北亚热带—暖温带过渡区典型森林生态系统土壤呼吸特征研究
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摘要
为揭示北亚热带-暖温带过渡区典型森林生态系统的土壤总呼吸及各组分呼吸的变异与碳排放特征，促进对森林生态系统碳输出过程的深入了解。本研究针对目前土壤呼吸研究中的热点与难点问题，以宝天曼森林生态观测站为依托，以土壤总呼吸组分分离为突破口，应用国内外认可的呼吸速率测定及分离方法对分布面积占该区域森林总面积95％以上的5个典型森林生态系统的土壤总呼吸(R_T)、土壤呼吸(R_S)、根呼吸(R_A)、土壤异养呼吸(R_H)及凋落物呼吸(R_L)在5个森林类型土壤温度、土壤含水量及凋落物含水量变幅分别为26.60～26.94℃、0.19～0.29 kg·kg~(-1)与1.85～1.96 kg·kg~(-1)条件下的变化过程及相关的环境、生物因子进行了为期一年的观测，并设置了土壤温湿度室内培养与呼吸速率测定试验，对更大温度范围内锐齿栎幼林的土壤呼吸特征进行了研究，经统计分析与模型模拟，结果表明：
(1)该区域森林生态系统土壤总呼吸与各组分呼吸有3种昼夜变化模式，第一种发生在春季阔叶林展叶前与秋季阔叶林落叶后，土壤总呼吸及各组分呼吸的变化均与土壤温度一致；第二种仅发生在夏季的幼林中，R_T、R_S与R_A与土壤温湿度的变化过程有短暂的解偶联现象，而R_H与R_L的变化与第一种相同：第三种模式发生在夏秋季的生长季中，仅有T_A与土壤温湿度有短暂的解偶联，而其它呼吸类型均与土壤温度的变化一致。R_T、R_S、R_H、R_A与R_L的最低值均出现在06：00，最高值出现在15：00～21：00。虽然土壤总呼吸与各呼吸组分在不同季节最低与最大值出现的时间存在差异，但昼夜变化均为单峰曲线。R_A与土壤温湿度的解偶联是否导致R_T与R_S与土壤温湿度解偶联与R_A在不同季节、不同林分中的绝对与相对大小有关；各呼吸在夏季的昼夜变幅＞秋季＞春季＞冬季，与水热变幅相同：随林龄增加，R_A与R_L昼夜变幅的季节性变异系数减小，而R_H的增大；R_T、R_S、R_H及R_A的季节性变化趋势均为单峰曲线，最小与最大值分别出现在1月与7月，总体上与土壤温度一致，含水量较低时对呼吸有明显制约，凋落物呼吸的季节性变化受其含水量的影响较大。
(2)温度与水分对呼吸影响的相对重要性存在季节性变化，土壤含水量与凋落物含水量分别低于0.20 kg·kg~(-1)与0.38 kg·kg~(-1)，而相对应地温与气温分别高于15℃与13℃的季节，含水量对呼吸有明显的制约作用；当土壤温度与气温分别低于15℃与13℃的季节，温度对呼吸的影响高于含水量：其它温湿度条件下，呼吸同时受到二者的明显影响：室内培养发现，锐齿栎幼林的土壤呼吸速率对土壤升温表现为正反馈，但土壤含水量过高(在0.35kg·kg~(-1)以上)时对二者的正反馈关系具有抑制作用。
(3)5个林分的土壤温度与R_T、R_S，R_A及R_H均呈显著(P＜0.05)或极显著(P＜0.01)的指数函数关系，可以分别解释上述呼吸昼夜与季节变化的80.62％～93.85％、86.84％～92.23％、79.05％～88.12％与82.63％～96.70％：昼夜与季节尺度上，土壤温度与凋落物呼吸分别呈显著的指数函数与二次函数关系，对其呼吸的昼夜与季节性变化的解释能力分别为68.10％～81.58％与34.78％～54.24％。各呼吸与含水量在昼夜与季节尺度上分别呈负相关与正相关。季节变化中，5种林分的R_T与其它4种林分(除锐齿栎老林外)的R_S、R_A及R_H与土壤含水量间有显著的一元一次函数关系，可分别解释上述呼吸季节性变化的24.8％～55.2％、23.8％～47.1％、20.45％～41.14％与23.06％～48.45，但室内培养试验发现二次函数对二者关系的解释能力更高；季节性变化中，5种林分凋落物含水量与凋落物呼吸具有显著的一元一次函数关系，能解释凋落物呼吸季节性变化的72.24％～79.50％。室内培养试验发现，只是在含水量较高或较低时，含水量的增加才对土壤呼吸速率有明显的抑制或促进作用。
(4)全年而言，5个林分的R_T、R_S、R_H的Q_(10)值分别为2.18～2.41(平均2.30)、2.30～2.44(平均为2.38)、2.09～2.35(平均为2.19)与2.49～2.82(平均为2.61)，上述各呼吸的温度敏感性指数间有显著差异(P＜0.05)。除根呼吸外，其它呼吸在冬季的Q_(10)值均＞春季＞秋季＞夏季，根呼吸的Q_(10)值在春季最高，其次为秋季，夏冬季较低。在土壤温度没有显著差异(P＞0.05)且含水量对土壤总呼吸没有明显抑制时，幼林土壤总呼吸的温度敏感性高于老林。昼夜与季节性变化中，每个林分土壤总呼吸的Q_(10)值均始终低于土壤呼吸。多数情况下，根呼吸的Q_(10)值＞土壤异养呼吸＞凋落物呼吸，但冬季时土壤异养呼吸的Q_(10)值＞凋落物呼吸＞根呼吸，且不同呼吸类型Q_(10)的相对大小在不同林分中也不相同。室内培养试验发现，温度对Q_(10)的影响较大，含水量的影响很小，含水量过高对Q_(10)有抑制作用。
(5)若考虑断根样方内细根分解对根呼吸与异养呼吸碳排放量估算的影响，锐齿栎幼林、锐齿栎老林、针-阔混交林、阔-阔混交林与栓皮栎林根呼吸的碳排放量应分别为724.52 gC·m~(-2)·a~(-1)、373.55 gC·m~(-2)·a~(-1)、441.56～441.87 gC·m~(-2)·a~(-1)、427.14 gC·m~(-2)·a~(-1)与484.43C·m~(-2)·a~(-1)，异养呼吸应分别为380.63 gC·m~(-2)·a~(-1)、405.57 gC·m~(-2)·a~(-1)、379.36～379.67 gC·m~(-2)·a~(-1)、485.05 gC·m~(-2)·a~(-1)与415.07 gC·m~(-2)·a~(-1)；凋落物呼吸的分别为：239.04 gC·m~(-2)·a~(-1)、119.79gC·m~(-2)·a~(-1)、218.60 gC·m~(-2)·a~(-1)、115.03 gC·m~(-2)·a~(-1)与277.29 gC·m~(-2)·a~(-1)；新鲜凋落物的分别为：86.59 gC·m~(-2)·a~(-1)、56.72 gC·m~(-2)·a~(-1)、65.92 gC·m~(-2)·a~(-1)、55.65 gC·m~(-2)·a~(-1)与98.84 gC·m~(-2)·a~(-1)；土壤呼吸的分别为：1105.15 gC·m~(-2)·a~(-1)、779.12 gC·m~(-2)·a~(-1)、821.23 gC·m~(-2)·a~(-1)、912.19gC·m~(-2)·a~(-1)和899.50 gC·m~(-2)·a~(-1)；总呼吸的分别为1345.19 gC·m~(-2)·a~(-1)、897.23 gC·m~(-2)·a~(-1)、1039.83 gC·m~(-2)·a~(-1)、1027.22 gC·m~(-2)·a~(-1)与1176.79 gC·m~(-2)·a~(-1)；每种林分中R_T、R_S、R_H和R_A的排放量均在1月最低，7月最高，变化趋势为单峰曲线；5种林分的R_S、R_H及R_A在夏秋两季的总排放量分别占全年的72.93％～76.55％、54.03％～59.50％与74.25％-82.21％。5种林分R_L凋落物呼吸碳排放量在夏季最高，其次为秋季、春季与冬季，夏秋两季排放量占全年的78.45％～83.67％。
