INFLUENCE OF INSTANTANEOUS AND INTEGRATED LIGHT-FLUX DENSITY ON LEAF ANATOMY AND PHOTOSYNTHESIS

Experiments were conducted to examine whether leaf adaptation to light in Fragaria virginiana (Rosaceae) was determined by peak photon-flux density or by total quanta received during the day. Leaf structure and apparent photosynthesis rates were similar under environments where total energy received was the same even though peak photon-flux density was different. When peak photon-flux density was held constant and total quanta varied, significant differences were noted in apparent photosynthesis, leaf thickness, specific leaf weight, mesophyll cell volume, and A^mes/A ratio. High total quanta produced high-light or sun-type leaves even at relatively low peak intensities. Thus, total light energy received during the day has a greater influence on leaf adaptation to light than does peak photon-flux density.     

THE INFLUENCE OF WATER STRESS AND TEMPERATURE ON LEAF PUBESCENCE DEVELOPMENT IN ENCELIA FARINOSA

The degree of leaf pubescence development in the arid land shrub Encelia farinosa Gray is affected by air temperature, leaf water potential, and previous history of the apical meristem during the current growing season. Changes in leaf pubescence levels change leaf spectral characteristics and affect both leaf temperature and photosynthesis. Decreasing leaf water potentials and increasing air temperatures both independently increase pubescence development as measured by decreased leaf absorptances. During any one growing season leaf absorptance may change reversibly coincident with air temperature changes, but with respect to water stress leaf absorptance only decreases as the season progresses. The ecological significance of regulation of the leaf spectral characteristics is discussed.     

Intraspecific variation of drought adaptation in brittlebush: leaf pubescence and timing of leaf loss vary with rainfall

Reflective leaf pubescence of the desert shrub Encelia farinosa (brittlebrush) reduces leaf temperature and plant water loss, and is considered adaptive in xeric environments. Yet, little is known about intraspecific variation in this trait. Among three populations in the northern range of E. farinosa, which span a very broad precipitation gradient, both leaf absorptance variation and differences in the timing of drought-induced leaf loss were broadly associated with climatic variability. Where mean annual rainfall was greatest, drought-induced leaf loss was earliest, but these plants also had higher population-level mean leaf absorptance values. Higher absorptance increases the relative dependence on latent heat transfer (transpirational cooling), but it also provides greater instantaneous carbon assimilation. Plants at the driest site reached lower leaf absorptance values and maintained leaves longer into the drought period. Lower leaf absorptance reduces water consumption, and extended leaf longevity may buffer against the unpredictability of growing conditions experienced in the driest site. These observations are consistent with a trade-off scenario in which plants from wetter regions might trade off water conservation for higher instantaneous carbon gain, whereas plants from drier regions reduce water consumption and extend leaf longevity to maintain photosynthetic activity in the face of unpredictable growing conditions. Received: 2 April 1997 / Accepted: 11 August 1997     

Pubescence and leaf spectral characteristics in a desert shrub,Encelia farinosa

Summary The effects of leaf hairs (pubescence) on leaf spectral characteristics were measured for the drought-deciduous desert shrub Encelia farinosa. Leaf absorptance to solar radiation is diminished by the presence of pubescence. The pubescence appears to be reflective only after the hairs have dried out. There are seasonal changes in leaf absorptance; leaves produced at the beginning of a growing season have high absorptances, whereas leaves produced during the growing season are more pubescent and have lower absorptances. The decrease in leaf absorptance is the result of an increase in pubescence density and thickness. Between 400 and 700 nm (visible wavelengths), pubescence serves as a blanket reflector. However, over the entire solar spectrum (400–3000 nm), the pubescence preferentially reflects near infrared radiation (700–3000 nm) over photosynthetically useful solar radiation (400–700 nm). Leaf absorptance to solar radiation (400–3000 nm) varies between 46 and 16%, depending on pubescence; whereas leaf absorptance to photosynthetically useful radiation (400–700 nm) may vary from 81 to 29%.C.I.W.-D.P.B. Publication No. 612     

Relation between Mesophyll Surface Area, Photosynthetic Rate, and Illumination Level during Development for Leaves of Plectranthus parviflorus Henckel

The influence of illumination level during leaf development on the mesophyll cell surface area per unit leaf area (A^mes/A), CO₂ resistances, and the photosynthetic rate was determined for leaves of Plectranthus parviflorus Henckel. The relative importance of A^mes/A versus CO₂ resistances in accounting for observed changes in photosynthesis was quantitatively evaluated using equations based on analogies to electrical circuits.     

