A stage-based study of drought response in Cryptantha flava (Boraginaceae): gas exchange, water use efficiency, and whole plant performance.

Models of global climate change predict an increase in the frequency of major droughts, yet we know little about the consequences of drought for the demography of natural populations. This study examined a population of the semi-desert perennial Cryptantha flava (Boraginaceae) to determine how plants of different developmental stages respond to drought through changes in leaf gas exchange, leaf water potential, water use efficiency, growth, and reproduction. In two of the four years, drought was applied using rainout shelters, and a severe natural drought occurred in another. Small, presumably younger, plants sometimes had lower rates of maximum photosynthesis, lower leaf water potentials, and lower instantaneous or integrated water-use efficiency than large plants. Small plants also had higher relative growth rates and lower reproductive effort. Large plants with evidence of shrinkage from a previously larger size often produced less growth and reproduction than large healthy plants, suggesting a decline in plant vigor with age. Drought depressed gas exchange and leaf water potentials equally in all plant stages. Thus, leaf-level physiological attributes provide no clues for why drought reduces growth more strongly in large plants. The results point to several additional avenues of research relevant to understanding stage-dependent or age-dependent plant performance under drought conditions.

Variation in carbon isotope discrimination in relation to plant performance in a natural population of Cryptantha flava.

Few studies of phenotypic selection have focused on physiological traits, especially in natural populations. The adaptive significance of plant water-use efficiency, the ratio of photosynthesis to water loss through transpiration, has rarely been examined. In this study, carbon isotopic discrimination, Delta, an integrated measure of water-use efficiency, was repeatedly measured in juveniles and adults in a natural population of the herbaceous desert perennial Cryptantha flava over a 4-year period and examined for plasticity in Delta, consistency between years in values of Delta, and evidence for selection on Delta phenotypes. There was significant concordance in Delta values among the 4 years for adult plants and significant correlations in Delta values measured in different years for juveniles and adults combined. The wettest year of the study, 1998, proved an exception because Delta values that year were not correlated with Delta values in any other year of the study. Consistency in Delta measured on the same plants in different years could indicate genotypic variation and/or consistency in the water status of the microhabitats the plants occupied. Two forms of plasticity in Delta were also evident; mean seasonal values were correlated with precipitation the preceding autumn, and Delta values also declined with plant size, indicating increasing water-use efficiency. Phenotypic selection was evident because in the first year of the study juvenile plants that would survive until year five averaged lower Delta values than did those that failed to survive. During the driest year, 2000, Delta was significantly negatively correlated with adult plant size, measured as the number of leaf rosettes, but the negative relationship between Delta and the number of flowering stalks, a more direct measure of fitness, was not significant. These results suggest that the direction of phenotypic selection on Delta changes as plants grow.

Effects of below- and aboveground competition from the vines Lonicera japonica and Parthenocissus quinquefolia on the growth of the tree host Liquidambar styraciflua.

Detrimental effects of vines on tree growth in successional environments have been frequently reported. Little is known, however, about the relative importance of below and aboveground competition from vines on tree growth. The objective of this study was to quantify and compare the growth responses of Liquidambar styraciflua saplings to below and/or aboveground competition with the exotic evergreen vine, Lonicera japonica (Japanese honeysuckle), and the native deciduous vine, Parthenocissus quinquefolia (Virginia creeper). Soil trenching and/or vine-trellising were used to control the type of vine competition experienced by trees. Comparisons among untrenched treatments tested for effects of belowground competition. Comparisons among trenched treatments tested for effects of aboveground competition. After two growing seasons, Lonicera japonica had a greater effect on the growth of L. styraciflua than did P. quinquefolia. This effect was largely due to root competition, as canopy competition only had a negative effect on tree growth when it occurred in combination with root competition. Leaf expansion was consistently and similarly affected by all treatments which involved belowground competition.

Water relations of stem succulent trees in north-central Baja California.

