Recruiment of juveniles into the population did not occur after 1985. Competition between shrubs could not be definitively demonstrated during the study, but only indirect tests for competition were used. Adult C. nauseosus and adults of all species combined (Sarcobatus vermiculatus with 9.6% cover, Tetradymia tetrameres with 0.8% cover and Chrysothamnus viscidiflorus with 0.02% cover) were randomly distributed, failing to support the hypothesis of competition if competition for water results in a uniform distribution of shrubs. Moreover, comparison of the spatial dispersion of C. nauseosus individuals before (1985) and after (1992) mortality did not reveal competitive thinning toward uniformity. On the other hand, larger individuals were farther from their nearest neighbor than were smaller individuals, an effect postulated to result from competitive thinning.     

Limited uptake of summer precipitation by bigtooth maple (Acer grandidentatum Nutt) and Gambel's oak (Quereus gambelii Nutt)

Winter and spring precipitation that saturates to deep soil layers precedes summer droughts in the Intermountain West. Occasional summer convection storms relieve summer drought, but are infrequent and unreliable from year to year, leading to the hypothesis that dominant tree species might not invest limited carbon reserves to surface roots to take up summer precipitation in these regions. We compared the hydrogen (D) and oxygen (¹⁸O) isotope ratios of winter, spring and summer precipitation to that of xylem sap water in Acer grandidentatum and Quercus gambelii, two dominant trees of this region. By this method we could identify water sources utilized throughout the growing season. Xylem D and ¹⁸O values changed significantly when each species leafed-out; this change was not associated with changes in either soil or plant water status (as measured by predawn and midday water potentials). This shift is apparently related increased transpirational flux, which may flush out residual stem water from the previous growing season. D values of xylem sap of both species matched winter precipitation input values throughout most of the summer, indicating a reliance on deep-soil moisture sources throughout the growing season. Mature Q. gambelii did not take up summer precipitation, whereas A. grandidentatum responded slightly to the largest summer rain event. Small trees of both species, particularly A. grandidentatum, showed a limited uptake of summer rains.     

Response of net ecosystem gas exchange to a simulated precipitation pulse in a semi-arid grassland: the role of native versus non-native grasses and soil texture

Physiological activity and structural dynamics in arid and semi-arid ecosystems are driven by discrete inputs or pulses of growing season precipitation. Here we describe the short-term dynamics of ecosystem physiology in experimental stands of native (Heteropogon contortus) and invasive (Eragrostis lehmanniana) grasses to an irrigation pulse across two geomorphic surfaces with distinctly different soils: a Pleistocene-aged surface with high clay content in a strongly horizonated soil, and a Holocene-aged surface with low clay content in homogenously structured soils. We evaluated whole-ecosystem and leaf-level CO₂ and H₂O exchange, soil CO₂ efflux, along with plant and soil water status to understand potential constraints on whole-ecosystem carbon exchange during the initiation of the summer monsoon season. Prior to the irrigation pulse, both invasive and native grasses had less negative pre-dawn water potentials ( _pd), greater leaf photosynthetic rates (A _net) and stomatal conductance (g _s), and greater rates of net ecosystem carbon exchange (NEE) on the Pleistocene surface than on the Holocene. Twenty-four hours following the experimental application of a 39 mm irrigation pulse, soil CO₂ efflux increased leading to all plots losing CO₂ to the atmosphere over the course of a day. Invasive species stands had greater evapotranspiration rates (ET) immediately following the precipitation pulse than did native stands, while maximum instantaneous NEE increased for both species and surfaces at roughly the same rate. The differential ET patterns through time were correlated with an earlier decline in NEE in the invasive species as compared to the native species plots. Plots with invasive species accumulated between 5% and 33% of the carbon that plots with the native species accumulated over the 15-day pulse period. Taken together, these results indicate that system CO₂ efflux (both the physical displacement of soil CO₂ by water along with plant and microbial respiration) strongly controls whole-ecosystem carbon exchange during precipitation pulses. Since CO₂ and H₂O loss to the atmosphere was partially driven by species effects on soil microclimate, understanding the mechanistic relationships between the soil characteristics, plant ecophysiological responses, and canopy structural dynamics will be important for understanding the effects of shifting precipitation and vegetation patterns in semi-arid environments.     

