Effect of CO2 on growth and toxicity of Alexandrium tamarense from the East China Sea,a major producer of paralytic shellfish toxins

In recent decades, the frequency and intensity of harmful algal blooms (HABs), as well as a profusion of toxic phytoplankton species, have significantly increased in coastal regions of China. Researchers attribute this to environmental changes such as rising atmospheric CO₂ levels. Such addition of carbon into the ocean ecosystem can lead to increased growth, enhanced metabolism, and altered toxicity of toxic phytoplankton communities resulting in serious human health concerns. In this study, the effects of elevated partial pressure of CO₂ (pCO₂) on the growth and toxicity of a strain of Alexandrium tamarense (ATDH) widespread in the East and South China Seas were investigated. Results of these studies showed a higher specific growth rate (0.31 ± 0.05 day⁻¹) when exposed to 1000 μatm CO₂, (experimental), with a corresponding density of (2.02 ± 0.19) × 10⁷ cells L⁻¹, that was significantly larger than cells under 395 μatm CO₂₍control). These data also revealed that elevated pCO₂ primarily affected the photosynthetic properties of cells in the exponential growth phase. Interestingly, measurement of the total toxin content per cell was reduced by half under elevated CO₂ conditions. The following individual toxins were measured in this study: C1, C2, GTX1, GTX2, GTX3, GTX4, GTX5, STX, dcGTX2, dcGTX3, and dcSTX. Cells grown in 1000 μatm CO₂ showed an overall decrease in the cellular concentrations of C1, C2, GTX2, GTX3, GTX5, STX, dcGTX2, dcGTX3, and dcSTX, but an increase in GTX1 and GTX4. Total cellular toxicity per cell was measured revealing an increase of nearly 60% toxicity in the presence of elevated CO₂ compared to controls. This unusual result was attributed to a significant increase in the cellular concentrations of the more toxic derivatives, GTX1 and GTX4.Taken together; these findings indicate that the A. tamarense strain ATDH isolated from the East China Sea significantly increased in growth and cellular toxicity under elevated pCO₂ levels. These data may provide vital information regarding future HABs and the corresponding harmful effects as a result of increasing atmospheric CO₂.     

海洋酸化生态学研究进展

工业革命以来,人类排放的大量二氧化碳引起温室效应的同时,也被海洋吸收使得全球海洋出现了严重的酸化。海洋酸化及伴随的海水碳酸盐化学体系的变化对海洋生物产生深远的影响。以海洋酸化对钙化作用和光合作用的影响为重点,总结了近年来关于海洋酸化的研究,介绍了海洋中不同生态系统对海洋酸化的响应。一方面,海水中CO23-浓度和碳酸钙饱和度的降低对海洋钙化生物造成严重损害,生活在高纬的冷水珊瑚和翼足目等文石生产者是最早的受害者;贝类和棘皮动物在钙化早期对海洋酸化尤其敏感,其幼体存活率受到海洋酸化的严重制约。另一方面,CO2浓度的增加能促进海洋植物的光合作用和生长,增加初级生产力,改变浮游植物的群落组成。此外,海洋酸化可以促进固氮和脱氮作用同时削弱硝化作用,改变溶氧浓度分布和金属的生物可利用性,从而对海洋生物产生间接影响。海洋酸化对海洋生态系统的影响机制复杂,影响程度深远。为了能准确的评估海洋酸化的生态学效应,需要更全面深入的研究。     

Phenotypic plasticity of coralline algae in a High CO2 world

It is important to understand how marine calcifying organisms may acclimatize to ocean acidification to assess their survival over the coming century. We cultured the cold water coralline algae, Lithothamnion glaciale, under elevated pCO₂ (408, 566, 770, and 1024 μatm) for 10 months. The results show that the cell (inter and intra) wall thickness is maintained, but there is a reduction in growth rate (linear extension) at all elevated pCO₂. Furthermore a decrease in Mg content at the two highest CO₂ treatments was observed. Comparison between our data and that at 3 months from the same long‐term experiment shows that the acclimation differs over time since at 3 months, the samples cultured under high pCO₂ showed a reduction in the cell (inter and intra) wall thickness but a maintained growth rate. This suggests a reallocation of the energy budget between 3 and 10 months and highlights the high degree plasticity that is present. This might provide a selective advantage in future high CO₂ world.     