(6)若考虑断根样方内细根分解对R_A与R_H贡献率估算的影响，林分1～林分5R_A的贡献率应依次分别为53.86％、41.63％、42.46％～42.49％、41.58％与41.17％；R_H的贡献率应依次分别为28.57％、45.21％、36.48％～36.51％、47.13％与35.27％；R_L的分别为17.77％、13.35％、21.01％、12.61％与23.56％；幼林、针阔混交林与栓皮栎林中根呼吸的年贡献率＞异养呼吸＞凋落物呼吸，而在其它2个林分中异养呼吸的年贡献率＞根呼吸＞凋落物呼吸，R_A贡献率的最小与最大值分别出现在15∶00以前与18∶00～21∶00，R_H的分别出现在18∶00～21∶00与15∶00以前，R_L的分别出现在15∶00～18∶00与12∶00之前；夏季根呼吸贡献率＞春季＞秋季＞冬季，R_H的贡献率在冬季最大，其次为春季，夏秋季较小；R_L贡献率在秋季最高，其次为夏季、春季与冬季，影响凡贡献率季节性变化的主导因子是凋落物含水量，其次为凋落物现存量，林分类型间凋落物现存量的差异解释了林分间R_L贡献率差异的62.60％～71.42％。
(7) 5个典型森林生态系统间的R_T、R_S及R_H均无显著差异(P＞0.05)，而林分间的R_A与R_L存在显著差异(P＜0.05)。在林分间温度、凋落物含水量没有显著差异(P＞0.05)，土壤含水量虽有极显著差异(P＜0.01)，但是解释能力很低的情况下，林分间活细根生物量的差异解释了林分间R_A差异的94.71％，凋落物现存量的差异解释了5种凋落物性质差异较大的林分类型间R_L差异的57.83％，解释了凋落物性质相似的4种阔叶林间R_L差异的99.9％。
In order to ascertain the carbon emission and variation characteristics of total soilrespiration and its components in typical forest ecosystems at the transitional area from thenorthern subtropics to warm temperate, and to understand their carbon output process clearly,the variation process of total soil respiration and its components and relative environmental andbiological factors under the condition that the variation range of soil temperature, soil watercontent and litter water content is 26.60～26.94℃, 0.19～4).29 kg·kg~(-1) and 1.85～1.96 kg·kg~(-1)respectively are simultaneously measured over the whole year in the 5 typical forestecosystems which account for 95% total forest area in this region with the support of BaoTianman forest ecosystem station and by using the approbatory methods of respirationmeasurement and respiration component partition. At the same time,the cultivation experimentof soil temperature and soil water content of Quercus aliena var.acuteserrata young stand wasset to understand the soil respiration characteristic at the larger scope of soil temperature. Thestatistical and simulated results show that:
(1) There are three diel variation patterns of total soil respiration and its components,the first which occurs before foliage spread in spring and after defoliation in fall is that thevariations of total soil respiration and its components are all consistent with soil temperature;the second which only occurs in summer in young-growth forest is that the variation of R_T, R_Sand R_A have short-term decoupling with soil temperature and soil water content, but thevariation of R_H and R_L is same as soil temperature; the third which occurs in growth seasons ofsummer and fall is that only the variation of R_A has short-term decoupling with soil temperatureand soil water content, the variations of the other respiration are all consistent with soiltemperature. The smallest and the biggest R_T, R_S, R_H ,R_A and R_L is at 06:00 and at 15:00～21:00respectively. The diel variation patterns for total soil respiration and its components are allsingle modal curve. Whether the decoupling of R_A with soil temperature and soil water contentcan lead to the decoupling of R_T and R_S with soil temperature and soil water content isdetermined by the absolute and relatively amount of root respiration in different stands andseasons.The diel variation range of respiration is highest in summer, followed by in spring and fall, and lowest in winter, its