Differences in water-use strategies along an aridity gradient between two coexisting desert shrubs (<Emphasis Type="Italic">Reaumuria soongorica</Emphasis> and <Emphasis Type="Italic">Nitraria sphaerocarpa</Emphasis>): isotopic approaches with physiological evidence

Background and aims

Understanding the responses of different plant species to changes in available water sources is critical for accurately modeling and predicting species dynamics. Our study aimed to explore whether there were differences in water-use strategies between the two coexisting shrubs (Reaumuria soongorica and Nitraria sphaerocarpa) in response to different amounts of summer precipitation.

Methods

We conducted 3 years of field observations at three sites along an aridity gradient from the middle to lower reaches of the Heihe River basin, northwestern China. Stable oxygen composition (δ¹⁸O) in plant xylem water, soil water and groundwater were analyzed concurrently with ecophysiological measurements at monthly intervals during the growing seasons.

Results

Water source for coexisting R. soongorica and N. sphaerocarpa did not differ at the sites with high precipitation, but significantly differed in more arid locations. The N. sphaerocarpa was more sensitive to summer precipitation than R. soongorica in terms of predawn water potential, stomatal conductance and foliage carbon-isotope discrimination.

Conclusions

The plants relying on groundwater maintained consistent water use strategies, but not plants that took up precipitation-derived water. We also found that the difference in water source uptake between the coexisting species was more apparent in more arid locations.

     

Plant N capture from pulses: effects of pulse size,growth rate,and other soil resources

In arid ecosystems, the ability to rapidly capture nitrogen (N) from brief pulses is expected to influence plant growth, survival, and competitive ability. Theory and data suggest that N capture from pulses should depend on plant growth rate and availability of other limiting resources. Theory also predicts trade-offs in plant stress tolerance and ability to capture N from different size pulses. We injected K¹⁵NO₃, to simulate small and large N pulses at three different times during the growing season into soil around the co-dominant Great Basin species Sarcobatus vermiculatus, Chrysothamnus nauseosus ssp. consimilis, and Distichlis spicata. Soils were amended with water and P in a partial factorial design. As predicted, all study species showed a comparable decline in N capture from large pulses through the season as growth rates slowed. Surprisingly, however, water and P availability differentially influenced the ability of these species to capture N from pulses. Distichlis N capture increased up to tenfold with water addition while Chrysothamnus N capture increased up to threefold with P addition. Sarcobatus N capture was not affected by water or P availability. Opposite to our prediction, Sarcobatus, the most stress tolerant species, captured less N from small pulses but more N from large pulses relative to the other species. These observations suggest that variation in N pulse timing and size can interact with variable soil water and P supply to determine how N is partitioned among co-existing Great Basin species.     

Differential uptake of summer precipitation among co-occurring trees and shrubs in a pinyon-juniper woodland

Measurements of the ratio of deuterium to hydrogen (D/H) in stem xylem water were used to determine the relative uptake of summer precipitation by four co-occurring plant species in southern Utah. The species compared included two trees, Juniperus osteosperma and Pinus edulis, and two shrubs, Artemisia tridentata and Chrysothamnus nauseousus. There were significant differences among species in the relative use of summer precipitation. Chrysothamnus nauseosus had stem water D/H ratios in May through August 1990 that were not significantly different from that of groundwater. In contrast, the other three species had stem water D/H ratios that were intermediate between the groundwater value and summer precipitation values, indicating that a mixture of both precipitation and groundwater was being used by these species. The two tree species generally had higher D/H values than did A. tridentata indicating a higher average uptake of summer precipitation, although the roots of J. osteosperma and P. edulis may not be as responsive to small precipitation events as A. tridentata. There was a strong negative correlation between stem water D/H ratios and predawn water potential, which suggests a relationship between plant rooting pattern and water source use. In addition, water-use efficiency during photosynthetic gas exchange, calculated from leaf carbon isotope composition, differed among species and was strongly correlated with differences in the relative uptake of summer precipitation.     

