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Notes: The frequency and duration of water on leaf surfaces have important consequences for plant growth and photosynthetic gas exchange. The objective of the present study was to compare the frequency and duration of leaf wetness under natural field conditions among species and to identify variation in structural features of leaves that may reduce surface wetness. During June–September 1992 in the central Rocky Mountains (USA), natural leaf wetting due to rain and dewfall was observed on 79 of 89 nights in open meadow habitats compared to only 29 of 89 nights in the understorey. Dew formation occurred at relative humidities that were often well below 100% because of radiational heat exchange with cold night skies and low wind speeds (〈 0.5 m s−1). A survey of 50 subalpine/montane species showed that structural characteristics associated with the occurrence and duration of leaf surface wetness differed among species and habitats. Both adaxial and abaxial surfaces accumulated moisture during rain and dewfall events. Leaf surfaces of open-meadow species were less wettable (P= 0.008), and had lower droplet retention (P= 0.015) and more stomata P= 0.017) than adjacent understorey species. Also, leaf trichomes reduced the area of leaf surface covered by moisture. Ecophysiological importance is suggested by the high frequency of leaf wetting events in open microsites, influences on growth and gas exchange, and correspondence between leaf surface wettability and habitat.

Notes: In the alpine-treeline ecotone of the Snowy Range in Wyoming, USA, microsite sky exposure of Englemann spruce (Picea englemannii) and subalpine fir (Abies lasiocarpa) seedlings (〈 5 years) was associated with the avoidance of low-nocturnal temperatures and high insolation, factors which appeared to result in low-temperature photoinhibition. In a field experiment, light-saturated photosynthesis (Asat) in current-year seedlings (newly germinated) of fir increased significantly (approximately seven-fold) in response to increased long-wave irradiance at night (warming), solar shading (approximately five-fold), and the combination of the two treatments (approximately eight-fold). Asat in current-year spruce remained unchanged in response to all treatments, but was over four-times higher than fir in control plots. These results indicated substantial low-temperature photoinhibition, and were supported by similar Asat trends in natural seedlings. Increased needle inclination and clustering in more exposed microsites for both species implicates the possible role of structural adaptations for decreased sky exposure and warmer leaf temperatures at night.

Notes: The potential contribution of intercellular light reflectance to photosynthesis was investigated by infiltrating shade leaves with mineral oil. Infiltration of leaves of Hydrophyllum canadense and Asarum canadense with mineral oil decreased adaxial leaf reflectance but increased transmittance. As a result of the large increase in transmittance, infiltration caused a decrease in absorptance of 25% and 30% at 550 and 750 nm, respectively. Thus, intercellular reflectance increased absorptance in these species by this amount. In a comparison of sun and shade leaves of Acer saccharum and Parthenocissus quinquefolia, oil infiltration decreased absorptance more in shade than in sun leaves. This difference suggests that the higher proportion of spongy mesophyll in shade leaves may increase internal light scattering and thus absorptance. The importance of the spongy mesophyll in increasing internal reflectance was also evident in comparisons of the optics of Populus leaves and in the fluorescence yield of oil-infiltrated leaves of several sun and shade species. Oil infiltration decreased the quantum yield of fluorescence (Fo) by 39–52% for shade leaves but only 21–25% for sun leaves. We conclude that the greater proportion of spongy parenchyma in shade leaves increased intercellular light scattering and thus absorptance. Direct measurements with fibre-optic light probes of the distribution of light inside leaves of Hydrophyllum canadense confirmed that oil infiltration decreased the amount of back-scattered light and that most of the light scattering for this species occurred from the middle of the palisade layer to the middle of the spongy mesophyll. We were not, however, able to assess the potential contribution of reflectance from the internal abaxial epidermis to total internal light scattering in these experiments. Using a mathematical model to compare the response of net photosynthesis (O2, flux) to incident irradiance for control leaves of H. canadense and theoretical leaves with no intercellular reflectance, we calculated that intercellular reflectance caused a 1.97-fold increase in photosynthesis at 20 μmol m−2s−1 (incident photon flux density). This enhancement of absorption and photosynthesis by inter-cellular reflectance, without additional production and maintenance of photosynthetic pigments, may maintain shade leaves above the photosynthetic light compensation point between sunflecks and maintain the light induction state during protracted periods of low diffuse light.

