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.     

EFFECTS OF CANOPY POSITION AND IRRADIANCE ON THE LEAF PHYSIOLOGY AND MORPHOLOGY OF PENTACLETHRA MACROLOBA (MIMOSACEAE)

Pentaclethra macroloba (Willd.) Kuntze (Mimosaceae) is a dominant late-successional tree species in the Atlantic lowland forests of Costa Rica. Leaves of P. macroloba from three heights in the forest canopy were compared with leaves of seedlings grown in controlled environment chambers under four different irradiance levels. Changes in leaf characteristics along the canopy gradient paralleled changes resulting from the light gradient under controlled conditions. The effect of light or canopy position on light-saturated photosynthesis was small, with maximum photosynthesis increasing from 5 to 6.5 μmol m^−-2 s^−-1 from understory to canopy. Both chamber grown and field leaves showed large adjustments in photosynthetic efficiency at low light via reductions in dark respiration rates and increases in apparent quantum yields. Light saturation of all leaves occurred at or below 500 μmol m^−-2 s^−-1. Leaf thickness, specific leaf weight, and stomatal density increased to a greater extent than saturated photosynthesis with higher irradiance during growth or height in the canopy. As a result, there was a poor correspondence between leaf thickness and light-saturated photosynthesis on an area basis. It is concluded that Pentaclethra macroloba possesses the characteristics of a typical shade-tolerant species.     

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis

Vertical structure of plant stands and canopies may change under conditions of elevated CO₂ due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO₂ on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO₂-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 μl l⁻¹) and elevated (610 μl l⁻¹) CO₂. Plants in each stand were grown in a common soil as closed “artificial ecosystems” with a ground area of 6.7 m². LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO₂ treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO₂ the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO₂. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO₂. CO₂ enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO₂ at the stand-level. We suggest that, under elevated CO₂, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.     

Variability of stomatal conductance, leaf anatomy, and seasonal leaf wettability of young and adult European beech leaves along a vertical canopy gradient

This study assessed the variation of leaf anatomy, chlorophyll content index (CCI), maximal stomatal conductance (g _s ^max ) and leaf wettability within the canopy of an adult European beech tree (Fagus sylvatica L.) and for beech saplings placed along the vertical gradient in the canopy. At the top canopy level (CL_28m) of the adult beech, CCI and leaf anatomy reflected higher light stress, while g _s ^max increased with height, reflecting the importance of gas exchange in the upper canopy layer. Leaf wettability, measured as drop contact angle, decreased from 85.5°?±?1.6° (summer) to 57.5°?±?2.8° (autumn) at CL_28m of the adult tree. At CL_22m, adult beech leaves seemed to be better optimized for photosynthesis than the CL_28m leaves because of a large leaf thickness with less protective and impregnated substances, and a higher CCI. The beech saplings, in contrast, did not adapt their stomatal characteristics and leaf anatomy according to the same strategy as the adult beech leaves. Consequently, care is needed when scaling up experimental results from seedlings to adult trees.     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

Seasonal changes in photosynthesis and nitrogen allocation in leaves of different ages in evergreen understory shrub <Emphasis Type="Italic">Daphniphyllum humile</Emphasis>

Seasonal changes in photosynthetic capacity, leaf nitrogen (N) content, leaf chlorophyll (Chl) content and leaf N allocation patterns in leaves of different ages in the evergreen understory shrub, Daphniphyllum humile Maxim, growing at a forest border and an understory site were studied. In current-year leaves at the understory site, the N and Rubisco contents increased from spring to autumn although their light-saturated photosynthetic rate at 22°C (P _max²²) remained stable, indicating that their mesophyll conductance rates declined as they completed their development and/or that they invested increasing amounts of their resources in photosynthetic enzymes during this period. In contrast, seasonal changes in P _max²² in current-year leaves at the forest border site were correlated with changes in Rubisco content. In 1-year old leaves at the understory site, P _max²² and contents of Chl, leaf N, and Rubisco remained stable from spring to autumn, while these parameters decreased in 1-year-old forest border leaves, indicating that N may have been remobilized from shaded 1-year-old leaves to sunlit current-year leaves. When leaves senesced at the forest border site the Rubisco content decreased more rapidly than that of light-harvesting proteins such as LHCII, suggesting that N remobilization from Rubisco may be more efficient, possibly because Rubisco has greater N costs and is soluble, whereas the light-harvesting proteins are membrane components.     