Sensitivity of water relations and photosynthesis to summer precipitation pulses for <Emphasis Type="Italic">Artemisia tridentata</Emphasis> and <Emphasis Type="Italic">Purshia tridentata</Emphasis>

For much of the western USA, precipitation occurs in pulses, the nature of which determine soil water potential and plant physiological performance. This research utilized three experiments to examine the sensitivity of photosynthesis and water relations for two widespread Great Basin Desert shrub species, Artemisia tridentata (which has both deep and shallow roots) and Purshia tridentata (which reportedly has only deep roots), to (1) variation in pulse magnitude size, (2) the kinetics of responses to pulses, and (3) the relationship between pulse-size and antecedent soil water content. At the study site in the southwestern Great Basin Desert, USA, summer rainfall exhibits a greater frequency of larger-sized events, and longer inter-pulse intervals, compared to annual patterns. Compared to pre-watering values, stem water potential initially increased by about 2.00 MPa for A. tridentata and 1.00 MPa for P. tridentata following watering to simulate an 11.5 mm rainfall pulse. For the same water addition, stomatal conductance increased by 0.3 mol m⁻² s⁻¹ and photosynthetic CO₂ assimilation increased 8-fold for A. tridentata and 6-fold for P. tridentata. Water potential and photosynthetic gas exchange were maximal for both species 2–3 days following a pulse addition. In comparison to P. tridentata, the increase in photosynthesis for A. tridentata was more pronounced for plants treated incrementally with several small pulses compared to plants treated with one pulse of an equivalent total volume. The results indicate that both species can respond to a range of summer rainfall pulse magnitudes within about 2 days, with A. tridentata generally exhibiting larger responses in comparison to the co-dominant shrub species P. tridentata, which at this study site does indeed have shallow roots. In a future climate, the timing and magnitude of summer rainfall pulses will determine the extent to which these two species undergo changes in water status and photosynthetic carbon uptake, with implications for their fitness.     

Responses of restored grassland ditch vegetation to hydrological changes, 1989–1992

The impact of hydrological manipulation of an unfertilised, Dutch peat grassland area on plant species composition on ditch banks and in ditchwater was studied. The hydrological manipulation involved raising the groundwater level by admitting nutrient-poor, Ca-rich groundwater in one compartment, and by retaining precipitation in another compartment. A third compartment served as control.The plant species composition showed significant correlations with the following hydrochemical parameters: in bank vegetation with K⁺ concentration, and winter and summer groundwater levels, water depth and elevation; in aquatic vegetation with pH, the concentrations of Cl^- organic-C and NH₄ ⁺ water temperature and elevation.The number of terrestrial plant species increased after compartmentation from 97 to 122; 16 submerged and floating species were found. Most new terrestrial species probably emerged in response to cessation of fertilization and biomass removal, since they showed no preference for any compartment. Five species showed preference for the groundwater compartment and two for the rainwater compartment. Of the new terrestrial plants, seven were relatively rare: Carex panicea, C. oederi, C. pallescens, C. vesicaria, Galium uliginosum, Juncus acutiflorus and Stellaria uliginosa. Of the aquatic plants, eight were relatively rare: Chara globularis, two Callitriche spp., four potamogetonaceae and Ranunculus circinatus.This study indicates that hydrological manipulation of grassland systems in which fertilisation has ceased has profound effects on the vegetation in the ditches of these systems. Bank vegetation responds more slowly and to other hydrochemical factors than aquatic vegetation. Short-term responses in terms of increase in diversity of vegetation pattern and species richness are promising. Long-term responses are not yet known.     

