川西亚高山森林凋落物分解初期土壤动物对红桦凋落叶质量损失的贡献

2010年10月26日-2011年4月18日在川西亚高山地区季节性冻融期间,选择典型的红桦-岷江冷杉林,采用凋落物分解袋法调查了不同网孔(0.02、0.125、1和3 mm)凋落物分解袋内的凋落物质量损失,分析微型、中型和大型土壤动物对红桦凋落叶分解的贡献.结果表明:在季节性冻融期间,0.02、0.125、1和3 mm分解袋内的红桦凋落叶质量损失率分别为11.8％、13.2％、15.4％和19.5％,不同体径土壤动物对红桦凋落叶质量损失的贡献率为39.5％;不同孔径凋落物袋内土壤动物的类群和个体相对密度与凋落叶的质量损失率的变化趋势相对一致.在季节性冻融的初期、深冻期和融化期,不同土壤动物对红桦凋落叶质量损失的贡献率为大型土壤动物(22.7％)＞中型土壤动物(11.9％)＞微型土壤动物(7.9％).季节性冻融期间土壤动物活动是影响川西亚高山森林凋落物分解的重要因素之一.     

川西高山林线交错带土壤动物对岷江冷杉和高山杜鹃凋落物分解的贡献

土壤动物是调控凋落物分解的重要生物因素.为了探究川西高山林线交错带土壤动物对两个优势物种岷江冷杉和高山杜鹃凋落物分解的贡献,在3个海拔梯度(针叶林-林线-高山草甸)采用凋落物分解袋试验,通过不同孔径的网袋(0.04 mm,基本排除土壤动物;3 mm,允许土壤动物通过),研究了分解554 d(2013年5月—2014年11月)土壤动物对凋落物的影响.结果表明: 在整个林线交错带上,岷江冷杉的分解速率(k)为0.209～0.243,高山杜鹃的k为0.173～0.189,岷江冷杉的k大于高山杜鹃.土壤动物的参与显著加速了两种凋落叶分解,同时土壤动物对两种凋落物分解的作用和贡献随海拔升高而降低.自针叶林、高山林线至高山草甸,土壤动物对岷江冷杉分解的质量损失率为15.2%、13.2%、9.8%,对高山杜鹃分解的质量损失率为20.1%、17.5%、12.4%;土壤动物对岷江冷杉分解的平均日贡献率为0.17%、0.13%、0.12%,对高山杜鹃分解的平均日贡献率为0.26%、0.25%、0.23%,土壤动物对高山杜鹃的分解影响相对较大.海拔、凋落物自身性质及其交互作用对土壤动物作用下凋落物的质量损失率和贡献率均表现出显著影响.土壤动物的作用于岷江冷杉和高山杜鹃分解的平均日贡献率在当年生长季(0.25%和0.44%)和次年生长季(0.10%和0.19%)均高于雪被期(0.07%和0.12%).回归分析表明,环境因子(日平均气温、冻融循环次数以及雪被厚度)可以解释土壤动物作用于岷江冷杉和高山杜鹃质量损失率的42.7%和50.9%,贡献率的43.2%和55.6%,这对了解土壤动物在凋落物分解中的作用和深入认识高山生态系统物质循环具有重要意义.     

土壤动物对川西高山/亚高山森林凋落物分解的贡献

采用凋落物分解袋法,研究了土壤动物对川西高山/亚高山森林代表性植物康定柳、方枝柏、红桦和岷江冷杉凋落物在分解第一年(2011年11月-2012年10月)不同关键时期质量损失的贡献.结果表明: 在凋落物第一年的分解过程中, 不同物种凋落物的分解速率大小依次为康定柳＞红桦＞岷江冷杉＞方枝柏,且均为生长季节大于冻融季节.土壤动物对凋落物分解的贡献率(P_fau)为方枝柏(26.7%)＞岷江冷杉(18.8%)＞红桦(15.7%)＞康定柳(13.2%),其中康定柳和方枝柏的P_fau在生长季节大于冻融季节,而红桦和岷江冷杉的P_fau为冻融季节大于生长季节.冻融季节土壤动物的作用与凋落物初始C、P和N/P显著相关,而生长季节则与N、C/N、木质素、木质素/纤维素显著相关.     

