Effect of water availability on leaf water isotopic enrichment in beech seedlings shows limitations of current fractionation models

Current models of leaf water enrichment predict that the differences between isotopic enrichment of water at the site of evaporation (Δ_e) and mean lamina leaf water enrichment (Δ_L) depend on transpiration rates ( E ), modulated by the scaled effective length ( L ) of water isotope movement in the leaf. However, variations in leaf parameters in response to changing environmental conditions might cause changes in the water path and thus L . We measured the diel course of Δ_L for ¹⁸O and ²H in beech seedlings under well-watered and water-limited conditions. We applied evaporative enrichment models of increasing complexity to predict Δ_e and Δ_L, and estimated L from model fits. Water-limited plants showed moderate drought stress, with lower stomatal conductance, E and stem water potential than the control. Despite having double E , the divergence between Δ_e and Δ_L was lower in well-watered than in water-limited plants, and thus, L should have changed to counteract differences in E . Indeed, L was about threefold higher in water-limited plants, regardless of the models used. We conclude that L changes with plant water status far beyond the variations explained by water content and other measured variables, thus limiting the use of current evaporative models under changing environmental conditions.     

Tracing carbon and oxygen isotope signals from newly assimilated sugars in the leaves to the tree-ring archive

The analysis of δ ¹³C and δ ¹⁸O in tree-ring archives offers retrospective insights into environmental conditions and ecophysiological processes. While photosynthetic carbon isotope discrimination and evaporative oxygen isotope enrichment are well understood, we lack information on how the isotope signal is altered by downstream metabolic processes.
In Pinus sylvestris , we traced the isotopic signals from their origin in the leaf water ( δ ¹⁸O) or the newly assimilated carbon ( δ ¹³C), via phloem sugars to the tree-ring, over a time-scale that ranges from hours to a growing season.
Seasonally, variable ¹³C enrichment of sugars related to phloem loading and transport did lead to uncoupling between δ ¹³C in the tree-ring, and the c _i/ c _a ratio at the leaf level. In contrast, the oxygen isotope signal was transferred from the leaf water to the tree-ring with an expected enrichment of 27‰, with time-lags of approximately 2 weeks and with a 40% exchange between organic oxygen and xylem water oxygen during cellulose synthesis.
This integrated overview of the fate of carbon and oxygen isotope signals within the model tree species P. sylvestris provides a novel physiological basis for the interpretation of δ ¹³C and δ ¹⁸O in tree-ring ecology.     

Physiological and isotopic (δ13C and δ18O) responses of three tropical tree species to water and nutrient availability

Water-use efficiency and stable isotope composition were studied in three tropical tree species. Seedlings of Tectona grandis , Swietenia macrophylla and Platymiscium pinnatum were grown at either high or low water supply, and with or without added fertilizer. These three species previously exhibited low, intermediate and high whole-plant water-use efficiency ( TE ) when grown at high water supply in unfertilized soil. Responses of TE to water and nutrient availability varied among species. The TE was calculated as experiment-long dry matter production divided by cumulative water use. Species-specific offsets were observed in relationships between TE and whole-plant ¹³C discrimination (Δ¹³C_p). These offsets could be attributed to a breakdown in the relationship between Δ¹³C_p and the ratio of intercellular to ambient CO₂ partial pressures ( c _i/ c _a) in P. pinnatum , and to variation among species in the leaf-to-air vapour pressure difference ( v ). Thus, a plot of v · TE against c _i/ c _a showed a general relationship among species. Relationships between δ ¹⁸O of stem dry matter and stomatal conductance ranged from strongly negative for S. macrophylla to no relationship for T. grandis . Results suggest inter-specific variation among tropical tree species in relationships between stable isotope ratios ( δ ¹³C and δ ¹⁸O) and the gas exchange processes thought to affect them.     