DIFFERENCES IN LEAF STRUCTURE,CHLOROPHYLL, AND NUTRIENTS FOR THE UNDERSTORY TREE ASIMINA TRILOBA

To investigate differences in leaf structure, chlorophyll and nutrients on terminal branches of the understory tree Asimina triloba, the first (proximal) and the last (distal) leaves to develop in the spring were compared. Proximal leaf expansion was completed before the overstory canopy was fully closed but distal leaf expansion occurred during and after the development of the overstory canopy. Fully expanded proximal leaves were 76% smaller in area, were 18% thicker and had 36% more stomates per m of leaf area when compared to distal leaves. In addition, maximum stomatal conductance to water vapor was greater (150 vs. 120 mmol m^−-2s^−-1) and the minimum PPFD required for maximum conductance was higher (200 vs. 150 μmol m^−-2s^−-1) for the proximal leaves. Chlorophyll content was also greater for proximal leaves, but nitrogen and phosphorus contents were lower throughout the entire summer. Seasonal measurements indicated an increase in chlorophyll a content and reductions in nitrogen content throughout the summer growth period for leaves from both positions. The results suggest that distal and proximal leaves differed physiologically and that the measured differences were related to the changing irradiance environment during leaf development. The time of leaf expansion, as indicated by leaf position on the branch, may be an important consideration when examining the water and photosynthetic relations of understory trees.     

Phospholipid synthesis and exchange in castor bean endosperm homogenates

Leaves on a bush of Hyptis emoryi Torr. varied in length from less than 1 cm when development occurred in full sunlight (e.g. 40 Mjoules m⁻²) to over 7 cm when the total daily solar irradiance was less than 3 Mjoules m⁻². The 1-cm sun leaves were 3-fold higher than the 7-cm shade leaves in chlorophyll per unit area, mesophyll thickness, and the internal to external leaf area ratio (A^mes/A). The higher A^mes/A caused a 1.2-cm leaf to have a 3-fold lower CO₂ liquid phase resistance than did a 7.1-cm leaf. Large thin shade leaves captured photosynthetically active radiation effectively (less than 7% passed through), but were not adapted to full sunlight. Specifically, when a 6.9-cm leaf was placed at 910 w m⁻² for 30 min, its temperature exceeded that of the air by nearly 8 C. For the common daytime air temperatures above 30 C for this desert shrub, large shade leaves would have temperatures far in excess of that optimum for photosynthesis for H. emoryi, 29 to 32 C.     

Changes in Morphology,Anatomy, and Photosynthetic Capacity of Needles of Japanese Larch (Larix kaempferi) Seedlings Grown in High CO2 Concentrations

Photosynthetic traits of two-year-old Japanese larch seedlings (Larix kaempferi Carr.) grown at elevated CO₂ concentrations were studied in relation to structural changes in the needles. Seedlings were grown at two CO₂ concentrations, 360 (AC) and 720 (EC) mol mol^–1 at high and low nutrient supply rates, high N (HN) and low N (LN). The photosynthetic capacity fell significantly in EC+LN, but increased significantly in EC+HN. Since the mesophyll surface area exposed to intercellular space per unit leaf area (A^mes/A) is correlated with the photosynthetic rate, we measured A^mes/A for larch needles growing in EC. Changes of A^mes/A in both EC+HN and EC+LN were very similar to the changes in photosynthetic capacity. This suggests that the changes of A^mes/A in EC probably caused the changes in the photosynthetic capacity. The changes of A^mes/A in EC were attributed to changes in the mesophyll cell size and mesophyll cell number. The photosynthetic capacity in EC can be explained by taking morphological and structural adaptations into account as well as biochemical factors.     

