Serious complications in experiments in which UV doses are effected by using different lamp heights.

Many experiments examining plant responses to enhanced ultraviolet-B radiation (280-315nm) simply compare an enhanced UV-B treatment with ambient UV-B (or no UV-B radiation in most greenhouse and controlled-environment studies). Some more detailed experiments utilize multiple levels of UV-B radiation. A number of different techniques have been used to adjust the UV dose. One common technique is to place racks of fluorescent UV-emitting lamps at different heights above the plant canopy. However, the lamps and associated support structure cast shadows on the plant bed below. We calculated one example of the sequence of shade intervals for two common heights of lamp racks and show the patterns and duration of shade which the plants receive is distributed differently over the course of the day for different heights of the lamp racks. We also conducted a greenhouse experiment with plants (canola, sunflower and maize) grown under unenergized lamp racks suspended at the same two heights above the canopy. Growth characteristics differed in unpredictable ways between plants grown under the two heights of lamp racks. These differences could enhance or obscure potential UV-B effects. Also, differences in leaf mass per unit foliage area, which were observed in this experiment, could contribute to differences in plant UV-B sensitivity. We recommend the use of other techniques for achieving multiple doses of UV-B radiation. These range from simple and inexpensive approaches (e.g., wrapping individual fluorescent tubes in layers of a neutral-density filter such as cheese cloth) to more technical and expensive alternatives (e.g., electronically modulated lamp control systems). These choices should be determined according to the goals of the particular experiment.

Root turnover and relocation in the soil profile in response to seasonal soil water variation in a natural stand of Utah juniper (Juniperus osteosperma).

Juniper species are noted for long-lived foliage, low and persistent gas exchange activity and drought tolerance. Because leaves and roots of the same species are thought to be similar in structure and life history, we hypothesized that Juniperus osteosperma (Torr.) Little (Utah juniper) fine roots would reflect the persistent aboveground foliage characteristic of this species. We monitored fine roots, less than 1 mm in diameter, by minirhizotron imaging to a depth of 150 cm over two growing seasons from April 2002 to December 2003. We measured fine root numbers, lengths and diameters, and noted the time of birth and death of root segments. We correlated our root data with soil water potential measured by thermocouple psychrometry and ecosystem evapotranspiration measured by ecosystem eddy flux. Median fine root lifespan, determined by the Kaplan-Meier product-limit method, was about one year, much less than foliage lifespan estimates of more than five years. Yet, roots of juniper live much longer than those of other Great Basin species. The median survivorship of shallow and deep roots was 144 and 448 days, respectively. Production of new roots was observed during periods of favorable soil water potential and there was a seasonal progression of increased new roots and root length during the warm season toward lower soil depths with root loss in the upper soil layers. This was also reflected in water extraction which progressed to greater soil depths later in the warm season. Aboveground, rates of ecosystem evapotranspiration decreased with decreasing soil water potentials in a similar manner in both 2002 and 2003, reflecting the relocation of roots to available water at depth. Juniper exhibited a flexible root depth distribution throughout the 20 months of this study, indicating the potential to respond to shifting soil water resources despite long fine root lifespans.

Fine root distribution and persistence under field conditions of three co-occurring Great Basin species of different life form.

Fine roots of an annual grass, a perennial grass and a perennial shrub were examined. Based on life histories and tissue composition, we expected the greatest root persistence for the shrub and shortest for the annual grass. Roots were observed with minirhizotrons over 2 yr for number, length and diameter changes. A Cox proportional hazard regression correlated root persistence with soil water, depth, diameter and date of production. In 2001, grass roots had similar persistence times, but shrub roots had the shortest. In 2002, the annual had the longest median root persistence, the perennial grass intermediate and the perennial shrub had the shortest. All species responded similarly to the magnitude of seasonal precipitation; root numbers increased with favorable soil moisture and disappeared with drying; fewer, thinner roots at greater soil depths were found in the drier year (2001). Root persistence increased with soil moisture, diameter and earlier appearance in the spring. Plasticity in root morphology and placement was influenced by water availability, yet persistence was surprisingly contrary to expectations.

Plant responses to current solar ultraviolet-B radiation and to supplemented solar ultraviolet-B radiation simulating ozone depletion: an experimental comparison.