Oxygen isotope enrichment (Δ18O) reflects yield potential and drought resistance in maize

Measurement of stable isotopes in plant dry matter is a useful phenotypic tool for speeding up breeding advance in C₃ crops exposed to different water regimes. However, the situation in C₄ crops is far from resolved, since their photosynthetic metabolism precludes (at least in maize) the use of carbon isotope discrimination. This paper investigates the use of oxygen isotope enrichment (Δ¹⁸O) as a new secondary trait for yield potential and drought resistance in maize ( Zea mays L). A set of tropical maize hybrids developed by the International Maize and Wheat Improvement Center was grown under three contrasting water regimes in field conditions. Water regimes clearly affected plant growth and yield. In accordance with the current theory, a decrease in water input was translated into large decreases in stomatal conductance and increases in leaf temperature together with concomitant ¹⁸O enrichment of plant matter (leaves and kernels). In addition, kernel Δ¹⁸O correlated negatively with grain yield under well-watered and intermediate water stress conditions, while it correlated positively under severe water stress conditions. Therefore, genotypes showing lower kernel Δ¹⁸O under well-watered and intermediate water stress had higher yields in these environments, while the opposite trend was found under severe water stress conditions. This illustrates the usefulness of Δ¹⁸O for selecting the genotypes best suited to differing water conditions.     

Relationships of grain δ13C and δ18O with wheat phenology and yield under water-limited conditions

Stable carbon isotope composition (δ¹³C) of dry matter has been widely investigated as a selection tool in cereal breeding programmes. However, reports on the possibilities of using stable oxygen isotope composition (δ¹⁸O) as a yield predictor are very scarce and only in the absence of water stress. Indeed, it remains to be tested whether changes in phenology and stomatal conductance in response to water stress overrule the use of either δ¹³C or δ¹⁸O when water is limited. To answer this question, a set of 24 genotypes of bread wheat ( Triticum aestivum ) were assayed in two trials with different levels of deficit irrigation and a third trial under rainfed conditions in a Mediterranean climate (northwest Syria). Grain yield (GY) and phenology (duration from planting to anthesis and from anthesis to maturity) were recorded, and the δ¹³C and δ¹⁸O of grains were analysed to assess their suitability as GY predictors. Both δ¹³C and δ¹⁸O showed higher broad-sense heritabilities ( H ²) than GY. Genotype means of GY across trials were negatively correlated with δ¹³C, as previously reported, but not with δ¹⁸O. Both isotopes were correlated with grain filling duration, whereas δ¹⁸O was also strongly affected by crop duration from planting to anthesis. We concluded that δ¹⁸O of grains is not a proper physiological trait to breed for suboptimal water conditions, as its variability is almost entirely determined by crop phenology. In contrast, δ¹³C of grains, despite being also affected by phenology, still provides complementary information associated with GY.     

Bulk leaf δ18O and δ13C reflect the intensity of intraspecific competition for water in a semi-arid tussock grassland

We investigated the extent to which plant water and nutrient status are affected by intraspecific competition intensity and microsite quality in a monodominant tussock grassland. Leaf gas exchange and stable isotope measurements were used to assess the water relations of Stipa tenacissima tussocks growing along a gradient of plant cover and soil depth in a semi-arid catchment of Southeast Spain. Stomatal conductance and photosynthetic rate decreased with increasing intensity of competition during the wet growing season, leading to foliar δ ¹⁸O and δ ¹³C enrichment. A high potential for runoff interception by upslope neighbours exerted strong detrimental effects on the water and phosphorus status of downslope S. tenacissima tussocks. Foliar δ ¹⁵N values became more enriched with increasing soil depth. Multiple stepwise regression showed that competition potential and/or rhizosphere soil depth accounted for large proportions of variance in foliar δ ¹³C, δ ¹⁸O and δ ¹⁵N among target tussocks (57, 37 and 64%, respectively). The results presented here highlight the key role that spatial redistribution of resources (water and nutrients) by runoff plays in semi-arid ecosystems. It is concluded that combined measurement of δ ¹³C, δ ¹⁸O and nutrient concentrations in bulk leaf tissue can provide insight into the intensity of competitive interactions occurring in natural plant communities.     

