季节性冻融对高山森林土壤微生物与生化特性的影响
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摘要
受季节性冻融影响明显的高山/亚高山地区是全球变化的敏感区域。作为不争的事实,全球气候变暖必然改变高山/亚高山地区的季节性冻融过程,直接或间接作用于生态系统地下/地上过程,从而对陆地生态系统过程施加强烈影响。土壤微生物是生态系统物质循环与能量转换的积极参与者,也是对环境变化的敏感响应者。这意味着,全球变暖可能通过对冬季土壤微生物的影响而深刻影响到高山/亚高山森林生态系统的物质循环与能量转换过程。但迄今为止,有关高山/亚高山森林土壤微生物对季节性冻融及暖冬响应的研究还鲜见报道,这不仅限制了对高山/亚高山森林生态系统过程的完整理解,也不利于更好的预测气候变暖背景下土壤生态过程可能发生的变化。因此,本文以受季节性冻融影响明显的川西高寒森林土壤为研究对象,(1)通过野外定位监测,同步研究季节性冻融不同关键时期下,岷江冷杉原始林(PF)、岷江冷杉红桦混交林(MF)及岷江冷杉次生林(SF)土壤有机层(OL)和矿质土壤层(MS)中的土壤微生物与生化特性；(2)通过自然环境梯度实验,将于高山森林(N T,海拔3580m)采集的土柱分别置于原位培养、海拔3300m(T2,理论增温2℃)和3000m(T4,理论增温4℃)的林地,以研究气温增加对高山森林上壤有机层(OL)和矿质土壤层(MS)微生物与生化特性的影响。以期为深入理解高寒森林生态系统对季节性冻融及其变化响应提供证据。主要研究结果如下：
1.随土壤的冻结-融化,3个森林群落土壤有机层(OL)和矿质土壤层(MS)的微生物生物量碳(MBC)、生物量氮(MBN)、生物量磷(MBP)含量及三者的比例(MBC:MBN、 MBC:MBP和MBN:MBP)均表现出先降低后升高的变化趋势。其中,MBC和MBP含量及MBC:MBN以冻结期最低,MBN含量,MBC:MBP和MBN:MBP以深冻期最低。
2.3个森林群落OL和MS的细菌和真菌rDNA基因拷贝数也随土壤冻结-融化过程表现出先降低后升高的变化趋势,以土壤冻结后最低。OL中古菌的rDNA基因拷贝数也表现出相似的变化,但冷杉原始林(PF)和冷杉次生林(sF)的古菌的rDNA基因拷贝数在MS中表现出先升高后降低的趋势,均在土壤冻结期最高。同时,各森林OL的细菌rDNA基因拷贝数显著高于MS,但OL的古菌rDNA基因拷贝数在土壤冻结期和深冻期则显著低于MS,而OL的真菌rDNA基因拷贝数从生长季节到深冻期均低于MS。此外,冻结过程显著降低了OL和MS中细菌群落的多样性,提高了优势度;但随后的融化过程又增加了细菌群落多样性,降低了优势度。OL中古菌群落多样性表现出与细菌相似的变化动态。然而,冻结过程使原始冷杉林(PF)的真菌群落的多样性呈上升趋势。
3.随土壤冻结-融化过程,sF和MF群落两个土层的铵态氮含量均呈降低趋势,PF群落呈先升高后降低。3个森林群落OL和MS的硝态氮含量均呈先升高后降低的趋势,且OL以深冻期最高,MS以融冻期最高。各森林群落OL的脲酶活性显著高于MS,且随土壤季节性冻融过程表现出先降低后升高的趋势,以冻结期最低。同时,各森林群落两个土层的氨氧化微生物amoA基因拷贝数也呈先降低后升高的趋势,氨氧化细菌(AOB)的amoA基因拷贝数深冻期最低,氨氧化古菌数量(AOA)的amoA基因拷贝数则以冻结期最低。此外,PF群落的AOA数量多高于：AOB,而sF群落的AOA多低于AOB。
4.与对照(NT)相比,气温增加2℃处理(T2)使两个土层的月平均温度明显下降,而气温增加4℃处理(T4)则月平均温度明显上升。T2处理使OL和MS的土壤冻融循环次数分别减少了日次和8次,T4处理则使L壤冻融循环次数分别减少了1次和3次。但土壤冻融期间内,各处理土壤温度明显波动,以T4处理的最为明显。在土壤冻结期,T4、T2和NT处理OL的平均上壤温度分别为一0.17±1.21℃、—1.17±0.90℃、—0.65±0.97℃；MS的平均土壤温度分别为：一0.06±0.99℃、—0.52±0.76℃、—0.08±0.72。C。在土壤融化期,T4、T2和NT处理OL的平均土壤温度分别为2.04+2.11℃、0.35+0.98℃、0.04+0.57℃；MS的平均土壤温度为1.87+1.89℃、—0.11+0.27℃、0.8士1.33℃。
5.与对照相比,在不同冻融时期内,模拟增温均降低了OL层的MBC、MBN、 MBP含量,增加了MS层的MBN含量。同时,模拟增温总体上使两个土层的MBC:MBN表现出降低趋势,而MBC:MBP和MBN:MBP表现出上升趋势。土壤细菌rDNA基因拷贝数对模拟增温响应的规律不明显。模拟增温明显增加了OL和MS的古菌、真菌rDNA基因拷贝数。与对照相比,模拟增温提高了OL细菌群落的多样性,降低其优势度,但对MS细菌群落的多样性影响不明显。增温也使OL和MS古菌群落、真菌群落的多样性增加、优势度降低。模拟增温还明显提高了OL和MS中：真菌：细菌、古菌：细菌和古菌：真菌数量比。