Field experiments assessing UV-B effects on plants have been conducted using two contrasting techniques: supplementation of solar UV-B with radiation from fluorescent UV lamps and the exclusion of solar UV-B with filters. We compared these two approaches by growing lettuce and oat simultaneously under three conditions: UV-B exclusion, near-ambient UV-B (control) and UV-B supplementation (simulating a 30% ozone depletion). This permitted computation of "solar UV-B" and "supplemental UV-B" effects. Microclimate and photosynthetically active radiation were the same under the two treatments and the control. Excluding UV-B changed total UV-B radiation more than did supplementing UV-B, but the UV-B supplementation contained more "biologically effective" shortwave radiation. For oat, solar UV-B had a greater effect than supplemental UV-B on main shoot leaf area and main shoot mass, but supplemental UV-B had a greater effect on leaf and tiller number and UV-B-absorbing compounds. For lettuce, growth and stomatal density generally responded similarly to both solar UV-B and supplemented UV-B radiation, but UV-absorbing compounds responded more to supplemental UV-B, as in oat. Because of the marked spectral differences between the techniques, experiments using UV-B exclusion are most suited to assessing effects of present-day UV-B radiation, whereas UV-B supplementation experiments are most appropriate for addressing the ozone depletion issue.

Reduction of solar UV-B mediates changes in the Sphagnum capitulum microenvironment and the peatland microfungal community.

The influence of near-ambient and reduced solar UV-B radiation on a peatland microfungal community was assessed by exposing experimental plots to UV-selective filtration. Replicate plots were covered with special plastic films to effect treatments of near-ambient and attenuated solar UV-B. The microfungal community from the top 1 cm of Sphagnum capitulum in a Tierra del Fuego peatland was censused throughout three growing seasons, between 1999 and 2002. Sphagnum capitula under near-ambient UV-B were more compressed and held more water than capitula under reduced UV-B. This water had a greater conductivity and was more acidic under near-ambient UV-B, as would be expected with increased leaching from the Sphagnum leaves. Nine regularly occurring hyphal fungi from the peatland were identified, at least to genus. Over three field seasons, no treatment effect on total fungal colony abundance was recorded, but individual species abundance was increased (Mortierella alpina), decreased (Penicillium frequentans), or was unaffected (P. thomii, Aureobasidium) by near-ambient UV-B. Species richness was also slightly lower under near-ambient UV-B. These treatment differences were smaller than seasonal or inter-annual fluctuations in abundance and species richness. In a growth chamber experiment, lamp UV-B treatments indicated that realistic fluxes of UV-B can inhibit fungal growth in some species. In addition to this direct UV-B effect, we suggest that changes in the peatland fungal community under near-ambient solar UV-B may also result from increased nutrient and moisture availability in the Sphagnum capitulum. The subtle nature of the responses of peatland fungi to solar UV-B suggests that most fungal species we encountered are well adapted to current solar UV-B fluxes in Tierra del Fuego.

Field testing of biological spectral weighting functions for induction of UV-absorbing compounds in higher plants.

Action spectra are typically used as biological spectral weighting functions (BSWF) in biological research on the stratospheric ozone depletion issue. Despite their critical role in determining the amount of UV supplied in experiments, there has been only limited testing of different functions under realistic field conditions. Here, we calculate effective radiation according to five published BSWF and evaluate the appropriateness of these BSWF in representing the induction of UV-absorbing compounds. Experiments were carried out in the field using both ultraviolet-B radiation (280-320 nm) supplementation and selective filtering of solar UV radiation. For the four species tested, BSWF that extend into the ultraviolet-A radiation (320-400 nm) (UV-A) with moderate effectiveness best represented the observed results. When compared with the commonly used generalized plant response, these BSWF suggest that simulations of ozone depletion will require more radiation than in the past experiments. However, they imply lower radiation supplements than a new plant growth BSWF that has a greater emphasis on UV-A wavelengths.

Root responses and nitrogen acquisition by Artemisia tridentata and Agropyron desertorum following small summer rainfall events.

