Osmotic and elastic adjustments in cold desert shrubs differing in rooting depth: coping with drought and subzero temperatures

Physiological adjustments to enhance tolerance or avoidance of summer drought and winter freezing were studied in shallow- to deep-rooted Patagonian cold desert shrubs. We measured leaf water potential (Ψ_L), osmotic potential, tissue elasticity, stem hydraulic characteristics, and stomatal conductance (g _S) across species throughout the year, and assessed tissue damage by subzero temperatures during winter. Species behavior was highly dependent on rooting depth. Substantial osmotic adjustment (up to 1.2?MPa) was observed in deep-rooted species exhibiting relatively small seasonal variations in Ψ_L and with access to a more stable water source, but having a large difference between predawn and midday Ψ_L. On the other hand, shallow-rooted species exposed to large seasonal changes in Ψ_L showed limited osmotic adjustment and incomplete stomatal closure, resulting in turgor loss during periods of drought. The bulk leaf tissue elastic modulus (ε) was lower in species with relatively shallow roots. Daily variation in g _S was larger in shallow-rooted species (more than 50?% of its maximum) and was negatively associated with the difference between Ψ_L at the turgor loss point and minimum Ψ_L (safety margin for turgor maintenance). All species increased ε by about 10?MPa during winter. Species with rigid tissue walls exhibited low leaf tissue damage at ?20?°C. Our results suggest that osmotic adjustment was the main water relationship adaptation to cope with drought during summer and spring, particularly in deep-rooted plants, and that adjustments in cell wall rigidity during the winter helped to enhance freezing tolerance.     

Evaluation of Restoration Techniques for the Succulent Karoo, South Africa

Abstract Possible constraints on the passive recovery of bare areas in the Karoo, a semiarid region in South Africa, include inadequate supply of seed, availability of suitable microsites for plant establishment, altered soil properties, and the truncation of key soil biotic processes. Here we investigate the possibility of initiating the restoration of bare areas by soil surface treatments with gypsum (CaSO₄) and/or organic mulch. We also apply an exogenous seed source to test the hypothesis that seed availability limits autogenic recovery. Both gypsum and mulch improved rain water infiltration, gypsum more so than mulch, and both treatments resulted in significantly higher numbers of reseeded seedlings compared with controls. Gypsum also improved the survival of the cohorts of seedlings of the larger seeded Tripteris sinuata. Tripteris showed the highest number of seedlings (maximum count of 150 seedlings/1,000 viable seeds sown) and surviving plants of the three reseeded species, which included two small‐seeded species, Ruschia spinosa and Chaetobromus dregeanus. Throughout the study period significantly higher plant volumes of naturally seeded annuals and perennials were recorded in the gypsum and/or mulch treatments compared with the controls. Germination and emergence of reseeded and naturally seeded plants appears to be determined by the availability of cool season (autumn to spring ) soil moisture, whereas follow‐up rainfall during this time is important for plant survival. Mulching of bare areas in the Succulent Karoo has the potential to re‐create vegetated areas that will further capture and conserve water, soil, and nutrients. Gypsum also showed positive results but might not be a cost‐effective option because of transport costs to these remote arid areas.     

Deuterium labelling of roots provides evidence of deep water access and hydraulic lift by Pinus nigra in a Mediterranean forest of NE Spain

We studied the tree access to deep water sources and the possibility of hydraulic lift from the deep roots of a Pinus nigra tree to the shallow soil layers in a Mediterranean forest of NE Spain. We also studied the use of hydraulically lifted water by neighboring trees, shrubs, and sprouts. We enriched the roots of a large P. nigra (10 m tall) with deuterium by accessing them from a below ground cave. During the next 3 days we measured stable deuterium isotopic composition of xylem sap, shoot predawn and midday water potentials, and the leaf δ¹³C and δ¹⁵N of the P. nigra tree, neighboring Quercus ilex ballota trees and sprouts, and Juniperus oxycedrus shrubs. The study was conducted both in dry summer and in wet spring. In summer, deuterated water absorbed by deep roots of P. nigra appeared in the stem water of neighboring plants and in surface soil. The most δD-enriched plant xylem sap was found in the enriched P. nigra tree, followed by the Q. ilex sprouts, the small Q. ilex trees and the surface soil (15 cm). All these trends disappeared in the wet spring season, when HDO only slightly appeared in the surface soil. The results show that the studied P. nigra tree accesses deep water source and conducts hydraulic lift in this Mediterranean forest in dry summer but not necessarily in wet spring.     