seasonal trend is same as that of water content and temperature;The seasonal variation coefficients of diel range of root respiration and litter respiration aredecreasing with stand age increasing, but that of heterotrophic is increasing with stand agesincreasing; The seasonal variation in total soil respiration, soil respiration, root respiration andheterotrophic respiration is single modal curve, which is same as that of soil temperature, thelargest and lowest respiration is in January and July respectively, but the respiration is clearlyrestricted by lower water content; The seasonal variation in litter respiration is more affectedby litter water content than by temperature.
(2) The relative importance of the effects of water and temperature on respiration hasseasonal variation, The respiration rate is largely controlled by temperature when soiltemperature and air temperature is below 15℃and 13℃respectively, while it is clearlylimited by water content when soil temperature is above 15℃and air temperature is above13℃respectively and soil water content and litter water content is below 0.20 kg·kg~(-1) and 0.38kg·kg~(-1) respectively. However, it is simultaneously affected by both temperature and watercontent under other condition. The soil water and temperature cultivation experiment indicatedthat the soil respiration is positively feedback with soil temperature increase, but the positivelyfeedback is restricted by water content which is over 0.35kg·kg~(-1) in Quercus alienavar.acuteserrata young stand.
(3) The relationship of soil temperature with total respiration, Soil respiration, rootrespiration and soil heterotrophie respiration is significant exponent function (P＜0.05) , soiltemperature can account for 80.62%～93.85%, 86.84%～92.23%, 79.05％～88.12% and82.63%～96.70% of diel and seasonal variation in above mentioned respiration respectively.The relationship of soil temperature with litter respiration is exponent function at diel scale butis quadratic equation at seasonal scale, soil temperature can account for 68.10%～81.58% ofdiel variation and 34.78%～54.24% seasonal variation in litter respiration. Respiration ispositively and negatively with water content at die and seasonal scale respectively, the simpleequation between soil water content and respiration indicates that soil water content canaccount for 24.8%～55.2% of seasonal variation in total soil respiration for 5 stands andaccount for 23.8%～47.1%, 20.45%～41.14% and 23.06%～48.45 of seasonal variation in soilrespiration, root respiration and soil heterotrophic respiration respectively for 4 stands (except for Quercus aliena var.acuteserrata old stand), but the cultivation experiment indicates therelationship between soil water content and respiration is quadratic equation. The relationshipbetween litter water content with litter respiration is significant single equation for 5 stands andlitter water content can account for 72.24%～79.5% of seasonal variation in litter respiration;The cultivation experiment indicates that soil water content increase can promote the soilrespiration when soil water content is very low and that soil water content increase can restrainsoil respiration when soil water is very high.
