Non-steady state effects in diurnal 180 discrimination by Picea sitchensis branches in the field

We report diurnal variations in 18O discrimination (18 delta) during photosynthesis (18 delta A) and respiration (18 delta R) of Picea sitchensis branches measured in branch chambers in the field. These observations were compared with predicted 18 delta (18 delta pred) based on concurrent measurements of branch gas exchange to evaluate steady state and non-steady state (NSS) models of foliage water 18O enrichment for predicting the impact of this ecosystem on the Delta 18O of atmospheric CO2. The non-steady state approach substantially improved the agreement between 18 delta pred and observed 18 delta (18 delta obs) compared with the assumption of isotopic steady state (ISS) for the Delta 18O signature of foliage water. In addition, we found direct observational evidence for NSS effects: extremely high apparent 18 delta values at dusk, dawn and during nocturnal respiration. Our experiments also show the importance of bidirectional foliage gas exchange at night (isotopic equilibration in addition to the net flux). Taken together, neglecting these effects leads to an underestimation of daily net canopy isofluxes from this forest by up to 30%. We expect NSS effects to be most pronounced in species with high specific leaf water content such as conifers and when stomata are open at night or when there is high relative humidity, and we suggest modifications to ecosystem and global models of delta 18O of CO2.     

Diurnal, seasonal and interannual variability of carbon isotope discrimination at the canopy level in response to environmental factors in a boreal forest ecosystem

Accurate estimation of temporal and spatial variations in photosynthetic discrimination of 13C is critical to carbon cycle research. In this study, a combined ecosystem-boundary layer isotope model, which was satisfactorily validated against intensive campaign data, was used to explore the temporal variability of carbon discrimination in response to environmental driving factors in a boreal ecosystem in the vicinity of Fraserdale Tower, Ontario, Canada (49 degrees 52'30'N, 81 degrees 34'12'W). A 14 year (1990-1996 and 1998-2004) hourly CO2 concentration and meteorological record measured on this tower was used for this purpose. The 14 year mean yearly diurnal amplitude of canopy-level discrimination Delta(canopy) was computed to be 2.8 +/- 0.5 per thousand, and the overall diurnal cycle showed that the greatest Delta(canopy) values occurred at dawn and dusk, while the minima generally appeared in mid-afternoon. The average annual Delta(canopy) varied from 18.3 to 19.7 per thousand with the 14 year average of 19 +/- 0.4 per thousand. The overall seasonality of Delta(canopy) showed a gradually increasing trend from leaf emergence in May-September and with a slight decrease at the end of the growing season in October. Delta(canopy) was negatively correlated to vapour pressure deficit and air temperature across hourly to decadal timescales. A strong climatic control on stomatal regulation of ecosystem isotope discrimination was found in this study.     

马占相思人工林净二氧化碳交换和碳同位素通量

应用稳定碳同位素技术,对马占相思人工林冠层受光和遮荫叶片的碳同化率(A_net)和叶面积指数(L)进行加权,将叶片水平的¹³C甄别率(Δ_i)扩展至冠层光合甄别率(Δ_canopy),测定光合固定和呼吸释放的碳同位素通量及其净交换通量.结果表明：Δ_canopy的日变化明显,日出前和中午出现较低值(18.47‰和19.87‰),而日落前达到最大(21.21‰);秋季末期(11月)至翌年夏季,Δ_canopy逐步升高,年平均为(20.37±0.29)‰.不同季节自养呼吸(日间叶片呼吸除外)和异养呼吸释放CO₂的碳同位素比率(δ¹³C)平均值分别为(-28.70±0.75)‰和(-26.75±1.3)‰,春季林冠夜间呼吸CO₂的δ¹³C最低(-30.14‰),秋季末期最高(-28.01‰).马占相思林与大气的CO₂碳同位素通量在春季和夏季中午时峰值分别为178.5和217 μmol·m^-2 ·s^-1·‰,日均值分别为638.4 和873.2 μmol·m^-2·s^-1·‰.冠层叶片吸收CO₂的碳同位素通量较呼吸释出CO₂的碳同位素通量高1.6～2.5倍,表明马占相思林日间吸收大量CO₂,降低空气CO₂浓度,具有改善环境的良好生态服务功能.     