开放式空气CO2浓度增高条件下旱地土壤气体CO2浓度廓线测定

设计了一套适合于FACE(free airCO2 enrichment)平台的旱地土壤气体CO2 浓度廓线测定方法 ,并将其应用于田间实验 .在江苏省无锡市郊区具有太湖地区典型水稻土的稻麦轮作农田 ,对FACE和对照麦田以及裸土 0～ 30cm土层的土壤气体CO2 浓度廓线进行了观测研究 .结果表明 ,所采用的方法满足进行旱地农田土壤气体CO2 浓度廓线研究的要求 ;在 0～ 30cm土层中 ,上层土壤气体中的CO2 向上垂直扩散要比下层土壤快 ;在作物旺盛生长期 ,大气CO2 浓度升高 2 0 0± 4 0 μmol·mol-1使 0～ 30cm土层的土壤气体CO2 浓度显著提高 14 %± 5 % (t 检验P <0 .0 0 1) .     

木本植物对CO2浓度和温度升高的相互作用的响应

CO2浓度和温度是影响木本植物生长和发育的两个关键因子,二者在全球变化中的相互作用对木本植物生长和发育具有显著的影响。大多数研究表明：CO2浓度增加和温度升高的相互作用可能影响木本植物的生长发育,促进光合作用;呼吸作用对CO2浓度增加和温度升高的相互作用存在长期和短期响应差异;二者的相互作用促进生物量增加和生产力的增长。木本植物对CO2浓度和温度升高的相互作用的响应程度因植物种类而异。     

Corynosoma spp. may be useful biological tags for saithe in the northern North Sea

The acanthocephalan parasites Corynosoma spp. could be useful biological tags for estimating the proportions of saithe recruiting from nursery areas in Scottish and Norwegian coastal waters to the central northern North Sea.     

A study on the inactivation of micro-organisms and enzymes by high pressure CO2

This study addresses some microbial inactivation phenomena induced by high pressure CO2 over micro-organisms and enzymes. The activity of four selected enzymes was measured before and after treatment with CO2 under pressure in both buffer solutions and natural cellular environment (E. coli cells and tomato paste). Results are reported for acid phosphatase, alkaline phosphatase, ATPase, and pectinase at different conditions of temperature, CO2 pressure, and treatment time (32-40 degrees C, 85-150 bar, 30-70 min). The results obtained show that the high pressure CO2 treatment induces an inactivation of cellular enzymatic activity higher than the one caused on the same enzymes in solution. However, the measured activity difference is not caused by a damage at the enzymes molecular level but is a consequence of the permeabilization of the cellular envelopes which leads to a release of unmodified enzymes from the cells with simultaneous drop of enzymatic cellular activity. The reported data suggest that the bacterial cell death is probably due not to a selective effect of high pressure CO2 treatment but to simultaneous detrimental action of CO2 on cellular membrane and cell wall.     

Modified future diurnal variability of the global surface ocean CO2 system

Our understanding of how increasing atmospheric CO₂ and climate change influences the marine CO₂ system and in turn ecosystems has increasingly focused on perturbations to carbonate chemistry variability. This variability can affect ocean-climate feedbacks and has been shown to influence marine ecosystems. The seasonal variability of the ocean CO₂ system has already changed, with enhanced seasonal variations in the surface ocean pCO₂ over recent decades and further amplification projected by models over the 21st century. Mesocosm studies and CO₂ vent sites indicate that diurnal variability of the CO₂ system, the amplitude of which in extreme events can exceed that of mean seasonal variability, is also likely to be altered by climate change. Here, we modified a global ocean biogeochemical model to resolve physically and biologically driven diurnal variability of the ocean CO₂ system. Forcing the model with 3-h atmospheric outputs derived from an Earth system model, we explore how surface ocean diurnal variability responds to historical changes and project how it changes under two contrasting 21st-century emission scenarios. Compared to preindustrial values, the global mean diurnal amplitude of pCO₂ increases by 4.8 μatm (+226%) in the high-emission scenario but only 1.2 μatm (+55%) in the high-mitigation scenario. The probability of extreme diurnal amplitudes of pCO₂ and [H⁺] is also affected, with 30- to 60-fold increases relative to the preindustrial under high 21st-century emissions. The main driver of heightened pCO₂ diurnal variability is the enhanced sensitivity of pCO₂ to changes in temperature as the ocean absorbs atmospheric CO₂. Our projections suggest that organisms in the future ocean will be exposed to enhanced diurnal variability in pCO₂ and [H⁺], with likely increases in the associated metabolic cost that such variability imposes.     