(4) The Q_(10) value of total respiration, soil respiration, soil heterotrophie respiration androot respiration is 2.18～2.41 (mean is 2.30) , 2.30～2.44(mean is 2.38) , 2.09～2.35 (mean is2.19)与2.49～2.82 (mean is 2.61) respectively at the annual scale, the Q_(10)value of abovementioned respiration is significant (P＜0.05) ; The Q_(10)value of the other respiration typesexcept for root respiration is highest in winter, followed by in spring and fall, and lowest insummer, but the Q_(10) value of root respiration is highest in spring, followed by in fall, andlowest in summer and winter; The higher Q_(10) value for total soil respiration in Quercus alienavar.acuteserrata young stand is than old stand under the condition that there is no significantdifference of the soil temperature between the 2 stands and that ther is no water contentlimitation; The Q_(10) value of root respiration is highest, followed by that of soil heterotrophicrespiration, that of litter respiration is lowest in many occasions, but the Q_(10) value of soilheterotrophie respiration is highest, followed by that of litter respiration, that of rootrespiration is lowest in winter; the Q_(10) value of total soil respiration is higer than that of soilrespiration regardless of diel and seasonal scale in 5 stands ; The order of Q_(10) values ofdifferent respiration in different stand has differences.The cultivation experiment indicates thatthe effects of soil temperature on Q_(10) value ,of soil respiration is very big ,but the effects of soilwater content on Q_(10) value of soil respiration is very small, the Q_(10) value of soil respiration isrestrained by very high soil water content too.
(5) The carbon release of root respiration in Quercus aliena var.acuteserrata youngstand, Quercus aliena var.acuteserrata old stand, broadleaf/coniferous mixed stand, broadleafmixed forest and Quercus variabilis stand is 724.52 gC·m~(-2)·a~(-1), 373.55 gC·m~(-2)·a~(-1),441.56～441.87 gC·m~(-2)·a~(-1), 427.14 gC·m~(-2)·a~(-1) and 484.43 C·m~(-2)·a~(-1) respectively; Their carbonrelease of soil heterotrophic respiration is 380.63 gC·m~(-2)·a~(-1), 405.57 gC·m~(-2)·a~(-1), 379.36～379.67 gC·m~(-2)·a~(-1), 485.05 gC·m~(-2)·a~(-1) and 415.07 gC·m~(-2)·a~(-1) respectively if taking account of fine rootdecomposition in root-excluded quadrates; Their carbon release of litter respiration is 239.04gC·m~(-2)·a~(-1), 119.79 gC·m~(-2)·a~(-1), 218.60 gC·m~(-2)·a~(-1), 115.03 gC·m~(-2)·a~(-1) and 277.29 gC·m~(-2)·a~(-1)respectively; Their carbon release of fresh litter respiration is 86.59 gC·m~(-2)·a~(-1), 56.72 gC·m~(-2)·a~(-1),65.92 gC·m~(-2)·a~(-1), 55.65 gC·m~(-2)·a~(-1) and 98.84 gC·m~(-2)·a~(-1) respectively; Their carbon release ofsoil respiration is 1105.15 gC·m~(-2)·a~(-1), 779.12 gC·m~(-2)·a~(-1), 821.23 gC·m~(-2)·a~(-1), 912.19 gC·m~(-2)·a~(-1)and 899.50 gC·m~(-2)·a~(-1) respectively; Their carbon release of total soil respiration is 1345.19gC·m~(-2)·a~(-1), 897.23 gC·m~(-2)·a~(-1), 1039.83 gC·m~(-2)·a~(-1), 1027.22 gC·m~(-2)·a~(-1)and 1176.79 gC·m~(-2)·a~(-1).The seasonal variation trend of soil respiration , soil heterotrophie respiration and rootrespiration is single modal curve, their smallest and biggest value is in January and July, Thepercentage of the carbon release of soil respiration, soil heterotrophic respiration and rootrespiration in summer and fall is 72.93%～76.55%, 54.03%～59.50% and 74.25%-82.21% ofannual release respectively for 5 stands; The carbon release of litter respiration is highest insummer, followed by in fall, lowest in spring and winter, the percentage of carbon release insummer and fall is 78.45%～83.67% of its annual release respectively for 5 stands
(6) The annual contribution rate of root respiration for stand 1～stand 5 is 53.86%,41.63%, 42.46%～42.49%, 41.58% and 41.17% respectively; Their annual contribution rate ofsoil heterotrophie respiration is 28.57%, 45.21%, 36.48%～36.51%, 47.13% and 35.27%respectively if taking account of fine root decomposition in root-excluded quadrates; Thecontribution rate of litter respiration is 17.77%, 13.35%, 21.01%, 12.61% and 23.56%; Thecontribution rate of root respiration is biggest, followed by soil heterotrophic respiration andthat of litter respiration is the lowest for Quercus aliena var.acuteserrata young stand,broadleaf/coniferous mixed stand and Quercus variabilis stand; But the contribution rate of soilheterotrophie respiration is biggest, followed by root respiration and that of litter respiration isthe lowest for the other two stands; The biggest and smallest contribution rate of rootrespiration is before 15:00 and at 18:00～21:00 respectively, the biggest and smallestcontribution rate of soil heterotrophic respiration is at 18:00～21:00 and before 15:00respectively and the biggest and smallest contribution rate of litter respiration is at 15:00～18:00and beforel2:00; The contribution rate of root respiration in summer is the biggest, followedby in spring and fall, lowest in winter, that of soil heterotrophic in winter is the biggest, followed by in spring and lowest in summer and fall, that of litter respiration in fall is biggestfollowed by summer, and lowest in spring and winter; The most important dominant factorscontrolling the seasonal variation in contribution rate of litter respiration is litter water content,the second factor is existing litter amount, which can accounts for 62.60%～71.42% of thedifference of contribution rate of litter respiration among the five stands.