Importance of short-term dynamics in carbon isotope ratios of ecosystem respiration (delta13C(R)) in a Mediterranean oak woodland and linkage to environmental factors

Temporal dynamics in carbon isotope ratios of ecosystem respiration (delta13C(R)) were evaluated on hourly, daily and annual timescales in a Mediterranean woodland. Emphasis was given to the periods of transition from wet to dry season and vice versa, when the system turns from a net carbon sink to a source. The constancy of nocturnal delta13C(R) was tested. The relationship between delta13C(R) (determined through Keeling plots) and environmental factors was evaluated through time-lag analysis. Delta13C(R) exhibited high annual variation (> 7). During the transition periods, delta13C(R) correlated significantly with factors influencing photosynthetic discrimination, soil respiration, and whole-canopy conductance. Time-lags differed between below- and above-ground variables, and between seasons. A shift in regression parameters with environmental factors indicated seasonal differences in ecosystem responsiveness (e.g. temperature acclimation). Delta13C(R) exhibited substantial nocturnal enrichment (> 4) from dusk to dawn. These data indicate pronounced short-term dynamics in delta13C(R) at hourly to daily timescales and a modulated response to environmental drivers. Substantial short-term changes in nocturnal delta13C(R) may have important implications for the sampling protocols of nocturnal Keeling plots.     

Natural entrainment without dawn and dusk: the case of the European ground squirrel (Spermophilus citellus).

Observational data collected in the field and in enclosures show that diurnal, burrow-dwelling European ground squirrels (Spermophilus citellus) never were above ground during twilight at dawn or at dusk. The animals emerged on average 4.02 h (SD = 0.45) after civil twilight at dawn and retreated in their burrows on average 2.87 h (SD = 0.47) before civil twilight at dusk. Daily patterns of light perceived by these burrowing mammals were measured with light-sensitive radio collar transmitters in an enclosure (the Netherlands) and in the field (Hungary). The observational data are corroborated by the telemetry data, which show clear daily patterns of timing of light perception including light perceived from the burrow entrances. The first light was observed by the animals on average 3.54 h (enclosure, SD = 0.45) and 3.60 h (field, SD = 0.31) after civil twilight at dawn, whereas the final observed light was on average 3.04 h (enclosure, SD = 0.64) and 2.02 h (field, SD = 0.72) before civil twilight at dusk. Thus, the animals do not perceive the rapid natural light-dark (LD) transitions that occur at civil twilight. Instead, they generate their own pattern of exposure to light within the natural LD cycle. The classical phase response model for entrainment by light or dark pulses cannot explain how the circadian system of this species remains entrained to the external, natural LD cycle while the major LD transitions are created by its own behavior.     

C4 photosynthetic isotope exchange in NAD-ME- and NADP-ME-type grasses

Monitoring photosynthetic isotope exchange is an important tool for predicting the influence of plant communities on the global carbon cycle in response to climate change. C(4) grasses play an important role in the global carbon cycle, but their contribution to the isotopic composition of atmospheric CO(2) is not well understood. Instantaneous measurements of (13)CO(2) (Delta(13)C) and C(18)OO (Delta(18)O) isotope exchange in five NAD-ME and seven NADP-ME C(4) grasses have been conducted to investigate the difference in photosynthetic CO(2) isotopic fractionation in these subgroups. As previously reported, the isotope composition of the leaf material (delta(13)C) was depleted in (13)C in the NAD-ME compared with the NADP-ME grasses. However, Delta(13)C was not different between subtypes at high light, and, although Delta(13)C increased at low light, it did so similarly in both subtypes. This suggests that differences in leaf delta(13)C between the C(4) subtypes are not caused by photosynthetic isotope fractionation and leaf delta(13)C is not a good indicator of bundle sheath leakiness. Additionally, low carbonic anhydrase (CA) in C(4) grasses may influences Delta(13)C and should be considered when estimating the contribution of C(4) grasses to the global isotopic signature of atmospheric CO(2). It was found that measured Delta(18)O values were lower than those predicted from leaf CA activities and Delta(18)O was similar in all species measured. The Delta(18)O in these C(4) grasses is similar to low Delta(18)O previously measured in C(4) dicots which contain 2.5 times the leaf CA activity, suggesting that leaf CA activity is not a predictor of Delta(18)O in C(4) plants.     