Resources in the Great Basin of western North America often occur in pulses, and plant species must rapidly respond to temporary increases in water and nutrients during the growing season. A field study was conducted to evaluate below ground responses of Artemisia tridentata and Agropyron desertorum, common Great Basin shrub and grass species, respectively, to simulated 5-mm (typical summer rain) and 15-mm (large summer rain) summer rainfall events. The simulated rainfall was labeled with K(15)NO(3) so that timing of plant nitrogen uptake could be monitored. In addition, soil NH(4)(+) and NO(3)(-) concentrations and physiological uptake capacities for NO(3)(-) and NH(4)(+) were determined before and after the rainfall events. Root growth in the top 15 cm of soil was monitored using a minirhizotron system. Surprisingly, there was no difference in the amount of labeled N acquired in response to the two rainfall amounts by either species during the 7-day sample period. However, there were differences between species in the timing of labeled N uptake. The N label was detected in above ground tissue of Agropyron within 1 h of the simulated rainfall events, but not until 24 h after the rainfall in Artemisia. For both Agropyron and Artemisia, root uptake capacity was similarly affected by the 5-mm and 15-mm rainfall. There was, however, a greater increase in uptake capacity for NH(4)(+) than for NO(3)(-), and the 15-mm event resulted in a longer response. No root growth occurred in either species in response to either rainfall event during this 8-day period. The results of this study indicate that these species are capable of utilizing nitrogen pulses following even small summer rainfall events during the most stressful period of the summer and further emphasize the importance of small precipitation events in arid systems.

Terrestrial ecosystems, increased solar ultraviolet radiation and interactions with other climatic change factors.

Based on research to date, we can state some expectations about terrestrial ecosystem response as several elements of global climate change develop in coming decades. Higher plant species will vary considerably in their response to elevated UV-B radiation, but the most common general effects are reductions in height of plants, decreased shoot mass if ozone reduction is severe, increased quantities of some phenolics in plant tissues and, perhaps, reductions in foliage area. In some cases, the common growth responses may be lessened by increasing CO2 concentrations. However, changes in chemistry of plant tissues will generally not be reversed by elevated CO2. Among other things, changes in plant tissue chemistry induced by enhanced UV-B may reduce consumption of plant tissues by insects and other herbivores, although occasionally consumption may be increased. Pathogen attack on plants may be increased or decreased as a consequence of elevated UV-B, in combination with other climatic changes. This may be affected both by alterations in plant chemistry and direct damage to some pathogens. Water limitation may decrease the sensitivity of some agricultural plants to UV-B, but for vegetation in other habitats, this may not apply. With global warming, the repair of some types of UV damage may be improved, but several other interactions between warming and enhanced UV-B may occur. For example, even though warming may lead to fewer killing frosts, with enhanced UV-B and elevated CO2 levels, some plant species may have increased sensitivity to frost damage.

Six years of solar UV-B manipulations affect growth of Sphagnum and vascular plants in a Tierra del Fuego peatland.

• Tierra del Fuego is subject to increases in solar UV-B radiation in the austral spring and summer due to ozone depletion. • Plastic films were used to filter solar UV-B radiation over peatland plots through six field seasons, resulting in near-ambient (c. 90%) and reduced (c. 17%) solar UV-B treatments. • As in the first three field seasons of treatments, near-ambient UV-B caused reduced height growth but had no effect on biomass production of the moss Sphagnum magellanicum. It reduced leaf and rhizome growth of Tetroncium magellanicum. Height growth and morphology of Empetrum rubrum and Nothofagus antarctica were only affected by solar UV-B during the fourth to sixth field seasons. There was also a decrease in Tetroncium leaf nitrogen under near-ambient UV-B. • Growth of Sphagnum was less affected than that of most emergent vascular plants. This enabled the Sphagnum mat to engulf more Nothofagus, and limit the escape of Empetrum under near-ambient UV-B. Yet, differences in the response of species to solar UV-B were not expressed as changes in plant community composition.

Carbon acquisition and water use in a Northern Utah Juniperus osteosperma (Utah juniper) population.

