Seasonal changes in photosynthetic characteristics of Pachysandra terminalis (Buxaceae), an evergreen woodland chamaephyte,in the cool temperate regions of Japan

Summary Seasonal changes in photosynthetic capacity, and photosynthetic responses to intercellular CO₂ concentration and irradiance were investigated under laboratory conditions on intact leaves of Pachysandra terminalis. Photosynthetic capacity and stomatal conductance under saturating light intensity and constant water vapor pressure deficit showed almost the same seasonal trend. They increased from early June just after the expansion of leaves, reached the maximum in late-Septemer, and then decreased to winter. In over-wintering leaves they recovered and increased immediately after snow-melting, reached a first maximum in late April, and then decreased to early July in response to the reduction of light intensity on the forest floor. There-after, they increased from mid August, reached a second maximum in late September, and then decreased to winter. The parallel changes of photosynthesis and stomatal conductane indicate a more or less constant intercellular CO₂ concentration throughout the year. The calculated values of relative stomatal limitation of photosynthesis were nearly constant throughout the year, irrespective of leaf age. The results indicate that the seasonal changes in light-saturated photosynthetic capacity are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Indeed, carboxylation efficiency assessed by the inital slope of the Ci-photosynthesis curve changed in proportion to seasonal changes of the photosynthetic capacity in both current-year and over-wintered leaves. High photosynthetic capacity in current-year leaves as compared with one-year-old leaves was also due to the high photosynthetic capacity of mesophyll. Nevertheless, stomatal conductance changed in proportion to photosynthetic capacity, indicating that stomatal conductance is regulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations are maintained constant. The quantum yield also changed seasonally parallel with that in the photosynthetic capacity.Contribution No. 2893 from the Institute of Low Temperature Science     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

Differences in the Activation State of Ribulose-1,5-bisphosphate Carboxylase/Oxygenase in Barley,Pea, and Wheat at Two Altitudes

Activation state of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) is an important parameter determining the rate of net photosynthesis (P _N) in situ for which no information is available with reference to altitude. We analyzed activation state along with P _N in three plant species and their cultivars grown at low (LA, 1 300 m) and high (HA, 4 200 m) altitudes. No significant change in P _N and the initial activity of RuBPCO was obtained with reference to altitude. However, activation state of RuBPCO was reduced significantly in the HA plants as compared to the LA ones. Hence low partial pressure of CO₂ prevailing at HA might be responsible for the lower activation state of RuBPCO.     

CO2 and water vapour exchange in four alpine herbs at two altitudes and under varying light and temperature conditions

CO₂ and water vapour exchange rates of four alpine herbs namely: Rheum emodi, R. moorcroftianum, Megacarpaea polyandra and Rumex nepalensis were studied under field conditions at 3600 m (natural habitat) and 550 m altitudes. The effect of light and temperature on CO₂ and water vapour exchange was studied in the plants grown at lower altitude. In R. moorcroftianum and R. nepalensis, the average photosynthesis rates were found to be about three times higher at 550 m as compared to that under their natural habitat. However, in M. polyandra, the CO₂ exchange rates were two times higher at 3600 m than at 550 m but in R. emodi, there were virtually no differences at the two altitudes. These results indicate the variations in the CO₂ exchange rates are species specific. The change in growth altitude does not affect this process uniformly.The transpiration rates in R. emodi and M. polyandra were found to be very high at 3600 m compared to 550 m and are attributed to overall higher stomatal conductance in plants of these species, grown at higher altitude. The mid-day closure of stomata and therefore, restriction of transpirational losses of water were observed in all the species at 550 m altitude. In addition to the effect of temperature and relative humidity, the data also indicate some endogenous rhythmic control of stomatal conductance.The temperature optima for photosynthesis was close to 30°C in M. polyandra and around 20°C in the rest of the three species. High temperature and high light intensity, as well as low temperature and high light intensity, adversely affect the net rate of photosynthesis in these species.Both light compensation point and dark respiration rate increased with increasing temperature.The effect of light was more prominent on photosynthesis than the effect of temperature, however, on transpiration the effect of temperature was more prominent than the effect of light intensity.No definite trends were found in stomatal conductance with respect to light and temperature. Generally, the stomatal conductance was highest at 20°C.The study reveals that all these species can easily be cultivated at relatively lower altitudes. However, proper agronomical methodology will need to be developed for better yields.     