Soil water availability and rooting depth as determinants of hydraulic architecture of Patagonian woody species

Adaptations of species to capture limiting resources is central for understanding structure and function of ecosystems. We studied the water economy of nine woody species differing in rooting depth in a Patagonian shrub steppe from southern Argentina to understand how soil water availability and rooting depth determine their hydraulic architecture. Soil water content and potentials, leaf water potentials (Ψ_Leaf), hydraulic conductivity, wood density (ρ_w), rooting depth, and specific leaf area (SLA) were measured during two summers. Water potentials in the upper soil layers during a summer drought ranged from −2.3 to −3.6 MPa, increasing to −0.05 MPa below 150 cm. Predawn Ψ_Leaf was used as a surrogate of weighted mean soil water potential because no statistical differences in Ψ_Leaf were observed between exposed and covered leaves. Species-specific differences in predawn Ψ_Leaf were consistent with rooting depths. Predawn Ψ_Leaf ranged from −4.0 MPa for shallow rooted shrubs to −1.0 MPa for deep-rooted shrubs, suggesting that the roots of the latter have access to abundant moisture, whereas shallow-rooted shrubs are adapted to use water deposited mainly by small rainfall events. Wood density was a good predictor of hydraulic conductivity and SLA. Overall, we found that shallow rooted species had efficient water transport in terms of high specific and leaf specific hydraulic conductivity, low ρ_w, high SLA and a low minimum Ψ_Leaf that exhibited strong seasonal changes, whereas deeply rooted shrubs maintained similar minimum Ψ_Leaf throughout the year, had stems with high ρ_w and low hydraulic conductivity and leaves with low SLA. These two hydraulic syndromes were the extremes of a continuum with several species occupying different portions of a gradient in hydraulic characteristics. It appears that the marginal cost of having an extensive root system (e.g., high ρ_w and root hydraulic resistance) contributes to low growth rates of the deeply rooted species.     

Plant functional types and ecological strategies in Patagonian forbs

Abstract. We identified four major functional types of forbs in the Patagonian steppe, taking into account phenological and morphological traits: (1) shallow-rooted mesophytic species (annuals), (2) shallow-rooted non-mesophytic species, (3) deep-rooted evergreen species, and (4) deep-rooted deciduous species. The major attributes differentiating these groups were the date at which seasonal growth ended, rooting depth, sprouting depth, distance between shoots of the same plant, and degree of ‘mesophytism’. We used Cluster and Principal Components Analyses to identify the groups, and the attributes determining them. Late-growth-cycle types had deep roots and/or high ‘xerophytism’. Late-cycle-xerophytic types had a great sprouting depth, and late-cycle-deep-rooted types had a great distance between shoots of the same plant. On the basis of current knowledge of the structure and functioning of the Patagonian steppe, we suggested three explanations to account for these correlations. 1. Late-cycle forbs survive summer water deficit if they have xerophytic characteristics that reduce transpiration water losses, and/or they have deep roots that increase water uptake. 2. Sprouting depth results from the shift of active buds to dormant buds at the end of the cycle. Summer forbs have a great sprouting depth because only buds which are located deep in the soil survive hot and dry summers. 3. Distant shoots of summer forbs allow them simultaneously to use the high protection against desiccating winds provided by shrubs, and the ample water availability of bare soil patches. All the functional types of forbs depend on winter water recharge to begin their cycles, but each one completes its cycle by using a different portion of the water resources available in spring and summer.     