(7) There is no significant differences in total soil respiration rate, soil respiration rateand the heterotrophic respiration rate among the five stands (P＞0.05), but the significantdifferences occur in the autotrophic respiration rate and litter respiration rote (P＜0.05). The livefine root biomass accounts for 94.71%of the difference of autotrophie respiration rates amongthe five stands, the existing litter amount accounts for 57.83% of the difference of litterrespiration among 5 stands in which the litter quality has obvious difference, but it canaccounts for 99.9% of the difference of litter respiration among 4 broadleaf stands in whichthe litter quality has no obvious difference.
(1)该区域森林生态系统土壤总呼吸与各组分呼吸有3种昼夜变化模式，第一种发生在春季阔叶林展叶前与秋季阔叶林落叶后，土壤总呼吸及各组分呼吸的变化均与土壤温度一致；第二种仅发生在夏季的幼林中，R_T、R_S与R_A与土壤温湿度的变化过程有短暂的解偶联现象，而R_H与R_L的变化与第一种相同：第三种模式发生在夏秋季的生长季中，仅有T_A与土壤温湿度有短暂的解偶联，而其它呼吸类型均与土壤温度的变化一致。R_T、R_S、R_H、R_A与R_L的最低值均出现在06：00，最高值出现在15：00～21：00。虽然土壤总呼吸与各呼吸组分在不同季节最低与最大值出现的时间存在差异，但昼夜变化均为单峰曲线。R_A与土壤温湿度的解偶联是否导致R_T与R_S与土壤温湿度解偶联与R_A在不同季节、不同林分中的绝对与相对大小有关；各呼吸在夏季的昼夜变幅＞秋季＞春季＞冬季，与水热变幅相同：随林龄增加，R_A与R_L昼夜变幅的季节性变异系数减小，而R_H的增大；R_T、R_S、R_H及R_A的季节性变化趋势均为单峰曲线，最小与最大值分别出现在1月与7月，总体上与土壤温度一致，含水量较低时对呼吸有明显制约，凋落物呼吸的季节性变化受其含水量的影响较大。
(2)温度与水分对呼吸影响的相对重要性存在季节性变化，土壤含水量与凋落物含水量分别低于0.20 kg·kg~(-1)与0.38 kg·kg~(-1)，而相对应地温与气温分别高于15℃与13℃的季节，含水量对呼吸有明显的制约作用；当土壤温度与气温分别低于15℃与13℃的季节，温度对呼吸的影响高于含水量：其它温湿度条件下，呼吸同时受到二者的明显影响：室内培养发现，锐齿栎幼林的土壤呼吸速率对土壤升温表现为正反馈，但土壤含水量过高(在0.35kg·kg~(-1)以上)时对二者的正反馈关系具有抑制作用。
(3)5个林分的土壤温度与R_T、R_S，R_A及R_H均呈显著(P＜0.05)或极显著(P＜0.01)的指数函数关系，可以分别解释上述呼吸昼夜与季节变化的80.62％～93.85％、86.84％～92.23％、79.05％～88.12％与82.63％～96.70％：昼夜与季节尺度上，土壤温度与凋落物呼吸分别呈显著的指数函数与二次函数关系，对其呼吸的昼夜与季节性变化的解释能力分别为68.10％～81.58％与34.78％～54.24％。各呼吸与含水量在昼夜与季节尺度上分别呈负相关与正相关。季节变化中，5种林分的R_T与其它4种林分(除锐齿栎老林外)的R_S、R_A及R_H与土壤含水量间有显著的一元一次函数关系，可分别解释上述呼吸季节性变化的24.8％～55.2％、23.8％～47.1％、20.45％～41.14％与23.06％～48.45，但室内培养试验发现二次函数对二者关系的解释能力更高；季节性变化中，5种林分凋落物含水量与凋落物呼吸具有显著的一元一次函数关系，能解释凋落物呼吸季节性变化的72.24％～79.50％。室内培养试验发现，只是在含水量较高或较低时，含水量的增加才对土壤呼吸速率有明显的抑制或促进作用。
(4)全年而言，5个林分的R_T、R_S、R_H的Q_(10)值分别为2.18～2.41(平均2.30)、2.30～2.44(平均为2.38)、2.09～2.35(平均为2.19)与2.49～2.82(平均为2.61)，上述各呼吸的温度敏感性指数间有显著差异(P＜0.05)。除根呼吸外，其它呼吸在冬季的Q_(10)值均＞春季＞秋季＞夏季，根呼吸的Q_(10)值在春季最高，其次为秋季，夏冬季较低。在土壤温度没有显著差异(P＞0.05)且含水量对土壤总呼吸没有明显抑制时，幼林土壤总呼吸的温度敏感性高于老林。昼夜与季节性变化中，每个林分土壤总呼吸的Q_(10)值均始终低于土壤呼吸。多数情况下，根呼吸的Q_(10)值＞土壤异养呼吸＞凋落物呼吸，但冬季时土壤异养呼吸的Q_(10)值＞凋落物呼吸＞根呼吸，且不同呼吸类型Q_(10)的相对大小在不同林分中也不相同。室内培养试验发现，温度对Q_(10)的影响较大，含水量的影响很小，含水量过高对Q_(10)有抑制作用。
(5)若考虑断根样方内细根分解对根呼吸与异养呼吸碳排放量估算的影响，锐齿栎幼林、锐齿栎老林、针-阔混交林、阔-阔混交林与栓皮栎林根呼吸的碳排放量应分别为724.52 gC·m~(-2)·a~(-1)、373.55 gC·m~(-2)·a~(-1)、441.56～441.87 gC·m~(-2)·a~(-1)、427.14 gC·m~(-2)·a~(-1)与484.43C·m~(-2)·a~(-1)，异养呼吸应分别为380.63 gC·m~(-2)·a~(-1)、405.57 gC·m~(-2)·a~(-1)、379.36～379.67 gC·m~(-2)·a~(-1)、485.05 gC·m~(-2)·a~(-1)与415.07 gC·m~(-2)·a~(-1)；凋落物呼吸的分别为：239.04 gC·m~(-2)·a~(-1)、119.79gC·m~(-2)·a~(-1)、218.60 gC·m~(-2)·a~(-1)、115.03 gC·m~(-2)·a~(-1)与277.29 gC·m~(-2)·a~(-1)；新鲜凋落物的分别为：86.59 gC·m~(-2)·a~(-1)、56.72 gC·m~(-2)·a~(-1)、65.92 gC·m~(-2)·a~(-1)、55.65 gC·m~(-2)·a~(-1)与98.84 gC·m~(-2)·a~(-1)；土壤呼吸的分别为：1105.15 gC·m~(-2)·a~(-1)、779.12 gC·m~(-2)·a~(-1)、821.23 gC·m~(-2)·a~(-1)、912.19gC·m~(-2)·a~(-1)和899.50 gC·m~(-2)·a~(-1)；总呼吸的分别为1345.19 gC·m~(-2)·a~(-1)、897.23 gC·m~(-2)·a~(-1)、1039.83 gC·m~(-2)·a~(-1)、1027.22 gC·m~(-2)·a~(-1)与1176.79 gC·m~(-2)·a~(-1)；每种林分中R_T、R_S、R_H和R_A的排放量均在1月最低，7月最高，变化趋势为单峰曲线；5种林分的R_S、R_H及R_A在夏秋两季的总排放量分别占全年的72.93％～76.55％、54.03％～59.50％与74.25％-82.21％。5种林分R_L凋落物呼吸碳排放量在夏季最高，其次为秋季、春季与冬季，夏秋两季排放量占全年的78.45％～83.67％。
(6)若考虑断根样方内细根分解对R_A与R_H贡献率估算的影响，林分1～林分5R_A的贡献率应依次分别为53.86％、41.63％、42.46％～42.49％、41.58％与41.17％；R_H的贡献率应依次分别为28.57％、45.21％、36.48％～36.51％、47.13％与35.27％；R_L的分别为17.77％、13.35％、21.01％、12.61％与23.56％；幼林、针阔混交林与栓皮栎林中根呼吸的年贡献率＞异养呼吸＞凋落物呼吸，而在其它2个林分中异养呼吸的年贡献率＞根呼吸＞凋落物呼吸，R_A贡献率的最小与最大值分别出现在15∶00以前与18∶00～21∶00，R_H的分别出现在18∶00～21∶00与15∶00以前，R_L的分别出现在15∶00～18∶00与12∶00之前；夏季根呼吸贡献率＞春季＞秋季＞冬季，R_H的贡献率在冬季最大，其次为春季，夏秋季较小；R_L贡献率在秋季最高，其次为夏季、春季与冬季，影响凡贡献率季节性变化的主导因子是凋落物含水量，其次为凋落物现存量，林分类型间凋落物现存量的差异解释了林分间R_L贡献率差异的62.60％～71.42％。