Water use and carbon acquisition were examined in a northern Utah population of Juniperus osteosperma (Torr.) Little. Leaf-level carbon assimilation, which was greatest in the spring and autumn, was limited by soil water availability. Gas exchange, plant water potential and tissue hydrogen stable isotopic ratio (deltaD) data suggested that plants responded rapidly to summer rain events. Based on a leaf area index of 1.4, leaf-level water use and carbon acquisition scaled to canopy-level means of 0.59 mm day(-1) and 0.13 mol m(-2) ground surface day(-1), respectively. Patterns of soil water potential indicated that J. osteosperma dries the soil from the surface downward to a depth of about 1 m. Hydraulic redistribution is a significant process in soil water dynamics. Eddy covariance data indicated a mean evapotranspiration rate of 0.85 mm day(-1) from March to October 2001, during which period the juniper population at the eddy flux site was a net source of CO2 (3.9 mol m(-2) ground area). We discuss these results in relation to the rapid range expansion of juniper species during the past century.

A meta-analysis of plant field studies simulating stratospheric ozone depletion.

The potential effects of increased ultraviolet-B radiation (UV-B, 280-320 nm) simulating stratospheric ozone depletion in field studies with vascular plants have previously been summarized only in narrative literature reviews. In this quantitative synthesis, we have assessed the significance of solar UV-B enhancement for ten commonly measured variables involving leaf pigmentation, plant growth and morphology, and photosynthesis using meta-analytic statistical methods. Of 103 papers published between 1976 and mid-1999 from field studies, more than 450 reports from 62 papers were included in the database. Effects of UV-B were most apparent for the case of UV-B-absorbing compounds with an average increase of approximately 10% across all studies when comparing the ambient solar UV-B control to the treatment (involving ambient UV-B plus a UV-B supplement from special UV lamps). Some morphological parameters such as plant height and leaf mass per area showed little or no response to enhanced UV-B. Leaf photosynthetic processes (leaf gas exchange and chlorophyll fluorescence) and the concentration of photosynthetic pigments (total chlorophylls and carotenoids) were also not affected. Shoot biomass and leaf area showed modest decreases under UV-B enhancement. The reduction in shoot biomass occurred only under very high levels of simulated ozone depletion and leaf area was affected only when studies inappropriately used the plant (i.e., the subreplicate) rather than the plot as the experimental replicate. To the best of our knowledge, this review provides the first quantitative estimates of UV-B effects in field-based studies using all suitable published studies as a database.

Solar ultraviolet-B radiation affects plant-insect interactions in a natural ecosystem of Tierra del Fuego (southern Argentina).

We examined the effects of solar ultraviolet-B radiation (UVB) on plant-herbivore interactions in native ecosystems of the Tierra del Fuego National Park (southern Argentina), an area of the globe that is frequently under the Antarctic "ozone hole" in early spring. We found that filtering out solar UVB from the sunlight received by naturally-occurring plants of Gunnera magellanica, a creeping perennial herb, significantly increased the number of leaf lesions caused by chewing insects. Field surveys suggested that early-season herbivory was principally due to the activity of moth larvae (Lepidoptera: Noctuidae). Manipulative field experiments showed that exposure to solar UVB changes the attractiveness of G. magellanica leaf tissue to natural grazers. In a laboratory experiment, locally caught moth caterpillars tended to eat more tissue from leaves grown without UVB than from leaves exposed to natural UVB during development; however, the difference between treatments was not significant. Leaves grown under solar UVB had slightly higher N levels than leaves not exposed to UVB; no differences between UVB treatments in specific leaf mass, relative water content, and total methanol-soluble phenolics were detected. Our results show that insect herbivory in a natural ecosystem is influenced by solar UVB, and that this influence could not be predicted from crude measurements of leaf physical and chemical characteristics and a common laboratory bioassay.

Hydraulic lift: consequences of water efflux from the roots of plants.

Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψs) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψs gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species.

Species interactions at the level of fine roots in the field: influence of soil nutrient heterogeneity and plant size.

