华北落叶松与阔叶树种混合凋落叶分解过程中养分释放和酶活性变化

森林凋落物是森林土壤的重要组成部分,凋落物分解在调控森林生态系统养分循环中发挥了关键作用。采用凋落物分解袋法,研究河北塞罕坝地区华北落叶松与白桦,华北落叶松与蒙古栎,华北落叶松、白桦和蒙古栎混合凋落叶及纯华北落叶松凋落叶分解过程中分解速率、养分释放和酶活性的变化。结果表明: 经过近2年的分解,混合凋落叶分解速率均显著高于纯华北落叶松凋落物叶;在所有处理中,华北落叶松与白桦混合凋落叶分解速率最高。在凋落叶分解过程中,不同处理养分含量变化一致,凋落叶N、P含量呈上升趋势,C、K含量和C/N呈下降趋势;相对纯华北落叶松凋落叶,各混合凋落叶分解可以促进凋落叶C、K的释放,但对N、P的释放有一定的抑制作用。在凋落叶分解过程中,不同处理凋落叶过氧化氢酶、脲酶、酸性磷酸酶活性呈上升趋势,蔗糖酶活性呈下降趋势;凋落叶分解速率与凋落叶过氧化氢酶、脲酶、酸性磷酸酶活性呈正相关,与蔗糖酶活性呈负相关。总体来看,华北落叶松和白桦、蒙古栎凋落叶混合可以促进华北落叶松凋落叶的分解,且凋落叶中酶活性动态变化与凋落叶的分解密切相关。     

Leaf litter decomposition characteristics and controlling factors across two contrasting forest types

两种不同森林类型叶凋落物分解特征及影响因子研究 叶凋落物分解为森林生态系统提供了重要的能量和养分来源。除传统的环境因素外,叶凋落物的降解过程也受到绿叶功能性状和叶凋落物基质质量的影响。然而,在群落水平上,绿叶功能性状和叶凋落物基质质量对不同森林群落叶凋落物分解的相对重要性仍不清楚。因此,本研究以北京东灵山地区7种典型森林群落类型的混合叶凋落物为研究对象,利用分解袋法通过360天的野外相似环境分解实验对叶凋落物的分解过程进行了研究。这些森林群落包括6种分别以胡桃楸(Juglans mandshurica)、青杨(Populus cathayana)、棘皮桦(Betula dahurica)、白桦(Betula platyphylla)、油松(Pinus tabuliformis) 和华北落叶松(Larix gmelinii var. principis-rupprechtii) 为优势种的单优种群落,以及一种以大叶白蜡(Fraxinus rhynchophylla)、蒙古栎(Quercus mongolica)和蒙椴(Tilia mongolica)为优势种的共优种群落。研究结果表明,不同森林群落之间叶凋落物分解速率存在显著差异。群落聚合的植物功能性状和叶凋落物基质质量分别解释了群落叶凋落物分解速率变异的35.60%和9.05%,两者交互作用解释率为23.37%,表明群落聚合的植物功能性状及其与叶凋落物基质质量的共同作用是影响群落叶凋落物分解速率变异的主要因素。通过冗余分析发现,叶片氮含量、叶干物质含量、叶片单宁含量和比叶面积能显著影响群落叶凋落物分解速率的变异。因此,在对群落水平上叶凋落物分解的研究应该关注群落聚合的绿叶功能性状对分解的影响。     

川西亚高山森林凋落物不同分解阶段碳氮磷化学计量特征及种间差异

为了了解亚高山森林凋落物在不同分解阶段的化学计量特征,采用空间代替时间的方法,以自然状态下凋落物的3个层次--新鲜凋落物层（L）、发酵层（F）、腐殖质层（H））模拟凋落物分解的不同阶段,对川西亚高山不同林分类型（岷江冷杉天然林、粗枝云杉人工林、白桦天然林、杜鹃矮曲林）凋落物的碳氮磷（C、N、P）及可溶性碳氮磷（DOC、SN、SP）含量进行研究。结果表明：林分类型及分解阶段将显著影响凋落物分解过程中的碳氮磷含量及其化学计量比。亚高山森林凋落物可溶性有机碳、水溶性磷含量均随着分解过程的不断进行而降低,分解初期快速淋溶,而分解中后期释放变缓。4种林分比较而言,水溶性碳氮磷含量表现为：白桦 > 杜鹃 > 冷杉 > 云杉,阔叶树种凋落物的可溶性碳氮磷普遍高于针叶树种,尤其在分解初期。针叶树种凋落物SN在分解过程中呈现释放模式,而阔叶树种SN呈现先富集后释放模式。凋落物C含量随着分解的不断进行而降低,冷杉、白桦及杜鹃N含量呈现先富集后释放的趋势,分解阶段对云杉与白桦各层P含量影响不显著,但冷杉却呈现先降低后升高的现象,而杜鹃则是在分解后期P含量显著降低。从总体来看,亚高山森林凋落物C/P和N/P均显著小于全球平均水平,凋落物C/N、C/P、N/P、DOC/C、SN/N、SP/P均随着分解的不断进行呈现降低的趋势。分解初期白桦和杜鹃DOC/C显著降低,而冷杉则在分解后期显著降低。冷杉N/P先升高后降低,杜鹃N/P随着分解的不断进行呈现升高趋势。这些研究结果为深入理解亚高山森林生态系统的凋落物分解进程和养分循环提供了依据。     