6.整个冻融期间,模拟增温对土壤OL和MS的脲酶活性、转化酶活性的影响并未表现出较为一致的变化规律。同时,与对照相比,T2和T4处理增加了OL中AOB的amoA基因拷贝数。T2处理同样增加了OL中AOA的amoA基因拷贝数和氨氧化古菌：细菌数量比,但T4处理使OL中AOA的amoA基因拷贝数降低氨氧化古菌：细菌数量比。然而,模拟增温对MS的AOB和AOA的amoA基因拷贝数的影响也没有表现出一致的规律。
7.模拟增温在不同的冻融时期对土壤OL和MS的有机碳(TOC)、可溶性有机碳(DOC)、全氮(TN)及铵态氮含量的影响并没有表现出较为一致的变化规律。与对照相比,经过为期一年的不同增温处理后,OL和MS的TOC含量略微降低,DOC含量显著降低。然而,模拟增温使土壤OL和MS的可溶性有机氮(DON).硝态氮含、矿质氮含量增加。同时,在OL中,各处理生长季节均表现为氮素固持,而非生长季节多表现为氮素矿化(NT除外),土壤净氮矿化量和矿化速率随模拟增温幅度增大而增大。在MS中,生长季节和非生长季节总体上表现为氮素固持(T4除外),生长季节内的土壤净氮矿化量、矿化速率也随模拟增温幅度增大而增大,但非生长季节内的土壤净氮矿化量和矿化速率随模拟增温幅度增大而减小。
The subalpine and alpine region, as affected by seasonal freezing and thawing, is a sensitive area to climate change. As an indisputable fact, global warming can alter the pattern of seasonal freezing and thawing, and directly and indirectly influence the processes of above-and below-ground ecology, and in turn give strong effects on the terrestrial ecosystem process. Soil microorganism not only plays their roles as driving force in material cycling and energy flow, but also acts as a sensitive bio-indicator to environmental change. The ongoing winter warming under climate change scenarios is suggested to have great effects on the pattern of seasonal snow cover and seasonal freeze-thaw and significantly change the structure and function of soil microorganism, subsequent have certain consequences for biogeochemical processes in the high latitudinal and altitudinal ecosystems. As yet, more attentions have concerned about the effect of experimental warming on the structure and function of soil microbial community during the growing season in the high latitudinal regions, and little information is available on the microbial response to experimental winter warming. Therefore, it is difficult for us to deeply understand the ecological process in the high-frigid ecosystem and predict the potential change of soil process under climate change scenarios.