End-of-season soil water depletion in relation to growth of herbaceous vegetation in a sub-humid Mediterranean dwarf-shrub community on two contrasting soils

Dwarf-shrub communities of Sarcopoterium spinosum dominate large areas of the landscape on hilly, eastern Mediterranean rangelands. Colonisation of new areas depends on the establishment of seedlings that must compete for water with the ubiquitous annual herbaceous species during the spring-winter growing season and also survive the first hot, dry summer. The present study investigated the role of the herbaceous vegetation patches growing between S. spinosum shrubs on the depletion of soil water during the critical transition period between the cool, rainy season and the dry summer. Dense and sparse herbaceous vegetation stands were established in S. spinosum dwarf-shrub communities by differential use of fertiliser on two contrasting soil types – a terra rossa overlying hard limestone where seedling establishment is low and a pale rendzina overlying a soft chalk substrate where seedling establishment is high. Soil water in the main root zone of the herbaceous vegetation between the shrubs was monitored with protected gypsum block sensors permanently placed at two depths (10 and 33 cm). Soil water depletion during the transition from the wet to the dry season was significantly more rapid under dense vegetation only on the terra rossa soil where the herbaceous vegetation also matured more rapidly than on the rendzina soil. However, in both habitats and under both dense and sparse vegetation, soil water depletion during the transition period left very little available water in the rooting zone of the herbaceous vegetation to maintain shrub seedlings throughout the summer. It was concluded that the difference in shrub seedling establishment success in the two habitats mainly reflects the differences in accessibility of water below the rooting zone of the herbaceous vegetation growing on the two contrasting soil types.     

Effect of deep and shallow root systems on the dynamics of soil inorganic N during 3-year crop rotations

Unused inorganic nitrogen (N_inorg) left in agricultural soils will typically leach to deeper soil layers. If it moves below the root zone it will be lost from the system, but the depth of the root zone depends on the crop species grown. In this experiment we studied the effect of 3-year crop sequences, with different combinations of deep-rooted and shallow-rooted crops, on soil N_inorg dynamics to 2.5 m soil depth and the possibility of crop utilization of N leached to deep soil layers. We grew ten different crop sequences for 3 years. The crops and catch crops grown were selected to allow different sequences of deep-rooted and shallow-rooted crops. Very different rooting depths were obtained, from only 0.5 m (leek), to ∼1.0 m (ryegrass and barley), 1.5 m (red beet), 2.0 m (fodder radish and white cabbage) and more than 2.5 m by the chicory catch crop. The results showed a significant retention of N_inorg within the 2.5 m soil profile from one year to the next, but the retained N had leached to deeper parts of the profile during the winter season. Only little N_inorg was retained over two winter seasons. The retention in the deeper soil layers allowed N_inorg to be taken up by succeeding deep-rooted main crops or catch crops. The effects of crop rooting depth on N_inorg in the subsoil layers from 1.0 to 2.5 m were striking. White cabbage reduced N_inorg below 1.0 m with up to 113 kg N ha^-1 during its growth. Grown after catch crops, leek and red beet left on average 60 kg N ha⁻¹ less below 1.0 m than leek and red beet grown without a preceding catch crop. We conclude that it is possible to design crop rotations with improved nitrogen use efficiency by using the differences in crop rooting patterns; deep-rooted crops or catch crops can be used to recover N_inorg leached after previous crops, and catch crops can be grown before shallow-rooted crops to lift the deep N_inorg up to layers where these crops have their roots.     

Seasonal water use patterns of woody species growing on the continuous dolostone outcrops and nearby thin soils in subtropical China

