The Influence of Previous Irradiance on Photosynthetic Induction in Three Species Grown in the Gap and Understory of a Fagus Crenata Forest

Photosynthetic induction responses to a sudden increase in photosynthetic photon flux density (PPFD) from lower background PPFD (0, 25, 50, and 100 mol m^–2 s^–1) to 1 000 mol m^–2 s^–1 were measured in leaves of Fagus crenata, Acer rufinerve Siebold & Zucc., and Viburnum furcatum growing in a gap and understory of a F. crenata forest in the Naeba mountains. In the gap, A. rufinerve exhibited more than 1.2-fold higher maximum net photosynthetic rate (P _Nmax) than F. crenata and V. furcatum. Meanwhile, in the understory F. crenata exhibited the highest P _Nmax among the three species. The photosynthetic induction period required to reach P _Nmax was 3–41 min. The photosynthetic responses to increase in PPFD depended on the background PPFD before increase in PPFD. The induction period required to reach P _Nmax was 2.5–6.5-fold longer when PPFD increased from darkness than when PPFD increased from 100 mol m^–2 s^–1. The induction period was correlated with initial P _N and stomatal conductance (g _s) relative to maximum values before increase in PPFD. The relationship was similar between the gap and the understory. As the background PPFD increased, the initial P _N and g _s increased, indicating that the degrees of biochemical and stomata limitations to dynamic photosynthetic performance decreased. Therefore, photosynthetic induction responses to increase in PPFD became faster with the increasing background PPFD. The differences in time required to reach induction between species, as well as between gap and understory, were mainly due to the varying of relative initial induction states in P _N and g _s at the same background PPFD.     

Environmental controls on the photosynthesis and respiration of a boreal lichen woodland: a growing season of whole-ecosystem exchange measurements by eddy correlation

Measurements of net ecosystem CO₂ exchange by eddy correlation, incident photosynthetically active photon flux density (PPFD), soil temperature, air temperature, and air humidity were made in a black spruce (Picea mariana) boreal woodland near Schefferville, Quebec, Canada, from June through August 1990. Nighttime respiration was between 0.5 and 1.5 kg C ha^–1 h^–1, increasing with temperature. Net uptake of carbon during the day peaked at 3 kg C ha^–1 h^–1, and the daily net uptake over the experiment was 12 kg C ha^–1 day^–1. Photosynthesis dropped substantially at leaf-to-air vapor pressure deficit (VPD) greater than 7 mb, presumably as a result of stomatal closure. The response of ecosystem photosynthesis to incident PPFD was markedly non-linear, with an abrupt saturation at 600 mol m^–2 s^–1. This sharp saturation reflected the geometry of the spruce canopy (isolated conical crowns), the frequently overcast conditions, and an increase in VPD coincident with high radiation. The ecosystem light-use efficiency increased markedly during overcast periods as a result of a more even distribution of light across the forest surface. A mechanistic model of forest photosynthesis, parameterized with observations of leaf density and nitrogen content from a nearby stand, provided accurate predictions of forest photosynthesis. The observations and model results indicated that ecosystem carbon balance at the site is highly sensitive to temperature, and relatively insensitive to cloudiness.     

Roles of dominant understory Sasa bamboo in carbon and nitrogen dynamics following canopy tree removal in a cool‐temperate forest in northern Japan

To clarify the role of dense understory vegetation in the stand structure, and in carbon (C) and nitrogen (N) dynamics of forest ecosystems with various conditions of overstory trees, we: (i) quantified the above‐ and below‐ground biomasses of understory dwarf bamboo (Sasa senanensis) at the old canopy‐gap area and the closed‐canopy area and compared the stand‐level biomasses of S. senanensis with that of overstory trees; (ii) determined the N leaching, soil respiration rates, fine‐root dynamics, plant area index (PAI) of S. senanensis, and soil temperature and moisture at the tree‐cut patches (cut) and the intact closed‐canopy patches (control). The biomass of S. senanensis in the canopy‐gap area was twice that at the closed‐canopy area. It equated to 12% of total biomass above ground but 41% below ground in the stand. The concentrations of NO₃^? and NH₄⁺ in the soil solution and soil respiration rates did not significantly change between cut and control plots, indicating that gap creation did not affect the C or N dynamics in the soil. Root‐length density and PAI of S. senanensis were significantly greater at the cut plots, suggesting the promotion of S. senanensis growth following tree cutting. The levels of soil temperature and soil moisture were not changed following tree cutting. These results show that S. senanensis is a key component species in this cool‐temperate forest ecosystem and plays significant roles in mitigating the loss of N and C from the soil following tree cutting by increasing its leaf and root biomass and stabilizing the soil environment.     