The subalpine and alpine forest of western Sichuan located in the eastern edge of Qinghai-Tibet plateau and the upper reaches of the Yangtze River is a principal part of the second biggest forest region (southwest forest region) in China, which play important roles in conserving water, nursing biodiversity and indicating regional climate change. Hence, in this study, we conducted two experiments consisting of a field investigation and an experimental warming to assess the effects of global warming on the biogeochemical processes in the high latitudinal and altitudinal ecosystems.(1) From August2009to August2010, the diversity of soil microorganism and their biochemical properties were simultaneously measured in primary fir (Abies faxoniana) forest (PF), fir and birch (Betula albosinensis) mixed forest (MF) and secondary fir forest (SF) which are three representative forests in the study region, employing the method of field sampling in combination with laboratory analysis in different critical periods, especially to the winter.(2) From May2010to August2011, untouched soil cores that sampled from the primary fir forest were respectively laid in three different altitudes (3600,3300and3000) to simulate experimental winter warming with increase amplitude of air temperature0℃(3600),2℃(3300) and4℃(3000). The soil microbial properties, soil enzyme activities, and soil carbon and nutrients content were simultaneously measured during the winter and growing season.
1. Season freeze-thaw significantly influenced the dynamic of soil microbial biomass in the subalpine and alpine forest. The freezing process decreased the microbial biomass C (MBC) and microbial biomass N (MBN) in both the soil organic layer (OL) and mineral soil layer (MS), but increased the microbial biomass P (MBP) in the two layers. In contrast, the thawing process increased the MBC, MBN and MBP in the two soil layers. The MBC/MBN, MBC/MBP and MBN/MBP showed a decrease and then increase trend as the soil sampled time. The lowest values of MBC/MBP and MBN/MBP were observed in the deeply frost stage (except MBC/MBP in the MF), and the lowest values of MBC/MBN were observed in the frozen stage.
2. As the soil freezing and thawing processes, the bacterial and fungal rDNA copy numbers showed a decrease and then increase trend in the OL and MS, and the lowest values were observed in the soil freezing stage. The archaeal rDNA copy numbers in the OL also showed a similar change as the bacteria and fungi, but that in the MS showed an increase and then decrease trend in the PF and SF, and the highest values were observed in the soil freezing stage. Moreover, the bacterial rDNA copy numbers in the OL were higher than that in the MS. However, the fungal rDNA copy numbers in the OL were lower than that in the MS from growing season to deeply frozen stage, and the archaea rDNA copy numbers in the OL also were lower than that in the MS in the soil freezing stage and deeply frozen stage. In addition, season freeze-thaw also obviously affected the diversity of bacteria, archaea, and fungi in both the OL and MS. For the bacteria, the freezing process decreased the diversity index (H') and increased the dominance index (C') in the two soil layer, but followed thawing process increased the diversity index (H') and decreased the dominance index (C). Although similar dynamic of the diversity indices were observed in the OL for the archaea, reverse change trend was observed in the MS. Moreover, the diversity of fungi showed increase trend as seasonal freeze-thaw.
3. Season freeze-thaw greatly influenced the extractable inorganic N (ammonium and nitrate), soil urease activity, and soil ammonia-oxidizing microorganism. The freezing process significantly decreased the soil urease activity, the quantity of the soil ammonia-oxidizing bacteria and soil ammonia-oxidizing archaea. After that, these indices significantly increased. The ammonium and extractable N showed a decrease and then increase trend in the MF and SF as affected by seasonal freeze-thaw. However, the nitrate showed an increase and then decrease trend. Moreover, the concentration of ammonium was significantly higher than that of nitrate.
4. As compared with air temperature increased0℃(NT), air temperature increased2℃(T2) obviously decreased the monthly mean soil temperature in both the OL and MS, but air temperature increased4℃(T4) significantly increased the monthly mean soil temperature in both the OL and MS. The freeze-thaw cycles in the OL and MS were decreased by11and8in the T2treatment, respectively, and these were decreased by1and3in the T4treatment. The soil temperature showed obvious fluctuation during the freezing and thawing period, and the amplitude of fluctuation in the T4was higher than that in the NT and T2. During soil freezing stage, the mean soil temperature of NT, T2, and T4in the OL was-0.17±1.21℃,-1.17±0.90℃-0.65±0.97℃, respectively, and that in the MS was-0.06±0.99℃,-0.52±0.76℃,-0.08±0.72℃, respectively. During soil thawing stage, the mean soil temperature of NT, T2, and T4in the OL2.04±2.11℃,0.35±0.98℃,0.04±0.57℃, respectively, and that in the MS was1.87±1.89℃、-0.11±0.27℃、0.8±1.33℃, respectively.