In karst regions, forests often grow on bedrock outcrops, however the water sources used by the forest vegetation are not known. This study aimed at investigating whether there were seasonal shifts (dry/wet season) of water sources for plants growing on the continuous dolostone outcrops, and comparing their differences with those growing on nearby thin soils in karst areas of southwest China. Rainwater, soil water within 0–30 cm depths, spring water (as a reflection of local deep water sources) and plant xylem water were sampled in March (late dry season) and July (mid rainy season) 2009, respectively. A direct inference approach and the IsoSource mixing model were used to estimate the contributions of different sources to the plant xylem water. On the outcrops, the deciduous tree species Radermachera sinica mainly used deep water sources during the dry season and a mixture of rainwater and deep water sources during the wet season. By contrast, the deciduous small shrub Alchornea trewioides largely relied on recent rainwater during both dry and wet seasons. Three non-deciduous species (Sterculia euosma, Schefflera octophylla and Ficus orthoneura) appear to rely on deep water sources during the wet seasons. In nearby thin soils, R. sinica mainly utilized deep water in the dry season and a mixture of soil water and deep water in the wet season. A. trewioides relied on the same water sources (rainwater-derived soil water) in the different seasons. The above results indicate that inter-specific differences in rooting patterns and leaf phenologies may lead to the differences in the sources of water used by coexisting plant species in karst regions.     

Stable isotopic observation of water use sources of Pinus sylvestris var. mongolica in Horqin Sandy Land, China

Since the late 1950s, Mongolian pine (Pinus sylvestris var. mongolica) has been widely planted for vegetation restoration in arid and semi-arid areas in North China. We used stable isotope signals from precipitation, soil water, and xylem water of Mongolian pine trees, which were planted in early 1980s on sand dunes in the east-southern margin of Horqin Sandy Land, to identify water uptake sources of this tree. (1) Stable ¹⁸O isotope composition of the xylem water exhibited little seasonality, suggesting that the trees use a relatively stable water source; (2) the water source of the pine trees primarily came from a soil depth of 20–60 cm (sampling depth up to 60 cm in this study) and the trees might use groundwater when soil moisture became extremely low; and (3) there was not much difference in water sources used by the pine trees grown at the top of the fixed dune and in the inter dune lowland, although these two sites had 3–8 m elevation difference. This study suggests that it is critical for Mongolian pine trees to access relatively reliable and stable water sources to grow in sandy land habitats, and timely recharging of rainwater to the trees’ rooting depth is requisite for avoiding and/or reducing their degradation caused by water shortage.     

天山林区4种主要灌木夏季水分来源差异

天山林区群落结构相对简单、木本植物种类较少,但天山林区灌木群落中主要木本植物间的水分竞争模式尚不明确,水分利用动态缺乏定量分析。运用稳定同位素技术,对天山林区灌木群落4种主要灌木的茎杆水分及各潜在水源的氢氧稳定同位素组成进行测定,运用IsoSource模型定量分析4种灌木在夏季对各潜在水源的相对利用比例,探讨天山林区灌木群落主要灌木树种水分来源差异及动态变化。结果发现:7月,当浅层土壤含水量充足时,密刺蔷薇、黑果栒子和金丝桃叶绣线菊均大幅度吸收利用浅层土壤水,相对利用比例高于89.3%,异果小檗则相反,即吸收利用各潜在水源(浅层土壤水30.7%、中层土壤水29.4%、深层土壤水25.7%、溪水14.2%,下同);8月,当浅层土壤含水量降低时,密刺蔷薇转移至60-100 cm深层土壤水和溪水,相对利用比例分别为64.8%和27%,黑果栒子和金丝桃叶绣线菊以相似比例吸收利用各潜在水源(33.8%和36.8%、30.9%和29.7%、23.5%和22.3%、11.8%和11.2%),异果小檗则表现出可能吸收利用80-100 cm以下更稳定的深层土壤水;9月,当浅层土壤含水量升高时,4种灌木均大量吸收利用浅层土壤水,相对利用比例高于72.2%。这表明,天山林区灌木群落主要树种可通过可塑性转换水分来源来应对环境水分变异,在时间和空间上有效分割灌丛水源从而减缓对水分资源的竞争压力,从而通过在水分资源利用上的生态位分化促进物种间的共存。     

Microtensiometer technique for in situ measurement of soil matric potential and root water extraction from a sandy soil