Carbon isotope discrimination and stomatal responses of mature Pinus sylvestris L. trees exposed in situ for three years to elevated CO2 and temperature

The Climate Change Experiment (CLIMEX) is a unique large scale facility in which an entire undisturbed catchment of boreal vegetation has been exposed to elevated CO₂ (560 ppm) and temperature (+3°C summer, +5°C winter) for the past three years with all the soil-plant-atmosphere linkages intact. Here, carbon isotope composition and stomatal density have been analysed from sequential year classes of needles of mature Scots pine trees (Pinus sylvestris L.) to investigate the response of time-integrated water-use efficiency (UWE) and stomatal density to CO₂ enrichment and climate change. Carbon isotope discrimination decreased and WUE increased in cohorts of needles developing under increased CO₂ and temperature, compared to needles on the same trees developing in pretreatment years. Mid-season instantaneous gas exchange, measured on the same trees for the past four years, indicated that these responses resulted from higher needle photosynthetic rates and reduced stomatal conductance. Needles of P. sylvestris developing under increased CO₂ and temperature had consistently lower stomatal densities than their ambient grown counterparts on the same trees. The stomatal density of P. sylvestris needles was inversely correlated with δ¹³C-derived WUE, implying some effect of this morphological response on leaf gas exchange. Future atmospheric CO₂ and temperature increases are therefore likely to improve the water economy of P. sylvestris, at least at the scale of individual needles, by affecting stomatal density and gas exchange processes.     

Different photosynthetic responses of wild and cultivated plants to high irradiance

In addition to other factors, high altitude (HA) environment is characterized by high photosynthetic photon flux density (PPFD). Photosynthetic characteristics of wild and cultivated plants were studied at different irradiances at Losar, India (altitude 4 200 m). Wild plants were tolerant to high PPFDs. Slopes of curve between net photosynthetic rate (P _N) and intercellular CO₂ concentration (C _i) or stomatal conductance (g _s) increased with increase in irradiance suggesting insensitivity or tolerance of these plants to higher PPFD. Cultivated plants, however, were sensitive to higher PPFD, their slopes of curves between P _N and C _i or g _s decreased with increased PPFD. Tolerance or insensitivity to higher PPFD was an important parameter affecting plant performance at HA.     

The Photosynthetic Response to Elevated CO2 in High Altitude Potato Species (Solanum curtilobum)

Plants of Solanum curtilobum (from high altitude) and Solanum tuberosum (from low altitude) were grown in open-top chambers in a greenhouse at either ambient (AC, 360 µmol mol^–1) or ca. twice ambient (EC, 720 µmol mol^–1) CO₂ concentrations for 30 d. CO₂ treatments started at the reproductive stage of the plants. There were similar patterns in the physiological response to CO₂ enrichment in the two species. Stomatal conductance was reduced by 59 % in S. tuberosum and by 55 % in S. curtilobum, but such a reduction did not limit the net photosynthetic rate (P _N), which was increased by approximately 56 % in S. curtilobum and 53 % in S. tuberosum. The transpiration rate was reduced by 16 % in both potato species while instantaneous transpiration efficiency increased by 80 % in S. tuberosum and 90 % in S. curtilobum. Plants grown under EC showed 36 and 66 % increment in total dry biomass, whereas yields (dry mass of tubers) were increased by 40 and 85 % in S. tuberosum and S. curtilobum, respectively. EC promoted productivity by increasing P _N. Thus S. tuberosum, cultivated around the world at low altitudes, and S. curtilobum, endemic of the highland Andes, respond positively to EC during the tuberisation stage.     