5. As compared with the NT, the MBC, MBN, and MBP in the OL were decreased with the increase of simulated warming amplitude, while MBN in the MS was increased with the increase of simulated warming amplitude. The ratio of MBC/MBN in OL was increased in growing season, but decreased in non-growing season. The ratio of MBC/MBN in MS was decreased. The ratio of MBC/MBP and MBN/MBP in OL and MS were both increased, only except for thawing period. Moreover, Bacterial, archaeal and fungal rDNA numbers were both increased in compared with control plot after annual warming treatment. The archaeal and fungal rDNA numbers has a positive feedback to the increase of simulated warming amplitude. The relative number of archaea and fungal was both increased by warming treatment. In addition, soil bacterial and archaeal community structure was sensitive to warming treatment, while fungal community was more consistent. The diversity of bacteria in OL and diversity of archaeal and fungal in OL and MS were both increased by warming treatment.
6. The warming decreased the activity of urease in all studied periods. But the effect of warming to invertase activity was depended on different periods. Warming increased the numbers of ammonia-oxidizing bacteria, especially during thawing period. But the numbers of ammonia-oxidizing archaea in OL was decreased by warming, while ammonia-oxidizing archaea in MS was change a little. In addition, the Log ratio of AOA to AOB in OL was decreased by warming, while this ratio in MS did not change with obviously tendency.
7. After annual different warming treatment, total organic carbon (TC) content was decreased by seasonal freeze-thaw. The TC content in OL was increased by warming in the end of studied period. Dissolved organic carbon (DOC) content was decreased by seasonal freeze-thaw. Total nitrogen (TN) content was decreased by seasonal freeze-thaw. Dissolved organic nitrogen (DON) content was decreased by seasonal freeze-thaw. The DON content was mainly increased by warming. NH4+-N content of OL was mainly increased by seasonal freeze-thaw, which in MS was decreased. The NH4+-N content was mainly decreased by warming. The NO3--N content was decreased by seasonal freeze thaw. The NO3--N content was mainly increased by warming. The mineral nitrogen content was decreased by seasonal freeze thaw. But the mineral nitrogen content was mainly increased by warming. But the effect of warming on TC, DOC, TN, NH4+-N, was depended on different periods. In addition, the soil nitrogen net mineralization rate and the amount of soil mineralized nitrogen was increased by warming amplitude n different treatment of OL and MS. However, the soil inorganic nitrogen was obviously immobilized in growing season, but it was minerlization in non-growing (freeze-thaw) season. The soil inorganic nitrogen in MS was mainly immobilized in all studied periods (except for T4treatment).