Photosynthetic and leaf morphological characteristics in Leucaena leucocephala as affected by growth under different neutral shade levels

Morphological and physiological measurements on individual leaves of Leucaena leucocephala seedlings were used to study acclimation to neutral shading. The light-saturated photosynthetic rate (P_{n max}) ranged from 19.6 to 6.5 mol CO₂ m^–2 s^–1 as photosynthetic photon flux density (PPFD) during growth decreased from 27 to 1.6 mol m^–2 s^–1. Stomatal density varied from 144 mm^–2 in plants grown in high PPFD to 84 mm^–2 in plants grown in low PPFD. Average maximal stomatal conductance for H₂O was 1.1 in plants grown in high PPFD and 0.3 for plants grown in low PPFD. Plants grown in low PPFD had a greater total chlorophyll content than plants grown in high PPFD (7.2 vs 2.9 mg g^–1 on a unit fresh weight basis, and 4.3 vs 3.7 mg dm^–2 on a unit leaf area basis). Leaf area was largest when plants were grown under the intermediate PPFDs. Leaf density thickness was largest when plants were grown under the largest PPFDs. It is concluded that L. leucocephala shows extensive ability to acclimate to neutral shade, and could be considered a facultative shade plant.Abbreviations the initial slope of the photosynthesis vs PPFD curve - P_{n max} the light-saturated photosynthetic rate - PPFD photosynthetic photon flux density     

Light environment under Rhododendron maximum thickets and estimated carbon gain of regenerating forest tree seedlings

Canopy tree recruitment is inhibited by evergreen shrubs in many forests. In the southern Appalachian mountains of the USA, thickets of Rhododendron maximum L. restrict dominant canopy tree seedling survival and persistence. Using R. maximum as a model system, we examined available light under the thickets and the photosynthetic responses of seedlings of canopy tree species. We tested the hypothesis that the additional shading from under R. maximum drives carbon gain in seedlings below the threshold for growth and survival. A reduction in light under the thicket was found where canopy openness (derived from canopy photographs) under R. maximum was half the amount measured in forest without R. maximum. R.␣maximum also reduced direct radiation by 50% and diffuse radiation by 12–29% compared to forest without the shrub layer. Mean mid-day PPFD (photosynthetically active photon flux density between 1000 and 1400 h) under R. maximum (obtained from quantum sensors) was below 10 mol m⁻² s⁻¹ on both clear and overcast days and the amount of sunflecks greater than 10 mol m⁻² s⁻¹ PPFD was only 0–20 min per day. In contrast, forest without R. maximum received a mean PPFD of 18–25 mol m⁻² s⁻¹ on clear days and a cumulative sunfleck duration of 100–220 min per day in all sky conditions. Consistent with light availability between the sites, daily carbon gain in Quercus rubra L. seedlings was lower in forest with R. maximum compared to forest where the shrub was absent. The presence of the shrub layer also significantly suppressed average mid-day photosynthesis of both Q. rubra and Prunus serotina Ehrt. seedlings on 8 out of 11 measurement dates. However, parameters derived from light response curves between seedlings growing in forest sites with or without a thicket of R. maximum was significantly different only in A _max (maximum photosynthetic rate), indicating a lack of further acclimation to the deeper shade under R. maximum. While the additional shade cast by R. maximum is sufficient to prevent the regeneration of tree seedlings under this shrub, there was sufficient heterogeneity in light under the thicket to imply that deep shade only partially explains the complete inhibition of regenerating canopy trees under R. maximum.     

