Martyn M. Caldwell

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

Abstract 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.

UV-B as an environmental factor in plant life: stress and regulation.

Recent studies indicate that increasing solar UV-B is not merely an environmental stress for plants. Solar UV-B can cause plant morphogenetic effects, which can, in turn, modify the architecture of plants and the structure of a vegetation, In addition, UV-B radiation affect the production of various secondary metabolites (such as flavonoids, tannins and lignin) with important physiological and ecological consequences.

Hydraulic lift: Substantial nocturnal water transport between soil layers by Artemisia tridentata roots

SummaryDiel soil water potential fluctuations reflected daytime depletion and nocturnal resupply of water in upper soil layers. Transpiration suppression experiments demonstrated that water absorption by roots caused the daytime depletion. The soil water potential data and experimental results suggest that at night water absorbed from moist soil by deeper roots is transported to and lost from roots into drier upper soil layers. The deeper roots appear to absorb and transport water both day and night. Implications for the efficiency of deep roots and water storage, nutrient uptake and water parasitism in upper soil layers are discussed.

Changes in biologically active ultraviolet radiation reaching the Earth's surface

Since publication of the 1998 UNEP Assessment, there has been continued rapid expansion of the literature on UV-B radiation. Many measurements have demonstrated the inverse relationship between column ozone amount and UV radiation, and in a few cases long-term increases due to ozone decreases have been identified. The quantity, quality and availability of ground-based UV measurements relevant to assessing the environmental impacts of ozone changes continue to improve. Recent studies have contributed to delineating regional and temporal differences due to aerosols, clouds, and ozone. Improvements in radiative transfer modelling capability now enable more accurate characterization of clouds, snow-cover, and topographical effects. A standardized scale for reporting UV to the public has gained wide acceptance. There has been increased use of satellite data to estimate geographic variability and trends in UV. Progress has been made in assessing the utility of satellite retrievals of UV radiation by comparison with measurements at the Earths surface. Global climatologies of UV radiation are now available on the Internet. Anthropogenic aerosols play a more important role in attenuating UV irradiances than has been assumed previously, and this will have implications for the accuracy of UV retrievals from satellite data. Progress has been made inferring historical levels of UV radiation using measurements of ozone (from satellites or from ground-based networks) in conjunction with measurements of total solar radiation obtained from extensive meteorological networks. We cannot yet be sure whether global ozone has reached a minimum. Atmospheric chlorine concentrations are beginning to decrease. However, bromine concentrations are still increasing. While these halogen concentrations remain high, the ozone layer remains vulnerable to further depletion from events such as volcanic eruptions that inject material into the stratosphere. Interactions between global warming and ozone depletion could delay ozone recovery by several years, and this topic remains an area of intense research interest. Future changes in greenhouse gases will affect the future evolution of ozone through chemical, radiative, and dynamic processes In this highly coupled system, an evaluation of the relative importance of these processes is difficult: studies are ongoing. A reliable assessment of these effects on total column ozone is limited by uncertainties in lower stratospheric response to these changes. At several sites, changes in UV differ from those expected from ozone changes alone, possibly as a result of long-term changes in aerosols, snow cover, or clouds. This indicates a possible interaction between climate change and UV radiation. Cloud reflectance measured by satellite has shown a long-term increase at some locations, especially in the Antarctic region, but also in Central Europe, which would tend to reduce the UV radiation. Even with the expected decreases in atmospheric chlorine, it will be several years before the beginning of an ozone recovery can be unambiguously identified at individual locations. Because UV-B is more variable than ozone, any identification of its recovery would be further delayed.