(7) 5个典型森林生态系统间的R_T、R_S及R_H均无显著差异(P＞0.05)，而林分间的R_A与R_L存在显著差异(P＜0.05)。在林分间温度、凋落物含水量没有显著差异(P＞0.05)，土壤含水量虽有极显著差异(P＜0.01)，但是解释能力很低的情况下，林分间活细根生物量的差异解释了林分间R_A差异的94.71％，凋落物现存量的差异解释了5种凋落物性质差异较大的林分类型间R_L差异的57.83％，解释了凋落物性质相似的4种阔叶林间R_L差异的99.9％。
In order to ascertain the carbon emission and variation characteristics of total soilrespiration and its components in typical forest ecosystems at the transitional area from thenorthern subtropics to warm temperate, and to understand their carbon output process clearly,the variation process of total soil respiration and its components and relative environmental andbiological factors under the condition that the variation range of soil temperature, soil watercontent and litter water content is 26.60～26.94℃, 0.19～4).29 kg·kg~(-1) and 1.85～1.96 kg·kg~(-1)respectively are simultaneously measured over the whole year in the 5 typical forestecosystems which account for 95% total forest area in this region with the support of BaoTianman forest ecosystem station and by using the approbatory methods of respirationmeasurement and respiration component partition. At the same time,the cultivation experimentof soil temperature and soil water content of Quercus aliena var.acuteserrata young stand wasset to understand the soil respiration characteristic at the larger scope of soil temperature. Thestatistical and simulated results show that:
(1) There are three diel variation patterns of total soil respiration and its components,the first which occurs before foliage spread in spring and after defoliation in fall is that thevariations of total soil respiration and its components are all consistent with soil temperature;the second which only occurs in summer in young-growth forest is that the variation of R_T, R_Sand R_A have short-term decoupling with soil temperature and soil water content, but thevariation of R_H and R_L is same as soil temperature; the third which occurs in growth seasons ofsummer and fall is that only the variation of R_A has short-term decoupling with soil temperatureand soil water content, the variations of the other respiration are all consistent with soiltemperature. The smallest and the biggest R_T, R_S, R_H ,R_A and R_L is at 06:00 and at 15:00～21:00respectively. The diel variation patterns for total soil respiration and its components are allsingle modal curve. Whether the decoupling of R_A with soil temperature and soil water contentcan lead to the decoupling of R_T and R_S with soil temperature and soil water content isdetermined by the absolute and relatively amount of root respiration in different stands andseasons.The diel variation range of respiration is highest in summer, followed by in spring and fall, and lowest in winter, its seasonal trend is same as that of water content and temperature;The seasonal variation coefficients of diel range of root respiration and litter respiration aredecreasing with stand age increasing, but that of heterotrophic is increasing with stand agesincreasing; The seasonal variation in total soil respiration, soil respiration, root respiration andheterotrophic respiration is single modal curve, which is same as that of soil temperature, thelargest and lowest respiration is in January and July respectively, but the respiration is clearlyrestricted by lower water content; The seasonal variation in litter respiration is more affectedby litter water content than by temperature.