樟树人工林凋落物养分含量及归还量对氮沉降的响应

氮沉降的持续增加对陆地生态系统的健康发展构成严重威胁,森林是陆地生态系统中重要的组成部分,大量的氮沉降对其结构和功能造成严重影响。凋落物是森林生态系统养分循环的重要组成部分,它对土壤肥力、森林生态系统养分循环等方面具有重要作用。为了探讨亚热带常绿阔叶森林凋落物对氮沉降增加的响应,在湖南省森林植物园以樟树人工林为研究对象进行模拟氮沉降的实验,实验设置4种氮添加水平CK(0g N m~(-2)a~(-1),对照)、LN(5g N m~(-2)a~(-1)),MN(15g N m~(-2)a~(-1)),HN(30g N m~(-2)a~(-1)),研究氮沉降对樟树林年凋落物量、凋落物养分含量以及归还量的影响。结果表明:不同施氮水平下(CK、LN、MN、HN),樟树林凋落物的年凋落量分别为(4.53±0.32)t hm~(-2)a~(-1)、(3.95±0.28)t hm~(-2)a~(-1)、(3.56±0.41)t hm~(-2)a~(-1)、(4.46±0.48)t hm~(-2)a~(-1),施氮抑制了樟树林的凋落量,且低、中氮处理下差异显著(P0.05);施氮处理后凋落物的养分含量大小顺序为:CNCaKMg,凋落物的碳含量没有显著变化,但氮含量都有所增加,因此,施氮降低了樟树凋落物各组分的C/N比;凋落物中元素的年归还量大小顺序表现为:CNCaKMg,施氮处理对凋落物C、K、Ca、Mg归还量有抑制作用,但对凋落物N归还量表现为促进作用。     

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。     

神农架常绿落叶阔叶混交林凋落物养分特征

凋落物是联结陆地生态系统植物与土壤养分的重要媒介,了解凋落物养分特征有助于理解陆地生态系统物质循环的机理。该研究于2015年收集了神农架地区常绿落叶阔叶混交林的新鲜凋落物及现存凋落物,测定其不同器官中大量元素(C、N、P、K、Ca、Mg)的含量,据此分析其养分含量、养分归还量、养分储量及化学计量比的特征。结果发现:该常绿落叶阔叶混交林新鲜凋落物的C、K养分含量显著高于现存凋落物,N、P、Ca、Mg养分含量显著低于现存凋落物;其凋落物大量元素的养分归还量及养分储量大小顺序均为C Ca N Mg K P,分别为1569.84、52.44、34.82、6.24、5.24、1.30 kg hm~(-2) a~(-1)及1835.29、87.87、51.17、12.12、3.90、1.95 kg hm~(-2) a~(-1);其新鲜凋落物及现存凋落物的C∶N∶P分别为1307.33∶27.73∶1及976.48∶26.77∶1,新鲜凋落物的C∶N、C∶P显著高于现存凋落物,N∶P无显著区别。研究表明,新鲜凋落物与现存凋落物养分含量之间的差异与不同元素在分解过程中的可淋溶性及生物固持等因素有关。该地区常绿落叶阔叶混交林凋落物养分归还量及养分储量相对于亚热带阔叶林平均水平较低;且显著低于喀斯特地区同类型森林,主要与其凋落物产量、降水量及植被类型有关。该森林生态系统新鲜凋落叶与中国及全球范围内阔叶树种凋落叶相比C∶N较低,C∶P、N∶P较高,这可能是由于该地区N沉降及P限制现象较为严重所致。     