Coping with Herbivory: Photosynthetic Capacity and Resource Allocation in Two Semiarid Agropyron Bunchgrasses,

SummaryAgropyron desertorum, a grazing-tolerant bunchgrass introduced to the western U.S. from Eurasia, and Agropyron spicatum, a grazing-sensitive bunchgrass native to North America, were examined in the field for photosynthetic capacity, growth, resource allocation, and tiller dynamics. These observations allowed identification of physiological characteristics that may contribute to grazing tolerance in semiarid environments. A uniform matrix of sagebrush, Artemisia tridentata, provided an ecologically relevant competitive environment for both bunch-grass species. Physiological activity, growth, and allocation were also followed during recovery from a severe defoliation treatment and were correlated with tiller dynamics.Potential photosynthetic carbon uptake of both species was dominated by stems and leaf sheaths during June, when maximum uptake rates occurred. For both species, water use efficiency of stems and sheaths was similar to that of leaf blades, but nitrogen investment per photosynthetic surface area was less than in blades. In addition, soluble carbohydrates in stems and sheaths of both species constituted the major labile carbon pools in control plants. Contrary to current theory, these findings suggest that culms from which leaf blades have been removed should be of considerable value to defoliated bunchgrasses, and in the case of partial defoliation could provide important supplies of organic nutrients for regrowth. These interpretations, based on total pool sizes, differ markedly from previous interpretations based on carbohydrate concentrations alone, which suggested that crowns contain large carbohydrate reserves. In this study, crowns of both species contained a minor component of the total plant carbohydrate pool.Following defoliation, A. desertorum plants rapidly reestablished a canopy with 3 to 5 times the photosynthetic surface of A. spicatum plants. This difference was primarily due to the greater number of quickly growing new tillers produced following defoliation. Agropyron spicatum produced few new tillers following defoliation despite adequate moisture, and carbohydrate pools that were equivalent to those in A. desertorum.Leaf blades of regrowing tillers had higher photosynthetic capacity than blades on unclipped plants of both species, but the relative increase, considered on a unit mass, area, or nitrogen basis, was greater for A. desertorum than for A. spicatum. Agropyron desertorum also had lower investment of nitrogen and biomass per unit area of photosynthetic tissues, more tillers and leaves per bunch, and shorter lived stems, all of which can contribute to greater tolerance of partial defoliation.Greater flexibility of resource allocation following defoliation was demonstrated by A. desertorum for both nitrogen and carbohydrates. Relatively more allocation to the shoot system and curtailed root growth in A. desertorum resulted in more rapid approach to the preclipping balance between the root and shoot systems, whereas root growth in A. spicatum continued unabated following defoliation. Nitrogen required for regrowth in both species was apparently supplied by uptake rather than reserve depletion. Carbohydrate pools in the shoot system of both species remained very low following severe defoliation and were approximately equivalent to carbon fixed in one day by photosynthesis of the whole canopy.

Geostatistical patterns of soil heterogeneity around individual perennial plants

1) Univariate, multivariate, and geostatistical techniques were used to quantify the scale and degree of soil variability around individual perennial sagebrush (Artemisia tridentata ssp. vaseyana) and bluebunch wheatgrass (Pseudoroegneria spicata ssp. spicata) plants. This variability was then compared to that found across the larger sagebrush-steppe site. Samples were taken every metre in a 10-m × 12-m grid and every 12.5 cm in nine nested 0.5 m × 0.5 m grids containing at least one Artemisia shrub or Pseudoroegneria tussock (362 total samples)