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

Summary 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 mol 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.     

SEASONAL PATTERNS OF LEAF PHOTOSYNTHETIC CAPACITY IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Seasonal patterns of leaf photosynthetic capacity and conductance were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan. Leaves of bigtooth aspen and red oak at the top of the canopy had higher maximum CO₂ Exchange Rate (CER) (10–15 μmol m ² s ¹) than leaves of sugar maple, red maple, red oak, and beech growing in the understory (4–5 μmol m ² s ¹). In all leaves, CER measured at light-saturation increased to a maximum near the completion of leaf expansion in early June, was constant until mid-September, and then rapidly declined until leaf death. A similar pattern was seen for CER measured in low light (1.5% full sun). Respiration rate in the dark was highest in young leaves and decreased during leaf expansion; a relatively constant rate was then maintained for the rest of leaf lifespan. The seasonal pattern of the initial slope of the light response of CER paralleled the pattern of light-saturated CER. The initial slope in midsummer ranged from values of 37 to 44 μmol/mol for species in the understory to 51 and 56 μmol/mol for red oak and bigtooth aspen, respectively, at the top of the canopy. Leaf conductance was constant throughout most of leaf lifespan, with some decline occurring in autumn. Leaves at the top of the canopy had higher conductances for water vapor (2–5 mm/s) than leaves in the understory (1–2 mm/s). All species maintained leaf intercellular CO, mole fractions (c,) near 200 uML/L until autumn, when c, increased during leaf senescence.     

Xanthophyll-cycle pigments and photosynthetic capacity in tropical forest species: a comparative field study on canopy,gap and understory plants

Xanthophyll-cycle pigments and photosynthetic capacity (PS_max) were analyzed in 25 species from different light environments (canopy, gap, understory) within a Panamanian tropical forest. (1) Sun-exposed leaves of canopy tree species showed the highest photosynthetic capacities and largest xanthophyll-cycle pools (violaxanthin, antheraxanthin, zeaxanthin) of about 87 mmol mol^-1 chlorophyll with only small amounts of -carotene [about 7 mmol mol^-1 chlorophyll = 8% of total (+) carotene pool]. Under high natural photon flux densities (PFDs) canopy leaves rapidly converted up to 96% of the xanthophyll-cycle pool into zeaxanthin. The back reaction to violaxanthin occurred much faster in low light than in complete darkness. At the end of the night, zeaxanthin still accounted for, on average, 14% of the total xanthophyll-cycle pigments. (2) Leaves of gap plants had intermediate values of PS_max and a 43% lower total carotenoid content than canopy leaves. The average size of the xanthophyll-cycle pool was 35 mmol mol^-1 chlorophyll, and -carotene accounted for up to 66% of the total (+) carotene pool. Under high light conditions gap plants converted, on average, 86% of the xanthophyll-cycle pigments into zeaxanthin. The back reaction, following a decrease in ambient PFD, was slower than the forward reaction. At the end of the night, zeaxanthin accounted for, on average, 7% of the xanthophyll-cycle pigments in gap plants. (3) Understory plants showed the lowest values of PS_max and the smallest xanthophyll-cycle pool of about 22 mmol mol^-1 chlorophyll. -Carotene accounted for up to 70% of total carotene. The conversion of xanthophyll-cycle pigments into zeaxanthin was negligible during short sunflecks of 1–2 min duration and PFDs up to about 400 mol m^-2 s^-1. At predawn, leaves of understory plants rarely contained any detectable zeaxanthin. Aechmea magdalenae, an understory CAM plant, showed exceptionally high rates of PS_max per unit leaf area compared to sympatric C₃ understory species.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. I. Leaf model parametrization

In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m^?2 s^?1) than in sun leaves (11–5μmol m^?2 s^?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).     

