Foliage of oaks grown under elevated CO2 reduces performance of Antheraea polyphemus (Lepidoptera: Saturniidae)

To understand how the increase in atmospheric CO2 from human activity may affect leaf damage by forest insects, we examined host plant preference and larval performance of a generalist herbivore, Antheraea polyphemus Cram., that consumed foliage developed under ambient or elevated CO2. Larvae were fed leaves from Quercus alba L. and Quercus velutina Lam. grown under ambient or plus 200 microl/liter CO2 using free air carbon dioxide enrichment (FACE). Lower digestibility of foliage, greater protein precipitation capacity in frass, and lower nitrogen concentration of larvae indicate that growth under elevated CO2 reduced the food quality of oak leaves for caterpillars. Consuming leaves of either oak species grown under elevated CO2 slowed the rate of development of A. polyphemus larvae. When given a choice, A. polyphemus larvae preferred Q. velutina leaves grown under ambient CO2; feeding on foliage of this species grown under elevated CO2 led to reduced consumption, slower growth, and greater mortality. Larvae compensated for the lower digestibility of Q. alba leaves grown under elevated CO2 by increasing the efficiency of conversion of ingested food into larval mass. Despite equivalent consumption rates, larvae grew larger when they consumed Q. alba leaves grown under elevated compared with ambient CO2. Reduced consumption, slower growth rates, and increased mortality of insect larvae may explain lower total leaf damage observed previously in plots in this forest exposed to elevated CO2. By subtly altering aspects of leaf chemistry, the ever-increasing concentration of CO2 in the atmosphere will change the trophic dynamics in forest ecosystems.     

Olfactory preferences of Popillia japonica, Vanessa cardui, and Aphis glycines for Glycine max grown under elevated CO2

Levels of atmospheric CO(2) have been increasing steadily over the last century and are projected to increase even more dramatically in the future. Soybeans (Glycine max L.) grown under elevated levels of CO(2) have larger herbivore populations than soybeans grown under ambient levels of CO(2). Increased abundance could reflect the fact that these herbivores are drawn in by increased amounts of volatiles or changes in the composition of volatiles released by plants grown under elevated CO(2) conditions. To determine impacts of elevated CO(2) on olfactory preferences, Japanese beetles (Popillia japonica Newman) and soybean aphids (Aphis glycines Matsumura) were placed in Y-tube olfactometers with a choice between ambient levels of CO(2) gas versus elevated levels of CO(2) gas or damaged and undamaged leaves and plants grown under ambient levels of CO(2) versus damaged and undamaged plants grown under elevated levels of CO(2). All plants had been grown from seeds under ambient or elevated levels of CO(2). Painted lady butterflies (Vanessa cardui L.) were placed in an oviposition chamber with a choice between plants grown under ambient and elevated levels of CO(2). A. glycines and V. cardui showed no significant preference for plants in either treatment. P. japonica showed no significant preference between ambient levels and elevated levels of CO(2) gas. There was a significant P. japonica preference for damaged plants grown under ambient CO(2) versus undamaged plants but no preference for damaged plants grown under elevated CO(2) versus undamaged plants. P. japonica also preferred damaged plants grown under elevated levels of CO(2) versus damaged plants grown under ambient levels of CO(2). This lack of preference for damaged plants grown under elevated CO(2) versus undamaged plants could be the result of the identical elevated levels of a green leaf volatile (2-hexenal) present in all foliage grown under elevated CO(2) regardless of damage status. Green leaf volatiles are typically released from damaged leaves and are used as kairomones by many herbivorous insects for host plant location. An increase in production of volatiles in soybeans grown under elevated CO(2) conditions may lead to larger herbivore outbreaks in the future.     

Does elevated CO2 ameliorate the impact of O3 on chlorophyll content and photosynthesis in potato (Solanum tuberosum)?