Water movement through Quercus rubra I. Leaf water potential and conductance during polycyclic growth

Two experiments examined simultaneous changes in leaf area (A_L), root length (L_r), stomatal conductance (g_s), leaf water potential (Ψ_L), transpiration and hydraulic plant conductance per unit leaf area (G) during the first three shoot cycles of northern red oak (Quercus rubra L.) grown under favourable and controlled conditions. Each shoot cycle consisted of bud swell, stem elongation, leaf expansion and rest; roots grew almost continuously. The g_s of all leaves decreased substantially while leaves of the newest flush were expanding and increased modestly when seedling leaf area remained constant. Overall, g_s decreased. The Ψ_L of mature leaves decreased during leaf expansion and increased by an equivalent amount during intervening periods. Possible explanations for the paired changes in g_s and Ψ_L are considered. Changes in G closely paralleled those of canopy g_s. These parallel changes during polycyclic seedling growth should act to keep seedling Ψ_L relatively constant as plant size increases and thereby help prevent Ψ_L from dropping to levels that would cause runaway embolism.     

RELATIONSHIPS BETWEEN PHOTOSYNTHETIC CAPACITY AND LEAF STRUCTURE IN SEVERAL SHADE PLANTS

Leaf structure, photosynthetic characteristics and related physiological parameters have been studied in three ornamental shade species: Fatsia japonica, Cissus rhombifolia (relatively light-tolerant plants), and Philodendron scandens (obligate shade plant). Species were grown in a shadehouse. Maximum photosynthetic photon flux density was 470 μmol m^-2 s^-1. Net rate of CO₂ uptake at light saturation (maximum Pn) in Fatsia was 6.90 ± 1.27 μmol m^-2 s^-1. In Cissus and Philodendron values were about 30% and 63% less respectively, than those measured in Fatsia. The nitrogen content, relative dry wt, specific leaf dry wt (SLDW), chlorophyll a/b ratio, and nitrogen to chlorophyll ratio were lower in Philodendron. However, leaf thickness in Philodendron (296 ± 17 μm) was about 54% and 160% higher, respectively, than in Fatsia and Cissus, and the ratio between mesophyll cell area and leaf surface area (A^mes/A) was nearly similar in the three species. However Philodendron exhibited a percentage of palisade parenchyma about three times lower than that observed in the two other species. The chloroplast number per mm of cell wall in transverse sections (chloroplast density) in the palisade parenchyma was fairly constant (about 65), irrespective of species. The “chloroplast density” in the spongy parenchyma of Philodendron was about 53% and 63%, respectively, of Fatsia and Cissus values. In Fatsia and Cissus chloroplast ultrastructure seems to change gradually and continuously from sun to shade type with the depth from the adaxial to abaxial surface. Special emphasis was given in order to determine the structural parameters best correlated with maximum Pn between the different species. In this way chloroplast number in transverse sections (chloroplast number) and the ratio between chloroplast area and leaf surface area (A^chl/A) were the parameters best correlated with maximum Pn, and stomatal frequency was also a good determinant of maximum Pn. However, leaf thickness, SLDW, and even A^mes/A ratio were weakly correlated with maximum Pn.     

Water potential in soil and Atriplex nummularia (phytoremediator halophyte) under drought and salt stresses

Atriplex nummularia is a halophyte widely employed to recover saline soils and was used as a model to evaluate the water potentials in the soil-plant system under drought and salt stresses. Potted plants grown under 70 and 37% of field capacity irrigated with solutions of NaCl and of a mixture of NaCl, KCl, MgCl₂ and CaCl₂ reproducing six electrical conductivity (EC): 0, 5, 10, 20, 30, and 40 dS m^?1. After 100 days, total water (Ψ_{w, plant}) and osmotic (Ψ_{o, plant}) potentials at predawn and midday and Ψ_{o, soil}, matric potential (Ψ_{m, soil}) and Ψ_{w, soil} were determined. The type of ion in the irrigation water did not influence the soil potential, but was altered by EC. The soil Ψ_o component was the largest contributor to Ψ_{w, soil}. Atriplex is surviving ECs close to 40 dS m^?1 due to the decrease in the Ψ_w. The plants reached a Ψ_w of approximately ?8 MPa. The water potentials determined for different moisture levels, EC levels and salt types showed huge importance for the management of this species in semiarid regions and can be used to recover salt affected soils.     