天童国家森林公园常绿阔叶林不同演替阶段群落光环境特征比较

常绿阔叶林是我国中亚热带东部典型植被类型,根据野外踏查和固定样地调查发现,天童国家森林公园内发育着常绿阔叶林一个完整的演替系列,包含着6个不同演替阶段。应用W inScanopy For C anopy A na lys is软件对研究区内不同演替阶段群落冠层进行分析,得到不同群落冠层和林下的光环境特征指标:PPFD(光合光量子通量密度)和相关的冠层结构形态学指标G ap fraction(空隙度)、LA I(叶面积指数)、M LA(平均叶倾角),通过对这些指标的分析比较,得到的基本规律大致是林冠层的光合有效光量子通量密度随演替逐渐降低,林冠下面的光合有效光量子通量密度随着群落演替的进展变化更为明显。马尾松林的林冠空隙度明显高于其他阶段的群落,总的趋势是随群落演替的进展而降低。叶面积指数随演替的进展而呈增加趋势。平均叶倾角随演替的进展先增大而后减小。这些结果反映了常绿阔叶林不同演替阶段群落由于不同树种树冠形态学结构的差异和微环境的不同,形成了特定群落内的特定光环境。     

Influence of krummholz mat microclimate on needle physiology and survival

Summary Microclimate and photosynthesis of krummholz mat growth forms of Picea engelmanii (Parry) and Abies lasiocarpa [Hook.] Nutt. were investigated to determine structural features which may aid survival in alpine environments. The structure of krummholz mats was described in terms of the vertical distribution of leaf area index and leaf area density, which exceeded 50 m^-1 (based on total leaf surface area) near the canopy surface and approached zero below 30 cm from the surface in both species. Photosynthetic photon flux density (PPFD, 0.4–0.7 m wavelengths) and wind decreased by an average of 6 and 50-fold, respectively, between 1 m above and 10 cm below mat surfaces in both species. Needle temperatures on a P. engelmannii krummholz mat during July averaged about 2°C above air temperature during the day, with a maximum overtemperature of greater than 20°C above T _air during one sunlit period. At night, needle temperatures averaged 3–4°C below T _air.Net photosynthesis in year-old P. engelmannii shoots reached a maximum at 15–20°C during July and August. Surface shoots were light saturated at near 1200 moles m^-2s^-1 PPFD, and had higher photosynthetic rates than subsurface, predominantly shaded shoots above 800 moles m^-2s^-1. Shade shoots had higher photosynthetic rates when PPFD was below 600 moles m^-2s^-1, and at 250 moles m^-2s^-1 shade shoots maintained about 50% of the net photosynthetic rate of sun shoots at light saturation. Shade shoots appeared capable of benefitting photosynthetically from elevated temperatures within krummholz mats despite relatively low light levels. Especially rapid photosynthesis may occur when canopy needles are illuminated by sunflecks and needle temperatures rise by 10° C or more.Snow cover appears crucial for the survival of needles during winter. Snow accumulated within krummholz needle canopies before the sub-canopy zone of unfoliated branches became filled. The concentrated needle growth in the krummholz canopy captured snow in early autumn without support from ground-level snowpack. Early snow cover in both species prevented cuticle abrasion and resulted in high winter needle water contents and viabilities for subsurface compared to surface needles which became abraded, severely dehydrated, and had high mortality between December and February, especially on windward sides of shoots.Extremely high concentrations of needles within krummholz mat canopies created an aerodynamic structure which elevated needle temperatures to more optimal photosynthetic levels in summer and resulted in more efficient snow accumulation in winter. These factors appear crucial for winter needle survival. Thus, krummholz mats appear to be an important adaptation in growth form which provides survival benefits in both summer and winter.     