Effects of increased solar ultraviolet radiation on terrestrial ecosystems

Elevated solar UV-B radiation associated with stratospheric ozone reduction may exert effects on terrestrial ecosystems through actions on plants, microbes, and perhaps on some animals. At the ecosystem level, the effects are less well understood than at the molecular and organismal levels. Many of the most important, yet less predictable, consequences will be indirect effects of elevated UV-B acting through changes in the chemical composition and form of plants and through changes in the abiotic environment. These indirect effects include changes in the susceptibility of plants to attack by insects and pathogens in both agricultural and natural ecosystems; the direction of these changes can result in either a decrease or an increase in susceptibility. Other indirect effects of elevated UV-B include changes in competitive balance of plants and nutrient cycling. The direct UV-B action on plants that results in changes in form or function of plants appears to occur more often through altered gene activity rather than damage. The yield of some crop varieties can be decreased by elevated UV-B, but other varieties are not affected. Plant breeding and genetic engineering efforts should be able to cope with the potential threats to crop productivity due to elevated UV-B. For forest trees, this may be more difficult if effects of elevated UV-B accumulate over several years. All effects of elevated UV-B radiation must be considered in the context of other climate changes such as increased temperature and levels of carbon dioxide, which may alter the UV-B responses, especially for plants. The actions of elevated carbon dioxide and UV-B appear to be largely independent, but interactions occur between changes in UV-B and other factors. Other ecosystem-level consequences of elevated UV-B radiation are emerging and their magnitude and direction will not be easily predicted.

The changing solar ultraviolet climate and the ecological consequences for higher plants

There is compelling evidence that a general erosion of the global ozone layer is occurring. Since ozone in the stratosphere absorbs much of the shortwave solar ultraviolet radiation (UV-B), diminished ozone means that more UV-B of a very specific wavelength composition will be received at the earths surface. Evaluating the implications for vegetation involves consideration of the wavelength specificity of biological photochemical reactions and their sensitivity to the extant and future solar spectrum. Recent research suggests the occurrence of direct damaging reactions and of indirect morphological and chemical responses with implications at the community and ecosystem levels.

Stratospheric Ozone Reduction, Solar UV-B Radiation and Terrestrial Ecosystems

Stratospheric ozone reduction is occurring and will continue to increase in magnitude into the next century. Yet, the consequences for terrestrial ecosystems of the increased solar UV-B (280–320 nm) radiation resulting from total column ozone reduction are not understood. Based on studies of higher plant response to UV-B, several possible consequences for ecosystems include decreased primary production, altered plant species composition, and altered secondary chemistry with implications for herbivory, litter decomposition and biogeochemical cycles. However, like the assessment of increased atmospheric CO2, extrapolation from studies with isolated plants to ecosystem function is very tenuous at best. Very few UV-B studies have dealt with multispecies systems. Most of the UV-B research in the past two decades (since the first suggestions of ozone reduction) has been conducted as short-term experiments in growth chambers and greenhouses where the unnatural spectral balance of radiation can lead to unrealistic conclusions. Technical difficulties in suitable measurement and manipulation of UV-B radiation also complicate the conduct of reliable experiments. This essay surveys and categorizes some 300 papers from the past 20 years on this subject, draws general conclusions from the research and offers some recommendations with respect to ecosystem consequences.

The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials

SummaryRoot proliferation in nutrient-rich soil patches is an important mechanism facilitating nutrient capture by plants. Although the phenomenon of root proliferation is well documented, the specific timing of this proliferation has not been investigated. We studied the timing and degree of root proliferation for three perennial species common to the Great Basin region of North America: a shrub, Artemisia tridentata, a native tussock grass, Agropyron spicatum, and an introduced tussock grass, Agropyron desertorum. One day after we applied nutrient solution to small soil patches, the mean relative growth rate of Agropyron desertorum roots in these soil patches was two to four times greater than for roots of the same plants in soil patches reated with distilled water. Most of the increased root growth came from thin, laterally branching roots within the patches. This rapid and striking root proliferation by Agropyron desertorum occurred in response to N-P-K enrichment as well as to P or N enrichment alone. A less competitive bunchgrass, Agrophyron spicatum, showed no tendency to proliferate roots in enriched soil patches during these two-week experiments. The shrub Artemisia tridentata proliferated roots within one day of initial solution injection in the N-enrichment experiment, but root proliferation of this species was more gradual and less consistent in the N-P-K and P-enrichment experiments, respectively. The ability of Agropyron desertorum to proliferate roots rapidly may partly explain both its general competitive success and its superior ability to exploit soil nutrients compared to Agropyron spicatum in Great Basin rangelands of North America.