This study examined the impact of season-long exposure to elevated carbon dioxide (CO2) and ozone (O3), individually and in combination, on leaf chlorophyll content and gas exchange characteristics in potato (Solanum tuberosum L. cv. Bintje). Plants grown in open-top chambers were exposed to three CO2 (ambient, 550 and 680 μmol mol-1) and two O3 treatments (ambient and elevated; 25 and 65 nmol mol-1, 8 h day-1 means, respectively) between crop emergence and maturity; plants were also grown in unchambered field plots. Non-destructive measurements of chlorophyll content and visible foliar injury were made for all treatments at 2-week intervals between 43 and 95 days after emergence. Gas exchange measurements were made for all except the intermediate 550 μmol mol-1 CO2 treatment. Season-long exposure to elevated O3 under ambient CO2 reduced chlorophyll content and induced extensive visible foliar damage, but had little effect on net assimilation rate or stomatal conductance. Elevated CO2 had no significant effect on chlorophyll content, but greatly reduced the damaging impact of O3 on chlorophyll content and visible foliar damage. Light-saturated assimilation rates for leaves grown under elevated CO2 were consistently lower when measured under either elevated or ambient CO2 than in equivalent leaves grown under ambient CO2. Analysis of CO2 response curves revealed that CO2-saturated assimilation rate, maximum rates of carboxylation and electron transport and respiration decreased with time. CO2-saturated assimilation rate was reduced by elevated O3 during the early stages of the season, while respiration was significantly greater under elevated CO2 as the crop approached maturity. The physiological origins of these responses and their implications for the performance of potato in a changing climate are discussed.     

Impact of elevated CO2 and increased temperature on Japanese beetle herbivory

To examine how the major elements of global change affect herbivory in agroecosystems, a multifactorial experiment was conducted where soybeans were grown at two levels of carbon dioxide and temperature, including those predicted for 2050, under otherwise normal field conditions. Japanese beetles (Popillia japonica Newman) were enclosed on foliage for 24 h, after which the beetle survivorship, total and per capita leaf consumption, and leaf protease inhibitor activity were measured. The direct effect of temperature on beetle consumption and survivorship also was measured under controlled environmental conditions. No differences in total foliage consumption were observed; however, beetles forced to feed at elevated temperature in the field demonstrated greater per capita consumption and reduced survivorship compared to beetles feeding at ambient temperature. Survivorship was also greater for beetles that consumed foliage grown under elevated CO₂, but there were no interactive effects of CO₂ and temperature, and no differences in leaf chemistry were resolved. Leaf consumption by beetles increased strongly with increasing temperature up to ～37° C, above which increased mortality caused a precipitous decrease in consumption. An empirical model based on the temperature dependence of leaf consumption and flight suggests that the 3.5°C increase in temperature predicted for 2050 will increase the optimal feeding window for the Japanese beetle by 290%. Elevated temperature and CO₂ operating independently have the potential to greatly increase foliage damage to soybean by chewing insects, such as Popillia japonica, potentially affecting crop yields.     

Evidence that carbon dioxide enrichment alleviates ureide-induced decline of nodule nitrogenase activity

The hypothesis that elevated [CO(2)] alleviates ureide inhibition of N(2)-fixation was tested. Short-term responses of the acetylene reduction assay (ARA), ureide accumulation and total non-structural carbohydrate (TNC) levels were measured following addition of ureide to the nutrient solution of hydroponically grown soybean. The plants were exposed to ambient (360 micromol mol(-1)) or elevated (700 micromol mol(-1)) [CO(2)]. Addition of 5 and 10 mM ureide to the nutrient solution inhibited N(2)-fixation activity under both ambient and elevated [CO(2)] conditions. However, the percentage inhibition following ureide treatment was significantly greater under ambient [CO(2)] as compared with that under elevated [CO(2)]. Under ambient [CO(2)] conditions, ARA was less than that under elevated [CO(2)] 1 d after ureide treatment. Under ambient [CO(2)], the application of ureide resulted in a significant accumulation of ureide in all plant tissues, with the highest concentration increases in the leaves. However, application of exogenous ureide to plants subjected to elevated [CO(2)] did not result in increased ureide concentration in any tissues. TNC concentrations were consistently higher under elevated [CO(2)] compared with those under ambient [CO(2)]. For both [CO(2)] treatments, the application of ureide induced a significant decrease of TNC concentrations in the leaves and nodules. For both leaves and nodules, a negative correlation was observed between TNC and ureide levels. Results indicate that product(s) of ureide catabolism rather than tissue ureide concentration itself are critical in the regulation of N(2)-fixation.     