Modelling environmental controls on ecosystem photosynthesis and the carbon isotope composition of ecosystem-respired CO2 in a coastal Douglas-fir forest

We developed and applied an ecosystem-scale model that calculated leaf CO₂ assimilation, stomatal conductance, chloroplast CO₂ concentration and the carbon isotope composition of carbohydrate formed during photosynthesis separately for sunlit and shaded leaves within multiple canopy layers. The ecosystem photosynthesis model was validated by comparison to leaf-level gas exchange measurements and estimates of ecosystem-scale photosynthesis from eddy covariance measurements made in a coastal Douglas-fir forest on Vancouver Island. A good agreement was also observed between modelled and measured δ ¹³C values of ecosystem-respired CO₂ ( δ _R). The modelled δ _R values showed strong responses to variation in photosynthetic photon flux density (PPFD), air temperature, vapour pressure deficit (VPD) and available soil moisture in a manner consistent with leaf-level studies of photosynthetic ¹³C discrimination. Sensitivity tests were conducted to evaluate the effect of (1) changes in the lag between the time of CO₂ fixation and the conversion of organic matter back to CO₂; (2) shifts in the proportion of autotrophic and heterotrophic respiration; (3) isotope fractionation during respiration; and (4) environmentally induced changes in mesophyll conductance, on modelled δ _R values. Our results indicated that δ _R is a good proxy for canopy-level C _c/ C _a and ¹³C discrimination during photosynthetic gas exchange, and therefore has several applications in ecosystem physiology.     

Life form-specific variations in leaf water oxygen-18 enrichment in Amazonian vegetation

Leaf water (18)O enrichment (Delta(o)) influences the isotopic composition of both gas exchange and organic matter, with Delta(o) values responding to changes in atmospheric parameters. In order to examine possible influences of plant parameters on Delta(o) dynamics, we measured oxygen isotope ratios (delta(18)O) of leaf and stem water on plant species representing different life forms in Amazonia forest and pasture ecosystems. We conducted two field experiments: one in March (wet season) and another in September (dry season) 2004. In each experiment, leaf and stem samples were collected at 2-h intervals at night and hourly during the day for 50 h from eight species including upper-canopy forest trees, upper-canopy forest lianas, and lower-canopy forest trees, a C(4) pasture grass and a C(3) pasture shrub. Significant life form-related differences were detected in (18)O leaf water values. Initial modeling efforts to explain these observations over-predicted nighttime Delta(o) values by as much as 10 per thousand. Across all species, errors associated with measured values of the delta(18)O of atmospheric water vapor (delta(v)) appeared to be largely responsible for the over-predictions of nighttime Delta(o) observations. We could not eliminate collection or storage of water vapor samples as a possible error and therefore developed an alternative, plant-based method for estimating the daily average delta(v) value in the absence of direct (reliable) measurements. This approach differs from the common assumption that isotopic equilibrium exists between water vapor and precipitation water, by including transpiration-based contributions from local vegetation through (18)O measurements of bulk leaf water. Inclusion of both modified delta(v) and non-steady state features resulted in model predictions that more reliably predicted both the magnitude and temporal patterns observed in the data. The influence of life form-specific patterns of Delta(o) was incorporated through changes in the effective path length, an important but little known parameter associated with the Péclet effect.     

Rhythms in the biting behaviour of a mosquito Armigeres subalbatus

Summary The biting cycle of Armigeres subalbatus is distinctly crepuscular, exhibiting two peaks of activity, a smaller one at dawn and a larger one at dusk. The biting cycle is entrained to natural light-dark cycles and the time interval from dawn to dawn or dusk to dusk peaks is exactly 24 h and from dawn to dusk or dusk to dawn is about 12 h measured at 50% level. This rhythm manifests itself day after day without any marked qualitative change.The rate of change of light intensity may determine the onset of crepuscular biting. The sudden increase (up to ca. 17 lx) or decrease (down to ca. 4 lx) in the intensity of ambient light at the time of sunrise or sunset coincides with the peak of the biting activity.The density of the population of the host-seeking females fluctuates in relation to the phases of the moon, increasing with the full moon phase and decreasing with the new moon phase.Even though the density of the population is greater outdoors than indoors both at ground levels and in the first floor, the peak of activity occurs at the same time in all the places. A vertical stratification of biting activity was also noticed.     