Interference at the level of fine roots in the field was studied by detailed examination of fine root distribution in small soil patches. To capture roots as they occur in natural three-dimensional soil space, we used a freezing and slicing technique for microscale root mapping. The location of individual roots intersecting a sliced soil core surface was digitized and the identity of shrub and grass roots was established by a chemical technique. Soil patches were created midway between the shrub, Artemisia tridentata, and one of two tussock grasses, Pseudoroegneria spicata or Agropyron desertorum. Some soil patches were enriched with nutrients and others given only deionized water (control); in addition, patches were located between plants of different size combination (large shrubs with small tussock grasses and small shrubs with large tussock grasses). The abundance of shrub and grass roots sharing soil patches and the inter-root distances of individual fine roots were measured. Total average rooting density in patches varied among these different treatment combinations by only a factor of 2, but the proportion of shrub and grass roots in the patches varied sixfold. For the shrub, the species of grass roots sharing the patches had a pronounced influence on shrub root density; shrub roots were more abundant if the patch was shared with Pseudoroegneria roots than if shared with Agropyron roots. The relative size of plants whose roots shared the soil patches also influenced the proportion of shrub and grass roots; larger plants were able to place more roots in the patches than were the smaller plants. In the nutrient-enriched patches, these influences of grass species and size combination were amplified. At the millimeter- to centimeter-scale within patches, shrub and grass roots tended to segregate, i.e., avoid each other, based on nearest-neighbor distances. At this scale, there was no indication that the species-specific interactions were the result of resource competition, since there were no obvious patterns between the proportion of shrub and grass roots of the two species combinations with microsite nutrient concentrations. Other potential mechanisms are discussed. Interference at the fine-root level, and its species-specific character, is likely an influential component of competitive success, but one that is not easily assessed.

The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field.

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.

Effects of soil temperature and nitrogen status on kinetics of 15 NO3 - uptake by roots of field-grown Agropyron desertorum (Fisch. ex Link) Schult.

Plant NO8 - acquisition is largely determined by root uptake capacity. Although root uptake capacity has been shown to be sensitive to both root temperature and previous nitrogen (N) supply in hydroponic systems, the uptake capacity response to similar environmental factors under field conditions has not been investigated. Using 15 NO3 - , root uptake capacities were determined in excised roots of Agropyron desertorum (Fisch. ex Link) Schult grown in the field at two soil temperatures and two N fertilization treatments. Variation in soil and root temperatures was achieved by application of clear plastic film or insulating mulch to the soil immediately around the target plants. Uptake rates were measured at six different assay solution concentrations (from 1 to 1000 µM external 15 NO3 - concentration range). Two months after the imposition of soil N and temperature treatments, a biphasic transport system (a high-affinity) saturable phase and a low-affinity transport phase) was apparent in low N-treated plants. Nitrate uptake capacity in the low-concentration range (1-500µM) was significantly reduced in N-fertilized plants compared with unfertilized control plants and the effect was more pronounced at high (27 °C) than low (17 °C) soil and assay temperatures. Furthermore, high soil N status inhibited the expression of a low-affinity NO3 - transport system which was clearly apparent at external NO3 - concentration ranges between 500 and 1000 mM in plants grown at low soil N. Prior soil N and temperature history may ultimately determine root ability to exploit NO3 - flushes which can result from changes in soil environmental conditions.

Competitive interaction in plant populations exposed to supplementary ultraviolet-B radiation.

Changes in plant growth and competitive balance between pairs of competing species were documented as a result of supplementary ultraviolet-B radiation (principally in the 290-315 nm waveband) under field conditions. This component of the terrestrial solar spectrum would be intensified if the atmospheric ozone layer were reduced. A method for calculating and statistically analyzing relative crowding coefficients was developed and used to evaluate the competitive status of the species pairs sown in a modified replacement series. The effect of the supplementary UV-B irradiance was generally detrimental to plant growth, and was reflected in decreased leaf area, biomass, height and density as well as changes in competitive balance for various species. For some species, interspecific competition apparently accentuated the effect of the UV-B radiation, while more intense intraspecific competition may have had the same effect for other species. A few species when grown in a situation of more severe mutual interspecific competition exhibited enhanced growth under the UV-B radiation treatment. This, however, was usually associated with a detrimental effect of the radiation on its competitor and thus was likely the result of its improved competitive circumstance rather than a beneficial physiological effect of the radiation.

Leaf epidermal transmittance of ultraviolet radiation and its implications for plant sensitivity to ulraviolet-radiation induced injury.