Photosynthetic Response to Irradiance in Valeriana jatamansi Jones,a Threatened Understorey Medicinal Herb of Western Himalaya

Net photosynthetic rate (P _N) of Valeriana jatamansi plants, grown under nylon net shade or under different tree canopies, was saturated with photons at 1 000 mol m^–2 s^–1 photosynthetic photon-flux-density (PPFD), whereas open-grown plants were able to photosynthesise even at higher PPFD, e.g. of 2 000 mol m^–2 s^–1. Plants grown under net shade had higher total chlorophyll (Chl) content per unit area of leaf surface. However, Chl a/b ratio was maximal in open-grown plants, but remained unchanged in plants grown in nylon net shade and under different tree canopies. Sun-grown plants had thicker leaves (higher leaf mass per leaf area unit), higher wax content, and higher P _N than shade grown plants. Thus V. jatamansi is able to acclimate to high PPFD and therefore this Himalayan species may be cultivated in open habitat to meet the ever-increasing industrial demand.     

Influence of Etherel and Gibberellic Acid on Carbon Metabolism,Growth, and Essential Oil Accumulation in Spearmint (Mentha Spicata)

Controlled environment chamber and glasshouse studies were conducted on six herbaceous annual species grown at 350 (AC) and 700 (EC) mol(CO₂) mol^-1 to determine whether growth at EC resulted in acclimation of the apparent quantum yield of photosynthesis (QY) measured at limiting photosynthetic photon flux density (PPFD), or in acclimation of net photosynthetic rate (P _N) measured at saturating PPFD. It was also determined whether acclimation in P _N at limiting PPFD was correlated with acclimation of carboxylation efficiency or ribulose-1,5-bisphosphate (RuBP) regeneration rate measured at saturating PPFD. Growth at EC reduced both the QY and P _N at limiting PPFD in three of the six species. The occurrence of photosynthetic acclimation measured at a rate limiting PPFD was independent of whether photosynthetic acclimation was apparent at saturating measurement PPFD. At saturating measurement PPFD, acclimation to EC in the apparent carboxylation efficiency and RuBP regeneration capacity also occurred independently. Thus at least three components of the photosynthetic system may adjust independently when leaves are grown at EC. Estimates of photosynthetic acclimation at both high and low PPFD are necessary to accurately predict photosynthesis at the whole plant or canopy level as [CO₂] increases.     

Long-term kinetics of the light-dependent regulation of ribulose-1,5-bisphosphate carboxylase/oxygenase activity in plants with and without 2-carboxyarabinitol 1-phosphate

The light-dependent modulation of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity was studied in two species: Phaseolus vulgaris L., which has high levels of the inhibitor of Rubisco activity, carboxyarabinitol 1-phosphate (CA1P), in the dark, and Chenopodium album L., which has little CA1P. In both species, the ratio of initial to fully-activated Rubisco activity declined by 40–50% within 60 min of a reduction in light from high a photosynthetic photon flux density (PPFD; >700 mol · m^–2 · s^–1) to a low PPFD (65 ± 15 mol · m^–2 · s^–1) or to darkness, indicating that decarbamylation of Rubisco is substantially involved in the initial regulatory response of Rubisco to a reduction in PPFD, even in species with potentially extensive CA1P inhibition. Total Rubisco activity was unaffected by PPFD in C. album, and prolonged exposure (2–6 h) to low light or darkness was accompanied by a slow decline in the activity ratio of this species. This indicates that the carbamylation state of Rubisco from C. album gradually declines for hours after the large initial drop in the first 60 min following light reduction. In P. vulgaris, the total activity of Rubisco declined by 10–30% within 1 h after a reduction in PPFD to below 100 mol · m^–2 · s^–1, indicating CA1P-binding contributes significantly to the reduction of Rubisco capacity during this period, but to a lesser extent than decarbamylation. With continued exposure of P. vulgaris leaves to very low PPFDs (< 30 mol · m^–2 · s^–1), the total activity of Rubisco declined steadily so that after 6–6.5 h of exposure to very low light or darkness, it was only 10–20% of the high-light value. These results indicate that while decarbamylation is more prominent in the initial regulatory response of Rubisco to a reduction in PPFD in P. vulgaris, binding of CA1P increases over time and after a few hours dominates the regulation of Rubisco activity in darkness and at very low PPFDs.Abbreviations CA1P 2-carboxyarabinitol 1-phosphate - CABP 2-carboxyarabinitol 1,5-bisphosphate - k_cat substrate-saturated turnover rate of fully carbamylated enzyme - PPFD photosynthetically active photon flux density (400–700 nm) - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP ribulose-1,5-bisphosphate     