Interactive Effects of Elevated CO2 and Ozone on Leaf Thermotolerance in Field-grown Glycine max

Humans are increasing atmospheric CO2, ground-level ozone (O3), and mean and acute high temperatures. Laboratory studies show that elevated CO2 can increase thermotolerance of photosynthesis in C3 plants. O3-related oxidative stress may offset benefits of elevated CO2 during heat-waves. We determined effects of elevated CO2 and O3 on leaf thermotolerance of field-grown Glycine max (soybean, C3). Photosynthetic electron transport (φet) was measured in attached leaves heated in situ and detached leaves heated under ambient CO2 and O3. Heating decreased φet, which O3 exacerbated. Elevated CO2 prevented O3-related decreases during heating, but only increased φet under ambient O3 in the field. Heating decreased chlorophyll and carotenoids, especially under elevated CO2. Neither CO2 nor O3 affected heat-shock proteins. Heating increased catalase (except in high O3) and CulZn-superoxide dismutase (SOD), but not MnSOD; CO2 and O3 decreased catalase but neither SOD. Soluble carbohydrates were unaffected by heating, but increased in elevated CO2. Thus, protection of photosynthesis during heat stress by elevated CO2 occurs in field-grown soybean under ambient O3, as in the lab, and high CO2 limits heat damage under elevated O3, but this protection is likely from decreased photorespiration and stomatal conductance rather than production of heat-stress adaptations.     

Effects of elevated CO2 and temperature on development and consumption rates of Octotoma championi and O. scabripennis feeding on Lantana camara

We carried out a factorial experiment to explore the effect of doubled CO₂ concentration and a 3 °C temperature increase on the development of a complete generation of the beetles Octotoma championi Baly and O. scabripennis Guérin‐Méneville (Coleoptera: Chrysomelidae). These species are biological control agents of Lantana camara L. (Verbenaceae), with a leaf‐mining larval phase and free‐living, leaf‐chewing adults. Plants grown at elevated CO₂ had enhanced above‐ground biomass, thicker leaves, reduced nitrogen concentration, and increased C:N ratios. Under the high temperature treatment, plants grown at ambient CO₂ suffered wilting and premature leaf loss, despite daily watering; this effect was ameliorated at elevated CO₂. The wilting of plants in the ambient CO₂/high temperature treatment reduced the emergence success of the beetles, particularly O. championi. Development time was accelerated by approximately 10–13 days at the higher temperature, but was not affected by CO₂. Neither CO₂ nor temperature affected adult beetle weight. Consumption rates of free‐living beetles were not affected by either CO₂ or temperature. By contrast, in the short‐term trials using excised foliage, beetles given no choice between ambient and elevated CO₂‐grown foliage, consumed more from ambient plants. When beetles were offered a choice between foliage grown at the two CO₂ levels, O. championi did not display a significant preference but O. scabripennis consumed more ambient CO₂‐grown foliage when feeding at the lower temperature. This study indicates that under future conditions of higher temperatures, amelioration of water stress in host plants growing in elevated CO₂ may benefit some endophagous insects by reducing premature leaf loss. Under some circumstances, this benefit may outweigh the deleterious effects of lower leaf nitrogen. Our results also indicate that foliage consumption under elevated CO₂ by mobile, adult insects on whole plants may not be significantly increased, as was previously indicated by short‐term experiments using excised foliage.     

Role of cysteine proteinase inhibitors in preference of Japanese beetles (<Emphasis Type="Italic">Popillia japonica</Emphasis>) for soybean (<Emphasis Type="Italic">Glycine max</Emphasis>) leaves of different ages and grown under elevated CO<Subscript>2</Subscript>

Elevated levels of CO₂, equivalent to those projected to occur under global climate change scenarios, increase the susceptibility of soybean foliage to herbivores by down-regulating the expression of genes related to the defense hormones jasmonic acid and ethylene; these in turn decrease the gene expression and activity of cysteine proteinase inhibitors (CystPIs), the principal antiherbivore defenses in foliage. To examine the effects of elevated CO₂ on the preference of Japanese beetle (JB; Popillia japonica) for leaves of different ages within the plant, soybeans were grown at the SoyFACE facility at the University of Illinois at Urbana-Champaign. When given a choice, JB consistently inflicted greater levels of damage on older leaves than on younger leaves, and there was a trend for a greater preference for young leaves grown under elevated CO₂ compared to those grown under ambient CO₂. More heavily damaged older leaves and those grown under elevated CO₂ had reduced CystPI activity, and JB that consumed leaves with lower CystPI activity had correspondingly greater gut proteinase activity. Younger leaves with higher CystPI activity and photosynthetic rates may contribute disproportionately to plant fitness and are more protected against herbivore attack than older foliage. Cysteine proteinase inhibitors are potent defenses against JB, and the effectiveness of this defense is modulated by growth under elevated CO₂ as well as leaf position.     