1.随土壤的冻结-融化,3个森林群落土壤有机层(OL)和矿质土壤层(MS)的微生物生物量碳(MBC)、生物量氮(MBN)、生物量磷(MBP)含量及三者的比例(MBC:MBN、 MBC:MBP和MBN:MBP)均表现出先降低后升高的变化趋势。其中,MBC和MBP含量及MBC:MBN以冻结期最低,MBN含量,MBC:MBP和MBN:MBP以深冻期最低。
2.3个森林群落OL和MS的细菌和真菌rDNA基因拷贝数也随土壤冻结-融化过程表现出先降低后升高的变化趋势,以土壤冻结后最低。OL中古菌的rDNA基因拷贝数也表现出相似的变化,但冷杉原始林(PF)和冷杉次生林(sF)的古菌的rDNA基因拷贝数在MS中表现出先升高后降低的趋势,均在土壤冻结期最高。同时,各森林OL的细菌rDNA基因拷贝数显著高于MS,但OL的古菌rDNA基因拷贝数在土壤冻结期和深冻期则显著低于MS,而OL的真菌rDNA基因拷贝数从生长季节到深冻期均低于MS。此外,冻结过程显著降低了OL和MS中细菌群落的多样性,提高了优势度;但随后的融化过程又增加了细菌群落多样性,降低了优势度。OL中古菌群落多样性表现出与细菌相似的变化动态。然而,冻结过程使原始冷杉林(PF)的真菌群落的多样性呈上升趋势。
3.随土壤冻结-融化过程,sF和MF群落两个土层的铵态氮含量均呈降低趋势,PF群落呈先升高后降低。3个森林群落OL和MS的硝态氮含量均呈先升高后降低的趋势,且OL以深冻期最高,MS以融冻期最高。各森林群落OL的脲酶活性显著高于MS,且随土壤季节性冻融过程表现出先降低后升高的趋势,以冻结期最低。同时,各森林群落两个土层的氨氧化微生物amoA基因拷贝数也呈先降低后升高的趋势,氨氧化细菌(AOB)的amoA基因拷贝数深冻期最低,氨氧化古菌数量(AOA)的amoA基因拷贝数则以冻结期最低。此外,PF群落的AOA数量多高于：AOB,而sF群落的AOA多低于AOB。
4.与对照(NT)相比,气温增加2℃处理(T2)使两个土层的月平均温度明显下降,而气温增加4℃处理(T4)则月平均温度明显上升。T2处理使OL和MS的土壤冻融循环次数分别减少了日次和8次,T4处理则使L壤冻融循环次数分别减少了1次和3次。但土壤冻融期间内,各处理土壤温度明显波动,以T4处理的最为明显。在土壤冻结期,T4、T2和NT处理OL的平均上壤温度分别为一0.17±1.21℃、—1.17±0.90℃、—0.65±0.97℃；MS的平均土壤温度分别为：一0.06±0.99℃、—0.52±0.76℃、—0.08±0.72。C。在土壤融化期,T4、T2和NT处理OL的平均土壤温度分别为2.04+2.11℃、0.35+0.98℃、0.04+0.57℃；MS的平均土壤温度为1.87+1.89℃、—0.11+0.27℃、0.8士1.33℃。
5.与对照相比,在不同冻融时期内,模拟增温均降低了OL层的MBC、MBN、 MBP含量,增加了MS层的MBN含量。同时,模拟增温总体上使两个土层的MBC:MBN表现出降低趋势,而MBC:MBP和MBN:MBP表现出上升趋势。土壤细菌rDNA基因拷贝数对模拟增温响应的规律不明显。模拟增温明显增加了OL和MS的古菌、真菌rDNA基因拷贝数。与对照相比,模拟增温提高了OL细菌群落的多样性,降低其优势度,但对MS细菌群落的多样性影响不明显。增温也使OL和MS古菌群落、真菌群落的多样性增加、优势度降低。模拟增温还明显提高了OL和MS中：真菌：细菌、古菌：细菌和古菌：真菌数量比。
6.整个冻融期间,模拟增温对土壤OL和MS的脲酶活性、转化酶活性的影响并未表现出较为一致的变化规律。同时,与对照相比,T2和T4处理增加了OL中AOB的amoA基因拷贝数。T2处理同样增加了OL中AOA的amoA基因拷贝数和氨氧化古菌：细菌数量比,但T4处理使OL中AOA的amoA基因拷贝数降低氨氧化古菌：细菌数量比。然而,模拟增温对MS的AOB和AOA的amoA基因拷贝数的影响也没有表现出一致的规律。
7.模拟增温在不同的冻融时期对土壤OL和MS的有机碳(TOC)、可溶性有机碳(DOC)、全氮(TN)及铵态氮含量的影响并没有表现出较为一致的变化规律。与对照相比,经过为期一年的不同增温处理后,OL和MS的TOC含量略微降低,DOC含量显著降低。然而,模拟增温使土壤OL和MS的可溶性有机氮(DON).硝态氮含、矿质氮含量增加。同时,在OL中,各处理生长季节均表现为氮素固持,而非生长季节多表现为氮素矿化(NT除外),土壤净氮矿化量和矿化速率随模拟增温幅度增大而增大。在MS中,生长季节和非生长季节总体上表现为氮素固持(T4除外),生长季节内的土壤净氮矿化量、矿化速率也随模拟增温幅度增大而增大,但非生长季节内的土壤净氮矿化量和矿化速率随模拟增温幅度增大而减小。
The subalpine and alpine region, as affected by seasonal freezing and thawing, is a sensitive area to climate change. As an indisputable fact, global warming can alter the pattern of seasonal freezing and thawing, and directly and indirectly influence the processes of above-and below-ground ecology, and in turn give strong effects on the terrestrial ecosystem process. Soil microorganism not only plays their roles as driving force in material cycling and energy flow, but also acts as a sensitive bio-indicator to environmental change. The ongoing winter warming under climate change scenarios is suggested to have great effects on the pattern of seasonal snow cover and seasonal freeze-thaw and significantly change the structure and function of soil microorganism, subsequent have certain consequences for biogeochemical processes in the high latitudinal and altitudinal ecosystems. As yet, more attentions have concerned about the effect of experimental warming on the structure and function of soil microbial community during the growing season in the high latitudinal regions, and little information is available on the microbial response to experimental winter warming. Therefore, it is difficult for us to deeply understand the ecological process in the high-frigid ecosystem and predict the potential change of soil process under climate change scenarios.