Gas exchange response of barley and pea cultivars to altitude variation in Himalaya

Leaf stomatal density (SD), net photosynthetic rates (P _N), and stomatal conductance (g _s) of Hordeum vulgare and Pisum sativum cultivars in Himalaya increased with altitude. Higher P _N and leaf temperature under low CO₂ partial pressure at high altitudes could evoke a higher g _s and SD to allow sufficient influx of CO₂ as well as more efficient leaf cooling through transpiration.     

长白山不同海拔白桦幼苗移栽至同一生境的光合及反射光谱特性

环境是影响植物生理性状的主要因素之一,通过改变植物的生长环境来研究植物生理性状的改变是目前研究植物生理生态的热点,而高海拔地区多样的环境为研究物种适应环境变化提供了良好的实验条件。本研究通过移栽的技术手段,将分布在海拔750 m、海拔1200 m和海拔1400 m的3~5年生白桦幼苗移植到海拔750 m相同的林下环境条件下,将3种白桦移栽幼苗和野生白桦幼苗的光响应参数、光谱反射率以及光谱反射指数进行对比研究,分析移栽初期和移栽一年后植物生理性状的异同,探索白桦幼苗迁入新环境的适应性。研究结果表明:移栽初期,通过对比不同海拔来源的白桦移栽幼苗彼此之间各项生理性状以及与原海拔野生白桦幼苗的生理性状发现,包括最大净光合速率(A_(max))、气孔导度(G_s)、胞间CO_2浓度(C_i)在内的生理性状差异显著(P0.05),幼苗光谱反射指数差异性显著(P0.05)。经过一年的适应,不同海拔来源的白桦幼苗与移栽地海拔750 m野生白桦幼苗的生理性状之间差异不显著(P0.05),幼苗光谱反射指数差异性不显著(P0.05),而与原海拔野生幼苗生理性状之间差异显著(P0.05),幼苗光谱反射指数差异显著(P0.05)。在移栽初期,移栽幼苗的各项生理指标已经因环境的改变而发生变化,但仍保留原海拔野生幼苗的生理特性;移栽一年后,移栽幼苗逐渐适应环境,其相关生理性状发生改变,并趋同于移栽地野生白桦幼苗的生理性状。对比一个生长季前后白桦移栽幼苗生理性状的变化可以发现,白桦作为先锋种植物,在新迁入环境后具有良好的环境适应性,能通过调整生理性状适应新环境。研究对幼苗在林下的演替以及森林的更新具有借鉴意义。     

High SO2 resistance ofClethra barbinervis established in a smoke-polluted area of Ashio,Tochigi Prefecture,Japan

The effect of SO₂ on the photosynthesis ofClethra barbinervis collected from a smoke-polluted area near the Ashio copper smelter in Tochigi Prefecture was compared withC. barbinervis collected from a nonpolluted district in Tsukuba, Ibaraki Prefecture andQuercus mongolica var.grosseserrata grown in a nonpolluted field in Nagano Prefecture. The plants were exposed to 0.5–1.5 p.p.m. SO₂ for 90 min (short-term) and to 0.3 p.p.m. SO₂ for 31–39 days (long-term). TheClethra plants from both sites had a lower intrinsic stomatal conductance and photosynthetic rate thanQuercus plants. Short-and long-term fumigation caused stomatal closure inQuercus plants, but had little effect on the stomatal conductance ofClethra plants. Under short-term fumigation, nonstomatal photosynthetic inhibition per unit of absorbed SO₂ was smallest inClethra plants from Ashio. Long-term fumigation caused photosynthetic decline and visible foliar injury toQuercus plants, but had no effect onClethra plants from Ashio. Consequently,Clethra plants from Ashio had a higher photosynthetic rate thanQuercus plants after long-term fumigation. These results suggest thatC. barbinervis populations in the smoke-polluted area of Ashio had evolved high SO₂ resistance connected with SO₂ detoxification ability in mesophyll cells.     

Why does photosynthesis decrease with needle age in Pinus pinaster?