How does elevated CO2 or ozone affect the leaf-area index of soybean when applied independently?

Changes in leaf-area index (LAI) may alter ecosystem productivity in elevated [CO2] or [O3]. By increasing the apparent quantum yield of photosynthesis (phi(c,max)), elevated [CO2] may increase maximum LAI. However, [O3] when elevated independently accelerates senescence and may reduce LAI. Large plots (20 m diameter) of soybean (Glycine max) were exposed to ambient (approx. 370 micromol mol(-1)) or elevated (approx. 550 micromol mol(-1)) CO2 or 1.2 times ambient [O3] using soybean free-air concentration enrichment (SoyFACE). In 2001 elevated CO2 had no detectable effect on maximum LAI, but in 2002 maximum LAI increased by 10% relative to ambient air. Elevated [CO2] also increased the phi(c,max) of shade leaves in both years. Elevated [CO2] delayed LAI loss to senescence by approx. 54% and also increased leaf-area duration. Elevated [O3] accelerated senescence, reducing LAI by 40% near the end of the growing season. No effect of elevated [O3] on photosynthesis was detected. Elevated [CO2] or [O3] affected LAI primarily by altering the rate of senescence; knowledge of this may aid in optimizing future soybean productivity.     

水稻田稗草叶片光合作用对开放式空气CO2浓度增高(FACE)的适应

于分蘖、拔节和抽穗 3个时期在空气CO2 浓度 (380 μmol·mol-1)下测定稻田中稗草叶片的净光合速率 (Pn) ,发现在开放式CO2 浓度增高 (FACE)条件下生长的稗草叶片后 2个时期的Pn显著低于普通空气中生长的对照 ,比对照下降约 2 0 % ,说明FACE条件下稗草叶片光合作用对高CO2 浓度发生了明显的适应 .同时 ,叶片的气孔导度 (Gs)和胞间CO2 浓度 (Ci)的下降更为明显 .与对照相比 ,叶片可溶性蛋白含量明显降低 ,拔节期只有对照的 6 2 .4 % ;高CO2 浓度下生长的稗草叶片Rubisco含量也降低 ,分蘖期和拔节期分别为对照的 87%和 84 % ,但其差异未达到显著水平 .可以认为 ,长期生长在高CO2 浓度下的C4植物稗草叶片光合作用的适应是叶片气孔部分关闭和可溶性蛋白含量下降的结果 .     

水稻叶片光合作用对开放式空气CO2浓度增高(FACE)的响应与适应

利用便携式光合气体分析系统 (LI 6 4 0 0 ) ,比较测定了高CO2 浓度 (FACE ,free airCO2 enrich ment)和普通空气CO2 浓度下生长的水稻叶片的净光合速率、水分利用率、表观量子效率和RuBP羧化效率等光合参数 .在各自生长CO2 浓度 (380vs 5 80 μmol·mol-1)下测定时 ,高CO2 浓度 (5 80 μmol·mol-1)下生长的水稻叶片的净光合速率、碳同化的表观量子效率和水分利用率明显高于普通空气 (380 μmol·mol-1)下生长的水稻叶片 .但是 ,随着FACE处理时间的延长 ,高CO2 浓度对净光合速率的促进作用逐渐减小 .在相同CO2 浓度下测定时 ,FACE条件下生长的水稻叶片净光合速率和羧化效率明显比普通空气下生长的对照低 .尽管高CO2 浓度下生长的水稻叶片的气孔导度明显低于普通空气中生长的水稻叶片 ,但两者胞间CO2 浓度差异不显著 ,因此高CO2 浓度下生长的水稻叶片光合下调似乎不是由气孔导度降低造成的 .     

