Seasonal Acclimation of Stem Photosynthesis in Woody Legume Species from the Mojave and Sonoran Deserts of California.

Photosynthesis (Pn) was measured in stems of two desert legumes, Caesalpinia virgata at a low elevation site (118 m) in the Sonoran Desert and Senna armata at a higher elevation (950 m) in the Mojave Desert. The lower elevation site experienced higher spring and summer temperatures than the higher elevation site, but the air vapor pressure, irradiance, and rainfall patterns were similar. Mid-morning maximum stem Pn was highest in May for C. virgata (7.8 [mu]mol m-2 s-1) and in July for S. armata (5.8 [mu]mol m-2 s-1). The seasonal variation in maximum stem Pn was not associated with changes in bulk tissue water potential or chlorenchyma tissue nitrogen concentration. The main environmental regulators of seasonal stem Pn were temperature and leaf to air vapor pressure gradient. Light-response curves indicated no major differences in apparent quantum yield or light compensation point between the spring and summer, but light-saturated stem Pn at ambient temperature decreased for C. virgata between these seasons. The optimal temperature for stem Pn remained the same for both species between the spring and the summer. However, stem Pn of both species increased at all temperatures between the spring and summer. Potential stem Pn under optimal conditions and CO2-saturated stem Pn increased for both species between spring and summer. The increase in stem Pn potential allowed these species to maintain stem Pn during the summer even though stem Pn responses to temperature and vapor pressure did not acclimate to seasonal climatic conditions.

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST- AND SOUTHWEST-FACING SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. II. WATER RELATIONS.

Three species of shrubs (Ericaceae) were found to segregate upon the northeast and southwest slopes of spur ridges on Brush Mountain, in southwestern Virginia. Rhododendron maximum was found only in valleys and lower northeast slopes, Rhododendron periclymenoides = R. nudiflorum) was found on northeast slopes while Kalmia latifolia was most abundant on southwest slopes. Previous vegetation studies indicated that these partially segregated distributions were related to irradiance and water availability. In field studies of water potential, R. periclymenoides had the lowest diurnal leaf water potentials and the largest seasonal variation in midday leaf water potential. Kalmia latifolia had the highest leaf conductance in field and phytotron experiments. Rhododendron maximum had the greatest seasonal osmotic adjustment followed by R. periclymenoides and K. latifolia. In phytotron experiments, the photosynthetic capacity of R. maximum was the most sensitive to water stress followed by R. periclymenoides and K. latifolia. Kalmia latifolia was able to modify its conductance rates to reduce water loss and maintain constant leaf water potential minimizing photosynthetic inhibition. Rhododendron periclymenoides showed extreme luxury spending of water indicated by high conductance and low photosynthesis. The ecophysiological responses to water and irradiance provided an explanation for the distributions of the three species. For example, R. maximum leaves are sensitive to elevated irradiance, and carbon gain is strongly influenced by water stress. Thus, R. maximum will perform best in low irradiance environments with ample water, such as valley sites. Each species had a unique set of adaptations for performing best in their optimum habitat.

Effects of manipulation of water and nitrogen regime on the water relations of the desert shrub Larrea tridentata.

Water and nitrogen regimes of Larrea tridentata shrubs growing in the field were manipulated during an annual cycle. Patterns of leaf water status, leaf water relations characteristics, and stomatal behavior were followed concurrently. Large variations in leaf water status in both irrigated and nonirrigated individuals were observed. Predawn and midday leaf water potentials of nonirrigated shrubs were lowest except when measurements had been preceded by significant rainfall. Despite the large seasonal variation in leaf water status, reasonably constant, high levels of turgor were maintained. Pressure-volume curve analysis suggested that changes in the bulk leaf osmotic potential at full turgor were small and that nearly all of the turgor adjustment was due to tissue elastic adjustment. The increase in tissue elasticity with increasing water deficit manifested itself as a decrease in the relative water content at zero turgor and as a decrease in the tissue bulk elastic modulus. Because of large hydration-induced displacement in the osmotic potential and relative water content at zero turgor, it was necessary to use shoots in their natural state of hydration for pressure-volume curve determinations. Large diurnal and seasonal differences in maximum stomatal conductance were observed, but could not easily be attributed to variations in leaf water potential or leaf water relations characteristics such as the turgor loss point. The single factor which seemed to account for most of the diurnal and seasonal differences in maximum stomatal conductance between individual shrubs was an index of soil/root/ shoot hydraulic resistance. Daily maximum stomatal conductance was found to decrease with increasing soil/root/ shoot hydraulic resistance. This pattern was most consistent if the hydraulic resistance calculation was based on an estimate of total canopy transpiration rather than the more commonly used transpiration per unit leaf area. The reasons for this are discussed. It is suggested that while stomatal aperture necessarily represents a major physical resistance controlling transpiration, plant hydraulic resistance may represent the functional resistance through its effects on stomatal aperture.

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.

Influences of microclimatic conditions and water relations on seasonal leaf dimorphism of Prosopis glandulosa var. torreyana in the Sonoran Desert, California.