Rates of photosynthesis vary with foliage age and typically decline from full-leaf expansion until senescence occurs. This age-related decline in photosynthesis is especially important in species that retain foliage for several years, yet it is not known whether the internal conductance to CO₂ movement (g _i) plays any role. More generally, g _i has been measured in only a few conifers and has never been measured in leaves or needles older than 1 year. The effect of ageing on g _i was investigated in Pinus pinaster, a species that retains needle for 4 or more years. Measurements were made in autumn when trees were not water limited and after leaf expansion was complete. Rates of net photosynthesis decreased with needle age, from 8 μmol m⁻² s⁻¹ in fully expanded current-year needles to 4.4 μmol m⁻² s⁻¹ in 3-year-old needles. The relative limitation due to internal conductance (0.24–0.35 out of 1) was in all cases larger than that due to stomatal conductance (0.13–0.19 out of 1). Internal conductance and stomatal conductance approximately scaled with rates of photosynthesis. Hence, there was no difference among year-classes in the relative limitations posed by internal and stomatal conductance or evidence that they cause the age-related decline in photosynthesis. There was little evidence that the age-related decline in photosynthesis was due to decreases in contents of N or Rubisco. The decrease in rates of photosynthesis from current-year to older needles was instead related to a twofold decrease in rates of photosynthesis per unit nitrogen and V _cmax/Rubisco (i.e., in vivo specific activity).     

Effects of low and elevated CO2 on C3 and C4 annuals

Abutilon theophrasti (C₃) and Amaranthus retroflexus (C₄), were grown from seed at four partial pressures of CO₂: 15 Pa (below Pleistocene minimum), 27 Pa (pre-industrial), 35 Pa (current), and 70 Pa (future) in the Duke Phytotron under high light, high nutrient, and wellwatered conditions to evaluate their photosynthetic response to historic and future levels of CO₂. Net photosynthesis at growth CO₂ partial pressures increased with increasing CO₂ for C₃ plants, but not C₄ plants. Net photosynthesis of Abutilon at 15 Pa CO₂ was 70% less than that of plants grown at 35 Pa CO₂, due to greater stomatal and biochemical limitations at 15 Pa CO₂. Relative stomatal limitation (RSL) of Abutilon at 15 Pa CO₂ was nearly 3 times greater than at 35 Pa CO₂. A photosynthesis model was used to estimate ribulose-1,5-bisphosphate carboxylase (rubisco) activity (Vc_max), electron transport mediated RuBP regeneration capacity (J _max), and phosphate regeneration capacity (PiRC) in Abutilon from net photosynthesis versus intercellular CO₂ (A–C _i) curves. All three component processes decreased by approximately 25% in Abutilon grown at 15 Pa compared with 35 Pa CO₂. Abutilon grown at 15 Pa CO₂ had significant reductions in total rubisco activity (25%), rubisco content (30%), activation state (29%), chlorophyll content (39%), N content (32%), and starch content (68%) compared with plants grown at 35 Pa CO₂. Greater allocation to rubisco relative to light reaction components and concomitant decreases in J _max and PiRC suggest co-regulation of biochemical processes occurred in Abutilon grown at 15 Pa CO₂. There were no significant differences in photosynthesis or leaf properties in Abutilon grown at 27 Pa CO₂ compared with 35 Pa CO₂, suggesting that the rise in CO₂ since the beginning of the industrial age has had little effect on the photosynthetic performance of Abutilon. For Amaranthus, limitations of photosynthesis were balanced between stomatal and biochemical factors such that net photosynthesis was similar in all CO₂ treatments. Differences in photosynthetic response to growth over a wide range of CO₂ partial pressures suggest changes in the relative performance of C₃ and C₄ annuals as atmospheric CO₂ has fluctuated over geologic time.     