Growth under elevated CO(2) ameliorates defenses against photo-oxidative stress in poplar (Populus alba x tremula)

To test the hypothesis that growth-CO(2) concentrations affect stress susceptibility, leaves of poplar trees (Populus alba x tremula) grown under ambient or about twofold ambient CO(2) concentrations were subjected to chilling temperatures at high light intensities or were exposed to paraquat. Photosynthesis was less diminished and electrolyte leakage was lower in stressed leaves from poplar trees grown under elevated [CO(2)] as compared with those from ambient [CO(2)]. Severe stress caused pigment and protein degradation but to a lower extent in leaves from elevated as compared with those from ambient [CO(2)]. The protection was accompanied by rapid induction of superoxide dismutase activities (EC 1.15.1.1). Ascorbate and glutathione-related detoxification systems as well as catalase (EC 1.11.1.6) activities were less resistant than superoxide dismutases and declined in stress-exposed leaves from poplars grown under elevated [CO(2)] to a similar extent as in those from trees grown under ambient [CO(2)]. These results suggest that the CO(2)-mediated amelioration of stress was confined to SOD and limited since the destruction of H(2)O(2)-degrading systems was not prevented.     

二氧化碳和臭氧浓度升高对春小麦生长及次生代谢的影响

通过开顶式气室(OTCs)研究了OTC对照（自然CO₂浓度约342 μmol·mol^-1,O₃浓度约30 nmol·mol^-1）、高浓度CO₂（550 μmol·mol^-1）、高浓度O₃（浓度为80 nmol·mol^-1）及其交互作用（CO₂ 550 μmol ·mol^-1,O₃ 80 nmol·mol^-1）对春小麦不同发育时期生物量、总酚量、黄酮含量及成熟期产量性状的影响.结果表明：CO₂浓度增加条件下,春小麦生物量和产量性状都显著高于OTC对照（P<0.05）;而O₃浓度升高条件下,小麦生物量降低,株高、穗长、穗粒质量及千粒重也显著低于对照;CO₂和O₃交互作用下各项指标处于二者之间.说明CO₂可以缓解O₃对小麦的负效应,而O₃对CO₂的正效应具有削弱作用,但二者的作用并非简单的叠加.CO₂、O₃浓度增加及其交互作用显著增加了春小麦叶片中的总酚含量,其中两者交互作用的效应更大,但在小麦生长后期,总酚含量增加量比对照有所降低.在小麦生长前期,各处理总黄酮含量均低于对照;而在成熟期,各处理都显著高于对照.     

CO2浓度升高对长白山三种树木幼苗叶碳水化合物和氮含量的影响

研究了持续 3个生长季高浓度CO2 处理的长白山主要树种红松、长白赤松和水曲柳幼苗叶 (片 )的蔗糖、果糖、可溶性总糖、淀粉及全氮含量变化 .结果表明 ,前两个生长季 70 0 μmol·mol-1CO2 处理 ,促进了水曲柳和长白赤松幼苗的淀粉累积 ,70 0和 5 0 0 μmol·mol-1CO2 却使红松幼苗的全N含量明显降低 .第三个生长季高浓度CO2 处理的第一周和第二周 ,红松和水曲柳幼苗的淀粉含量增加 ,全N含量降低 ,第八周时 ,水曲柳仍维持原来的变化趋势 .第三个生长季高浓度CO2 处理 ,并未使长白赤松的C、N含量有明显的增减 .CO2 浓度影响了碳水化合物在叶 (片 )中的积累形式     

The effects of ethylene, depressed oxygen and elevated carbon dioxide on antioxidant profiles of senescing spinach leaves