Relative growth rate in relation to physiological and morphological traits for northern hardwood tree seedlings: species,light environment and ontogenetic considerations

The influence of ontogeny, light environment and species on relationships of relative growth rate (RGR) to physiological and morphological traits were examined for first-year northern hardwood tree seedlings. Three Betulaceae species (Betula papyrifera, Betula alleghaniensis and Ostrya virginiana) were grown in high and low light and Quercus rubra and Acer saccharum were grown only in high light. Plant traits were determined at four ages: 41, 62, 83 and 104 days after germination. In high light (610 mol m^–2 s^–1 PPFD), across species and ages, RGR was positively related to the proportion of the plant in leaves (leaf weight ratio, LWR; leaf area ratio, LAR), in situ rates of average canopy net photosynthesis (A) per unit mass (A_mass) and per unit area (A_area), and rates of leaf, stem and root respiration. In low light (127 mol m^–2 s^–1 PPFD), RGR was not correlated with A_mass and A_area whereas RGR was positively correlated with LAR, LWR, and rates of root and stem respiration. RGR was negatively correlated with leaf mass per area in both high and low light. Across light levels, relationships of CO₂ exchange and morphological characteristics with RGR were generally weaker than within light environments. Moreover, relationships were weaker for plant parameters containing a leaf area component (leaf mass per area, LAR and A_area), than those that were solely mass-based (respiration rates, LWR and A_mass). Across light environments, parameters incorporating the proportion of the plant in leaves and rates of photosynthesis explained a greater amount of variation in RGR (e.g. LWR^*A_mass, R²=0.64) than did any single parameter related to whole-plant carbon gain. RGR generally declined with age and mass, which were used as scalars of ontogeny. LWR (and LAR) also declined for seven of the eight species-light treatments and A declined in four of the five species in high light. Decreasing LWR and A with ontogeny may have been partially responsible for decreasing RGR. Declines in RGR were not due to increased respiration resulting from an increase in the proportion of solely respiring tissue (roots and stems). In general, although LWR declined with ontogeny, specific rates of leaf, stem, and root respiration also decreased. The net result was that whole-plant respiration rates per unit leaf mass decreased for all eight treatments. Identifying the major determinants of variation in growth (e.g. LWR^*A_mass) across light environments, species and ontogeny contributes to the establishment of a framework for exploring limits to productivity and the nature of ecological success as measured by growth. The generality of these relationships both across the sources of variation we explored here and across other sources of variation in RGR needs further study.     

Can light-saturated photosynthesis in lowland tropical forests be estimated by one light level?

Leaf-level net photosynthesis (A_n) estimates and associated photosynthetic parameters are crucial for accurately parameterizing photosynthesis models. For tropical forests, such data are poorly available and collected at variable light conditions. To avoid over- or underestimation of modeled photosynthesis, it is critical to know at which photosynthetic photon flux density (PPFD) photosynthesis becomes light-saturated. We studied the dependence of A_n on PPFD in two tropical forests in French Guiana. We estimated the light saturation range, including the lowest PPFD level at which A_sat (A_n at light saturation) is reached, as well as the PPFD range at which A_sat remained unaltered. The light saturation range was derived from photosynthetic light-response curves, and within-canopy and interspecific differences were studied. We observed wide light saturation ranges of A_n. Light saturation ranges differed among canopy heights, but a PPFD level of 1,000 µmol m⁻² s⁻¹ was common across all heights, except for pioneer trees species that did not reach light saturation below 2,000 µmol m⁻² s⁻¹. A light intensity of 1,000 µmol m⁻² s⁻¹ sufficed for measuring A_sat of climax species at our study sites, independent of the species or the canopy height. Because of the wide light saturation ranges, results from studies measuring A_sat at higher PPFD levels (for upper canopy leaves up to 1,600 µmol m⁻² s⁻¹) are comparable with studies measuring at 1,000 µmol m⁻² s⁻¹.     