Atmospheric CO2 concentration may directly affect leaf respiration measurement in tobacco,but not respiration itself*

When atmospheric CO₂ concentration increases, various consequences for plant metabolism have been suggested, such as changes in photosynthesis, photorespiration or respiration which can affect growth and carbon sequestration. In addition to long‐term (indirect) effects on respiration, short‐term (direct) effects of CO₂ concentration on the respiration of leaves, shoots and roots are described in the literature. In most cases, respiration is reported to be inhibited by increased CO₂ concentration, but the mechanism(s) are not yet understood. It has been shown previously that, when the respective technical problems and properties of a gas exchange system are fully considered, a short‐term increase in CO₂ (up to 4200 µmol mol^?1) had no effect on respiration of Phaseolus or Populus leaves (Jahnke, Plant, Cell and Environment 24, 1139–1151, 2001). However, in the present study, large (apparent) CO₂ effects were found with mature Nicotiana leaves whereas, in young leaves, the effect was absent. The experimental results clearly show that the observed direct CO₂ effect on dark CO₂ efflux in the mature tobacco leaves was caused by leakage of CO₂ inside the leaves (and the magnitude of the effect was dependent on the size of the leakage). Nicotiana leaves are, in contrast to Phaseolus and Populus leaves (which are heterobaric), characterized by a homobaric anatomy in which intercellular air spaces are not compartmented and provide a continuous system of open pores in the lateral (paradermal) direction of the leaves. Mesophyll porosity increases with leaf development, which explains the differences between young and mature tobacco leaves. When internal leakage was experimentally restricted, the CO₂ inhibition on CO₂ efflux was no longer observed. It is concluded that the measured direct CO₂ effect(s) on leaf CO₂ efflux in the dark are artefactual, and that a true direct CO₂ effect on leaf respiration does not exist.     

Nitrogen fertilization raises CO2 efflux from inorganic carbon: A global assessment

Nitrogen (N) fertilization is an indispensable agricultural practice worldwide, serving the survival of half of the global population. Nitrogen transformation (e.g., nitrification) in soil as well as plant N uptake releases protons and increases soil acidification. Neutralizing this acidity in carbonate‐containing soils (7.49 × 10⁹ ha; ca. 54% of the global land surface area) leads to a CO₂ release corresponding to 0.21 kg C per kg of applied N. We here for the first time raise this problem of acidification of carbonate‐containing soils and assess the global CO₂ release from pedogenic and geogenic carbonates in the upper 1 m soil depth. Based on a global N‐fertilization map and the distribution of soils containing CaCO₃, we calculated the CO₂ amount released annually from the acidification of such soils to be 7.48 × 10¹² g C/year. This level of continuous CO₂ release will remain constant at least until soils are fertilized by N. Moreover, we estimated that about 273 × 10¹² g CO₂‐C are released annually in the same process of CaCO₃ neutralization but involving liming of acid soils. These two CO₂ sources correspond to 3% of global CO₂ emissions by fossil fuel combustion or 30% of CO₂ by land‐use changes. Importantly, the duration of CO₂ release after land‐use changes usually lasts only 1–3 decades before a new C equilibrium is reached in soil. In contrast, the CO₂ released by CaCO₃ acidification cannot reach equilibrium, as long as N fertilizer is applied until it becomes completely neutralized. As the CaCO₃ amounts in soils, if present, are nearly unlimited, their complete dissolution and CO₂ release will take centuries or even millennia. This emphasizes the necessity of preventing soil acidification in N‐fertilized soils as an effective strategy to inhibit millennia of CO₂ efflux to the atmosphere. Hence, N fertilization should be strictly calculated based on plant‐demand, and overfertilization should be avoided not only because N is a source of local and regional eutrophication, but also because of the continuous CO₂ release by global acidification.     