Hydrogen peroxide production is an early event during bicarbonate induced stomatal closure in abaxial epidermis of <Emphasis Type="Italic">Arabidopsis</Emphasis>

The presence of 2 mM bicarbonate in the incubation medium induced stomatal closure in abaxial epidermis of Arabidopsis. Exposure to 2 mM bicarbonate elevated the levels of H₂O₂ in guard cells within 5 min, as indicated by the fluorescent probe, dichlorofluorescein diacetate (H₂DCF-DA). Bicarbonate-induced stomatal closure as well as H₂O₂ production were restricted by exogenous catalase or diphenylene iodonium (DPI, an inhibitor of NAD(P)H oxidase). The reduced sensitivity of stomata to bicarbonate and H₂O₂ production in homozygous atrbohD/F double mutant of Arabidopsis confirmed that NADP(H) oxidase is involved during bicarbonate induced ROS production in guard cells. The production of H₂O₂ was quicker and greater with ABA than that with bicarbonate. Such pattern of H₂O₂ production may be one of the reasons for ABA being more effective than bicarbonate, in promoting stomatal closure. Our results demonstrate that H₂O₂ is an essential secondary messenger during bicarbonate induced stomatal closure in Arabidopsis.     

No heat-stimulated germination found in herbaceous species from burned subtropical grassland

The inverse relationship between numbers of stomata (stomatal frequency) on tree leaves and ambient CO₂ concentration is increasingly applied for reconstructing past atmospheric CO₂ levels. The abundance of leaf remains of Quercus robur in Holocene peat and lake deposits in Europe makes this species potentially suitable for high-resolution stomatal frequency analysis. In order to quantify the CO₂ responsiveness of the species, the behavior of the stomatal index for Q. robur during the current anthropogenic CO₂ increase is determined on the basis of buried, herbarium and modern leaf material from the Netherlands. The stomatal index (SI), expressing the ratio of the number of stomata in a given area divided by the total number of stomata and other epidermal cells in that same area, is used in order to minimize influences on stomatal frequency of environmental conditions other than CO₂. The sigmoid SI response pattern recorded for Q. robur resembles that of the closely related species Q. petraea, although there is a difference in the timing of the response limitation of the two species to increasing atmospheric CO₂. For calibration purposes only the linear phase of the sigmoidal response curve is taken into consideration in the presented CO₂ response model, which allows confident combination of Q. robur and Q. petraea over the interval from 290 to 325 ppmv CO₂. The model is conservative in reconstructing past CO₂ mixing ratios outside the range of monitored response. As a result of the observed SI response limit, the model predicts CO₂ levels below 325 ppmv with a mean error of 10.2 ppmv, whereas higher CO₂ levels are underestimated.     

九寨沟两种常见藓类植物对模拟氮沉降的生理响应

为探讨氮沉降对九寨沟藓类植物的影响，该研究以当地优势藓类植物锦丝藓(Actinothuidium hookeri)和塔藓(Hylocomium splendens)为对象，以NH₄NO₃为氮源，设置对照(0 kg N·hm^-2·a^-1)、低浓度(20 kg N·hm^-2·a^-1)、高浓度(50 kg N·hm^-2·a^-1)3种处理，开展为期6个月的氮沉降模拟实验。结果表明：(1)氮沉降处理导致两种藓类植物的活性氧、丙二醛、叶绿素、脯氨酸和可溶性蛋白含量显著增加，同时锦丝藓过氧化氢酶、过氧化物酶、超氧化物歧化酶、抗坏血酸过氧化物酶活性增加。(2)对于生长旺期和生长末期的塔藓，氮沉降导致其过氧化物酶、过氧化氢酶、抗坏血酸过氧化物酶活性降低。(3)锦丝藓的综合隶属函数值随氮沉降浓度增大而增加，在生长旺期和生长末期，塔藓综合隶属函数值对氮沉降的响应存在差异。综上认为，两种藓类植物对氮沉降处理的生理响应存在差异，高浓度氮沉...     