Effects of water stress on oxygen, hydrogen and carbon isotope ratios in two species of cotton plants

Abstract. Two cotton species ( Gossypium hirsutum L. cv. SJ-2 and Gossypium barbadence cv. S-5) were grown under irrigated (wet) and non-irrigated (dry) conditions in the same field. Leaf water was enriched in ¹⁸O and deuterium in the dry treatment relative to the wet treatment for both species. Only in plants of S-5 was a similar enrichment observed in leaf cellulose. In both species, the isotopic composition of leaf cellulose must reflect the isotopic composition of the actual water pool involved in cellulose synthesis. Therefore, our observations indicate that one species (SJ-2) can maintain a relative isolation of this water pool from direct evapotranspirational effects. Such plant species will more faithfully record, in the isotopic composition of organic matter, the isotopic composition of ground water. In contrast, the isotopic composition of organic matter in plants such as S-5 could be used as an integrated signal reflecting humidity conditions during growth. Water use efficiency, based on seed-cotton yield and total water applied, correlated linearly with differences in carbon isotopic ratios between species in both the wet and dry treatments and between treatments in each species.     

Isotopic composition of transpiration and rates of change in leaf water isotopologue storage in response to environmental variables

During daylight hours, the isotope composition of leaf water generally approximates steady‐state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady‐state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of ¹⁸O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non‐steady state. We show that non–steady‐state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.     

Stable Isotopes and Carbon Cycle Processes in Forests and Grasslands

Abstract: Scaling and partitioning are frequently two difficult challenges facing ecology today. With regard to ecosystem carbon balance studies, ecologists and atmospheric scientists are often interested in asking how fluxes of carbon dioxide scale across the landscape, region and continent. Yet at the same time, physiological ecologists and ecosystem ecologists are interested in dissecting the net ecosystem CO₂ exchange between the biosphere and the atmosphere to achieve a better understanding of the balance between photosynthesis and respiration within a forest. In both of these multiple-scale ecological questions, stable isotope analyses of carbon dioxide can play a central role in influencing our understanding of the extent to which terrestrial ecosystems are carbon sinks. In this synthesis, we review the theory and present field evidence to address isotopic scaling of CO₂ fluxes. We first show that the ¹³C isotopic signal which ecosystems impart to the atmosphere does not remain constant over time at either temporal or spatial scales. The relative balances of different biological activities and plant responses to stress result in dynamic changes in the ¹³C isotopic exchange between the biosphere and atmosphere, with both seasonal and stand-age factors playing major roles influencing the ¹³C biosphere-atmosphere exchange. We then examine how stable isotopes are used to partition net ecosystem exchange fluxes in order to calculate shifts in the balance of photosynthesis and respiration. Lastly, we explore how fundamental differences in the ¹⁸O isotopic gas exchange of forest and grassland ecosystems can be used to further partition terrestrial fluxes.     

Diurnal and seasonal variation in the carbon isotope composition of leaf dark-respired CO2 in velvet mesquite (Prosopis velutina)

We evaluated diurnal and seasonal patterns of carbon isotope composition of leaf dark-respired CO₂ ( δ ¹³C_l) in the C₃ perennial shrub velvet mesquite ( Prosopis velutina ) across flood plain and upland savanna ecosystems in the south-western USA. δ ¹³C_l of darkened leaves increased to maximum values late during daytime periods and declined gradually over night-time periods to minimum values at pre-dawn. The magnitude of the diurnal shift in δ ¹³C_l was strongly influenced by seasonal and habitat-related differences in soil water availability and leaf surface vapour pressure deficit. δ ¹³C_l and the cumulative flux-weighted δ ¹³C value of photosynthates were positively correlated, suggesting that progressive ¹³C enrichment of the CO₂ evolved by darkened leaves during the daytime mainly resulted from short-term changes in photosynthetic ¹³C discrimination and associated shifts in the δ ¹³C signature of primary respiratory substrates. The ¹³C enrichment of dark-respired CO₂ relative to photosynthates across habitats and seasons was 4 to 6‰ at the end of the daytime period (1800 h), but progressively declined to 0‰ by pre-dawn (0300 h). The origin of night-time and daytime variations in δ ¹³C_l is discussed in terms of the carbon source(s) feeding respiration and the drought-induced changes in carbon metabolism.     