Strategic diel regulation of body mass in European robins

Stochastic dynamic programming (SDP) is a computational technique that has been used to model daily routines of foraging in small birds. A diurnal bird must build up its fat reserves towards dusk in order to avoid starvation during the night, when it cannot feed. However, as well as the benefits of avoiding starvation, storing fat imposes costs such as an increased predation risk and higher flight and metabolic costs. There is therefore an optimal level of fat reserves for a bird to reach at dusk in order to survive overnight without being left with excessive fat reserves at dawn. I tested a prediction common to all SDP models of daily foraging routines, that a bird will attempt to reach this level at dusk, regardless of its fat reserves the previous dawn. I provided supplementary food to manipulate the fat reserves at dawn of free-living European robins, Erithacus rubecula. Diurnal changes in body mass (a reliable estimate of fat reserves) were then monitored remotely. Robins provided with an ad libitum food supply reached almost exactly the same body mass at dusk, regardless of their body mass at dawn, supporting the prediction that birds attempt to reach a target level of reserves at dusk. Copyright 2000 The Association for the Study of Animal Behaviour.     

基于稳定碳同位素的北京西山侧柏林生态系统呼吸区分

基于稳定碳同位素对北京西山侧柏林生态系统呼吸进行定量拆分,能够为该地区森林生态系统碳交换研究奠定基础。本研究采用光谱技术对森林不同高度处CO₂浓度和δ¹³C值进行连续观测,同时采用土壤气室和枝条气室测定地下呼吸和地上呼吸的δ¹³C值,求得生态系统呼吸各组分比例。结合土壤呼吸通量的测定,实现对生态系呼吸的定量区分。结果表明: 森林生态系统各呼吸组分夜间δ¹³C值呈波动变化,植物地上呼吸δ¹³C值变化范围为-31.74‰～-23.33‰,土壤地下呼吸δ¹³C值变化范围为-32.11‰～-27.74‰,生态系统呼吸δ¹³C值介于二者之间。夜间平均土壤呼吸通量为1.70 μmol·m^-2·s^-1,占生态系统总呼吸的47%～91%;夜间地上呼吸量的平均值为0.72 μmol·m^-2·s^-1,对生态系统总呼吸的贡献较小。由同位素混合模型推算求得的日间呼吸变异大于温度响应模型的结果,其平均值分别为2.31和2.28 μmol·m^-2·s^-1。     

青海省都兰县沟里乡幼龄藏狐昼间行为节律

2007年3月6日～4月25日,采用焦点取样和瞬时扫描取样相结合的方法对3只幼龄藏狐(Vulpesferrilata)的昼间行为进行观察记录,并分析其节律及组成.结果表明,休息行为是幼龄藏狐日常活动的主要组成部分,其次为捕食行为和嬉戏行为.幼龄藏狐以晨昏活动为主,主要活动时间集中在8:00～9:00时和16:00～19:00时,其余时间以休息为主.成体藏狐出现后,幼龄藏狐嬉戏行为增加,而休息行为减少,并伴随有奔跑和亲昵行为.     

Carbon isotope fractionation during photorespiration and carboxylation in Senecio

The magnitude of fractionation during photorespiration and the effect on net photosynthetic (13)C discrimination (Delta) were investigated for three Senecio species, S. squalidus, S. cineraria, and S. greyii. We determined the contributions of different processes during photosynthesis to Delta by comparing observations (Delta(obs)) with discrimination predicted from gas-exchange measurements (Delta(pred)). Photorespiration rates were manipulated by altering the O(2) partial pressure (pO(2)) in the air surrounding the leaves. Contributions from (13)C-depleted photorespiratory CO(2) were largest at high pO(2). The parameters for photorespiratory fractionation (f), net fractionation during carboxylation by Rubisco and phosphoenolpyruvate carboxylase (b), and mesophyll conductance (g(i)) were determined simultaneously for all measurements. Instead of using Delta(obs) data to obtain g(i) and f successively, which requires that b is known, we treated b, f, and g(i) as unknowns. We propose this as an alternative approach to analyze measurements under field conditions when b and g(i) are not known or cannot be determined in separate experiments. Good agreement between modeled and observed Delta was achieved with f = 11.6 per thousand +/- 1.5 per thousand, b = 26.0 per thousand +/- 0.3 per thousand, and g(i) of 0.27 +/- 0.01, 0.25 +/- 0.01, and 0.22 +/- 0.01 mol m(-2) s(-1) for S. squalidus, S. cineraria, and S. greyii, respectively. We estimate that photorespiratory fractionation decreases Delta by about 1.2 per thousand on average under field conditions. In addition, diurnal changes in Delta are likely to reflect variations in photorespiration even at the canopy level. Our results emphasize that the effects of photorespiration must be taken into account when partitioning net CO(2) exchange of ecosystems into gross fluxes of photosynthesis and respiration.     