季节性冻融期间土壤动物对高山草甸两种凋落叶失重的贡献

季节性冻融期间高山草甸凋落叶的分解可为生长季节植物生长提供必要的养分,对于维持生态系统物质循环和养分平衡具有重要作用。然而,土壤动物对凋落叶分解是否具有明显的贡献仍然缺乏一致认识。因此,以高山草甸代表性植物黄花亚菊(Ajania nubigena)和黑褐苔草(Carex atrofusca)凋落叶为研究对象,采用不同孔径凋落叶袋排除土壤动物的方法,研究冬季不同冻融时期(冻结前期、冻结期和融化期)土壤动物对凋落叶失重的贡献。整个季节性冻融期间土壤动物对黄花亚菊和黑褐苔草两种凋落叶失重率的作用分别为12.07%和4.03%,总贡献率分别为46.39%和24.14%。土壤动物对两种凋落叶失重率的作用均在融化期最大,而土壤动物对黄花亚菊凋落叶失重率的作用在冻结初期最小,土壤动物对黑褐苔草凋落叶失重率的作用在冻结期最小。整个季节性冻融期,土壤动物对凋落叶失重率的作用和贡献率与正积温和凋落叶初始C、N浓度和C/N比均呈显著的正相关关系。因此,季节性冻融期间土壤动物对高山草甸凋落叶分解具有明显的贡献,但这些过程受冻融格局和凋落叶初始质量的调控。     

季节性冻融期间亚高山森林凋落物的质量变化

凋落物质量是影响凋落物分解的重要生物因子,其在季节性冻融期间的变化可能对亚高山森林生态系统过程产生显著的影响。因此,采用凋落物分解袋法,研究了岷江冷杉(Abies faxoniana)和白桦(Betula platyphylla)凋落物质量在一个季节性冻融期间(2006年10月至2007年4月)的变化。季节性冻融期间,岷江冷杉和白桦凋落物的木质素(L)和纤维素的降解率为全年降解的70%-75%,岷江冷杉和白桦凋落物的C/N、L/N和纤维素/N均显著增加,而纤维素/P均有所降低。岷江冷杉凋落物的C/P和L/P有所增加,但白桦凋落物的C/P和L/P有所降低。可见,季节性冻融期间,亚高山森林凋落物的质量发生了较为显著的变化,其显著影响了亚高山凋落物分解过程。     

高山森林林窗对凋落叶分解的影响

林窗对降水和光照等环境条件的再分配以及分解者群落的影响可能深刻作用于森林凋落物分解过程,但有关高山森林林窗大小对凋落物分解的影响尚无研究报道。采用凋落物分解袋法,研究了川西高山森林不同大小林窗对非生长季节和生长季节红桦(Betula albo-sinensis)和岷江冷杉(Abies faxoniana)凋落叶质量损失的影响。结果显示,经过1a的分解,不同生境下红桦和岷江冷杉凋落叶分别分解了27.25%—30.12%和27.04%—27.96%,其中非生长季节占53.83%—60.18%和50.23%—59.09%。林窗对红桦和岷江冷杉凋落叶质量损失的影响因物种不同而呈现季节差异。总体上,林窗加快了岷江冷杉凋落叶的分解而延缓了红桦凋落叶的分解。与郁闭林下相比,林窗显著增加了2种凋落叶非生长季节的质量损失速率,显著降低了生长季节2种凋落叶的质量损失速率;2种凋落叶质量损失速率在非生长季节随林窗面积增大而加快,在生长季节随林窗面积增大而减慢。林窗显著影响了初冻期、深冻期和融化期岷江冷杉凋落叶的质量损失率,但对红桦凋落叶质量损失率影响不显著。可见,高山森林凋落物分解过程受到林窗的显著影响,并且阔叶和针叶凋落叶在非生长季节和生长季节对林窗的响应具有明显差异。     