Inter- and intra-generic differences in growth,reproduction, and fitness of nine herbaceous annual species grown in elevated CO2 environments

In assessing the capacity of plants to adapt to rapidly changing global climate, we must elucidate the impacts of elevated carbon dioxide on reproduction, fitness and evolution. We investigated how elevated CO₂ influenced reproduction and growth of plants exhibiting a range of floral morphologies, the implications of shifts in allocation for fitness in these species, and whether related taxa would show similar patterns of response. Three herbaceous, annual species each of the genera Polygonum, Ipomoea, and Cassia were grown under 350 or 700 ppm CO₂. Vegetative growth and reproductive output were measured non-destructively throughout the full life span, and vegetative biomass was quantified for a subsample of plants in a harvest at first flowering. Viability and germination studies of seed progeny were conducted to characterize fitness precisely. Early vegetative growth was often enhanced in high-CO₂ grown plants of Polygonum and Cassia (but not Ipomoea). However, early vegetative growth was not a strong predictor of subsequent reproduction. Phenology and production of floral buds, flowers, unripe and abscised fruits differed between CO₂ treatments, and genera differed in their reproductive and fitness responses to elevated CO₂. Polygonum and Cassia species showed accelerated, enhanced reproduction, while Ipomoea species generally declined in reproductive output in elevated CO₂. Seed quality and fitness (in terms of viability and percentage germination) were not always directly correlated with quantity produced, indicating that output alone may not reliably indicate fitness or evolutionary potential. Species within genera typically responded more consistently to CO₂ than unrelated species. Cluster analyses were performed separately on suites of vegetative and reproductive characters. Some species assorted within genera when these reproductive responses were considered, but vegetative responses did not reflect taxonomic affinity in these plants. Congeners may respond similarly in terms of reproductive output under global change, but fitness and prognoses of population persistence and evolutionary performance can be inferred only rarely from examination of vegetative characters alone.     

Effects of CO2 elevation on canopy development in the stands of two co-occurring annuals

Elevated CO₂ may increase dry mass production of canopies directly through increasing net assimilation rate of leaves and also indirectly through increasing leaf area index (LAI). We studied the effects of CO₂ elevation on canopy productivity and development in monospecific and mixed (1:1) stands of two co-occurring C₃ annual species, Abutilon theophrasti, and Ambrosia artemisiifolia. The stands were established in the glasshouse with two CO₂ levels (360 and 700 l/l) under natural light conditions. The planting density was 100 per m² and LAI increased up to 2.6 in 53 days of growth. Root competition was excluded by growing each plant in an individual pot. However, interference was apparent in the amount of photons absorbed by the plants and in photon absorption per unit leaf area. Greater photon absorption by Abutilon in the mixed stand was due to different canopy structures: Abutilon distributed leaves in the upper layers in the canopy while Ambrosia distributed leaves more to the lower layers. CO₂ elevation did not affect the relative performance and light interception of the two species in mixed stands. Total aboveground dry mass was significantly increased with CO₂ elevation, while no significant effects on leaf area development were observed. CO₂ elevation increased dry mass production by 30–50%, which was mediated by 35–38% increase in the net assimilation rate (NAR) and 37–60% increase in the nitrogen use efficiency (NUE, net assimilation rate per unit leaf nitrogen). Since there was a strong overall correlation between LAI and aboveground nitrogen and no significant difference was found in the regression of LAI against aboveground nitrogen between the two CO₂ levels, we hypothesized that leaf area development was controlled by the amount of nitrogen taken up from the soil. This hypothesis suggests that the increased LAI with CO₂ elevation observed by several authors might be due to increased uptake of nitrogen with increased root growth.     

Improvement of growth conditions and gas exchange of<Emphasis Type="Italic"> Fagus sylvatica</Emphasis> L. seedlings planted below a recently thinned<Emphasis Type="Italic"> Pinus sylvestris</Emphasis> L. stand