It has been suggested that antioxidants play a role in regulating or modulating senescence dynamics of plant tissues. Ethylene has been shown to promote early plant senescence while controlled atmospheres (CA; reduced O2 levels and elevated CO2 levels) can delay its onset and/or severity. In order to examine the possible importance of various antioxidants in the regulation of senescence, detached spinach (Spinacia oleracea L.) leaves were stored for 35 d at 10 degrees C in one of three different atmospheres: (1) ambient air (0.3% CO2, 21.5% O2, 78.5% N2), (2) ambient air + 10 ppm ethylene to promote senescence, or (3) CA (10% CO2, 0.8% O2 and 89.2% N2) to delay senescence. At weekly intervals, material was assessed for activities of the antioxidant enzymes ascorbate peroxidase (ASPX; EC 1.11.1.11), catalase (CAT; EC 1.11.1.6), dehydroascorbate reductase (DHAR; EC 1.8.5.4), glutathione reductase (GR; EC 1.6.4.2), monodehydroascorbate reductase (MDHAR; EC 1.6.5.4), and superoxide dismutase (SOD; EC 1.15.1.1), and concentrations of the water-soluble antioxidant compounds ascorbate and glutathione. Indicators of the rate and severity of senescence (lipid peroxidation, chlorophyll, and soluble protein levels) were also determined. Results indicated that the rate and severity of senescence was similar between the leaves stored in ambient air or CA until day 35, at which point the ambient air-stored leaves exhibited a sharp increase in lipid peroxidation. Tissues under both storage regimes demonstrated significant declines only in levels of ASPX, CAT, and ascorbate. Glutathione content in the CA-stored tissue also significantly dropped, but only on day 35. In contrast, spinach leaves stored in ambient air + ethylene experienced a rapid decrease in levels of all the antioxidants assessed except SOD. Declines in levels of ASPX, CAT, and ascorbate over the 35 d storage period regardless of the composition of the storage atmosphere suggests that regulation of H2O2 levels plays an important role in both the dynamics and severity of post-harvest senescence of spinach.     

大气CO2浓度倍增对植物暗呼吸的影响

以长期生长于350和700μmolCO_2·mol~(-1)空气的开顶式培养室的杜仲(Eucommia ulmoides Oliv.)、紫花苜蓿(Medicago sativa L.)、玉米(Zea mays L.)等10种植物的离体成熟叶片或整株为材料,研究不同测定温度(15～35℃)下,CO_2浓度倍增对植物暗呼吸的影响。结果表明:在较低温度(15℃、20℃)下,CO_2浓度倍增对植物暗呼吸没有显著效应,在较高温度(30℃、35℃)下多数被测植物的暗呼吸显著增强。讨论了实验所得结果在未来全球气候变化中的可能的意义。     

大气CO2浓度升高对香蕉光合作用及光合碳循环过程中叶氮分配的影响

生长在高CC2浓度(700±56μl     

Responses of Plant Dark Respiration to Doubled CO2 Concentration

Ten species of plants were grown at ambient (350μmol CO2·mol-1 air) and doubled (700 μmol CO2·mol-1 air) CO2 concentrations at ambient temperature and illumination in order to examine changes of dark respiration of whole seedlings or detached leaves. Effects of CO2 on dark respiration were determined by brief exposure ( ≤ 5 min) to corresponding CO2 concentration and temperatures ( 15,20,25,30 and 35 ℃ ) with infrared CO2 analyzer. The reductions in dark respiration on a weight base for leaves of East-Liaoning oak (Quercus liaotungensis Koidz. ) at 15,20 and 25 ℃ and of soybean ( Glycine max L. ) at 20,25,30 and 35 ℃ and for whole seedlings of three- tcoloured amaranth (Amaranthus tricolor L. ) at 15 and 20 ℃ and cucumber ( Cucumis sativus L. ) at 15 cE measured at elevated concentration relative to the ambient CO2 concentration were observed. No significant difference in respiration responded was observed to elevated or ambient CO2 concentrations at 15 ℃ in maize (Zea mays L. ) seedlings and alfalfa (Medicago sativa L. ) leaves, at 35 ℃ in East-Liaoning oak leaves and at 20,25 and 30 ℃ in three-coloured amaranth seedlings. However CO2 efflux in leaves of weeping willow (Salix babylonica L. ), simon poplar (Populus simonii Carr. ) and eucommia (Eucommia ulmoides Oliv. ) at 15,20,25,30 and 35 ℃, alfalfa at 20,25,30 and 35 ℃, East-Liaoning oak at 30 ℃, maize at 15 ℃, seedlings of common buckwheat (Fagotrytum esculentum Moench) at 15,20,25,30 and 35 ℃, cucumber and maize at 20,25,30 and 35 ℃ and three-coloured amaranth at 35 ℃ showed an increase at elevated in contrast to ambient CO2 concentration. In general, at lower temperatures (i. e. 15, 20 ℃ ) there was no significant difference between elevated and ambient CO2 concentration for dark respiration, while at higher temperatures (i. e. 30,35 ℃ ) elevated CO2 concentration positively stimulate clark respiretion. It has not yet been described that double CO2 concentration could enhance plant dark respiration at 30 and 35 ℃. Impacts of the characteristics in dark respiration on the future changes of vegetation and its mechanism were discussed.