The suitability of microtensiometers to measure the spatial variation of soil matric potential and its diurnal change was tested in a pot experiment with pearl millet (Pennisetum americanum [L.] Leeke) in a sandy soil as the soil dried out.The temporal and spatial resolution of this technique allowed precise measurement of soil matric potential and thus estimation of soil water extraction from different compartments as well as from the whole rooting zone. The technique also allowed the measurement of rehydration of plants at night and root water uptake rate per unit soil volume or per unit root length. The precision of determination of root water uptake depended greatly on the accuracy of the estimate of hydraulic conductivity, which was derived from a bare soil and might be different for a cropped soil owing to aggregation induced by the root system. A linear relationship between root length and water uptake was found (r²=0.82), irrespective of variation in soil water content between compartments and despite the variation in root age, xylem differentiation and suberin formation expected to exist between different compartments of the rooting zone. As the experiment was carried out in a range of soil matric potentials between –4 and –30 kPa, drought stress did not occur. Further information at lower soil matric potentials are required, to address questions such as the importance of soil resistance for water uptake, or which portion of the root system has to be stressed to induce hormonal signals to the shoot. The microtensiometer technique can be applied to soil matric potentials up to –80 kPa.     

Increased snowfall weakens complementarity of summer water use by different plant functional groups

Winter snowfall is an important water source for plants during summer in semiarid regions. Snow, rain, soil water, and plant water were sampled for hydrogen and oxygen stable isotopes analyses under control and increased snowfall conditions in the temperate steppe of Inner Mongolia, China. Our study showed that the snowfall contribution to plant water uptake continued throughout the growing season and was detectable even in the late growing season. Snowfall versus rainfall accounted for 30% and 70%, respectively, of the water source for plants, on the basis of hydrogen stable isotope signature (δD) analysis, and accounted for 12% and 88%, respectively, on the basis of oxygen stable isotope signature (δ¹⁸O) analysis. Water use partitioning between topsoil and subsoil was found among species with different rooting depths. Increased snowfall weakened complementarity of plant water use during summer. Our study provides insights into the relationships between precipitation regimes and species interactions in semiarid regions.     

Sources of water used by riparian Eucalyptus camaldulensis overlying highly saline groundwater

Water sources of Eucalyptus camaldulensis Dehn. trees were investigated on a semiarid floodplain in south-eastern Australia. The trees investigated ranged in distance from 0.5 to 40 m from a stream, with electrical conductivity 0.8 dSm^–1, and grew over groundwater with electrical conductivity ranging from 30 to 50 dSm^–1. The sources of water being used by the trees were investigated using the naturally occurring stable isotopes of water and measurements of soil water potential. Xylem water potential and leaf conductance were also examined to identify the trees' response to using these sources of water. Trees at distances greater than about 15 m from the stream used no stream water. The trees used groundwater in summer and a combination of groundwater and rain-derived surface-soil water (0.05–0.15 m depth) in winter. In doing so they suffered water stress at electrical conductivities higher than approximately 40 dSm^–1 (equivalent to approximately –1.4 MPa). Trees adjacent to the stream used stream water directly in summer, but may have used stream water from the soil profile in winter, after the stream had risen and recharged the soil water. E. camaldulensis appeared to be partially opportunistic in the sources of water they used.     

Significant Difference in Hydrogen Isotope Composition Between Xylem and Tissue Water in Populus Euphratica

Deuterium depletions between stem water and source water have been observed in coastal halophyte plants and in multiple species under greenhouse conditions. However, the location(s) of the isotope fractionation is not clear yet and it is uncertain whether deuterium fractionation appears in other natural environments. In this study, through two extensive field campaigns utilizing a common dryland riparian tree species Populus euphratica Oliv., we showed that no significant δ¹⁸O differences were found between water source and various plant components, in accord with previous studies. We also found that no deuterium fractionation occurred during P. euphratica water uptake by comparing the deuterium composition (δD) of groundwater and xylem sap. However, remarkable δD differences (up to 26.4‰) between xylem sap and twig water, root water and core water provided direct evidence that deuterium fractionation occurred between xylem sap and root or stem tissue water. This study indicates that deuterium fractionation could be a common phenomenon in drylands, which has important implications in plant water source identification, palaeoclimate reconstruction based on wood cellulose and evapotranspiration partitioning using δD of stem water.     

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.     