Comparative ecophysiology of Encelia farinosa and Encelia frutescens

Summary Encelia farinosa and Encelia frutescens are drought-decidous shrubs whose distributions overlap throughout much of the Sonoran Desert. During hot and dry periods, leaves of E. farinosa utilize increased leaf reflectance to reduce leaf temperature, whereas leaves of E. frutescens have substantially higher leaf conductances and rely on increased transpirational cooling to reduce leaf temperature. E. farinosa is common on the dry slope microhabitats, whereas E. frutescens occurs only in wash microhabitats where greater soil moisture is available to provide the water necessary for transpirational cooling. E. farinosa tends not to persist in wash microhabitats because of its greater susceptibility to flashfloods. The consequences and significance of increased leaf reflectance versus increased transpirational cooling to leaf temperature regulation are discussed.     

Low moisture availability reduces the positive effect of increased soil temperature on biomass production of white birch (Betula papyrifera) seedlings in ambient and elevated carbon dioxide concentration

White birch (Betula papyrifera Marsh.) seedlings were grown under two carbon dioxide concentrations ([CO₂]) (360 vs 720 μmol mol^?1), three soil temperatures (T_soil) (5, 15, 25°C initially, increased to 7, 17, 27°C, respectively, one month later), and three moisture regimes (low: 30–40%, intermediate: 45–55%, high: 60–70% field water capacity) for four months in environment‐controlled greenhouses. The dry mass of stem, leaves, and roots was measured after 2 and 4 months of treatment. Low T_soil decreased stem, leaf and total biomass in both measurements, however, the decrease was significantly greater in the elevated than ambient [CO₂] after 4 months. Intermediate T_soil increased root biomass in both measurements. Low moisture reduced stem, leaf, root and total biomass after both 2 and 4 months of treatment. There was a significant T_soil‐moisture interactive effect on leaf, root, and total biomass after 4 months of treatment, suggesting that the magnitude of biomass enhancement in warmer T_soil was dependent on the moisture regime. For instance, the increase in total biomass from the low to high T_soil was 22, 50, and 47% under the low, intermediate and high moisture regimes, respectively. In contrast, the T_soil×moisture effect on stem biomass was significant after 2 months, but not after 4 months of treatment. High T_soil increased leaf mass ratio (LMR) after 4 months of treatment, but decreased both root mass ratio (RMR) after both 2 and 4 months, and root:shoot ratio (RSR) after 4 months of treatment. The low moisture regime decreased LMR after 2 and 4 months of treatment, but increased RSR after 4 months of treatment. There were no significant [CO₂] effects on biomass allocation or [CO₂]×T_soil×moisture interactions on biomass production/allocation.     

Internal Leaf Area and Cellular CO2 Resistance: Photosynthetic Implications of Variations with Growth Conditions and Plant Species

The influences of illumination, temperature, and soil water potential during development on leaf thickness, mesophyll cell wall area per unit leaf area (^Ames/A), and the cellular CO₂, resistance expressed on a mesophyll cell wall area basis (r_CO2^cell,) were examined for Plectranthus parviflorus Henckel. Although the ranges of all three growth conditions caused at least 9-fold variations in the leaf biomass produced in 4 weeks, only the illumination had a major effect on internal leaf morphology, e.g. the thickness went from 279 to 831 μm and A^mes/A from 10.5 to 34.8 as the photosynthetically active radiation was raised from 3 to 53 nEinsteins cm^?2 s^?1, while r_CO2^cell remained close to 154 s cm^?1. Variations in the growth temperature, soil water potential, and the nutritional status of the plant, affected photosynthesis mainly by changes in r_CO2^cell. To compare the influence of internal leaf area on photosynthesis for other plants, especially those with low A^mes/A values, the maximum rates of CO₂ uptake at light saturation and photosynthetically optimal temperatures were also determined for a moss, Mnium ciliare (C. Muell.) Schimp., and two ferns, Adiantum decorum Moore and Alsophila australe R. Br. As Ames/A went from 2.00 for the moss to 3.8, 7.5, 11.7, and 20.8 for the fens, the illumination at light saturation and the maximum rate of photosynthesis both progressively increased. The cellular CO₂ resistance, which theoretically might have a lower limit of 20 s cm^?1, ranged from 85 to 190 s cm^?1.     