西沙永兴岛抗风桐与海岸桐群落凋落叶分解及中型土壤动物的贡献

我国南海诸岛主要是珊瑚岛。植物凋落物分解是生态系统元素循环的关键环节,但目前关于南海珊瑚岛生态系统凋落物分解的研究还是空白。以我国西沙群岛的优势树种抗风桐（Pisonia grandis）和海岸桐（Guettarda speciosa）为研究对象,采用凋落物袋法,分别于分解期间的第3、6、9、13和15个月取样,探究中型土壤动物对两种植物群落中凋落物分解过程中质量损失和养分释放的影响。结果表明：与没有中型土壤动物存在的情况（0.1 mm凋落物袋）相比,分解开始后的6个月内,中型土壤动物存在（2 mm凋落物袋）使抗风桐和海岸桐凋落叶分解速率分别提高了12.3%和4.8%（P<0.05）;分解6-15个月期间,中型土壤动物存在使抗风桐和海岸桐凋落叶分解速率分别提高了33.0%和12.3%（P<0.05）。中型土壤动物排除显著影响了不同分解阶段凋落叶总碳（Total carbon,TC）、总氮（Total nitrogen,TN）、纤维素、木质素和半纤维素的残留率变化。中型土壤动物群落组成受土壤温度显著影响（P<0.05）,它们对凋落叶分解的贡献可能主要受优势类群如真螨目和寄螨目的影响。相较海岸桐,抗风桐凋落叶的分解周期更短,中型土壤动物对其的贡献更大;选用抗风桐作为南海珊瑚岛退化植被恢复或新建的先锋种对促进生态系统元素循环更有利。     

季节性雪被对高山森林凋落物分解的影响

季节性雪被可能对高山森林凋落物分解产生重要影响, 但一直没有深入的研究。该文采用凋落物分解袋法, 于2010-2012年雪被覆盖下几个关键时期(冻结初期、深冻期和融化期)以及生长季节, 研究了川西高山森林代表性树种岷江冷杉(Abies faxoniana)、红桦(Betula albosinensis)、四川红杉(Larix mastersiana)和方枝柏(Sabina saltuaria)凋落叶在不同厚度冬季雪被下的分解动态。经过两年的分解, 不同雪被覆盖下岷江冷杉凋落物分解率为33.98%-39.55%, 红桦为46.49%-48.22%, 四川红杉为42.30%-44.93%, 方枝柏为40.34%-43.84%。相对于无雪被覆盖环境, 厚型雪被覆盖均小幅提高了4种凋落物两年的失重率(1.57%-5.57%)。3个针叶树种(岷江冷杉、四川红杉和方枝柏) Olson凋落物分解系数k均以厚型雪被覆盖最大, 薄型雪被覆盖最小, 而阔叶树种红桦分解系数k则表现为无雪被>薄型雪被>较厚型雪被>厚型雪被>中型雪被。尽管在第二年生长季中雪被对红桦凋落物分解的促进作用不明显, 但雪被覆盖明显促进了两年各个关键时期岷江冷杉、四川红杉和方枝柏凋落物的分解。第一年雪被期凋落物分解对当年分解总量的贡献达42.5%-65.5%, 季节性雪被变化明显改变了凋落物冬季分解格局, 对深冻期凋落物分解过程影响尤为显著。综上所述, 当前气候变化情景下冬季雪被的减少可能减缓该区森林凋落物分解过程, 但相对于易分解的阔叶凋落物, 针叶凋落物的响应特征可能更为强烈。     