Root Architecture and Root:Shoot Allocation of Shrubs and Saplings in Two Lowland Tropical Forests: Implications for Life-Form Composition1

Entire root systems of saplings of five canopy species and of six shrub and treelet species growing in lowland mixed dipterocarp forest at Andulau, Brunei were excavated and measured. Referring to a prior study at Gigante, Panama, two-way, fixed-factor ANOVAs were used to compare life–forms and sites. Rooting depth and the proportion of root surface area in the upper 20 cm of soil did not differ significantly between life-forms because some treelets/shrubs at Andulau were deep-rooted; all saplings studied were deep-rooted. The root:leaf area ratios of both saplings and treelets/ shrubs at Andulau were significantly higher than those at Gigante. We attribute this strong difference to the lower soil content of available nutrients at Andulau where rainfall shortage is less severe and regular than at Gigante. Available data on life-form composition and mortality rates in large plots are consistent with our proposal that shallow-rooted shrubs and treelets are more vulnerable to drought than deep-rooted life-forms. We suggest that future studies of water use partitioning, wood anatomy, leaf morphology, and associations with neighboring plants would benefit from an explicit examination of their relation to rooting depth.     

Alleviation of soil acidity in Ultisol and Oxisol for corn growth

Malaysian Ultisols and Oxisols are characterized by low pH, high soil solution Al concentration and Ca and/or Mg deficiencies, which are limiting to corn growth. An experiment was conducted to determine the changes in solid and soil solution phase properties of a representative Ultisol and Oxisol following applications of ground magnesium limestone (GML), gypsum and their combinations, and their effects on corn growth. A plot of pAl against lime potential (pH-1/2 pCa) showed that the points were mostly positioned between the theoretical lines for kaolinite-quartz and gibbsite equilibrium, reflecting the kaolinitic-oxidic mineralogy of the Ultisol and Oxisol. Gypsum application increased Al concentration in the soil solutions of the Ultisol, but had no significant effect on that of the Oxisol. The increase in Al concentration in the Ultisol was due to an increase in ionic strength. Gypsum application increased soil solution pH of the Oxisol due to release of OH as a result of ligand exchange between SO₄ and OH ions on the oxides of Fe and/or Al. Exchangeable Al in both soils was reduced by gypsum application. The reduction was associated with solid phase immobilization through alunite formation; the soil solutions of soil samples treated with 2 and 4 t gypsum ha⁻¹ were supersaturated with respect to alunite. Application of GML at 2 t ha⁻¹ together with 1–2t gypsum ha⁻¹ gave high top weight of corn. Relative top weight of corn was positively correlated with a soil solution Mg and Ca/Al concentration ratio, but negatively correlated with soil solution Al concentration. Foliar Al corn was positively correlated with soil solution Al concentration. Soil solution Al and Mg concentrations, and Ca/Al concentration ratio can be used as indices of soil acidity in Ultisols and Oxisols. ei]{gnB E}{fnClothier}     

Water relations of the root hemiparasite Olax phyllanthi (Labill) R.Br. (Olacaceae) and its multiple hosts

Summary Water relations of the root hemiparasite Olax phyllanthi were compared with those of its major species of hosts in natural habitat in coastal heath near Denmark, SW Australia. Leaf water potentials of Olax during winter were 0.4 to 1.4 MPa lower (more negative) than those of all (29) non parasitic host species examined. During the dry summer months (January to March), shallow-rooted hosts developed water potentials up to 3 MPa lower than those of Olax, and were accordingly rated as no longer accessible as a source of water to the hemiparasite. By contrast, deep-rooted hosts, with access to the water table, showed water potentials less negative than Olax, and haustorial contacts retained with these apparently enabled continued extraction of water and nutrients throughout the summer. Three other species of root hemiparasites parasitized by Olax, but not themselves parasitizing Olax, showed leaf water potentials throughout the year very close to, and mostly slightly more negative than those of Olax. Nocturnal measurements of leaf water potential in winter (July and August) in soil at field capacity (water potential –0.006 MPa) showed maintenance of a 0.5–0.8 MPa potential difference between Olax and a range of common host species. By dawn most hosts had equilibrated with the water potential of the soil, whereas both exposed and bagged Olax plants recorded potentials of –0.8 MPa. Daytime rates of transpiration and photosynthesis of Olax were less than those of a range of common hosts, but water use efficiencies were not consistently different between hemiparasite and hosts. This was reflected in almost identical mean values for carbon isotope ratio (¹³C/¹²C) between Olax (mean value –27.0) and thirteen frequently exploited hosts ( value –27.1). The results are discussed in relation to published information on other angiosperm hemiparasites.     