Abstract Gas exchange and growth of beech seedlings planted in the understory of a recently thinned pinewood were recorded for 2 years. Relative irradiance was assessed by hemispherical photographs taken just after the thinning. Predawn water potential (_pd), daily gas exchange and chlorophyll fluorescence were measured several times during the two growing seasons. Maximum values of photosynthesis (A _max) and stomatal conductance to water vapour (g _wvmax) were established from daily data. Maximum quantum efficiency of PS II was recorded at dawn by taking the variable to maximum chlorophyll fluorescence ratio on dark adapted leaves (F _v/F _m). In the middle of each summer, leaf nitrogen content and leaf mass per area were evaluated, and height growth and basal area increment were recorded at the end of the season. The thinning treatment removed half the trees and generated around 10% more available relative irradiance (GLF). This was followed by an increase in net photosynthesis at saturating PPFD (A _sat) and in maximum stomatal conductance to water vapour (g _wvmax). Moreover, specific leaf mass (SLM) and mass based nitrogen content (N_m) showed higher values for seedlings in the thinned stand. In both years, a positive relationship was established between the area based nitrogen content (N_a) and maximum net photosynthesis (A _max). In 1998, a year with a dry summer, seedlings suffered a significant drop in daily A _max irrespective of the thinning regime. This was a response to an increase in stomatal limitation to net photosynthesis, g _wvmax reaching the lowest value on dates with the highest drought. A lack of decrease of Fv/Fm confirmed the absence of significant non-stomatal limitation to A as a consequence of photoinhibition after opening the pinewood. A higher maximum quantum efficiency of open PS II centres (Fv/Fm) was registered in seedlings in the thinned stand. The significance of the differences between the treatments was stronger in the second year after thinning. In 1999, a year with frequent summer storms, water availability increased for seedlings growing under the thinned pinewood. Overall, the reduced pine overstory had a positive effect on physiological responses of beech seedlings, which was translated into improved seedling growth.     

正常和环割条件下不同形态氮素添加对红椎幼苗光合特性的影响

采用盆栽试验和韧皮部环割方法研究了碳水化合物供应、氮素形态及其交互作用对红椎幼苗叶片光合特性的影响。无机氮源采用硝酸铵(AN)、铵态氮(NH_4~+-N)、硝态氮(NO_3~--N),有机氮源采用尿素(Urea)、精氨酸(Arg)和甘氨酸(Gly),氮素施用量均为10g N/m~2,处理时间为10 d。研究结果表明,环割、氮素形态及其交互作用均显著影响了红椎幼苗叶片的净光合速率(Pn);单一环割处理显著降低了红椎幼苗叶片Pn、气孔导度(Gs)、胞间CO_2浓度(Ci)、蒸腾速率(Tr)、相对叶绿素含量和CO2利用效率(CUE),但显著提高了叶片水分利用效率(WUE)和气孔限制值(Ls),其叶片Pn降低主要是由气孔限制和叶绿素含量降低所导致的。正常条件下,所有氮素形态处理均显著降低了红椎幼苗叶片的Pn、Gs、Ci和Tr,但显著升高了叶片Ls和WUE,其叶片Pn降低的主要原因是供氮过高引起的叶片气孔限制。正常条件下,除Arg处理外,其他氮素形态均显著升高了红椎幼苗叶片的CUE,其中以Gly处理的促进作用最大。环割条件下,3种有机态氮素的添加均显著缓解了单一环割处理对红椎幼苗叶片Pn的抑制作用,尤其以精氨酸最为明显,供应精氨酸的叶片Pn值从单一环割处理的强烈抑制中恢复到了对照水平,而无机态氮的供应均未显著改变这种抑制。结果表明短期碳水化合物供应的阻断会显著抑制红椎幼苗叶片的光合能力,适量供应精氨酸、甘氨酸和尿素等有机态氮会有效缓解这种抑制作用,其中以精氨酸的作用最为明显。     

新疆准噶尔盆地早侏罗世掌鳞杉科植物化石 及古大气CO2浓度

植物化石气孔参数分析是目前恢复古大气二氧化碳浓度较为精准的方法之一,银杏类和松柏类等是恢复古大气CO_2浓度常用的化石类群。本文利用新疆准噶尔盆地下侏罗统三工河组的松柏类掌鳞杉科Brachyphyllum(Hirmeriella?)sp.化石对早侏罗世大气CO_2浓度进行了重建,获得早侏罗世大气CO_2浓度为～1200ppm,丰富了早侏罗世大气CO_2浓度信息,进一步说明掌鳞杉科植物通过气孔比率法在重建侏罗纪大气CO_2浓度方面的可靠性。掌鳞杉科植物的旱生构造和较高的大气CO_2浓度表明早侏罗世Toarcian期大洋缺氧事件在陆地生态系统内可能产生了一定的响应。