Ecophysiology of selected tree species in different plant communities at the periphery of the Atlantic Forest of SE-Brazil II. Spatial and ontogenetic dynamics in Andira legalis, a deciduous legume tree

Andira legalis Vell. Toledo (Leguminosae, Papilionoideae) is a deciduous tree of the coastal vegetation of Brazil. It was studied in the restinga ecosystems of the Atlantic coast along a moisture gradient of open dry restinga (annual precipitation 800 mm, water table high at 0.9–1.2 m but partially saline), intermediate restinga (annual precipitation 1164 mm, water table low at 2–3 m) and wet restinga (annual precipitation 1000 mm, water table high at 0.5–1 m). In addition, a comparison of the ecophysiological performance was made of A. legalis in the wet restinga and a dune forest (water table low at 2–6 m). Plants of A. legalis at a given site and even within clonal stands show considerable phenological plasticity, where especially timing of leaf shedding and new bud break, leaf expansion and maturation varies between neighbouring individuals and all stages can be observed at the same time. We used analyses of soluble carbohydrate carbon compounds, soluble non-protein nitrogen compounds, stable isotope signatures (¹³C, ¹⁵N) and parameters of fluorescence of chlorophyll a for describing metabolic relations and photosynthetic behaviour and to deduce possible variations in ecophysiological strategies in response to site conditions. The phenological plasticity is described in relation to predominant metabolite levels in leaves, roots, xylem and phloem, but its ecological advantage—perhaps related to staggered resource acquisition and susceptibility to predators—remains an open question. For overall ecophysiological performance among the environmental factors root access to the ground water table appeared to be more essential than significantly different precipitation regimes, as plants in the intermediate restinga appeared to be more stressed than those in the other two open restingas. Shading in the dune forest was also an important environmental constraint.     

Summer rain pulses may stimulate a CO2 release rather than absorption in desert halophyte communities

Background and aims

The response of plants and soil to rain pulses determines seasonal variations in the exchange of materials and energy at the ecosystem scale in arid and semi-arid regions. We assessed how the ecosystem carbon exchange (NEE) of desert halophyte communities of different plant functional-types responds to summer precipitation pulses in Tamarix and Haloxylon communities.

Methods

Plant water status, photosynthetic gas exchange, soil respiration and net ecosystem carbon exchange were measured to test the hypothesis that high physiological sensitivity may induce a greater changes in NEE resulting from the summer precipitation pulses in Haloxylon community.

Results

Plant water status and photosynthetic assimilation did not differ before and after summer precipitation pulses in either community. In contrast, soil respiration and NEE responded strongly to summer precipitation events in both communities. At the ecosystem level, precipitation pulses induced a pulse of CO₂ release, rather than absorption. The NEE response to summer precipitation was less in the deep-rooted Tamarix community, compared to the shallow-rooted Haloxylon community, which was even converted into a carbon source after summer precipitation inputs. As a result, the effect of summer precipitation inputs on soil respiration was more important than the plant (carbon assimilation) response in determining the ecosystem response to episodic precipitation pulses.     

Limitations on Sphagnum growth and net primary production in the foothills of the Philip Smith Mountains,Alaska

Summary In the foothills of the Philip Smith Mountains, Brooks Range, Alaska, tussock tundra occurs on rolling hills and in valleys that were shaped by Pleistocene glaciations. During the 1986 and 1987 summer seasons, Sphagnum growth and production were determined in water tracks on tundra slopes that acted to channel water flow to the valley bottom stream and in intertrack tundra areas that were relatively homogeneous with respect to downslope drainage. Measurements were made under ambient environmental conditions and on mosses receiving supplemental irrigation in each area. Growth rate for Sphagnum spp. (cm shoot length increase/day) was low and relatively constant in intertrack tundra and highest but quite variable in water tracks. A strong negative correlation was found between Sphagnum spp. growth rate and solar irradiance in the shady environment below Salix canopies in the water tracks. Estimates of net annual dry weight (DW) production for Sphagnum spp. ranged from 0.10 g DW dm^-2 yr^-1 in intertrack tundra vegetation to 1.64 g DW dm^-2 yr^-1 in well-shaded water tracks. Experimental water additions had little effect on growth and production in intertrack tundra and well-developed water tracks, but significantly increased growth in a weakly-developed water track community. Low production over large areas of tundra slopes may occur due to presence of slow growing species resistant to dessication in intertrack tundra as opposed to rapidly growing less compact species within the limited extent of water tracks. We hypothesize that species capable of rapid growth occur also in weakly-developed water tracks, and that these are water-limited more often than plants occurring in well-developed water track situations. Where experienced, high light intensity may additionally limit growth due to photoinhibition.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

Isotopic and anatomical signals for interpreting fire-related responses in Pinus halepensis

Key message

Our study aims to define isotopic and anatomical responses to fires of P. halepensis . Main results: decrease in tree growth and relative conductivity and increase in water use efficiency.