(2) The relative importance of the effects of water and temperature on respiration hasseasonal variation, The respiration rate is largely controlled by temperature when soiltemperature and air temperature is below 15℃and 13℃respectively, while it is clearlylimited by water content when soil temperature is above 15℃and air temperature is above13℃respectively and soil water content and litter water content is below 0.20 kg·kg~(-1) and 0.38kg·kg~(-1) respectively. However, it is simultaneously affected by both temperature and watercontent under other condition. The soil water and temperature cultivation experiment indicatedthat the soil respiration is positively feedback with soil temperature increase, but the positivelyfeedback is restricted by water content which is over 0.35kg·kg~(-1) in Quercus alienavar.acuteserrata young stand.
(3) The relationship of soil temperature with total respiration, Soil respiration, rootrespiration and soil heterotrophie respiration is significant exponent function (P＜0.05) , soiltemperature can account for 80.62%～93.85%, 86.84%～92.23%, 79.05％～88.12% and82.63%～96.70% of diel and seasonal variation in above mentioned respiration respectively.The relationship of soil temperature with litter respiration is exponent function at diel scale butis quadratic equation at seasonal scale, soil temperature can account for 68.10%～81.58% ofdiel variation and 34.78%～54.24% seasonal variation in litter respiration. Respiration ispositively and negatively with water content at die and seasonal scale respectively, the simpleequation between soil water content and respiration indicates that soil water content canaccount for 24.8%～55.2% of seasonal variation in total soil respiration for 5 stands andaccount for 23.8%～47.1%, 20.45%～41.14% and 23.06%～48.45 of seasonal variation in soilrespiration, root respiration and soil heterotrophic respiration respectively for 4 stands (except for Quercus aliena var.acuteserrata old stand), but the cultivation experiment indicates therelationship between soil water content and respiration is quadratic equation. The relationshipbetween litter water content with litter respiration is significant single equation for 5 stands andlitter water content can account for 72.24%～79.5% of seasonal variation in litter respiration;The cultivation experiment indicates that soil water content increase can promote the soilrespiration when soil water content is very low and that soil water content increase can restrainsoil respiration when soil water is very high.
(4) The Q_(10) value of total respiration, soil respiration, soil heterotrophie respiration androot respiration is 2.18～2.41 (mean is 2.30) , 2.30～2.44(mean is 2.38) , 2.09～2.35 (mean is2.19)与2.49～2.82 (mean is 2.61) respectively at the annual scale, the Q_(10)value of abovementioned respiration is significant (P＜0.05) ; The Q_(10)value of the other respiration typesexcept for root respiration is highest in winter, followed by in spring and fall, and lowest insummer, but the Q_(10) value of root respiration is highest in spring, followed by in fall, andlowest in summer and winter; The higher Q_(10) value for total soil respiration in Quercus alienavar.acuteserrata young stand is than old stand under the condition that there is no significantdifference of the soil temperature between the 2 stands and that ther is no water contentlimitation; The Q_(10) value of root respiration is highest, followed by that of soil heterotrophicrespiration, that of litter respiration is lowest in many occasions, but the Q_(10) value of soilheterotrophie respiration is highest, followed by that of litter respiration, that of rootrespiration is lowest in winter; the Q_(10) value of total soil respiration is higer than that of soilrespiration regardless of diel and seasonal scale in 5 stands ; The order of Q_(10) values ofdifferent respiration in different stand has differences.The cultivation experiment indicates thatthe effects of soil temperature on Q_(10) value ,of soil respiration is very big ,but the effects of soilwater content on Q_(10) value of soil respiration is very small, the Q_(10) value of soil respiration isrestrained by very high soil water content too.