Leaf epidermal transmittance of ultraviolet radiation (280-400 nm) was examined in several plant species to determine the capability of the epidermis to attenuate solar ultraviolet radiation. Epidermal samples were mechanically isolated and examined with a spectroradiometer/integrating sphere for transmittance. A survey of 25 species exposed to natural insolation was conducted. Although the species differed in life form, habitat type, and epidermal characteristics, epidermal transmittance was generally less than 10%. Ultraviolet radiation was attenuated 95 to 99% in more than half of the species. In 16 species, flavonoid and related pigments in the epidermis accounted for 20 to 57% of the attenuation. Several species exposed to supplemental ultraviolet irradiation (288-315 nm) in a greenhouse exhibited significant (P≦0.05) depressions in epidermal transmittance of 31 to 47%, apparently resulting from an increase in ultraviolet-absorbing pigments.

Carbon balance, productivity, and water use of cold-winter desert shrub communities dominated by C3 and C4 species.

Common generalizations concerning the ecologic significance of C4 photosynthesis were tested in a study of plant gas exchange, productivity, carbon balance, and water use in monospecific communities of C3 and C4 salt desert shrubs. Contrary to expectations, few of the hypotheses concerning the performance of C4 species were supported. Like the C3 species, Ceratoides lanata, the C4 shrub, Atriplex confertifolia, initiated growth and photosynthetic activity in the cool spring months and also exhibited maximum photosynthetic rates at this time of year. To compete successfully with C3 species, Atriplex may have been forced to evolve the capacity for photosynthesis at low temperatures prevalent during the spring when moisture is most abundant. Maximum photosynthetic rates of Atriplex were lower than those of the C3 species. This was compensated by a prolonged period of low photosynthetic activity in the dry late summer months while Ceratoides became largely inactive. However, the annual photosynthetic carbon fixation per ground area was about the same in these two communities composed of C3 and C4 shrubs. The C4 species did not exhibit greater leaf diffusion resistance than the C3 species. The photosynthesis/transpiration ratios of the two species were about the same during the period of maximum photosynthetic rates in the spring. During the warm summer months the C4 species did have superior photosynthesis/transpiration ratios. Yet, since Ceratoides completed a somewhat greater proportion of its annual carbon fixation earlier in the season, the ratio of annual carbon fixation/transpiratory water loss in the two communities was about the same. Atriplex did incorporate a greater percentage of the annual carbon fixation into biomass production than did Ceratoides. However, this is considered to be a reflection of properties apart from the C4 photosynthetic pathway. Both species displayed a heavy commitment of carbon to the belowground system, and only about half of the annual moisture resource was utilized in both communities.

Gas exchange of four arctic and alpine tundra plant species in relation to atmospheric and soil moisture stress.

Gas exchange measurements of two arctic tundra plants, Dupontia fischeri and Carex aquatilis, and two alpine tundra species, Deschampsia caespitosa and Geum rossii, were conducted under a range of atmospheric and soil moisture stress conditions to determine if photosynthetic adaptations to water stress may play a role in the local distributions of these species. Under low soil moisture stress, the species which are normally restricted to wet sites, Dupontia and Deschampsia, exhibited higher net photosynthetic rates than Carex and Geum which are more widely distributed. However, photosynthetic of the wider ranging species was not so abruptly curtailed as that of the wet site species when the plants were exposed to increasing atmospheric or soil moisure stress. Although the depression of photosynthesis with water stress in these tundra species could be partially attributed to reduced stomatal aperture, with decreased soil water potential most of the decline of photosynthesis was due to a greater non-stomatal or residual resistance, indicating a direct impact of water stress on the photosynthetic apparatus. Dark respiration did not increase with enhanced water stress. Stomata of the wet site species did not appear to undergo a closing response until the bulk leaf water potential decreased. However, at high soil water potential reduced stomatal aperture of the more widely distributed species was noted before leaf water potential dropped. Therefore, stomata of these species may respond directly to the vapor pressure gradient between the leaf and the atmosphere when soil moisture potential is high. The wet site species typically exhibited higher photosynthesis/transpiration ratios at low soil moisture stress; however, as soil moisture stress increased, the photosynthesis/transpiration ratios of the wider ranging species generally exceeded those of the wet site species.