Abstract

We investigated the ecophysiological responses of a Pinus halepensis Mill. stand surviving two wildfires in southern France. Basal area, isotope composition and anatomical traits were analysed before and after fires, using tree rings to assess the ecological responses of trees to heat-related damage. The years were determined based on the presence of fire scars. Stable isotopes (δ¹³C and δ¹⁸O) were measured in tree rings before and after the “fire years”. Anatomical observations allowed qualitative analysis of the scar region and quantification of tracheid size in tree rings before and after the fire years. Relative and percentage conductivity of earlywood and latewood far from the woundwood were estimated. Results showed a decrease in tree growth after the fire events accompanied by an increase in ¹³C-derived water use efficiency (WUE_i) and a decrease in relative conductivity. The positive relationship between δ¹³C and δ¹⁸O suggested that both isotopic variations are mostly driven by changes in stomatal conductance following fire events. P. halepensis proved to be a strong isohydric species, able to survive frequent fires with temporary ecophysiological modifications and anatomical adaptations. Our findings afford new insights into post-fire survival strategies of this species in an environment where fires are predicted to increase in frequency during the twenty-first century.     

Monosoonal precipitation responses of shrubs in a cold desert community on the Colorado Plateau

South-eastern Utah forms a northern border for the region currently influenced by the Arizona monosoonal system, which feeds moisture and summer precipitation into western North America. One major consequence predicted by global climate change scenarios is an intensification of monosoonal (summer) precipitation in the aridland areas of the western United States. We examined the capacity of dominant perennial shrubs in a Colorado Plateau cold desert ecosystem of southern Utah, United States, to use summer moisture inputs. We simulated increases of 25 and 50 mm summer rain events on Atriplex canescens, Artemisia filifolia, Chrysothamnus nauseosus, Coleogyne ramosissima, and Vanclevea stylosa, in July and September with an isotopically enriched water (enriched in deuterium but not ¹⁸O). The uptake of this artificial water source was estimated by analyzing hydrogen and oxygen isotope ratios of stem water. The predawn and midday xylem water potentials and foliar carbon isotope discrimination were measured to estimate changes in water status and water-use efficiency. At. canescens and Ch. nauseosus showed little if any uptake of summer rains in either July or September. The predawn and midday xylem water potentials for control and treatment plants of these two species were not significantly different from each other. For A. filifolia and V. stylosa, up to 50% of xylem water was from the simulated summer rain, but the predawn and midday xylem water potentials were not significantly affected by the additional summer moisture input. In contrast, C. ramosissima showed significant uptake of the simulated summer rain (>50% of xylem water was from the artificial summer rain) and an increase in both predawn and midday water potentials. The percent uptake of simulated summer rain was greater when those rains were applied in September than in July, implying that high soil temperature in midsummer may in some way inhibit water uptake. Foliar carbon isotope discrimination increased significantly in the three shrubs taking up simulated summer rain, but pre-treatment differences in the absolute discrimination values were maintained among species. The ecological implications of our results are discussed in terms of the dynamics of this desert community in response to changes in the frequency and dependability of summer rains that might be associated with a northward shift in the Arizona monsoon boundary.     

Effects of plant size and water relations on gas exchange and growth of the desert shrub Larrea tridentata