箱式通量观测技术和方法的理论假设及其应用进展

碳(CO₂、CH₄)、氮(N₂O)和水汽(H₂O)等温室气体的交换通量是生态系统物质循环的核心, 是地圈-生物圈-大气圈相互作用的纽带。稳定同位素光谱和质谱技术和方法的进步使碳稳定同位素比值(δ ¹³C)和氧稳定同位素比值(δ ¹⁸O)(CO₂)、δ ¹³C (CH₄)、氮稳定同位素比值(δ ¹⁵N)和δ ¹⁸O (N₂O)、氢稳定同位素比值(δD)和δ ¹⁸O (H₂O)的观测成为可能, 与箱式通量观测技术和方法结合可以实现土壤、植物乃至生态系统尺度温室气体及其同位素通量观测研究。该综述以CO₂及其δ ¹³C通量的箱式观测技术和方法为例, 概述了箱式通量观测系统的基本原理及分类, 阐述了系统设计的理论要求和假设, 综述了从野外到室内土壤、植物叶-茎-根以及生态系统尺度箱式通量观测研究的应用进展及问题, 展望了气体分析精度和准确度、观测数据精度和准确度以及观测数据的代表性评价在箱式通量观测研究中的重要性。     

Temporal and spatial variations in the oxygen-18 content of leaf water in different plant species

Temporal variations in the δ¹⁸ oxygen (δ¹⁸O) content of water transpired by leaves during a simulated diurnal cycle fluctuated around the δ¹⁸O content of the source water. Reconstructed variations in the δ¹⁸O values of leaf water differed markedly from those predicted by conventional models. Even when transpiring leaves were maintained under constant conditions for at least 3 h, strict isotopic steady-state conditions of leaf water (equality of the ¹⁸O/¹⁶O ratios in the input and transpired water) were rarely attained in a variety of plant species (Citrus reticu-lata, Citrus paradisi, Gossypium hirsutum, Helianthus annuns, Musa musaceae and Nicotinia tabacum). Isotopic analysis of water transpired by leaves indicated that leaves approach the isotopic steady state in two stages. The first stage takes 10 to 35 min (with a rate of change of about 3–3%h^?1), while in the second stage further approach to the isotopic steady state is asymptotic (with a rate of change of about 0–4% h^?1), and under conditions of low transpiration leaves can last for many hours. Substantial spatial isotopic heterogeneity was maintained even when leaves were at or near isotopic steady state. An underlying pattern in this isotopic heterogeneity is often discerned with increasing ¹⁸O/¹⁶O ratios from base to tip, and from the centre to the edges of the leaves. It is also shown that tissue water along these spatial isotopic gradients, as well as the average leaf water, can have ¹⁸O/¹⁶O ratios both lower and higher than those predicted by the conventional Craig and Gordon model. We concluded, first, that at any given time during the diurnal cycle of relative humidity the attainment of an isotopic steady state in leaf water cannot be assumed a priori and, secondly, that the isotopic enrichment pattern of leaf water reflects gradual enrichment along the water-flow pathway (e.g. as in a string of pools), rather than a single-step enrichment from source water, as is normally assumed.     

Latent manganese deficiency increases transpiration in barley (Hordeum vulgare)

To investigate if latent manganese (Mn) deficiency leads to increased transpiration, barley plants were grown for 10 weeks in hydroponics with daily additions of Mn in the low n M range. The Mn-starved plants did not exhibit visual leaf symptoms of Mn deficiency, but Chl a fluorescence measurements revealed that the quantum yield efficiency of PSII (F_v/F_m) was reduced from 0.83 in Mn-sufficient control plants to below 0.5 in Mn-starved plants. Leaf Mn concentrations declined from 30 to 7 μg Mn g⁻¹ dry weight in control and Mn-starved plants, respectively. Mn-starved plants had up to four-fold higher transpiration than control plants. Stomatal closure and opening upon light/dark transitions took place at the same rate in both Mn treatments, but the nocturnal leaf conductance for water vapour was still twice as high in Mn-starved plants compared with the control. The observed increase in transpiration was substantiated by ¹³C-isotope discrimination analysis and gravimetric measurement of the water consumption, showing significantly lower water use efficiency in Mn-starved plants. The extractable wax content of leaves of Mn-starved plants was approximately 40% lower than that in control plants, and it is concluded that the increased leaf conductance and higher transpirational water loss are correlated with a reduction in the epicuticular wax layer under Mn deficiency.     