Linkage between seasonal gas exchange and hydraulic acclimation in the top canopy leaves of Fagus trees in a mesic forest in Japan

We investigated how leaf gas exchange and hydraulic properties acclimate to increasing evaporative demand in mature beech trees, Fagus crenata Blume and Fagus japonica Maxim., growing in their natural habitat. The measurements in the top canopy leaves were conducted using a 16-m-high scaffolding tower over two growing seasons. The daily maxima of net photosynthetic rate for the early growing season were close to the annual maximum value (11.9 mol m^–2 s^–1 in F. crenata and 7.7 mol m^–2 s^–1 in F. japonica). The daily maxima of water vapor stomatal conductance were highest in the summer, approximately 0.3 mol m^–2 s^–1 in F. crenata and 0.15 mol m^–2 s^–1 in F. japonica. From the early growing season to the summer season, the leaf-to-air vapor pressure deficit increased and the daily minima of leaf water potentials decreased. However, there was no loss of leaf turgor in the summer as a result of effective osmotic adjustment. Both the soil-to-leaf hydraulic conductance per unit leaf area and the twig hydraulic conductivity simultaneously increased in the summer, probably as a result of production of new vessels in the xylem. These results suggest that both osmotic adjustment and increased hydraulic conductance resulted in the largest diurnal maximum of stomatal conductance in the summer, resulting in the lowest relative stomatal limitation on net photosynthetic rate, although the leaf-to-air vapor pressure deficit was highest. These results indicate that even in a mesic forest, in which excessive hydraulic stress does not occur, the seasonal acclimation of hydraulic properties at both the single leaf and whole plant levels are important for plant carbon gain.     

Linking light attenuation,sunflecks, and canopy architecture in mesic shrub thickets

Expansion of shrubs into grasslands is often accompanied by a reduction in understory light and an associated reduction of shade-intolerant species. However, effects of specific canopy architectural characteristics on the light environment under shrub thickets are unknown. Our objective was to determine what characteristics of canopy architecture most influence understory light in monospecific shrub thickets. We quantified understory light and canopy architecture for five shrub species in the eastern United States that have a history of expansion, and we used multiple regression to determine which canopy characteristics best predicted light attenuation and relative contribution of sunflecks. Measurements included leaf angle, leaf azimuth, branch bifurcation ratio, leaf area index (LAI), canopy depth (the vertical distance from the bottommost leaf to the top of the canopy), and leaf area density (LAD) as well as understory photosynthetic photon flux density (PPFD). The best predictor of light attenuation and the occurrence of sunflecks for all species was canopy depth. Multiple leaf and plant-level traits were correlated with canopy depth but not with LAI or LAD. The invasive shrub Elaeagnus umbellata had the lowest understory light levels of the species examined although LAI values for Morella cerifera and Rhododendron maximum were higher. Branch bifurcation ratios for E. umbellata were significantly higher than for other species and this likely contributed to the differences in light attenuation and suppression of sunflecks. The potential of shrubs to intercept light is primarily dependent on vertical distribution of leaves in the canopy which is itself correlated with fine-scale, species-specific variations in leaf display.     

The role of leaf and canopy-level gas exchange in the replacement of Quercus virginiana (Fagaceae) by Juniperus ashei (Cupressaceae) in semiarid savannas

Photosynthesis, transpiration, and leaf area distribution were sampled in mature Quercus virginiana and Juniperus ashei trees to determine the impact of leaf position on canopy-level gas exchange, and how gas exchange patterns may affect the successful invasion of Quercus communities by J. ashei. Sampling was conducted monthly over a 2-yr period in 12 canopy locations (three canopy layers and four cardinal directions). Photosynthetic and transpiration rates of both species were greatest in the upper canopy and decreased with canopy depth. Leaf photosynthetic and transpiration rates were significantly higher for Q. virginiana (4.1–6.7 μmol CO₂·m⁻²·s⁻¹ and 1.1–2.1 mmol H₂O·m⁻²·s⁻¹) than for J. ashei (2.1–2.8 μmol CO₂·m⁻²·s⁻¹ and 0.7–1.0 mmol H₂O·m⁻²·s⁻¹) in every canopy level and direction. Leaves on the south and east sides of both species had higher gas exchange rates than leaves on the north and west sides. Although Quercus had a greater mean canopy diameter than Juniperus (31.3 vs. 27.7 m²), J. ashei had significantly greater leaf area (142 vs. 58 m²/tree). A simple model combining leaf area and gas exchange rates for different leaf positions demonstrated a significantly greater total canopy carbon dioxide uptake for J. ashei compared to Q. virginiana (831 vs. 612 g CO₂·tree⁻¹·d⁻¹, respectively). Total daily water loss was also greater for Juniperus (125 vs. 73 Ltree⁻¹·d⁻¹). Differences in leaf gas exchange rates were poor predictors of the relationship between the invasive J. ashei and the codominant Q. virginiana. Leaf area and leaf area distribution coupled with leaf gas exchange rates were necessary to demonstrate the higher overall competitive potential of J. ashei.     