季节性冻融对岷江冷杉和白桦凋落物酶活性的影响

为了了解季节性冻融对亚高山森林生态系统过程的影响,2006年10月—2007年10月,采用凋落物分解袋法,研究了冻融和非冻融处理下岷江冷杉和白桦凋落物的酶活性动态.结果表明：凋落物的酶活性均具有明显的月变化,但活性高峰与酶类有关;季节性冻融显著影响了冷杉和白桦凋落物分解过程中的转化酶、纤维素酶、脲酶和脱氢酶活性(P＜0.05).冻融处理使冷杉凋落物分解过程中的转化酶、纤维素酶和脲酶活性在生长季节内分别增加了66.1%、14.8%和76.3%,但脱氢酶活性降低了18.4%;冻融处理使白桦凋落物分解过程中的转化酶活性增加了39.4%,但脲酶、纤维素酶和脱氢酶活性分别降低了18.2%、28.7%和15.6%.冻融处理对两种凋落物的过氧化物酶和多酚氧化酶活性影响不显著.说明季节性冻融对亚高山森林凋落物分解过程具有不同程度的影响,其影响程度与凋落物种类和分解阶段有关.     

亚高山森林林窗大小对凋落叶木质素降解的影响

木质素降解是认识高寒森林凋落物分解过程的关键环节,可能受到林窗大小及其在不同季节水热环境的影响。采用分解袋法,研究了川西亚高山森林不同面积大小林窗下红桦(Betula albo-sinensis)和岷江冷杉(Abies faxoniana)凋落叶在初冻期、深冻期、融化期、生长季节初期、生长季节中期和生长季节后期的木质素分解动态特征。研究结果表明,采样时间和林窗面积大小对两种凋落叶的木质素降解均有显著影响。经历1a分解,红桦凋落叶的木质素降解了21.53%—27.65%,而岷江冷杉凋落叶的木质素富集了7.95%—19.40%。较大林窗促进了冬季岷江冷杉凋落叶和生长季节红桦凋落叶木质素的降解,抑制了冬季红桦凋落叶木质素的降解;而生长季节岷江冷杉凋落叶木质素富集速率则为林下大林窗中林窗小林窗。逐步回归分析表明,凋落叶木质素的降解过程在冬季主要受到负积温和土壤冻融循环次数的影响(木质素结构的物理破碎),而在生长季节则主要受到平均温度和正积温的影响(木质素的生物降解)。可见,川西亚高山森林木质素降解受林窗格局变化的显著影响,且林窗大小对凋落叶木质素降解的影响与物种和分解时期有关。     

Contribution of meso- and macro-fauna to nitrogen release in needle litter decomposition of Abies faxoniana during the freeze-thaw period

The litterbags with different mesh sizes (0.125, 1.000 and 3.000 mm) were placed on the forest floor in October 2010, and collected after each of the three stages in the freeze-thaw season (OF: the onset of freezing stage; DF: the deeply frozen stage; and TS: the thawing stage). Over the whole freeze-thaw period, 14.01% ± 1.26, 20.64% ± 1.61 and 30.89% ± 0.70 of N released from fir litter in the litterbags with 0.125, 1.000 and 3.000 mm, respectively. The contributions of meso- and macro-fauna to N releases were 21.45% ± 3.80 and 33.18% ± 5.39, respectively. Macro-fauna contributed more to N releases than meso-fauna regardless of decomposition stages in the freeze-thaw period. Compared with other stages in the freeze-thaw period, higher contributions of both macro- and meso-fauna were observed at DF stage, and the lower contributions at TS stage. The results suggested that soil fauna have a great contribution to N releases during litter decomposition in the freeze-thaw period, but the contribution was significantly affected by temperature dynamics and the body size of soil fauna.     