Effects of elevated CO2 and increased nitrogen deposition on photosynthesis and growth of understory plants in spruce model ecosystems

We studied the effects of atmospheric CO₂ enrichment (280, 420 and 560 l CO₂ l^–1) and increased N deposition (0,30 and 90 kg ha^–1 year^–1) on the spruce-forest understory species Oxalis acetosella, Homogyne alpina and Rubus hirtus. Clones of these species formed the ground cover in nine 0.7 m² model ecosystems with 5-year-old Picea abies trees (leaf area index of approx 2.2). Communities grew on natural forest soil in a simulated montane climate. Independently of N deposition, the rate of light-saturated net photosynthesis of leaves grown and measured at 420 l CO₂ l^–1 was higher in Oxalis and in Homogyne, but was not significantly different in Rubus compared to leaves grown and measured at the pre-industrial CO₂ concentration of 280 l l^–1. Remarkably, further CO₂ enrichment to 560 l l^–1 caused no additional increase of CO₂ uptake. With increasing CO₂ supply concentrations of non-structural carbohydrates in leaves increased and N concentrations decreased in all species, whereas N deposition had no significant effect on these traits. Above-ground biomass and leaf area production were not significantly affected by elevated CO₂ in the more vigorously growing species O. acetosella and R. hirtus, but the slow growing H. alpina produced almost twice as much biomass and 50% more leaf area per plant under 420 l CO₂ l^–1 compared to 280 l l^–1 (again no further stimulation at 560 l l^–1). In contrast, increased N addition stimulated growth in Oxalis and Rubus but had no effect on Homogyne. In Oxalis (only) biomass per plant was positively correlated with microhabitat quantum flux density at low CO₂, but not at high CO₂ indicating carbon saturation. On the other hand, the less shade-tolerant Homogyne profited from CO₂ enrichment at all understory light levels facilitating its spread into more shady micro-habitats under elevated CO₂. These species-specific responses to CO₂ and N deposition will affect community structure. The non-linear responses to elevated CO₂ of several of the traits studied here suggest that the largest responses to rising atmospheric CO₂ are under way now or have already occurred and possible future responses to further increases in CO₂ concentration are likely to be much smaller in these understory species.     

Responses of C4 grasses to atmospheric CO2 enrichment

Summary The growth and photosynethetic responses to atmospheric CO₂ enrichment of 4 species of C₄ grasses grown at two levels of irradiance were studied. We sought to determine whether CO₂ enrichment would yield proportionally greater growth enhancement in the C₄ grasses when they were grown at low irradiance than when grown at high irradiance. The species studied were Echinochloa crusgalli, Digitaria sanguinalis, Eleusine indica, and Setaria faberi. Plants were grown in controlled environment chambers at 350, 675 and 1,000 l 1^-1 CO₂ and 1,000 or 150 mol m^-2 s^-1 photosynthetic photon flux density (PPFD). An increase in CO₂ concentration and PPFD significantly affected net photosynthesis and total biomass production of all plants. Plants grown at low PPFD had significantly lower rates of photosynthesis, produced less biomass, and had reduced responses to increases in CO₂. Plants grown in CO₂-enriched atmosphere had lower photosynthetic capacity relative to the low CO₂ grown plants when exposed to lower CO₂ concentration at the time of measurement, but had greater rate of photosynthesis when exposed to increasing PPFD. The light level under which the plants were growing did not influence the CO₂ compensation point for photosynthesis.     