Water use by the desert cucurbit Citrullus colocynthis (L.) Schrad.

Summary The rates of water use and leaf surface conductance of Citrullus colocynthis (Cucurbitacea) were evaluated from measurements of the surface temperature and microenvironment of leaves. At desert sites in Saudi Arabia the transpiration rates reached 0.13–0.17 g m^-2 s^-1 and the leaf temperatures were always close to air temperature. Leaf models (dry) placed in the canopy were considerably warmer than the air. To investigate responses over a wider range of conditions, plants were grown in a controlled environment room. It was found that when conditions were made hotter than those that occurred in the desert, the stomatal conductance increased greatly. Transpiration rate attained 0.6 g m^-2 s^-1 and the leaves were up to seven degrees cooler than the air. The results suggest a finely-tuned control mechanism working like a switch when the leaves experience extreme conditions, and enabling the plant to avoid lethal temperatures.     

Photosynthesis of individual field-grown cotton leaves during ontogeny

Photosynthetic characteristics of field-grown cotton (Gossypium hirsutum L.) leaves were determined at several insertion levels within the canopy during the growing season. Single-leaf measurements of net photosynthesis (Pn), stomatal conductance to CO₂ (g_s·CO₂), substomatal CO₂, leaf area expansion, leaf nitrogen, and light intensity (PPFD) were recorded for undisturbed leaves within the crop canopy at 3–4 day intervals during the development of all leaves at main-stem nodes 8, 10, and 12. Patterns of Pn during leaf ontogeny exhibited three distinct phases; a rapid increase to maximum at 16–20 days after leaf unfolding, a relatively short plateau, and a period of linear decline to negligible Pn at 60–65 days. Analysis of the parameters which contributed to the rise and fall pattern of Pn with leaf age indicated the primary involvement of leaf area expansion, leaf nitrogen, PPFD, and g_s·CO₂ in this process. The response of Pn and g_s·CO₂ to incident PPFD conditions during canopy development was highly age dependent. For leaves less than 16 days old, the patterns of Pn and g_s·CO₂ were largely controlled by non-PPFD factors, while for older leaves Pn and g_s·CO₂ were more closely coupled to PPFD-mediated processes. Maximum values of Pn were not significantly different for any of the leaves monitored in this study, however, those leaves at main-stem node 8 did possess a significantly diminished photosynthetic capacity with age compared to upper canopy leaves. This accelerated decline in Pn could not be explained by age-related variations in g_s·CO₂ since all leaves showed similar changes in g_s·CO₂ with leaf age.Abbreviations g_s·CO₂ stomatal conductance to CO₂ - Pn net photosynthesis - PPFD photosynthetic photon flux density     

热带次生林不同林层植物叶片非结构性碳水化合物的季节变化及其对氮磷添加的响应

非结构性碳水化合物(Non-structural Carbohydrates, NSCs)是植物生长代谢过程中重要的能量来源。通过在华南热带次生林进行氮磷添加试验,探究不同林层植物叶片NSCs的季节变化及其对氮磷添加的响应,取样时间为2019年1月、4月、7月和10月。结果表明:1)植物叶片NSCs存在显著的种间差异,磷(P)添加对叶片淀粉和NSCs含量具有显著影响,且物种与磷添加的交互作用显著影响叶片淀粉含量。2)黑嘴蒲桃和紫玉盘叶片NSCs含量对氮(N)添加的响应较为敏感,而白车和竹节叶片NSCs含量对P添加的响应较为敏感,氮磷同时添加(+NP)对植物叶片NSCs的增效作用最好。3)植物叶片NSCs存在显著的季节性变化,且季节与林层间的交互作用对叶片可溶性糖和NSCs含量具有显著影响。4)不同林层植物对氮磷添加的响应不同,氮磷添加使林下层植物叶片可溶性糖含量增高,林冠层降低,在干季,N添加会使林下层植物叶片淀粉含量增高,林冠层降低。P添加的影响恰好与之相反。在湿季,氮磷添加使林下层和林冠层植物叶片的淀粉含量增加。5)林冠层植物叶片NSCs含量高于林下层,且林下层植物叶片NSCs含量...     