SPM1 encoding a vacuole-localized protease is required for infection-related autophagy of the rice blast fungus Magnaporthe oryzae

The chlorite dismutase (Cld) of Pseudomonas chloritidismutans was purified from the periplasmic fraction in one step by hydroxyapatite chromatography. The enzyme has a molecular mass of 110 kDa and consists of four 31-kDa subunits. Enzyme catalysis followed Michaelis–Menten kinetics, with V _max and K _m values of 443 U mg⁻¹ and 84 μM, respectively. A pyridine–NaOH–dithionite-reduced Cld revealed a Soret peak at 418 nm, indicative for protoheme IX. The spectral data indicate the presence of 1.5 mol  protoheme IX mol⁻¹ tetrameric enzyme while metal analysis revealed 2.2 mol iron mol⁻¹ tetrameric enzyme. High concentrations of chlorite resulted in the disappearance of the Soret peak, which coincided with loss in activity. Electron paramagnetic resonance analyses showed an axial high-spin ferric iron signal. Cld was inhibited by cyanide, azide, but not by hydroxylamine or 3-amino-1,2,3-triazole. Remarkably, the activity was drastically enhanced by kosmotropic salts, and chaotropic salts decreased the activity, in accordance with the Hofmeister series. Chlorite conversion in the presence of ¹⁸O-labeled water did not result in the formation of oxygen with a mass of 34 (¹⁶O–¹⁸O) or a mass of 36 (¹⁸O–¹⁸O), indicating that water is not a substrate in the reaction and that both oxygen atoms originate from chlorite.     

STUDY OF RNA IN SUBCELLULAR FRACTIONS FROM RAT BRAIN: SIMULTANEOUS INCORPORATION OF [14C]URIDINE AND [3H]METHYL-l-METHIONINE

Abstract— By using a combination of subcutaneous and intraventricular injections of [¹⁴C]uridine and [³H]methyl- l -methionine we have obtained maximum incorporation in about 40 min of both radioactive precursors into nuclear RNA from rat brain. In this nuclear fraction we found at least two different types of RNA that were rapidly labelled. One of them incorporated both [¹⁴C]uridine and [³H]methyl groups and seemed to correspond to species of rRNA and their precursors. The other RNA fraction was less methylated or non-methylated and exhibited sedimentation coefficients distributed along a continuous 8–30 % sucrose density gradient. At least part of the latter type of RNA very probably was mRNA, but much of it must conespond to a different RNA similar to that recently described in HeLa cells by P enman , V esco and P enman (1968).
We also found that labelled 185 and 285 rRNA components began leaving the nucleus for the cytoplasm within 24 to 33 min after the radioactive precursors had been injected, and, in the cytoplasmic fraction, the patterns of incorporation for [¹⁴C]uridine and [³H]-methyl groups were similar for the 18S and 28S rRNA components. We estimate that in this fraction of rat brain the 18S rRNA component was 1·4 times more methylated than the 28S component. We also detected a lower sedimentation coefficient for the non- or slightly methylated, species of soluble RNA found in the cytoplasmic fraction.     

Uptake of [3H]Dopamine into Dopaminergic and Noradrenergic Neurones of the Isolated Neurointermediate Lobe of the Rat Hypophysis. Effects of Desipramine and Nomifensine

Abstract: The isolated neurointermediate lobe (NIL) of the rat hypophysis accumulates [³H]dopamine from the incubation medium. Column chromatographic analysis showed that 92% of the tissue radioactivity was contained in the catecholamine fraction. [³H]Dopamine represented 70% and [³H]noradrenaline 30% of the [³H]catecholamines. Desipramine (1 μM) prevented the formation of [³H]noradrenaline without affecting the storage of [³H]dopamine. Nomifensine (10 μM) blocked the storage of [³H]dopamine and [³H]noradrenaline. Thus, in the NIL, [³H]dopamine is taken up into dopaminergic and noradrenergic neurones. In the latter, [³H]dopamine is converted to [³H]noradrenaline, indicating a significant dopamine β-hydroxylase activity in the NIL tissue. A selective labeling of the dopamine stores with [³H]dopamine can be achieved in the presence of desipramine.     