Steady-state and dynamic photosynthetic response of Adenocaulon bicolor (Asteraceae) in its redwood forest habitat

Summary The gas exchange characteristics under steadystate and transient light conditions were determined for a redwood forest understory herb Adenocaulon bicolor, that depends on use of sunflecks for a large fraction of its daily carbon gain. Measurements under steady-state conditions indicated that this species has photosynthetic characteristics that are typical for understory plants. The mean light-saturated assimilation rate was 5.26 mol CO₂ m^-2 s^-1; the light saturation and compensation occurred at 243 and 2 mol photons m^-2 s^-1, respectively. This light compensation point was much less than the photon flux density under diffuse light in the understory so that positive assimilation could be maintained throughout the day. When leaves that had been in diffuse light for at least 2 h were exposed to a sudden increase in PFD to saturating levels, 10–30 min were required for both assimilation and stomatal conductance to reach maximum values. Calculation of intercellular CO₂ pressures, however, suggest that for the first 10 min after the light increase, biochemical factors were responsible for most of the increase in assimilation. Thereafter stomatal opening caused a further increase in assimilation that was no more than 25% of the total. When fully induced leaves were returned to low light, induction was rapidly lost even though stomatal conductance decreased only slowly. This rapid loss of induction limited the capacity of A. bicolor to use sunflecks after low light periods that lasted longer than 1–2 min. However, during periods when sunflecks are more frequent there is probably little loss of induction. Under these conditions, sunflecks are used with high efficiency for assimilation.     

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2–3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 mol photons m^–2 s^–1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r ²=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F _v/F _m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO₂ flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.     

Ammonium and nitrate uptake in gap,generalist and understory species of the genus Piper

Summary We studied root net uptake of ammonium (NH ₄ ⁺ ) and nitrate (NO ₃ ^– ) in species of the genus Piper (Piperaceae) under high, intermediate and low photosynthetically active photon flux densities (PFD). Plants were grown hydroponically, and then transferred to temperature controlled (25° C) root cuvettes for nutrient uptake determinations. Uptake solutions provided NH ₄ ⁺ and NO ₃ ^– simultaneously (both) or separately (single). In the first experiment, seven species of Piper, from a broad range of rainforest light habitats ranging from gap to understory, were screened for mineral nitrogen preference (100 M NH ₄ ⁺ and/or 100 M NO ₃ ^– ) at intermediate PFD (100 mol m^–2 s^–1). Preference for NH ₄ ⁺ relative to NO ₃ ^– , defined as the ratio of NH ₄ ⁺ (both):NO ₃ ^– (both) net uptake, was higher in understory species than in gap species. Ammonium repression of NO ₃ ^– uptake, defined as the ratio of NO ₃ ^– (single): NO ₃ ^– (both) net uptake, was also higher in understory species as compared to gap species. In a second set of experiments, we examined the effect of nitrogen concentration (equimolar, 10 to 1000 M) on NH ₄ ⁺ preference and NH ₄ ⁺ repression of NO ₃ ^– net uptake at high (500 mol m^–2 s^–1) and low (50 mol m^–2 s^–1) PFD in a gap (P. auritum), generalist (P. hispidum) and understory species (P. aequale). All species exhibited negligible NH ₄ ⁺ repression of NO ₃ ^– net uptake at high PFD. At low PFD, NH ₄ ⁺ preference and repression of NO ₃ ^– net uptake occurred in all species (understory > generalist > gap), but only at intermediate nitrogen concentrations, i.e. between 10 and 200 M. Ammonium repression of net NO ₃ ^– uptake decreased or increased rapidly (in < 48 h) after transitions from low to high or from high to low PFD respectively. No significant diurnal patterns in NO ₃ ^– or NH ₄ ⁺ net uptake were observed.CIWDPB publication # 1130     

Effects of water vapour pressure deficit and stomatal conductance on photosynthesis,internal pressurization and convective flow in three emergent wetland plants