Effect of Irradiance on Chlorophyll Estimation with the Minolta SPAD-502 Leaf Chlorophyll Meter

Leaf chlorophyll content may be used as an indirect indicatorof crop nitrogen status. Chlorophyll meter values (SPAD values)taken with the Minolta SPAD-502 chlorophyll meter in the shadeplantOxalis acetosellaL. and in winter wheat (Triticum aestivumL.)varied by 15 and 8%, respectively, with variation in irradiance.The lowest SPAD-values were measured at high irradiance. Duringa natural night-day-night cycle SPAD values for winter wheatwere lowest in the middle of the day, highest at low irradianceat dusk and dawn and intermediate in darkness before dawn andafter dusk. The results indicate that irradiance during measurementshould be considered when using the Minolta SPAD-502 chlorophyllmeter for the estimation of crop N-status.Copyright 1998 Annalsof Botany Company Chlorophyll meter, nitrogen, irradiance,Oxalis acetosellaL.,Triticum aestivumL., winter wheat.     

Assessing environmental and physiological controls over water relations in a Scots pine (Pinus sylvestris L.) stand through analyses of stable isotope composition of water and organic matter

This study investigated the influence of meteorological, pedospheric and physiological factors on the water relations of Scots pine, as characterized by the origin of water taken up, by xylem transport as well as by carbon isotope discrimination (Delta13C) and oxygen isotope enrichment (Delta18O) of newly assimilated organic matter. For more than 1 year, we quantified delta2H and delta18O of potential water sources and xylem water as well as Delta13C and Delta18O in twig and trunk phloem organic matter biweekly, and related these values to continuously measured or modelled meteorological parameters, soil water content, stand transpiration (ST) and canopy stomatal conductance (G(s)). During the growing season, delta18O and delta2H of xylem water were generally in a range comparable to soil water from a depth of 2-20 cm. Long residence time of water in the tracheids uncoupled the isotopic signals of xylem and soil water in winter. Delta18O but not Delta13C in phloem organic matter was directly indicative of recent environmental conditions during the whole year. Delta18O could be described applying a model that included 18O fractionation associated with water exchange between leaf and atmosphere, and with the production of organic matter as well as the influence of transpiration. Phloem Delta13C was assumed to be concertedly influenced by G(s) and photosynthetically active radiation (PAR) (as a proxy for photosynthetic capacity). We conclude that isotope signatures can be used as effective tools (1) to characterize the seasonal dynamics in source and xylem water, and (2) to assess environmental effects on transpiration and G(s) of Scots pine, thus helping to understand and predict potential impacts of climate change on trees and forest ecosystems.     

Diurnal variation of the delta 13C of pine needle respired CO2 evolved in darkness

The delta 13C of pine needle CO2 evolved in darkness (delta 13Cr) for slash pine trees (Pinus elliottii) was determined by placing recently collected pine needles in darkness and collecting respired CO2 over a short time period (<15 min). Delta 13Cr measurements were made over several 24 h periods to test the hypothesis that significant variation in delta 13Cr would be observed during a diurnal cycle. The delta 13Cr measurements from the 24 h time series trials showed a consistent midday 13C-enrichment (5-10 per thousand) relative to bulk biomass. The delta 13Cr values became more 13C-depleted at night and following shading, and approached bulk-biomass delta 13C values by dawn. The effect of night-time respired 13C-enriched CO2 on the delta 13C value of the remaining assimilate is shown to be minimal (13C depleted by 0.22 per thousand) under field conditions for P. elliottii needles.     