Photosynthetic strategies of summergreen and evergreen understory herbs of the boreal mixedwood forest

Seasonal differences in photosynthesis and stomatal conductance of four herbaceous perennials from beneath a deciduous canopy was assessed at two light levels (60 and 400 μmol m⁻² s⁻¹ photosynthetic photon flux density, PPFD) and two leaf temperatures (7 and 20°C). Leaves of an evergreen, Pyrola asarifolia Michx., a wintergreen, Cornus canadensis L., and two summergreen species, Rubus pubescens Raf. and Aralia nudicaulis L., were collected at four times during the growing season. In addition, midsummer light response curves were obtained for one summergreen (A. nudicaulis) and one evergreen species (P. asarifolia) at both 7 and 20°C. Gas exchange measurements were made in the laboratory under controlled environmental conditions. For leaves collected in April, when insolation was high due to the leafless overstory, only P. asarifolia had green leaves, and there was no effect of temperature or light on this species' photosynthesis. P. asarifolia's net assimilation rate (NA) in April was about 30% of it's maximum in late summer. In early summer (June), A. nudicaulis and R. pubescens had higher NA at the higher temperature; at this time, these summergreen species also reached their maximum NA. Midsummer photosynthetic light response curves showed that the light-saturation point was higher and more responsive to leaf temperature in the summergreen A. nudicaulis than in the evergreen P. asarifolia. The summergreen species appear to have a photosystem which performs at high rates during early- and mid-summer, as well as a taller stature which allows them to intercept more light. The photosynthetic system of the ever/wintergreen species is adapted to the low ground-level light conditions in the summer and there does not appear to be an adjustment to take further advantage of the higher light in the spring and fall period. The adaptation of the evergreen and wintergreen understory species is tolerance to low temperatures, enabling them to photosynthesize into the fall till the first continuous frosts occur in the understory and also permitting the evergreen species to begin photosynthesis early in the spring. Received: 17 October 1996 / Accepted: 2 May 1997     

Gas exchange acclimation to elevated CO2 in upper-sunlit and lower-shaded canopy leaves in relation to nitrogen acquisition and partitioning in wheat grown in field chambers

Growth at elevated CO₂ often decreases photosynthetic capacity (acclimation) and leaf N concentrations. Lower-shaded canopy leaves may undergo both CO₂ and shade acclimation. The relationship of acclimatory responses of flag and lower-shaded canopy leaves of wheat (Triticum aestivum L.) to the N content, and possible factors affecting N gain and distribution within the plant were investigated in a wheat crop growing in field chambers set at ambient (360 μmol mol⁻¹) and elevated (700 μmol mol⁻¹) CO₂, and with two amounts of N fertilizer (none and 70 kg ha⁻¹ applied on 30 April). Photosynthesis, stomatal conductance and transpiration at a common measurement CO₂, chlorophyll and Rubisco levels of upper-sunlit (flag) and lower-shaded canopy leaves were significantly lower in elevated relative to ambient CO₂-grown plants. Both whole shoot N and leaf N per unit area decreased at elevated CO₂, and leaf N declined with canopy position. Acclimatory responses to elevated CO₂ were enhanced in N-deficient plants. With N supply, the acclimatory responses were less pronounced in lower canopy leaves relative to the flag leaf. Additional N did not increase the fraction of shoot N allocated to the flag and penultimate leaves. The decrease in photosynthetic capacity in both upper-sunlit and lower-shaded leaves in elevated CO₂ was associated with a decrease in N contents in above-ground organs and with lower N partitioning to leaves. A single relationship of N per unit leaf area to the transpiration rate accounted for a significant fraction of the variation among sun-lit and shaded leaves, growth CO₂ level and N supply. We conclude that reduced stomatal conductance and transpiration can decrease plant N, leading to acclimation to CO₂ enrichment.     