Internal pressurization and convective gas flow in emergent wetland plants is a function of the water vapour pressure deficit (WPD) and stomatal conductance (G _s) separating the external atmosphere from the internal aerenchyma. We have compared the effects of WPD and G _s under a range of light intensities on static pressures and convective flows in Phragmites australis, Typha orientalis and Baumea articulata. The capacity of the three species to generate flows per unit leaf area differed, being greatest in P. australisand lowest in B. articulata. In all three species, decreasing light intensity from full sunlight (2200 mol m^–2 s^–1 photosynthetically active photon flux density (PPFD)) to < 200 and < 10 mol m^–2 s^–1PPFD caused immediate decreases in photosynthetic assimilation, followed by more gradual decreases in transpiration and G _s. However, internal pressures and flows in the two low light intensities remained similar to values recorded in full sunlight. WPD was more significantly related to pressures and flows in P. australis and T. orientalis than G _s. In B. articulata, pressures increased at low G _s values but flow rates were unaffected, as predicted by earlier models describing pore size effects on pressures and flows. The data suggest that emergent macrophytes can maintain significant internal convection even at low light intensities, and this may be beneficial for nocturnal aeration, particularly in arid climates where the atmospheric humidity at night is low.     

Photosynthetic acclimation to light in juveniles of two cloud forest tree species

The photosynthetic response of juveniles of Decussocarpus rospigliosii, an emergent primary forest species and shade tolerant in its juvenile stages and Alchornea triplinervia, a gap-colonizing species of tropical cloud forest in Venezuela was studied. Daily courses of microenvironmental variables and gas exchange under contrasting light conditions (gap and understory) were carried out in their natural environment and transplanted to different light regimes (shade and sun) in the field. The photosynthetic response and some anatomical characteristics of plants from different treatments were analyzed in the laboratory. Photosynthetic rates were low for both species, and were negative during some diurnal periods, related to the low photosynthetically active radiation levels obtained at both gap (6% of total radiation) and understory (2%). A. triplinervia shows higher rates (1.5–3.0 molm^{-2 -1}) than D. rospigliosii (0.7–1.1 molm^-2s^-1). Both species showed increased photosynthetic rates when grown in gaps. A. triplinervia did not adjust its maximum photosynthetic rates to the prevailing light conditions. In contrast, D. rospigliosii responded to increased light levels. Both species showed low light compensation points when grown under total shade. There was a partial stomatal closure generally during midday in D. rospigliosii. A. triplinervia presented lower leaf conductances, transpiration rates and lesser stomatal control. Some leaf anatomical characteristics, in both species, were affected by variations in the light regime (i.e. increased leaf thickness, leaf specific weight and stomatal density). These results suggest that both species have the ability to respond to variations in their natural light environments, therefore maintaining a favorable carbon balance during the day.     

Contrasting traits,contrasting environments,and considerations on population dynamics under a changing climate: an ecophysiological field study of two co‐dominant tree species

The increase in light availability resulting from canopy changes or opening is not always beneficial and can inhibit photosynthesis of tree seedlings already under other environmental stress. Tree seedlings' responses to compounded abiotic stress depend on their life‐history traits, and understanding the variations of such responses is important for understanding population dynamics under a changing climate. In this study we investigate how the photosynthesis of juveniles of two canopy tree species with different life‐history traits, Abies sachalinensis and Betula ermanii, differs in two contrasting sites at a sub‐boreal forest in northern Japan—one under a deciduous canopy (Closed site) and the other at a wide canopy opening (Open site). Seedlings at the Open site had low F_v/F_m (quantum yield of photosystem II) for a longer period than those at the Closed site. Abies sachalinensis at the Closed site showed lower F_v/F_m in spring than those at the Open site, but recovered after the canopy's new leaves flushed, indicating its acclimation to the shaded condition. Mean P_max (light‐saturated photosynthetic rate at ambient CO₂ levels) of A. sachalinensis seedlings was affected by site and air temperature, while B. ermanii seedlings were also affected by precipitation. Only B. ermanii's seedlings presented growth in the period studied, in spite of observed mid‐day drops to F_v/F_m attributed to water‐deficit‐related photoprotection. Results suggest that the climate change predicted for the Hokkaido area may increase the competitive advantage of broad‐leaved deciduous species, such as B. ermanii, in relation to evergreen conifers like A. sachalinensis.