Contribution of Soil Fauna to Foliar Litter-Mass Loss in Winter in an Ecotone between Dry Valley and Montane Forest in the Upper Reaches of the Minjiang River

Litter decomposition during winter can provide essential nutrients for plant growth in the subsequent growing season, which plays important role in preventing the expansion of dry areas and maintaining the stability of ecotone ecosystems. However, limited information is currently available on the contributions of soil fauna to litter decomposition during winter in such ecosystems. Therefore, a field experiment that included litterbags with two different mesh sizes (0.04 mm and 3 mm) was conducted to investigate the contribution of soil fauna to the loss of foliar litter mass in winter from November 2013 to April 2014 along the upper reaches of the Minjiang River. Two litter types of the dominant species were selected in each ecosystem: cypress (Cupressus chengiana) and oak (Quercus baronii) in ecotone; cypress (Cupressus chengiana) and clovershrub (Campylotropis macrocarpa) in dry valley; and fir (Abies faxoniana) and birch (Betula albosinensis) in montane forest. Over one winter incubation, foliar litter lost 6.0%-16.1%, 11.4%-26.0%, and 6.4%-8.5% of initial mass in the ecotone, dry valley and montane forest, respectively. Soil fauna showed obvious contributions to the loss of foliar litter mass in all of the ecosystems. The highest contribution (48.5%-56.8%) was observed in the ecotone, and the lowest contribution (0.4%-25.8%) was observed in the montane forest. Compared with other winter periods, thawing period exhibited higher soil fauna contributions to litter mass loss in ecotone and dry valley, but both thawing period and freezing period displayed higher soil fauna contributions in montane forest. Statistical analysis demonstrated that the contribution of soil fauna was significantly correlated with temperature and soil moisture during the winter-long incubation. These results suggest that temperature might be the primary control factor in foliar litter decomposition, but more active soil fauna in the ecotone could contribute more in litter decomposition and its related ecological processes in this region.     

土壤动物对川西亚高山和高山森林凋落叶第一年不同分解时期N和P元素动态的影响

土壤动物对高寒森林凋落物养分元素动态具有重要影响, 但这种影响受控于凋落物质量及环境条件。为了解土壤动物对高寒森林凋落物不同分解时期凋落物中N和P元素动态的影响, 采用凋落物分解袋的方法, 于凋落物第一年分解的不同时期, 即冻结前期、冻结期、融化期、生长季节初期、生长季节中期和生长季节末期, 研究了3.00和0.04 mm孔径凋落物袋中川西亚高山和高山森林的代表性植物——康定柳(Salix paraplesia)、方枝柏(Sabina saltuaria)、红桦(Betula albosinensis)和岷江冷杉(Abies fargesii var. faxoniana)凋落物中的N和P元素动态特征。结果表明: 康定柳和红桦凋落物中的N元素呈现出释放—富集—释放的模式, 方枝柏、岷江冷杉凋落物中的N元素则表现为释放—富集模式; 凋落物P元素总体表现为释放模式, 但4种植物凋落物均在生长季节中期具有明显的富集过程; 从凋落物分解的第一年来看, 土壤动物明显促进了4种植物凋落物N的释放, 而抑制了P的释放; 不同时期土壤动物对凋落物中N和P释放量的影响存在显著差异, 且分别与正积温呈极显著正相关和极显著负相关关系; 相对于阔叶植物凋落物, 土壤动物对针叶植物凋落物中N和P元素动态的影响更为显著。这些结果为深入了解高寒森林生态系统土壤动物与凋落物分解等物质循环过程的相互联系具有重要意义。     

Soil Fauna Affects Dissolved Carbon and Nitrogen in Foliar Litter in Alpine Forest and Alpine Meadow