(5) The carbon release of root respiration in Quercus aliena var.acuteserrata youngstand, Quercus aliena var.acuteserrata old stand, broadleaf/coniferous mixed stand, broadleafmixed forest and Quercus variabilis stand is 724.52 gC·m~(-2)·a~(-1), 373.55 gC·m~(-2)·a~(-1),441.56～441.87 gC·m~(-2)·a~(-1), 427.14 gC·m~(-2)·a~(-1) and 484.43 C·m~(-2)·a~(-1) respectively; Their carbonrelease of soil heterotrophic respiration is 380.63 gC·m~(-2)·a~(-1), 405.57 gC·m~(-2)·a~(-1), 379.36～379.67 gC·m~(-2)·a~(-1), 485.05 gC·m~(-2)·a~(-1) and 415.07 gC·m~(-2)·a~(-1) respectively if taking account of fine rootdecomposition in root-excluded quadrates; Their carbon release of litter respiration is 239.04gC·m~(-2)·a~(-1), 119.79 gC·m~(-2)·a~(-1), 218.60 gC·m~(-2)·a~(-1), 115.03 gC·m~(-2)·a~(-1) and 277.29 gC·m~(-2)·a~(-1)respectively; Their carbon release of fresh litter respiration is 86.59 gC·m~(-2)·a~(-1), 56.72 gC·m~(-2)·a~(-1),65.92 gC·m~(-2)·a~(-1), 55.65 gC·m~(-2)·a~(-1) and 98.84 gC·m~(-2)·a~(-1) respectively; Their carbon release ofsoil respiration is 1105.15 gC·m~(-2)·a~(-1), 779.12 gC·m~(-2)·a~(-1), 821.23 gC·m~(-2)·a~(-1), 912.19 gC·m~(-2)·a~(-1)and 899.50 gC·m~(-2)·a~(-1) respectively; Their carbon release of total soil respiration is 1345.19gC·m~(-2)·a~(-1), 897.23 gC·m~(-2)·a~(-1), 1039.83 gC·m~(-2)·a~(-1), 1027.22 gC·m~(-2)·a~(-1)and 1176.79 gC·m~(-2)·a~(-1).The seasonal variation trend of soil respiration , soil heterotrophie respiration and rootrespiration is single modal curve, their smallest and biggest value is in January and July, Thepercentage of the carbon release of soil respiration, soil heterotrophic respiration and rootrespiration in summer and fall is 72.93%～76.55%, 54.03%～59.50% and 74.25%-82.21% ofannual release respectively for 5 stands; The carbon release of litter respiration is highest insummer, followed by in fall, lowest in spring and winter, the percentage of carbon release insummer and fall is 78.45%～83.67% of its annual release respectively for 5 stands
(6) The annual contribution rate of root respiration for stand 1～stand 5 is 53.86%,41.63%, 42.46%～42.49%, 41.58% and 41.17% respectively; Their annual contribution rate ofsoil heterotrophie respiration is 28.57%, 45.21%, 36.48%～36.51%, 47.13% and 35.27%respectively if taking account of fine root decomposition in root-excluded quadrates; Thecontribution rate of litter respiration is 17.77%, 13.35%, 21.01%, 12.61% and 23.56%; Thecontribution rate of root respiration is biggest, followed by soil heterotrophic respiration andthat of litter respiration is the lowest for Quercus aliena var.acuteserrata young stand,broadleaf/coniferous mixed stand and Quercus variabilis stand; But the contribution rate of soilheterotrophie respiration is biggest, followed by root respiration and that of litter respiration isthe lowest for the other two stands; The biggest and smallest contribution rate of rootrespiration is before 15:00 and at 18:00～21:00 respectively, the biggest and smallestcontribution rate of soil heterotrophic respiration is at 18:00～21:00 and before 15:00respectively and the biggest and smallest contribution rate of litter respiration is at 15:00～18:00and beforel2:00; The contribution rate of root respiration in summer is the biggest, followedby in spring and fall, lowest in winter, that of soil heterotrophic in winter is the biggest, followed by in spring and lowest in summer and fall, that of litter respiration in fall is biggestfollowed by summer, and lowest in spring and winter; The most important dominant factorscontrolling the seasonal variation in contribution rate of litter respiration is litter water content,the second factor is existing litter amount, which can accounts for 62.60%～71.42% of thedifference of contribution rate of litter respiration among the five stands.
(7) There is no significant differences in total soil respiration rate, soil respiration rateand the heterotrophic respiration rate among the five stands (P＞0.05), but the significantdifferences occur in the autotrophic respiration rate and litter respiration rote (P＜0.05). The livefine root biomass accounts for 94.71%of the difference of autotrophie respiration rates amongthe five stands, the existing litter amount accounts for 57.83% of the difference of litterrespiration among 5 stands in which the litter quality has obvious difference, but it canaccounts for 99.9% of the difference of litter respiration among 4 broadleaf stands in whichthe litter quality has no obvious difference.
引文
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