Differential water resource use by herbaceous and woody plant life-forms in a shortgrass steppe community

We conducted a study to test the predictions of Walter's two-layer model in the shortgrass steppe of northeastern Colorado. The model suggests that grasses and woody plants use water resources from different layers of the soil profile. Four plant removal treatments were applied in the spring of 1996 within a plant community codominated by Atriplex canescens (a C₄ shrub) and Bouteloua gracilis (a C₄ grass). During the subsequent growing season, soil water content was monitored to a depth of 180 cm. In addition, stem and leaf tissue of Atriplex, Bouteloua and the streamside tree Populus sargentii were collected monthly during the growing seasons of 1995 and 1996 for analysis of the δ¹⁸O value of plant stem water (for comparison with potential water sources) and the δ¹³C value of leaves (as an indicator of plant water status). Selective removal of shrubs did not significantly increase water storage at any depth in the measured soil profile. Selective removal of the herbaceous understory (mainly grasses) increased water storage in the top 60 cm of the soil. Some of this water gradually percolated to lower layers, where it was utilized by the shrubs. Based on stem water δ¹⁸O values, grasses were exclusively using spring and summer rain extracted from the uppermost soil layers. In contrast, trees were exclusively using groundwater, and the consistent δ¹³C values of tree leaves over the course of the summer indicated no seasonal changes in gas exchange and therefore minimal water stress in this life-form. Based on anecdotal rooting-depth information and initial measurements of stem water δ¹⁸O, shrubs may have also had access to groundwater. However, their overall δ¹⁸O values indicated that they mainly used water from spring and summer precipitation events, extracted from subsurface soil layers. These findings indicate that the diversity of life-forms found in this shortgrass steppe community may be a function of the spatial partitioning of soil water resources, and their differential use by grasses, shrubs, and trees. Consequently, our findings support the two-layer model in a broad sense, but indicate a relatively flexible strategy of water acquisition by shrubs. Received: 23 December 1997 / Accepted: 16 September 1998     

Seasonal water use characteristics of tall wheatgrass [Agropyron elongatum (Host) Beauv.] in a saline environment

It is essential to characterize the water use of plants that have potential for the stabilization of rising saline ground-water which could lead to increases in soil salinity. In this study, several techniques were used to determine the seasonal water use characteristics of the perennial grass Agropyron elongatum (tall wheatgrass) growing in a moderately saline, dryland environment with a fluctuating shallow groundwater table varying in electrical conductivity between 0 and 10 dS m^?1. Soil conditions were examined in terms of water potential measurements, plant water sources were identified using a stable isotope of water (deuterium, ²H) and evapotranspiration was estimated using hydrological and ventilated chamber methods. Seasonal changes in soil water potential were caused by salt accumulation and soil moisture leading to changes in plant water availability, particularly in the surface soil region over summer and autumn. Evapotranspiration in A. elongatum was high over summer during the peak period of growth (4 mm d^?1), with evidence of water use from groundwater and from specific regions of the soil profile. Evapotranspiration was low during the period that A. elongatum was partially senescent in autumn (< 0.5 mm d^?1) and the lowest leaf water potential of -3 MPa that was measured occurred during this period of moderate water stress. Intermediate levels of water use (1.5 mm d^?1) were measured during winter when the entire soil profile was available for water uptake. Based on physiological characteristics, including aspects of summer water use, root morphology and salt tolerance in A. elongatum, we conclude that this species is suitable for stabilizing the level of moderately saline groundwater in parts of southern Australia, which could reduce the potential for soil salinization.