Herbivory in canopy gaps created by a typhoon varies by understory plant leaf phenology

1. Availabilities of light and soil nitrogen for understory plants vary by extent of canopy gap formation through typhoon disturbance. We predicted that variation in resource availability and herbivore abundance in canopy gaps would affect herbivory through variation in leaf traits among plant species. We studied six understory species that expand their leaves before or after canopy closure in deciduous forests. We measured the availabilities of light, soil nitrogen, soil water content, and herbivore abundance in 20 canopy gaps (28.3–607.6 m²) formed by a typhoon and in four undisturbed stands. We also measured leaf traits and herbivory on understory plants. 2. The availabilities of light and soil nitrogen increased with increasing gap size. However, soil water content did not. The abundance of herbivorous insects (such as Lepidoptera and Orthoptera) increased with increasing gap size. 3. Concentrations of condensed tannins, total phenolics, and nitrogen in leaves and the leaf mass per area increased in late leaf expansion species with increasing gap size, whereas none of the leaf traits varied by gap size in early leaf expansion species. 4. Herbivory increased on early leaf expansion species with increasing gap size, but decreased on late leaf expansion species. In these late leaf expansion species, total phenolics and C : N ratio had negative relationships with herbivory. 5. These results suggested that after typhoon disturbance, increased herbivory on early leaf expansion species can be explained by increased herbivore abundance, whereas decreased herbivory on late leaf expansion species can be explained by variation in leaf traits.     

The effects of light acclimation during and after foliage expansion on photosynthesis ofAbies amabilis foliage within the canopy

Variation in the photosynthetic function ofAbies amabilis foliage within a canopy was examined and related to three different processes that affect foliage function: foliage aging, sun-shade acclimation that occurred while foliage was expanding, and reacclimation after expansion was complete. Foliage produced in the sun had higher photosynthesis at light saturation (A _max, mol·m^-2·s^-1), dark respiration (mol·m^-2·s^-1), nitrogen content (g·m^-2), chlorophyll content (g·m^-2), and chlorophylla:b ratio, and a lower chlorophyll to nitrogen ratio (chl:N), than foliage produced in the shade. As sun foliage becomes shaded, it becomes physiologically similar to shade foliage, even though it still retains a sun morphology. Shaded sun foliage exhibited lowerA _max, dark respiration, nitrogen content, and chlorophylla:b ratio, and a higher chl:N ratio than sun foliage of the same age remaining in the open. However, shaded sun foliage had a higher chlorophyll content than sun foliage remaining in the open, even though true shade foliage had a lower chlorophyll content than sun foliage. This anomaly arises because as sun foliage becomes shaded, it retains a higher nitrogen content than shade foliage in a similar light environment, but the two forms have similar chl:N ratios. Within the canopy, most physiological indicators were more strongly correlated with the current light environment than with foliage age or leaf thickness, with the exception of chlorophyll content.A _max decreased significantly with both decreasing current light environment of the foliage and increasing foliage age. The same trend with current light and age was found for the chlorophylla:b ratio. Foliage nitrogen content also decreased with a decrease in current light environment, but no distinct pattern was found with foliage age. Leaf thickness was also important for predicting leaf nitrogen content: thicker leaves had more nitrogen than thinner leaves regardless of light environment or age. The chl:N ratio had a strong negative correlation with the current light environment, and, as with nitrogen content, no distinct pattern was found with foliage age. Chlorophyll content of the foliage was not well correlated with any of the three predictor variables: current light environment, foliage age or leaf thickness. On the other hand, chlorophyll content was positively correlated with the amount of nitrogen in a leaf, and once nitrogen was considered, the current light environment was also highly significant in explaining the variation in chlorophyll content. It has been suggested that the redistribution of nitrogen both within and between leaves is a mechanism for photosynthetic acclimation to the current light environment. Within theseA. amabilis canopies, both leaf nitrogen and the chl:N ratio were strongly correlated with the current light environment, but only weakly with leaf age, supporting the idea that changing light is the driving force for the redistribution of nitrogen both within and between leaves. Thus, our results support previous theories on nitrogen distribution and partitioning. However,A _max was significantly affected by both foliage age and the current light environment, indicating that changes in light alone are not enough to explain changes inA _max with time.