细菌对五大连池火山森林凋落物早期分解的影响

为了解火山森林生态系统中微生物在凋落物分解和营养循环中的作用，以五大连池火山熔岩台地森林优势树种落叶松、白桦、山杨为研究对象，采用凋落物袋野外原位分解和高通量测序方法，分析细菌群落结构及多样性对凋落物分解的响应。结果表明：经过18个月的分解，3种植物叶片凋落物质量残留率为63.9%～68.1%,白桦叶片凋落物分解最快，分解前后的N含量、C∶N、N∶P存在显著差异。3种植物叶片凋落物的细菌物种数和多样性指数具有显著差异。变形菌门、放线菌门、拟杆菌门为优势菌门，根瘤菌属、鞘氨醇单胞菌属、假单胞菌属为优势菌属，且在3种凋落物间存在显著差异。分解18个月后，凋落物中优势细菌类群与火山熔岩台地土壤趋向一致，火山森林生态系统中凋落物细菌多样性和群落结构受到凋落物中P含量、C∶N和N∶P的影响。     

雪被期川西高山林线交错带凋落物可溶性有机碳的淋失特征

凋落物可溶性有机碳(dissolved organic carbon,DOC)是土壤DOC的主要来源之一,同时,凋落物DOC的淋失过程是生态系统碳循环的重要环节。本文采用凋落物分解袋法,于2012年10月—2013年5月,以川西高山林线交错带12种代表性植物的凋落物为材料,研究了冬季雪被期凋落物DOC在暗针叶林森林、高山林线和高山草甸3种分解生境中的淋失特征。结果表明:1)凋落物初始DOC含量为22.78~178.8 g·kg-1,物种间差异显著;2)经过一个雪被期(182 d),凋落物DOC淋失率为18.92%~62.33%,各物种DOC的淋失率对分解生境的响应差异较大;3)分解生境差异对凋落物DOC的淋失产生了显著影响,暗针叶林生境中凋落物DOC的淋失率显著高于林线和高山草甸生境;4)雪被是影响冬季高山林线交错带凋落物DOC淋失的主要环境因素。因此,高山林线交错上雪被期DOC淋失的空间变异性主要是景观尺度分布的雪被差异造成的,分解生境中较厚的雪被更利于凋落物DOC的淋失。     

核磁共振技术在森林凋落物分解研究中的应用

凋落物分解是连接森林生态系统地上和地下部分的关键过程,影响着森林生态系统的养分循环,对全球碳(C)循环和森林生态系统生产力具有重要的调控作用。特别是,在当前全球气候变化的背景下,研究森林凋落物不同有机C组分的分解动态对于了解土壤有机质的形成、稳定性及其对气候变化的反馈效应至关重要。核磁共振技术(NMR)具有无损和非侵入等特点,是研究有机化合物结构的有效手段,已在凋落物分解研究中得到广泛应用。本文概述了NMR技术应用于森林凋落物分解过程的最新研究进展,包括森林凋落物有机C基团的NMR特征,凋落物分解过程中不同C基团的变化特征及其机理、凋落物分解过程的NMR评价指标等,并分析了NMR技术在凋落物分解研究中存在的问题和未来研究方向。NMR技术将有助于了解凋落物有机C基团的分解特征及其对土壤有机质形成和稳定的影响机理。     

凋落物输入对三江平原弃耕农田土壤基础呼吸和活性碳组分的影响

土壤有机碳的积累主要由土壤有机质的输入与输出间的净平衡决定的,植被的恢复和凋落物质的大量输入是土壤恢复的先决条件,凋落物的输入在土壤恢复过程中起着至关重要的作用.通过对不同类型凋落物输入到三江平原弃耕农田后土壤的基础呼吸、溶解有机碳(DOC)和土壤微生物量碳(MBC)的研究表明:相同种类凋落物输入后,输入到土壤总有机碳(TOC)背景值低的凋落物被微生物降解的速率大于TOC背景值高的土壤,TOC较低的土壤能够加快微生物对输入凋落物的分解,不利于有机质的积累;不同类型凋落物的输入使土壤基础呼吸、DOC和MBC等活性组分的生成和降解产生差异,改变了凋落物的降解速率,在三江平原研究的4种主要植被类型中,人工林凋落物最容易降解,小叶章、大豆的降解能力次之,玉米是最难降解的凋落物.     