Flocking behaviour in the twilight ascents of Common Swifts Apus apus

Among the many unique flight behaviours of Common Swifts Apus apus, the most puzzling may be their ascents to high altitudes during both dusk and dawn. Twilight ascents have been hypothesized to be functionally related to information acquisition, including integration of celestial orientation cues, high‐altitude visual landmarks and sampling of weather conditions. However, their exact purpose remains unknown. We tracked Common Swifts with tracking radar at their breeding grounds in southern Sweden, and present evidence that during the dusk ascent and dawn descent they often occur in flocks, whereas during the dusk descent and dawn ascent phase they do not. This flocking behaviour suggests that swifts may benefit from conspecific interactions during twilight ascents and descents, possibly through more robust cue acquisition and information exchange in groups, or extending social behaviour also seen in screaming parties before dusk.     

Diurnal lighting patterns and habitat alter opsin expression and colour preferences in a killifish

Spatial variation in lighting environments frequently leads to population variation in colour patterns, colour preferences and visual systems. Yet lighting conditions also vary diurnally, and many aspects of visual systems and behaviour vary over this time scale. Here, we use the bluefin killifish (Lucania goodei) to compare how diurnal variation and habitat variation (clear versus tannin-stained water) affect opsin expression and the preference to peck at different-coloured objects. Opsin expression was generally lowest at midnight and dawn, and highest at midday and dusk, and this diurnal variation was many times greater than variation between habitats. Pecking preference was affected by both diurnal and habitat variation but did not correlate with opsin expression. Rather, pecking preference matched lighting conditions, with higher preferences for blue at noon and for red at dawn/dusk, when these wavelengths are comparatively scarce. Similarly, blue pecking preference was higher in tannin-stained water where blue wavelengths are reduced. In conclusion, L. goodei exhibits strong diurnal cycles of opsin expression, but these are not tightly correlated with light intensity or colour. Temporally variable pecking preferences probably result from lighting environment rather than from opsin production. These results may have implications for the colour pattern diversity observed in these fish.     

Fish drift in a Danube sidearm‐system: II. Seasonal and diurnal patterns

Seasonal and diurnal patterns of larval and juvenile fish drift were investigated in the Marchfeldkanal, a man‐made side branch of the Danube River near Vienna, Austria. A clear seasonal pattern with peak densities in mid‐June was found. Species composition varied over time, showed a site specific pattern and was dominated by tubenose goby Proterorhinus marmoratus . Water temperature was the main factor responsible for the increase of drift densities until the median drift date and repeated occurrence of early larval stages in drift indicated repeated spawning for many species. Significant differences in drift densities between different time periods of the day (day, dusk, night and dawn) were found for common bream Abramis brama , barbel Barbus barbus , chub Leuciscus cephalus , tubenose goby and roach Rutilus rutilus . The highest drift rates occurred at night (2200–0400 hours), with 86% of all larvae drifting during the hours of darkness. Fish larvae of different lengths drifting at different phases of the day were found for common bream, bleak Alburnus alburnus and chub, with largest larvae drifting during dusk (chub) and day (bleak and common bream). For bleak, all gudgeon species Gobio spp., tubenose goby, roach and for all cyprinid species combined, one 2 h night sample was found to be sufficient to predict the total 24 h drift.     

生态系统光合与呼吸拆分的同位素理论、方法和应用进展

生态系统光合和呼吸是构成净生态系统CO₂交换量(NEE)的重要组分。涡度相关技术可直接观测生态系统NEE,并通过建立温度回归或光响应曲线等函数将NEE统计拆分为生态系统光合和呼吸,但是存在自相关和高估白天呼吸等问题。稳定同位素红外光谱技术的进步使高时间分辨率大气CO₂及其稳定碳同位素组成(δ¹³C)的连续观测成为可能,与涡度相关技术观测的NEE数据相结合,可实现昼夜和季节尺度生态系统光合和呼吸拆分。本文系统阐述了生态系统光合与呼吸的同位素通量拆分方法的基本理论与假设,阐述了同位素通量观测技术的发展及其应用进展,综述了同位素通量拆分理论解析生态系统光合与呼吸过程的新机制认识,最后总结并展望了同位素通量拆分理论的不确定性以及开展多种拆分方法综合比较的必要性。