Notes: Summary The temperature and water relations of the largleafed, high-elevation species Frasera speciosa, Balsamorhiza sagittata, and Rumex densiflorus were evaluated in the Medicine Bow Mountains of southeast Wyoming (USA) to determine the influence of leaf size, orientation, and arrangement on transpiration. These species characteristically have low minimum stomatal resistances (〈60 s m-1) and high maximum transpiration rates (〉260 mg m-2s-1 for F. speciosa). Field measurements of leaf and microclimatic parameters were incorporated into a computer simulation using standard energy balance equations which predicted leaf temperature (T leaf) and transpiration for various leaf sizes. Whole-plant transpiration during a day was simulated using field measurements for plants with natural leaf sizes and compared to transpiration rates simulated for plants having identical, but hypothetically smaller (0.5 cm) leaves during a clear day and a typically cloudy day. Although clear-day transpiration for F. speciosa plants with natural size leaves was only 2.0% less per unit leaf area than that predicted for plants with much smaller leaves, daily transpiration of B. sagittata and R. densiflorus plants with natural leaf sizes was 16.1% and 21.1% less, respectively. The predicted influence of a larger leaf size on transpiration for the cloudy day was similar to clear-day results except that F. speciosa had much greater decreases in transpiration (12.7%). The different influences of leaf size on transpiration between the three species was primarily due to major differences in leaf absorptance to solar radiation, orientation, and arrangement which caused large differences in T leaf. Also, simulated increases in leaf size above natural sizes measured in the field resulted in only small additional decreases in predicted transpiration, indicating a leaf size that was nearly optimal for reducing transpiration. These results are discussed in terms of the possible evolution of a larger leaf size in combination with specific leaf absorptances, orientations and arrangements which could act to reduce transpiration for species growing in short-season habitats where the requirement for rapid carbon fixation might necessitate low stomatal resistances.

Notes: Summary The influence of variations in the boundary air layer thickness on transpirtion due to changes in leaf dimension or wind speed was evaluated at a given stomatal resistance (r s) for various combinations of air temperature (T a) and total absorbed solar energy expressed as a fraction of full sunlight (S ffs). Predicted transpiration was found to either increase or decrease for increases in leaf size depending on specific combinations of T a, S ffs, and r s. Major reductions in simulated transpiration with increasing leaf size occurred for shaded, highly reflective, or specially oriented leaves (S ffs=0.1) at relatively high T a when r s was below a critical value of near 500 s m-1. Increases in S ffs and decreases in T a lowered this critical resistance to below 50 s m-1 for S ffs=0.7 and T a=20°C. In contrast, when r s was above this critical value, an increase in leaf dimension (or less wind) resulted in increases in transpiration, especially at high T a and S ffs. For several combinations of T a, S ffs, and r s, transpiration was minimal for a specific leaf size. These theoretical results were compared to field measurements on common desert, alpine, and subalpine plants to evaluate the possible interactions of leaf and environmental parameters that may serve to reduce transpiration in xeric habitats.

Notes: Summary The daily temperature and water relations of 7 perennial subalpine, understory species (1 shrub, 1 subshrub, 5 herbs) were compared in the Rocky Mountains of southeastern Wyoming with an emphasis on the effects of natural sun and shade exposure. Field measurements of rainfall; leaf, air, and soil temperatures; stomatal conductance to water vapor diffusion; and plant and soil water potentials were supplemented with leaf and root morphological measurements to evaluate potential adaptive patterns in understory species. Morphologically, all 7 species had relatively broad leaves that were hypostomous and bicolored with the abaxial leaf surface lighter than the abaxial surface. Root systems tended to be shallow (〈20 cm), especially for the herbaceous species. Although soil water potentials from 4 to 40 cm depths remained relatively high throughout the summer (〉-1.0 MPa), plant xylem water potentials for sunlit plants decreased to below-2.0 MPa during midday. During these sunlit periods, leaf temperatures and conductances increased substantially, leading to severe wilting for 4 of the 5 herbaceous species. Stomatal conductance and density for 6 of the 7 species monitored were much greater on abaxial compared to adaxial leaf sides and substantial stomatal closure occurred when either leaf side was oriented to receive direct sunlight. Moreover, stomatal opening on abaxial leaf sides corresponded to the amount of sunlight incident upon the adaxial rather than abaxial leaf surfaces. The 2 shrubby species did not wilt during these periods and were characterized by the highest leaf temperatures (〉30°C). These 2 species also had consistantly lower xylem water potentials throughout the summer growth period. These results are discussed in terms of the possible adaptive significance of midday wilting, leaf hypostomy and bicoloration and stomatal behavior to the water and photosynthetic relations of understory species.

Notes: Summary Environmental and water relations parameters during fall were monitored for six conifer tree species common to the central Rocky Mountains growing naturally at the same location (Pinus contorta, Pinus ponderosa, Pinus flexilus, Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii). Subsequent to what appeared to be the beginning of seasonal stomatal closure, leaf conductance to water vapor declined sharply following the onset of freezing air temperatures at night. A coincident rapid decline in morning xylem pressure potentials (ψp) also occurred which resulted in values that were considerably below afternoon ψp. Continuing decreases in maximum leaf conductance during the day were highly correlated with corresponding decreases in minimum nocturnal air temperatures of the preceding night. By mid-December, morning ψp returned to values very near afternoon ψp and were only slightly lower than before the onset of subfreezing nights. A preliminary model is proposed which interprets the qualitative interaction between air and soil temperatures, soil and plant water potentials, and leaf conductance during seasonal stomatal closure in fall.