Dissolved organic carbon (DOC) and total dissolved nitrogen (TDN) are generally considered important active biogeochemical pools of total carbon and nitrogen. Many studies have documented the contributions of soil fauna to litter decomposition, but the effects of the soil fauna on labile substances (i.e., DOC and TDN) in litter during early decomposition are not completely clear. Therefore, a field litterbag experiment was carried out from 13th November 2013 to 23rd October 2014 in an alpine forest and an alpine meadow located on the eastern Tibetan Plateau. Litterbags with different mesh sizes were used to provide access to or prohibit the access of the soil fauna, and the concentrations of DOC and TDN in the foliar litter were measured during the winter (the onset of freezing, deep freezing and thawing stage) and the growing season (early and late). After one year of field incubation, the concentration of DOC in the litter significantly decreased, whereas the TDN concentration in the litter increased. Similar dynamic patterns were detected under the effects of the soil fauna on both DOC and TDN in the litter between the alpine forest and the alpine meadow. The soil fauna showed greater positive effects on decreasing DOC concentration in the litter in the winter than in the growing season. In contrast, the dynamics of TND in the litter were related to seasonal changes in environmental factors, rather than the soil fauna. In addition, the soil fauna promoted a decrease in litter DOC/TDN ratio in both the alpine forest and the alpine meadow throughout the first year of decomposition, except for in the late growing season. These results suggest that the soil fauna can promote decreases in DOC and TDN concentrations in litter, contributing to early litter decomposition in these cold biomes.     

Habitat patchiness affects decomposition and faunal diversity: a microcosm experiment on forest floor

Environmental heterogeneity has been intensively studied, but little is known about relationships between habitat patchiness and soil processes. The aim of this study was to investigate (1) the impact of patchiness of the litter layer on the decomposer community and litter decomposition rate, and (2) whether the impact of soil fauna on the rates of processes differs in relation to patchiness. An experiment was carried out in microcosms with coniferous forest humus and four kinds of litter with different C:N ratios or stages of decomposition, either separately (i.e. in patches) or mixed with each other. Microarthropod species diversity was better maintained in the patchy systems. In the absence of soil fauna, community respiration was higher in the patchy microcosms, but in the presence of fauna the opposite pattern was observed. The contribution of soil fauna to the rate of decomposition was clearly greater in the mixed litter systems. Based on the results, a hypothesis is presented that in the patchy litter layer the soil fungi can create connections between different materials located some centimeters apart, thus enhancing decomposition, while in the mixed litter the scale of millimeters is more appropriate for the soil fauna, known to accelerate the process rates. Received: 10 November 1997 / Accepted: 20 April 1998     

雪被斑块对高山森林两种灌木凋落叶质量损失的影响

高山/亚高山森林灌木层植物凋落物的分解对于系统物质循环等过程具有重要意义, 并可能受到冬季不同厚度雪被斑块下冻融格局的影响。该文采用凋落物分解袋法, 研究了高山森林典型灌层植物华西箭竹(Fargesia nitida)和康定柳(Salix paraplesia)凋落物在沿林窗-林下形成的冬季雪被厚度梯度(厚型雪被斑块、较厚型雪被斑块、中型雪被斑块、薄型雪被斑块、无雪被斑块)上在第一年不同关键时期(冻结初期、冻结期、融化期、生长季节初期和生长季节后期)的质量损失特征。在整个冻融季节, 华西箭竹和康定柳凋落叶的平均质量损失分别占全年的(48.78 ± 2.35)%和(46.60 ± 5.02)%。冻融季节雪被覆盖斑块下凋落叶的失重率表现出厚型雪被斑块大于薄型雪被斑块的趋势,而生长季节无雪被斑块的失重率明显较高。尽管如此, 华西箭竹凋落物第一年分解表现出随冬季雪被厚度增加而增加的趋势, 但康定柳凋落物第一年失重率以薄型雪被斑块最高, 而无雪被斑块最低。同时, 相关分析表明冻融季节凋落叶的失重率与平均温度和负积温呈极显著正相关, 生长季节凋落叶的失重率与所调查的温度因子并无显著相关关系, 但全年凋落物失重率与平均温度和正/负积温均显著相关。这些结果清晰地表明, 未来冬季变暖情境下高山森林冬季雪被格局的改变将显著影响灌层植物凋落物分解, 影响趋势随着物种的差异具有明显差异。