CO2倍增条件下长白赤松幼苗土壤CO2廓线的动态

采用固定在土壤中的气井系统,监测土壤剖面的CO2动态及其与长白赤松 (Pinus sylvestris var. sylvestriformis (Takenouchi) Cheng et C. D. Chu) 幼苗根系发展之间的关系.实验研究共设4种CO2处理,分别是环境CO2浓度,无苗;CO2为700 μmol/mol,无苗;环境CO2浓度,有苗;CO2为700 μmol/mol,有苗.通过对土壤剖面CO2气体的同步采集与分析表明:土壤CO2廓线与幼苗根系的生物活性密切相关.在土壤表面及壤土和沙土的边界层中,根系分布密集,根系的呼吸作用对那两个土层CO2贡献大;随着幼苗的季节生长,与环境CO2浓度比较,CO2倍增将导致土壤剖面上CO2浓度最大区域由表面向壤土和沙土边界层的转移.本文采用的气井系统提供了一种对土壤无破坏、经济、简单并且能够用于监测幼苗地下过程的廓线研究方法.     

Seaweed fails to prevent ocean acidification impact on foraminifera along a shallow‐water CO2 gradient

Ocean acidification causes biodiversity loss, alters ecosystems, and may impact food security, as shells of small organisms dissolve easily in corrosive waters. There is a suggestion that photosynthetic organisms could mitigate ocean acidification on a local scale, through seagrass protection or seaweed cultivation, as net ecosystem organic production raises the saturation state of calcium carbonate making seawater less corrosive. Here, we used a natural gradient in calcium carbonate saturation, caused by shallow‐water CO₂ seeps in the Mediterranean Sea, to assess whether seaweed that is resistant to acidification (Padina pavonica) could prevent adverse effects of acidification on epiphytic foraminifera. We found a reduction in the number of species of foraminifera as calcium carbonate saturation state fell and that the assemblage shifted from one dominated by calcareous species at reference sites (pH ~8.19) to one dominated by agglutinated foraminifera at elevated levels of CO₂ (pH ~7.71). It is expected that ocean acidification will result in changes in foraminiferal assemblage composition and agglutinated forms may become more prevalent. Although Padina did not prevent adverse effects of ocean acidification, high biomass stands of seagrass or seaweed farms might be more successful in protecting epiphytic foraminifera.     

长白山四种林分土壤CO2释放通量的研究

对长白山地区 4种典型林分下土壤释放CO2 通量的研究表明 ,在较短的时间尺度上 ,4种林型在CO2 释放通量上有显著的差别 ,各林型CO2 通量的释放表现出明显的空间异质性 ,其释放顺序为 :山杨白桦混交林 >原始阔叶红松林 >白桦林 >山杨林 ,其中山杨白桦混交林对CO2 释放的贡献最大 ,而山杨林最小。 4种林分土壤CO2 通量的全日变化过程并不一致 ,除山杨白桦混交林和气温有较一致性的规律 ,呈较明显的单峰型外 ,其余并无明显的变化规律。凋落物对林地CO2 释放通量的变化有显著的影响 ,有凋落物覆盖的林地土壤CO2 释放量明显大于无凋落物覆盖的林地 ,其平均通量为 2 6 2 86～ 5 5 6 5 5mgC·m-2 ·h-1,而无凋落物覆盖林地土壤CO2 的平均通量为 2 0 5 85～ 395 6 9mg·m-2 ·h-1。凋落物排放CO2 量占林地总排放量的14 %～ 30 % ,并且改变了林地CO2 释放通量大小的排序。     

Modelling assimilation and intercellular CO2 from measured conductance: a synthesis of approaches

A spectrum of models that estimate assimilation rate A from intercellular carbon dioxide concentration (C_i) and measured stomatal conductance to CO₂ (g_c) were investigated using leaf‐level gas exchange measurements. The gas exchange measurements were performed in a uniform loblolly pine stand (Pinus taeda L.) using the Free Air CO₂ Enrichment (FACE) facility under ambient and elevated atmospheric CO₂ for 3 years. These measurements were also used to test a newly proposed framework that combines basic properties of the A–C_i curve with a Fickian diffusion transport model to predict the relationship between C_i/C_a and g_c, where C_a is atmospheric carbon dioxide concentration. The widely used Ball–Berry model and five other models as well as the biochemical model proposed by Farquhar et al. (1980) were also reformulated to express variations in C_i/C_a as a function of their corresponding driving mechanisms. To assess the predictive capabilities of these approaches, their respective parameters were estimated from independent measurements of long‐term stable carbon isotope determinations (δ¹³C), meteorological variables, and ensemble A–C_i curves. All eight approaches reproduced the measured A reasonably well, in an ensemble sense, from measured water vapour conductance and modeled C_i/C_a. However, the scatter in the instantaneous A estimates was sufficiently large for both ambient and elevated C_a to suggest that other transient processes were not explicitly resolved by all eight parameterizations. An important finding from our analysis is that added physiological complexity in modeling C_i/C_a (when g_c is known) need not always translate to increased accuracy in predicting A. Finally, the broader utility of these approaches to estimate assimilation and net ecosystem exchange is discussed in relation to elevated atmospheric CO₂.     