EXTRACELLULAR POLYSACCHARIDE AND LEAF CAPACITANCE IN A HAWAIIAN BOG SPECIES,ARGYROXIPHIUM GRAYANUM (COMPOSITAE-MADIINAE)

Mature leaves of Argyroxiphium grayanum exhibit an unusual structure-function relationship. The leaves are traversed by several large channels filled with extracellular polysaccharide. Infrared spectral data suggest that the polysaccharide is mainly pectin, which is consistent with previous data based on its histochemical staining properties. The polysaccharide exhibits a very high water-holding capacity, which appears to result in a very high leaf capacitance at high water contents and water potentials. The volume-normalized leaf capacitance of A. grayanum is 0.67 MPa^–1 over the water potential range of –0.1 to –0.5 MPa. Coupled with the low time constant for water flux within the leaves (57 sec), the high leaf capacitance of A. grayanum may enable it to avoid rapid declines in leaf water potential. Thus, in its native bog habitat in Hawaii, the high leaf capacitance of A. grayanum may be of benefit in terms of minimizing the development of short-term leaf water deficits.     

LEAF OPTICAL PROPERTIES OF RAINFOREST SUN AND EXTREME SHADE PLANTS

The optical properties of the leaves of twelve tropical sun species and thirteen tropical extreme shade species were examined with an integrating sphere attached to a spectroradiometer. Measurements of diffuse reflectance and transmittance allowed calculations of absorptance, 350–1,100 nm. Although some shade species absorbed higher percentages of quantum flux densities for photosynthesis (400–700 nm, PPFD) than the mean for the sun species, the sun and shade species as groups were not significantly different from each other: 90.2, S.D. 3.6% for shade species and 88.6, S.D. 2.4% for the sun species. The groups of species did not differ in total absorptance of energy 350–1,100 nm. Furthermore, the sun and shade species were identical in their shift of absorptance at wavelengths between 650 and 750 nm. The anthocyanic coloration of the leaf undersurfaces of two species polymorphic for this characteristic (Trionela hirsuta and Ischnosciphon pruinosus) is correlated with increased absorptance at the upper end of the action spectrum of photosynthesis. Although sun and shade species have similar optical properties, the energy investment (as documented by dry wt per unit area of leaf surface) is much less for the shade species.     

AUXIN-INDUCED HYPONASTY OF THE LEAF BLADE OF PHASEOLUS VULGARIS

The lateral margins of immature primary leaf blades of Phaseolus vulgaris L. cv. ‘Pinto’ curve up and in toward the midrib when auxin is applied to the leaf. The leaves are most sensitive to auxin shortly after they first unfold and leaves which have grown to about 60 % or more of their ultimate area no longer give this hyponastic response. The response is specific for auxins and is inhibited by the anti-auxins, trans-cinnamic acid and para-chlorophenoxyisobutyric acid. Ethylene and ethylene-generating compounds failed to induce hyponasty, suggesting the response is due to a positive growth promotion by auxin. Measurements of the distance between the lateral margins of the leaf at its maximum width were used to provide quantitative estimates of the degree of hyponasty. Between 2 and 4 hr after auxin application a direct proportionality was found between the amount of curvature and the logarithm of the indoleacetic acid concentration over the range of 10⁻⁶ to 10⁻³ m. The relative sensitivity of the leaves to different auxins was qualitatively similar to that observed in many straight-growth bioassays. Similar responses were obtained when auxin was applied by a carborundum wounding procedure. Potential applications of this auxin bioassay for investigations of the role of auxin in the normal plagiotropic growth behavior of leaf lamina and of the role of auxin in the initiation of various plant diseases are suggested.     