The subalpine and alpine forest of western Sichuan located in the eastern edge of Qinghai-Tibet plateau and the upper reaches of the Yangtze River is a principal part of the second biggest forest region (southwest forest region) in China, which play important roles in conserving water, nursing biodiversity and indicating regional climate change. Hence, in this study, we conducted two experiments consisting of a field investigation and an experimental warming to assess the effects of global warming on the biogeochemical processes in the high latitudinal and altitudinal ecosystems.(1) From August2009to August2010, the diversity of soil microorganism and their biochemical properties were simultaneously measured in primary fir (Abies faxoniana) forest (PF), fir and birch (Betula albosinensis) mixed forest (MF) and secondary fir forest (SF) which are three representative forests in the study region, employing the method of field sampling in combination with laboratory analysis in different critical periods, especially to the winter.(2) From May2010to August2011, untouched soil cores that sampled from the primary fir forest were respectively laid in three different altitudes (3600,3300and3000) to simulate experimental winter warming with increase amplitude of air temperature0℃(3600),2℃(3300) and4℃(3000). The soil microbial properties, soil enzyme activities, and soil carbon and nutrients content were simultaneously measured during the winter and growing season.
1. Season freeze-thaw significantly influenced the dynamic of soil microbial biomass in the subalpine and alpine forest. The freezing process decreased the microbial biomass C (MBC) and microbial biomass N (MBN) in both the soil organic layer (OL) and mineral soil layer (MS), but increased the microbial biomass P (MBP) in the two layers. In contrast, the thawing process increased the MBC, MBN and MBP in the two soil layers. The MBC/MBN, MBC/MBP and MBN/MBP showed a decrease and then increase trend as the soil sampled time. The lowest values of MBC/MBP and MBN/MBP were observed in the deeply frost stage (except MBC/MBP in the MF), and the lowest values of MBC/MBN were observed in the frozen stage.
2. As the soil freezing and thawing processes, the bacterial and fungal rDNA copy numbers showed a decrease and then increase trend in the OL and MS, and the lowest values were observed in the soil freezing stage. The archaeal rDNA copy numbers in the OL also showed a similar change as the bacteria and fungi, but that in the MS showed an increase and then decrease trend in the PF and SF, and the highest values were observed in the soil freezing stage. Moreover, the bacterial rDNA copy numbers in the OL were higher than that in the MS. However, the fungal rDNA copy numbers in the OL were lower than that in the MS from growing season to deeply frozen stage, and the archaea rDNA copy numbers in the OL also were lower than that in the MS in the soil freezing stage and deeply frozen stage. In addition, season freeze-thaw also obviously affected the diversity of bacteria, archaea, and fungi in both the OL and MS. For the bacteria, the freezing process decreased the diversity index (H') and increased the dominance index (C') in the two soil layer, but followed thawing process increased the diversity index (H') and decreased the dominance index (C). Although similar dynamic of the diversity indices were observed in the OL for the archaea, reverse change trend was observed in the MS. Moreover, the diversity of fungi showed increase trend as seasonal freeze-thaw.