The use of positron emission tomography for studies of long-distance transport in plants: uptake and transport of 18F

Abstract. Positron emission tomography (PET) has been utilized to obtain dynamic images of long distance nutrient translocation in plants. Positron emitting ¹⁸F, produced by a Van de Graaff accelerator using the reaction ¹⁸O(p,n)¹⁸F, was fed in solution to excised stems of Glycine max positioned vertically in a large-aperture PET detector system. Images of tracer activity were recorded with a time resolution of 0.5 min and a spatial resolution of 4 mm. Maximum tracer activities at stem sites were obtained within 3 min of the pulse feed. A model is presented enabling evaluation of regional values for tracer flow, tracer binding, flow speed and flow volume. Analysis of data for one stem position yielded a flow volume of 2.1mm³ min⁻¹ and a flow speed of 36cm min⁻¹. Comparison with the distribution of ¹⁴C-inulin, which was simultaneously fed to the cut stems, indicates the ¹⁸F is suitable for use as an apoplastic tracer; 92% of the tracer activity accumulated in the leaves. The fraction of ¹⁸F that remained bound was most concentrated at stem nodal regions, an observation consistent with the existence of transfer cells at these sites. Advantages and limitations of PET applied to plant physiological investigations are discussed.     

Stable isotope evidence (δ13C, δ18O) for winter feeding on seaweed by Neolithic sheep of Scotland

The antiquity of the use of seaweed to feed domestic animals was investigated through carbon ( δ ¹³C) and oxygen ( δ ¹⁸O) isotope analysis of tooth enamel bioapatite. The analysis was performed on sheep and cattle teeth from two Neolithic sites in Orkney (Scotland). At the Knap of Howar, c . 3600 bc , carbon isotopes reflect grazing on terrestrial plants throughout the year for both sheep and cattle, with no contribution of seaweed to their diet. At the Holm of Papa Westray North (HPWN), c . 3000 bc , significant contribution of seaweed to the sheep diet during winter is indicated by bioapatite δ ¹³C values as high as −5.7‰, far outside of the range of values expected for the feeding on terrestrial C₃ plants, and δ ¹⁸O values higher than expected during winter, possibly caused by ingestion of oceanic water with seaweed. Ingestion of seaweed by sheep at HPWN might have been necessitated by severe reduction of pastures during winter. Results suggest that sheep ingested fresh seaweed rather than dry fodder, perhaps directly on the shore as sheep do nowadays on North Ronaldsay. A significant difference between the two populations is the exclusive reliance on seaweed by the North Ronaldsay sheep, which have developed physiological adaptations to this diet. Contribution of seaweed to the sheep winter diet at HPWN might have been a first step towards this adaptation.     

基于机器语言的岷江上游流域表层土壤氢氧稳定同位素空间分布模拟

为了探明利用机器学习方法对表层土壤氢氧稳定同位素组成(δ²H和δ¹⁸O)进行空间模拟的可行性,揭示δ²H和δ¹⁸O在岷江上游流域的大尺度分布变化规律,本研究在该区域共采集183个0~10 cm土层样本进行氢氧稳定同位素分析。通过变量筛选后,分别采用反向传播(BP)神经网络、随机森林(RF)和支持向量机(SVM)建立研究区浅层土壤δ²H和δ¹⁸O的模型并进行精度评价,通过结构方程模型(SEM)揭示各辅助变量与土壤水δ²H和δ¹⁸O之间的机理过程。结果表明: RF模型的预测精度最高,可分别解释研究区表层土壤δ²H和δ¹⁸O变异的75.0%和64.0%。在该模型中,土壤体积含水量是最重要的辅助变量,对δ²H和δ¹⁸O的贡献率分别达到48.9%和37.4%。植被因子对表层土壤δ²H和δ¹⁸O的影响比气候因子大,且气候因子对δ²H和δ¹⁸O的影响过程受到植被因子的干扰。在所有辅助变量中,降水氢氧同位素由于分馏作用对δ²H和δ¹⁸O的影响最小。该区域表层土壤δ²H和δ¹⁸O在生长季各月的变化显著,在生长季初期和末期主要受植被的影响分别呈大幅度上升和下降趋势,而气候变化则导致生长季中期小幅波动。