CAUSAL MECHANISMS OF LEAF DIMORPHISM IN THE AQUATIC ANGIOSPERM CALLITRICHE HETEROPHYLLA

This report investigates the physiological basis for the production of dimorphic leaves on the aquatic angiosperm Callitriche heterophylla. In nature, the leaf morphology of this plant depends on whether the shoot apex is submerged in or emergent from water. The water-form leaves that develop on submerged apices assume a long, linear shape in contrast to the short, obovate appearance of land forms on emergent apices. The parameters of length/width ratio and stomatal density were used as developmental indices to characterize how natural conditions, fluctuating water levels and other experimental treatments affect leaf shape. Transferring submerged and emergent shoots to the alternative culture conditions caused immature leaves to assume the characteristics appropriate to their new environment. Moreover, the treatments of 0.24 mol mannitol, high temperature (30 C) and 10^−-5 m abscisic acid induced submerged shoots to produce land-form leaves whereas 10^−-5 m gibberellic acid mediated the development of water-form leaves on emergent shoots. Water, osmotic and pressure potentials of immature leaves in the control and experimental treatments were determined by thermocouple psychrometry. Under natural conditions, growing water forms exhibited high turgors (3–5 bars) while developing land forms showed much lower turgors (0–1 bar). Similar correlations between turgor pressure and leaf morphology were observed in the case of the gibberellic acid and mannitol treatments. However, abscisic acid and high temperature caused the developing land-form leaves to exhibit high turgors without a concomitant change to the water-form morphology. Microscopic measurements of epidermal cells established that irrespective of the experimental conditions, water-form leaves had longer and narrower epidermal cells with less convoluted anticlinal walls than land forms. Cell counts indicated that the numbers of epidermal cells did not account for the observed differences in leaf morphology. The results are interpreted in terms of how cell expansion might regulate leaf morphology in aquatic angiosperms.     

MICROCLIMATIC EFFECTS ON WATER RELATIONS,LEAF TEMPERATURES,AND THE DISTRIBUTION OF HERACLEUM LANATUM AT HIGH ELEVATIONS

The small-scale distribution of an understory herb, Heracleum lanatum, was evaluated in terms of leaf temperature and water relations limitations due to a large leaf size (630 cm²). Diurnal variations in transpiration (4 to 60 mg m⁻² s⁻¹) were influenced by fluctuations in solar irradiance, wind speed, leaf temperature and stomatal conductance. Computer simulations indicated that leaf temperatures in a forest clearing would be > 12 C above air temperature, with maximum transpiration rates of 140 mg m⁻² s⁻¹, and daily water loss to be over 200% greater than values at natural understory locations. Simulations of nocturnal temperature relations indicated ~100 W m ⁻² less incident longwave irradiance in the forest clearing as compared to the understory (560 vs. 660 W m⁻² at 400 hr). This difference led to predicted leaf temperatures being as low as 6 C below air temperature in the forest clearing while measured leaf temperatures in the forest understory were within 1.5 C of air temperature throughout the night. Furthermore, minimum air temperatures were at or below 6 C on 36% of the nights during the summer growth period indicating that in open areas leaves of H. lanatum would frequently be below 0 C and subject to possible freeze damage. Heracleum lanatum may be more abundant in the shaded understory of the subalpine forest because exposure in open environments would result in high leaf temperatures and increased transpirational water loss during the day, as well as low leaf temperatures with the possibility of freeze damage at night.     

Characterization of a glabrate Encelia farinosa mutant: morphology,ecophysiology, and field observations

An individual Encelia farinosa (Asteraceae) from Death Valley, California is described that is completely lacking in the dense leaf pubescence covering characteristic of the species. While leaf absorptances to solar radiation of the mutant E. farinosa are much greater than those of the wild type, other morphological aspects and photosynthetic characteristics appear to be similar to those typical for E. farinosa. Leaf temperatures of mutant and wild type E. farinosa in the field are similar, but only because of steeper leaf angles and higher leaf conductances to water vapor in the mutant form. As a consequence of a greater water expenditure for transpirational cooling, the mutant E. farinosa becomes deciduous much earlier in the drought period than do the normal pubescent leaved E. farinosa.