3. Season freeze-thaw greatly influenced the extractable inorganic N (ammonium and nitrate), soil urease activity, and soil ammonia-oxidizing microorganism. The freezing process significantly decreased the soil urease activity, the quantity of the soil ammonia-oxidizing bacteria and soil ammonia-oxidizing archaea. After that, these indices significantly increased. The ammonium and extractable N showed a decrease and then increase trend in the MF and SF as affected by seasonal freeze-thaw. However, the nitrate showed an increase and then decrease trend. Moreover, the concentration of ammonium was significantly higher than that of nitrate.
4. As compared with air temperature increased0℃(NT), air temperature increased2℃(T2) obviously decreased the monthly mean soil temperature in both the OL and MS, but air temperature increased4℃(T4) significantly increased the monthly mean soil temperature in both the OL and MS. The freeze-thaw cycles in the OL and MS were decreased by11and8in the T2treatment, respectively, and these were decreased by1and3in the T4treatment. The soil temperature showed obvious fluctuation during the freezing and thawing period, and the amplitude of fluctuation in the T4was higher than that in the NT and T2. During soil freezing stage, the mean soil temperature of NT, T2, and T4in the OL was-0.17±1.21℃,-1.17±0.90℃-0.65±0.97℃, respectively, and that in the MS was-0.06±0.99℃,-0.52±0.76℃,-0.08±0.72℃, respectively. During soil thawing stage, the mean soil temperature of NT, T2, and T4in the OL2.04±2.11℃,0.35±0.98℃,0.04±0.57℃, respectively, and that in the MS was1.87±1.89℃、-0.11±0.27℃、0.8±1.33℃, respectively.
5. As compared with the NT, the MBC, MBN, and MBP in the OL were decreased with the increase of simulated warming amplitude, while MBN in the MS was increased with the increase of simulated warming amplitude. The ratio of MBC/MBN in OL was increased in growing season, but decreased in non-growing season. The ratio of MBC/MBN in MS was decreased. The ratio of MBC/MBP and MBN/MBP in OL and MS were both increased, only except for thawing period. Moreover, Bacterial, archaeal and fungal rDNA numbers were both increased in compared with control plot after annual warming treatment. The archaeal and fungal rDNA numbers has a positive feedback to the increase of simulated warming amplitude. The relative number of archaea and fungal was both increased by warming treatment. In addition, soil bacterial and archaeal community structure was sensitive to warming treatment, while fungal community was more consistent. The diversity of bacteria in OL and diversity of archaeal and fungal in OL and MS were both increased by warming treatment.
6. The warming decreased the activity of urease in all studied periods. But the effect of warming to invertase activity was depended on different periods. Warming increased the numbers of ammonia-oxidizing bacteria, especially during thawing period. But the numbers of ammonia-oxidizing archaea in OL was decreased by warming, while ammonia-oxidizing archaea in MS was change a little. In addition, the Log ratio of AOA to AOB in OL was decreased by warming, while this ratio in MS did not change with obviously tendency.
7. After annual different warming treatment, total organic carbon (TC) content was decreased by seasonal freeze-thaw. The TC content in OL was increased by warming in the end of studied period. Dissolved organic carbon (DOC) content was decreased by seasonal freeze-thaw. Total nitrogen (TN) content was decreased by seasonal freeze-thaw. Dissolved organic nitrogen (DON) content was decreased by seasonal freeze-thaw. The DON content was mainly increased by warming. NH4+-N content of OL was mainly increased by seasonal freeze-thaw, which in MS was decreased. The NH4+-N content was mainly decreased by warming. The NO3--N content was decreased by seasonal freeze thaw. The NO3--N content was mainly increased by warming. The mineral nitrogen content was decreased by seasonal freeze thaw. But the mineral nitrogen content was mainly increased by warming. But the effect of warming on TC, DOC, TN, NH4+-N, was depended on different periods. In addition, the soil nitrogen net mineralization rate and the amount of soil mineralized nitrogen was increased by warming amplitude n different treatment of OL and MS. However, the soil inorganic nitrogen was obviously immobilized in growing season, but it was minerlization in non-growing (freeze-thaw) season. The soil inorganic nitrogen in MS was mainly immobilized in all studied periods (except for T4treatment).
引文
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