Larrea tridentata is a xerophytic evergreen shrub, dominant in the arid regions of the southwestern United States. We examined relationships between gasexchange characteristics, plant and soil water relations, and growth responses of large versus small shrubs of L. tridentata over the course of a summer growing season in the Chihuahuan Desert of southern New Mexico, USA. The soil wetting front did not reach 0.6 m, and soils at depths of 0.6 and 0.9 m remained dry throughout the summer, suggesting that L. tridentata extracts water largely from soil near the surface. Surface soil layers (<0.3 m) were drier under large plants, but predawn xylem water potentials were similar for both plant sizes suggesting some access to deeper soil moisture reserves by large plants. Stem elongation rates were about 40% less in large, reproductively active shrubs than in small, reproductively inactive shrubs. Maximal net photosynthetic rates (P_max) occurred in early summer (21.3 mol m^-2 s^-1), when pre-dawn xylem water potential (XWP) reached ca. -1 MPa. Although both shrub sizes exhibited similar responses to environmental factors, small shrubs recovered faster from short-term drought, when pre-dawn XWP reached about -4.5 MPa and P_max decreased to only ca. 20% of unstressed levels. Gas exchange measurements yielded a strong relationship between stomatal conductance and photosynthesis, and the relationship between leaf-to-air vapor pressure deficit and stomatal conductance was found to be influenced by pre-dawn XWP. Our results indicate that stomatal responses to water stress and vapor pressure deficit are important in determining rates of carbon gain and water loss in L. tridentata.     

Exceptional Photosynthetic Performance of Capparis spinosa L. Under Adverse Conditions of Mediterranean summer

Diurnal and seasonal fluctuations in water potential (), stomatal conductance (g _s), transpiration rate (E), and net photosynthetic rate (P _N) were monitored in Capparis spinosa L., a Mediterranean plant growing during summer, i.e. at the period considered the most stressful for local plant life. In spite of the complete absence of rain, exhibited a modest drop at midday (–2.7 MPa), but was fully recovered overnight, indicating sufficient access to water sources. The stomata remained open throughout the day and season and the high E resulted in leaf temperatures up to 3.9 °C below air temperature. Additionally, P _N of the fully exposed leaves was higher than 25 mol m^–2 s^–1 for more than 10 h per day throughout the summer growth period. No symptoms of photooxidative stress were shown, as judged by maximum photosystem 2 photochemical efficiency (F_v/F_m) and the function of xanthophyll cycle. Indeed, diurnal inter-conversions of the xanthophyll cycle components were modest during the summer and a more intensive function of the cycle was only evident during leaf senescence in autumn. In comparison with a semi-deciduous and an evergreen sclerophyll co-existing in the same ecosystem, C. spinosa assimilated up to 3.4 times more CO₂ per m² during its growth period (May to October) and up to 1.8 times more on an annual basis.     

Is the bark of shining gum (Eucalyptus nitens) a sun or a shade leaf?

Photoinhibition and pigment composition of green stem tissues of field-grown adult Eucalyptus nitens were measured on clear spring days with low morning temperatures—conditions that cause photoinhibition in leaves of many plant species. The sun-exposed (north-facing) bark contained less chlorophyll a+b (531 vs 748 mol m^–2), neoxanthin (29 vs 41), and -carotene (54 vs 73), more xanthophyll cycle pigments per unit surface area and per unit chlorophyll (71 vs 53 mol m^–2 and 141 vs 66 mmol mol^–1 chlorophyll), and less lutein per unit chlorophyll (239 vs 190) than the shaded (southern) side. Maximum electron flow rates were 60 mol m^–2 s^–1 on the sun-exposed side, and about 10 mol m^–2 s^–1 on the shaded side. F_v/F_m was always lower than 0.8 on the sun-exposed side and the de-epoxidation state (DEPS) of the xanthophyll cycle was dominated by zeaxanthin in midday samples. F_v/F_m increased quickly after darkening, but DEPS recovered more slowly to 40% overnight. This suggested that rapidly reversible pH-dependent quenching was responsible for the bulk of changes in PS II efficiency. F_v/F_m remained below 0.8 overnight, which may well be indicative of photo-damage to PSII. In contrast, DEPS of the shaded side was lower, and F_v/F_m was higher, than for the sun-exposed side. We conclude that E. nitens chlorenchyma on the sun-exposed stem side has a photosynthetic pigment composition similar to sun leaves and it experiences significant photoinhibition in the field.     