Water relations of several stem succulent trees were measured in north-central Baja California in comparisons to other growth forms in the same habitat. Our research concentrated on three stem succulent species (Idria collumnaris, Pachycormus discolor and Bursera microphylla) each with a different succulent stem morphology. The stem succulent trees had 1 to 4 kg H2O/m3 of trunk while the other trees and shrubs in the same habitat had 0.6 to 0.8 kg H2O/m3. The diurnal and seasonal variation in leaf water potential was small for the stem succulent species in comparison to deciduous and evergreen species as a consequence of the stem-water, buffering capacity. In addition, the leaf conductance of the stem succulent species was low (60 mmol m-2 s-1) and yet, the leaf conductance decreased through the day similar to adjacent evergreen and deciduous species. The leaves of the stem succulent trees lost turgor at low saturated water deficits (0.06 to 0.14), had comparatively high osmotic potentials, and high values of elastic modulus in comparison to adjacent evergreen and deciduous species. The stem acts as an important buffering mechanism allowing for the maintenance of leaf turgor in these stem succulent trees. The low transpiration rates of the stem succulent trees may be a mechanism to minimize leaf saturated water deficit and extend leaf longevity.

Field water relations of three temperate vines.

A two year comparative field water relations study was conducted in central Maryland on three sympatric temperate lianas, Lonicera japonica, Vitis vulpina and Parthenocissus quinquefolia. Seasonal physiological activity was longer by approximately 9 weeks in the evergreen L. japonica, while peak rates of stem elongation were 4-10 fold higher in the two deciduous species. There were marked differences in vascular anatomy and water use patterns among the three species, however all three evidenced varying degrees of stomatal closure in response to decreasing soil water availability and increasing atmospheric evaporative demand. The range of leaf water potentials measured in these species was quite narrow in comparison to other temperate woody species. Two of the species showed no alterations in their tissue water release properties in response to decreased soil water availability, while V. vulpina showed a limited capacity in this regard. Most significant among the species differences in water relations were the conservative water use patterns of P. quinquefolia, and the midday maxima of transpirational water loss measured in L. japonica compared to the morning peaks in traspiration for the two deciduous species. The differences found in anatomy, leaf phenology, climbing mechanics, water relations and canopy development among these three sympatric vines implies a spatial and temporal partitioning of light and water resources and emphasizes the diversity of morphological-physiological suites of characters present among species co-occurring in the same macrohabitat.

Field photosynthesis, microclimate and water relations of an exotic temperate liana, Pueraria lobata, kudzu.

Kudzu occurs in a variety of habitats in the southeastern United States. It is most common in exposed, forest edge sites and road cuts where it forms an extensive ground canopy as well as a canopy overtopping nearby trees, but it can also be found in completely open fields and deeply shaded sites within a forest. Microclimate, stomatal conductance, leaf water potential and photosynthetic responses to light, temperature and humidity were measured in two contrasting microhabitats on Pueraria lobata, kudzu. Midsummer leaf temperatures and leaf-to-air water vapor deficits for plants growing in an exposed site were significantly greater than for those in a shaded site, exceeding 35° C and 50 mmol mol-1, respectively. Maximum stomatal conductance exceeded 400 mmol m-2 s-1 in exposed leaves during peak vegetative growth. Stomatal conductance in shaded leaves was approximately half the value measured in exposed leaves on any particular dya. Maximum photosynthetic carbon uptake was also higher in leaves growing in exposed sites compared to leaves in shaded sites, exceeding 18.7 and 14.0 µmol m-2 s-1, respectively. Photosynthesis, stomatal conductance and intercellular CO2 concentration decreased dramatically in response to increasing water vapor deficit for leaves from both sites. However, transpiration showed an initial increase at intermediate water vapor deficits, leveling off or even decreasing at higher values. Leaf water potential demonstrated marked diurnal variation, but remained constant over a wide range of transpirational water fluxes. This latter feature, combined with microenvironmental modification through rapid leaf orientation and pronounced stomatal responses to water vapor deficits may represent important adaptive responses in the exploitation of a diverse array of habitats by kudzu.

Photosynthetic characteristics of Sonoran Desert winter annuals.