Elevated CO2 alters belowground exoenzyme activities in tussock tundra

A three-year exposure to a CO2 concentration of 680 mol mol-1 altered the enzymic characteristics of root surfaces, associated ectomycorrhizae, and in soils surrounding roots in a tussock tundra ecosystem of north Alaska, USA. At elevated CO2, phosphatase activity was higher on Eriophorum vaginatum root surfaces, ectomycorrhizal rhizomorphs and mantles associated with Betula nana roots, and in Oe and Oi soil horizons associated with plant roots. Also, endocellulase and exocellulase activities at elevated CO2 were higher in ectomycorrhizal rhizomorphs and lower in Oe and Oi soil horizons associated with roots. These results suggest that arctic plants respond to raised CO2 by increasing activities associated with nutrient acquisition, e.g. higher phosphatase activities on surfaces of roots and ectomycorrhizae, and greater cellulase activity in ectomycorrhizae. Changes in enzyme activities of surrounding soils are consistent with an increase in carbon exudation from plant roots, which would be expected to inhibit cellulase activities and stimulate phosphatase activities of soil microflora. These data were used to modify existing simulation models describing tussock phosphatase activities and litter decay. Model projections suggest that observed increases in phosphatase activities at 680 mol mol-1 CO2 could augment total annual phosphorus release within tussocks by more than 40%, at present levels of root and ectomycorrhizae biomass. This includes a nearly three-fold increase in potential phosphatase activity of E. vaginatum roots, per unit of surface area. Observed reductions in cellulase activities could diminish cellulose turnover by 45% in soils within rooting zones, which could substantially increase mineral nitrogen availability in soils due to lowered microbial immobilization.     

Rewinding the tape: selection of algae adapted to high CO2 at current and pleistocene levels of CO2

Selective history is thought to constrain the extent and direction of future adaptation by limiting access to genotypes that are advantageous in a novel environment. Populations of Chlamydomonas previously selected at high CO2 were either backselected at ambient levels of CO2, or selected at levels of CO2 that last occurred during glaciation in the Pleistocene. There was no effect of selective history on adaptation to either level of CO2, and the high CO2 phenotypes were evolutionarily reversible such that fitness in ambient CO2 returned to values seen in controls. CO2 uptake affinity improved relative to the ancestor in both ambient and glacial CO2, although wild-type regulation of CO2 uptake, which deteriorated during previous selection at high CO2, was not restored by selection at lower levels of CO2. Trade-offs in both CO2 uptake affinity and growth were seen after selection at any given level of CO2. Adaptation to ambient and glacial-era levels of CO2 produced a range of phenotypes, suggesting that chance rather than selective history contributes to the divergence of replicate populations in this system.     

Yield response of Lolium perenne swards to free air CO2 enrichment increased over six years in a high N input system on fertile soil