Δ13C and tree-ring width reflect different drought responses in Quercus ilex and Pinus halepensis

Holm oak (Quercus ilex L.) and Aleppo pine (Pinus halepensis Mill) are representative of two different functional types of trees extensively found in the Mediterranean: evergreen sclerophyllous and drought-adapted conifers. The former is considered a partially drought-tolerant species, whereas the latter is a typically drought-avoiding, water-saving species. We postulated that contrasting strategies in response to water deficits in Q. ilex and P. halepensis would lead to a differential sensitivity to changes in water availability. To test this hypothesis, we compared the response of both species in growth rate (measured as radial increments) and intrinsic water use efficiency [WUE_i, as inferred from carbon isotope discrimination (¹³C) in wood samples] among sites from different provenance regions in NE Spain. We found significant differences in ¹³C and growth among provenance regions, partly explained by contrasting water availability. Wood ¹³C was positively related with precipitation and the ratio between precipitation and potential evapotranspiration (P / E). However, these relationships were stronger in P. halepensis (for P / E, r ²=0.67, P <0.001) than in Q. ilex (r ²=0.42, P <0.01). In addition, radial growth was positively related with precipitation and ¹³C in P. halepensis (r ²=0.32 and r ²=0.35, respectively, P <0.01), but not in Q. ilex. We concluded that P. halepensis was more sensitive than Q. ilex to water availability, showing faster increase in WUE_i in response to water stress. We also found that the effect of north/south aspect on ¹³C and growth was site-specific, and unrelated to climatic variables.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Response of understory vegetation to variable tree mortality following a mountain pine beetle epidemic in lodgepole pine stands in northern Utah

We analyzed the long-term dynamics of aboveground biomass ofLeymus chinense steppe in relation to interannual variation of precipitation and temperature during 1980–1989 at levels of community, growth form and species in the Xilin river basin, Inner Mongolia Autonomous Region, China. Annual aboveground net primary production (ANPP) varied from 154.00 g m^-2 yr^-1 in 1980 to 318.59 g m^-2 yr^-1 in 1988, with a mean of 248.63 g m^-2 yr^-1 and the coefficient of variation of 25%. ANPP was not significantly correlated to annual precipitation and total precipitation during April–September atp0.05 level, but precipitation in May and August accounted for 69% of interannual variation of ANPP. The means of rain use efficiency and water use efficiency ofL. chinense steppe were 8.1 kg DM ha^-1 mm^-1 yr^-1 and 0.89 mg DM g^-1 H₂O respectively. Aboveground biomass of various growth forms and species had different response patterns to interannual variation of precipitation and temperature. Monthly and seasonal distribution of precipitation and temperature were the key controls of aboveground biomass of species.     

Partitioning of water and nitrogen in co-occurring Mediterranean woody shrub species of different evolutionary history

We studied the interspecific and intraspecific variation in the development of water stress and in the use of different water and nitrogen sources during the spring (wet season) and summer (dry season) in a shrub community in NE Spain. We measured shoot water potentials, stable deuterium isotopic composition (D) of xylem sap, leaf mass per area, leaf N and C concentrations, gas exchange, leaf ¹³C, and leaf ¹⁵N of the dominant species (Quercus coccifera, Arbutus unedo, Pistacia lentiscus, Erica multiflora, Globularia alypum). The D, the ¹³C and the shoot water potential values showed diurnal, seasonal, intraspecific and interspecific variation in the source and use of water. There was also seasonal, intraspecific and interspecific variation in the foliar ¹⁵N and N concentrations. In summer, some species (A. unedo, P. lentiscus and E. multiflora) presented significantly different D values in morning and afternoon measurements likely indicating that they used different sources of water during the day, and a dual root system in these species. We conjecture that dew may be one of these water sources. Species predawn water potential was negatively correlated with species xylem water D. There was also a positive correlation between ¹³C and D in P. lentiscus, species for which we took additional samples from nearby sites. These results suggest that the access to water from greater depths allowed the maintenance of more favourable plant water supply. Multivariate principal component analysis based on the studied hydrological and isotope variables clearly separated the seasons (wet spring and dry summer) and the species. The species resulted separated according to their evolutionary history (Pre-Mediterranean and Mediterranean) and the associated root and functional traits. These results show water (and nitrogen) partitioning among coexisting species of the same functional type (Mediterranean woody shrubs). They also show the great intraspecific plasticity of responses to resource availability.