Seasonal measurements of microclimatic conditions were compared to seasonal indices of leaf structural components and plant water relations in Prosopis glandulosa var. torryana. P. glandulosa had two short periods of leaf production which resulted in two distinct even aged cohorts of leaves. The two leaf cohorts (summer, winter) were concurrent in the summer and fall, contrasting to previous studies on other species in which one leaf form replaces a previous leaf type. The structural characteristics of these two cohorts differed significantly in two replicate year cycles. The leaves of the spring cohort were larger in weight and area but similar to the summer cohort in specific leaf weight and leaflet number. The second growth period leaves constituted only a small proportion of the total plant leaf area. The dimorphism between the two cohorts was best associated with plant water relations and not energy load. Second growth period leaves maintained turgor to greater water deficits but lost turgor at higher leaf water potentials. Seasonal osmotic adjustment occurred for first growth period leaves but not second growth period leaves. The small leaves produced during the hot climate were most likely the result of low turgor potential during development rather than an adaptation to tolerate stressful environments.

Estimates of N2-fixation from variation in the natural abundance of 15N in Sonoran desert ecosystems.

The 15N abundance of tissues of five Prosopis specimens at our primary study site (a Prosopis woodland at Harper's Well in the Sonoran desert of Southern California) was determined over two growing seasons 1980 and 1981. The 15N abundance of soil and of tissues of presumed non-N2-fixing (control) plants was also measured. Prosopis tissues were significantly lower in 15N than either soil N or corresponding tissues of presumed non-N2-fixing plants which derive their N entirely from soil. Soil N was also significantly higher in 15N than atmospheric N2. We conclude that it is feasible to use variations in the natural abundance of 15N as an index of N2-fixation in this kind of ecosystem, and that N2-fixation is of considerable importance to Prosopis growing at this site.We also determined the 15N abundance of leaf tissue of presumed N2-fixing and control plants growing at the same site at six additional sites (five in the Sonoran desert of southern California and one in Baja California, Mexico near the town of Catavina). Four of these additional sites were dominated by Prosopis and two were mixed communities. There were statistically significant differences between the 15N abundances of the pooled legume population and control plants at all sites, although not every legume specimen exhibited this difference. From 15N abundance data we estimated the fractional contribution of biologically fixed N to the N economy of desert legumes. We concluded that N2-fixation is very important to Prosopis at six of seven sites in the Sonoran Desert. At the site where Prosopis did not appear to be fixing N2, N2-fixation was important only for legumes of the sub-family Papilionoideae, Lupinus, Dalea, Astragalus and Lotus.

Influence of water relations and temperature on leaf movements of rhododendron species.

Rhododendron maximum L. and R. Catawbiense L. are subcanopy evergreen shrubs of the eastern United States deciduous forest. Field measurements of climate factors and leaf movements of these species indicated a high correlation between leaf temperature and leaf curling; and between leaf water potential and leaf angle. Laboratory experiments were performed to isolate the influence of temperature and cellular water relations on leaf movements. Significant differences were found between the patterns of temperature induction of leaf curling in the two species. Leaves of the species which curled at higher temperatures (R. catawbiense) also froze at higher leaf temperatures. However, in both cases leaf curling occurred at leaf temperatures two to three degrees above the leaf freezing point. Pressure volume curves indicated that cellular turgor loss was associated with a maximum of 45% curling while 100% or more curling occurred in field leaves which still had positive cell turgor. Moisture release curves indicated that 70% curling requires a loss of greater than 60% of symplastic water which corresponds to leaf water potentials far below those experienced in field situations. Conversely, most laboratory induced changes in leaf angle could be related to leaf cell turgor loss.

Inhibition of seedling survival under Rhododendron maximum (Ericaceae): could allelopathy be a cause?

In the southern Appalachian mountains a subcanopy species, Rhododendron maximum, inhibits the establishment and survival of canopy tree seedlings. One of the mechanisms by which seedlings could be inhibited is an allelopathic effect of decomposing litter or leachate from the canopy of R. maximum (R.m.) on seed germination, root elongation, or mycorrhizal colonization. The potential for allelopathy by R.m. was tested with two bioassay species (lettuce and cress), with seeds from four native tree species, and with three ectomycorrhizal fungi. Inhibitory influences of throughfall, fresh litter, and decomposed litter (organic layer) from forest with R.m. (+R.m. sites) were compared to similar extractions made from forest without R.m. (-R.m. sites). Throughfall and leachates of the organic layer from both +R.m. and -R.m. sites stimulated germination of the bioassay species above that of the distilled water control, to a similar extent. There was an inhibitory effect of leachates of litter from +R.m. sites on seed germination and root elongation rate of both bioassay species compared with that of litter from -R.m. sites. Native tree seed stratified in forest floor material from both forest types had a slightly higher seed germination rate compared with the control. A 2-yr study of seed germination and seedling mortality of two tree species, Quercus rubra and Prunus serotina, in field plots showed no significant influence of litter or organic layer from either forest type. Incorporating R.m. leaf material into the growth medium in vitro depressed growth of one ectomycorrhizal species but did not affect two other species. Leaf material from other deciduous tree species depressed ectomycorrhizal growth to a similar or greater extent as leaf material from R.m. In conclusion, R.m. litter can have an allelopathic effect on seed germination and root elongation of bioassay species as well as some ectomycorrhizal species. However, this allelopathic affect is not manifest in field sites and is not likely to be an important cause for the inhibition of seedling survival within thickets of R.m.