After a step increase in the atmospheric partial pressure of CO₂ (pCO₂), the availability of mineral N may be insufficient to meet the plant's increased demand for N. Over time, however, the ecosystem may adapt to the new conditions, and a new equilibrium may be established in the fluxes of C and N. This would result in a higher dry mass (DM) yield response of the plants to elevated pCO₂. The effect of elevated atmospheric pCO₂ (60 Pa pCO₂) was studied in Lolium perenne L. swards with two N fertilization treatments (14 and 56 g m^?2 y^?1) in a six‐year FACE (Free Air Carbon dioxide Enrichment) experiment. In the high N treatment, the input of N with fertilizer considerably exceeded the export of N with the harvested plant material in both CO₂ treatments leading to an apparent net input of N into the ecosystem. Accordingly, the proportion of harvested N derived from ¹⁵N labelled fertilizer N, applied throughout the experiment (< 6 years), increased over the years. Under these high N conditions, the annual DM yield response of the Lolium perenne sward to elevated pCO₂ increased (from 7% in 1993 to 25% in 1998). In parallel, the response of N yield to elevated pCO₂ increased, and the initially negative effect of elevated pCO₂ on specific leaf area (SLA) disappeared. The high N input system seemed to overcome in part an initially limiting effect of N on the yield response to elevated pCO₂ within a few years. In contrast, there was no apparent net input of N into the ecosystem in the low N treatment, because N fertilization just compensated the export of N with the harvested plant material. Accordingly, the proportion of harvested N yield, derived from fertilizer N, which was applied throughout the experiment, remained low. At low N, the availability of mineral N strongly limited plant growth and yield production in both CO₂ treatments; the low yields of DM and N, the low concentration of N in the plant material, and the low SLA reflected this. Although the plants grew under the same environmental conditions and the same management treatment as plants in the high N treatment, the response of DM yields to elevated pCO₂ in the low N treatment remained weak throughout the experiment (5% in 1993 and 9% in 1998). The results are discussed in the context of the sizes of the different N pools in the soil, the allocation of N within the plant and the possible effects on temporal immobilization, and the availability of mineral N for yield production as affected by elevated pCO₂ and N fertilization.     

Winners always win: growth of a wide range of plant species from low to future high CO2

Evolutionary adaptation to variation in resource supply has resulted in plant strategies that are based on trade‐offs in functional traits. Here, we investigate, for the first time across multiple species, whether such trade‐offs are also apparent in growth and morphology responses to past low, current ambient, and future high CO₂ concentrations. We grew freshly germinated seedlings of up to 28 C₃ species (16 forbs, 6 woody, and 6 grasses) in climate chambers at 160 ppm, 450 ppm, and 750 ppm CO₂. We determined biomass, allocation, SLA (specific leaf area), LAR (leaf area ratio), and RGR (relative growth rate), thereby doubling the available data on these plant responses to low CO₂. High CO₂ increased RGR by 8%; low CO₂ decreased RGR by 23%. Fast growers at ambient CO₂ had the greatest reduction in RGR at low CO₂ as they lost the benefits of a fast‐growth morphology (decoupling of RGR and LAR [leaf area ratio]). Despite these shifts species ranking on biomass and RGR was unaffected by CO₂, winners continued to win, regardless of CO_2. Unlike for other plant resources we found no trade‐offs in morphological and growth responses to CO₂ variation, changes in morphological traits were unrelated to changes in growth at low or high CO₂. Thus, changes in physiology may be more important than morphological changes in response to CO₂ variation.     

开放式空气CO2浓度增高对水稻冠层微气候的影响

利用位于江苏省无锡市安镇的我国唯一的农田开放式空气CO2 浓度增高 (FACE)系统平台 ,于2 0 0 1年 8月 2 6日至 10月 13日 (水稻抽穗至成熟期 )进行水稻作物冠层微气候连续观测 ,以研究FACE对水稻冠层微气候特征的影响 .结果表明 ,FACE降低了水稻叶片的气孔导度 ,FACE与对照水稻叶片气孔导度的差异上层叶片大于下层叶片 ,生长前期大于生长后期 .FACE使白天水稻冠层和叶片温度升高 ,这种差异生长前期大于生长后期 ;但FACE对夜间水稻冠层温度的影响不明显 .在水稻旺盛生长的抽穗开花期 ,晴天正午前后FACE水稻冠层温度比对照高 1.2℃ ;从开花至成熟期 ,FACE水稻冠层白天平均温度比对照高 0 .4 3℃ .FACE对冠层空气温度也有影响 ,白天水稻冠层空气温度FACE高于对照 ,这种差异随太阳辐射增强而增大且冠层中部大于冠层顶部 ;冠层中部空气温度FACE与对照的差异 (Tface-Tambient)日最大值在 0 .4 7～ 1.2℃之间 ,而冠层顶部的Tface-Tambient日最大值在 0 .37～ 0 .8℃之间 .夜间水稻冠层空气温度FACE与对照差别不大 ,变化在± 0 .3℃之内 .而FACE对水稻冠层空气湿度无显著影响 ,表明FACE使水稻叶片气孔导度降低 ,从而削弱了植株的蒸腾降温作用 ,导致水稻冠层温度和冠层空气温度升高 ,改变了整个水稻冠层的温度环