Photosynthesis in Sonoran Desert winter annuals appeared to be similar to those observed in other C3 photosynthetic pathway herbs, although photosynthetic capacities ranged from 18 to 65 µmol CO1 m(-2) s(-1) under natural conditions. The higher photosynthetic capacities were associated with high leaf conductances to water vapor (up to 39 mm s(-1)). Leaf Kjeldahl nitrogen contents were high, ranging up to 44.9 mg g(-1). We suggest that the high photosynthetic capacities in several species may be related to resource availability and enable successful exploitation of the short, unpredictable growth periods experienced by these annuals. Although photosynthetic rates in desert winter annuals spanned a wide range, the relationship between leaf conductance and maximum photosynthesis appeared simiar to that of other C3 vascular plants. It is possible that the resulting constant intercellular, CO2 concentrations were related to minimizing excessive water loss, while not severely imposing limitations to photosynthetic gains.

Ecophysiology of two solar tracking desert winter annuals : III. Gas exchange responses to light, CO2 and VPD in relation to long-term drought.

The gas exchange responses of potted, outdoor and greenhouse grown plants of the Sonoran Desert annuals Lupinus arizonicus (Wats.) and Malvastrum rotundifolium (Gray) were examined. Light saturation of leaf photosynthetic rates did not occur in either species at quantum flux densities exceeding 2.0 mmol m-2 s-1. Decreasing water potentials due to long-term drought did not alter this pattern of light response, though it did lower both photosynthetic rates and leaf conductances. Absolute maximum net photosynthetic rates exceeded 60 µmol m-2 s-1 and 50 µmol m-2 s-1 for M. rotundifolium and L. arizonicus, respectively. Both species showed a two level control of leaf conductance, responding to bulk leaf water potential and vapor pressure deficit. There were non-stomatal effects of drought upon photosynthesis in each species. Leaves of M. rotundifolium exhibited a constant stomatal, inhibition of approximately 19%, while in leaves of L. arizonicus stomatal inhibition ranged from 12-40% with decreasing leaf water potentials. These physiological data lend support to previous reports on the divergent water use patterns of these co-occurring species.

Ecophysiology of two solar tracking desert winter annuals : IV. Effects of leaf orientation on calculated daily carbon gain and water use efficiency.

This paper represents an empirical study on the effect of different leaf orientations on the daily carbon gain and transpirational water loss of desert winter annuals. Laboratory physiological data on Malvastrum rotundifolium (Gray) and Lupinus arizonicus (Wats) were combined with energy budget concepts and field measurements of water relations and leaf movements to predict carbon gain patterns for horizontally oriented, diaheliotropic and paraheliotropic leaf movement types. The results showed contrasting patterns of carbon gain and water loss. L. arizonicus, which is capable of both dia- and paraheliotropic leaf movements, had the lowest rates of daily carbon gain and water loss. But these low rates resulted in the highest water use efficiencies under early season conditions and high water availability. M. rotundifolium, a diaheliotropic species, was predicted as having the highest rates of carbon gain and water loss on a daily basis over a wide range of environmental conditions and water availability. Despite possessing the highest rates for transpiration, its water use efficiency was higher in relation to other leaf movement types, under a variety of conditions. This result was extremely sensitive to soil water availability. The results were discussed in relation to the ecological ramifications of leaf movements in arid land annuals.

Ecophysiology of two solar tracking desert winter annuals : II. Leaf movements, water relations and microclimate.

The seasonal course of water relations in field populations of two leaf solar tracking desert winter annuals was examined. Measurements were made of leaf movements in relation to leaf conductance and water potential. Malvastrum rotundifolium maintained solar tracking movements up to the wilting point of the plant (-4 MPa). Lupinus arizonicus altered its morphology through paraheliotropic leaf movements as leaf water potentials declined to-1.8 MPa. Diurnal patterns of leaf conductance showed marked seasonal trends, with gas exchange activity being restricted to early morning hours as water availability declined. Studies of potted plants showed that L. arizonicus was not able to alter its osmotic potential in response to drought, while M. rotundifolium underwent a 1.86 MPa reduction in osmotic potential. The significance of the two contrasting patterns is discussed in terms of observed plant distribution and origin.