黄土丘陵区刺槐人工林林龄增加土壤微环境变化对凋落物分解的影响

凋落物所处的土壤微环境是影响其分解的关键因素之一,然而在黄土高原广泛栽植的刺槐人工林中,土壤微环境随林龄增加如何改变、其对凋落物分解过程的影响趋势尚不清楚。为明确上述问题,以油松凋落物(典型的难分解凋落物)和白三叶凋落物(易分解)为对象,分别在林龄为10、20、33 a和43 a的刺槐林地土壤表面进行为期592 d的模拟分解试验,检测凋落物分解特征以及地表土壤理化生物学性质随林龄增加的变化趋势,并分析凋落物分解速率与土壤微环境指标间的关系。结果表明:(1)随林龄增加,油松凋落物的分解速率呈先小幅降低后提高的趋势,白三叶凋落物的分解速率持续提高(P<0.05);(2)总体而言,随林龄增加林地表层土壤温度呈先降后增趋势,土壤湿度、有效磷含量和pH持续降低,而速效氮含量持续提高(P<0.05);(3)林龄增加显著改变了林地土壤微生物群落结构,特别是在各分解时间点时均导致真菌属的明显演替现象。土壤中9种凋落物分解酶的总酶活性和木质纤维素酶活性均在分解第108天时随林龄增加呈先降后增趋势,而在分解第389天和第592天时持续提高(P<0.05)。(4)油松凋落物分解速率仅与土壤总酶活性、真菌群落结构和铵态氮含量呈显著正相关,白三叶凋落物分解速率则与总酶和木质纤维素酶活性、细菌和真菌群落结构、温度和铵态氮含量显著正相关,而与土壤湿度和pH显著负相关(P<0.05)。综上所述,刺槐林龄增加引起的土壤理、化和生物微环境变化总体倾向于加速凋落物的分解过程。     

杉木和香樟酸雨酸解底物的分解格局

采用凋落物分解袋法,选取湘西地区两种人工林优势树种(香樟和杉木)的凋落叶作为分解材料,分析了两种凋落叶经酸解处理后凋落物分解及其微生物活性的变化。结果表明:酸解处理过程会使两种凋落叶损失一定的质量,随着酸解强度的增加质量损失增加,且酸解处理对香樟凋落物质量损失的影响较杉木凋落物大。不同物种凋落物对酸解强度的差异性反应产生了后续分解过程的差异格局:酸雨酸解作用的增强抑制了杉木凋落物分解过程中包括真菌生物量以及纤维素酶与木质素酶在内的微生物活性;而对于香樟凋落物分解过程,微生物活性对酸雨酸解的响应因变量不同、分解期不同而存在差异性。两物种凋落物的总失重率、木质素和纤维素分解率对酸解作用的响应及其在不同分解期的表现也存在差异性:对于杉木凋落物,在分解前期其失重率表现为T1T2T3,而在后期随酸解强度的增大而升高,即T3T2/T1;香樟凋落物在分解的前期(T1T2T3)与后期(T1T2T3)情况则正好与杉木凋落物相反。总之,酸雨酸解凋落物不仅使底物有机组成发生了变化,在一定程度上导致凋落物物理结构紧密程度改变,而且也可能相应地改变了凋落物定殖微生物群落,这些复合影响从不同程度上决定了凋落物分解及其微生物活性对凋落物底物酸解的响应。     

亚热带不同植被恢复阶段林地凋落物层现存量和养分特征

为揭示亚热带森林植被自然恢复过程中,凋落物层现存量及其养分元素储存能力的演变,采用空间代替时间的方法,在位于亚热带丘陵区的长沙县选取地域相邻、生境条件基本一致的檵木+南烛+杜鹃灌草丛（Loropetalum chinense+Vaccinium bracteatum +Rhododendron simsii scrub-grass-land,LVR）、檵木+杉木+白栎灌木林（L.chinense+Cunninghamia lanceolata+Quercus fabri shrubbery,LCQ）、马尾松+柯+檵木针阔混交林（Pinus massoniana +Lithocarpus glaber +L.chinense coniferous-broad leaved mixed forest,PLL）、柯+红淡比+青冈常绿阔叶林（L.glaber+Cleyera japonica+Cyclobalanopsis glauca evergreen broad-leaved forest,LAG）作为一个恢复序列,设置固定样地,采集未分解层（U层）、半分解层（S层）、已分解层（D层）凋落物样品,测定凋落物层现存量和主要养分元素含量、储量及其释放率,分析植物多样性指数与凋落物层现存量、养分元素含量的相关性。结果表明：1）凋落物层及各分解层凋落物现存量随着植被恢复而增加;同一恢复阶段D层凋落物现存量最高,占凋落物层现存量的41.59%-51.02%,不同分解层凋落物现存量的差异随着植被恢复而增大;各恢复阶段凋落物分解率为0.44-0.61,周转期为1.65-2.28 a。2）凋落物层及各分解层凋落物主要养分元素含量均表现为：N > Ca > Mg > K > P,随着植被恢复呈现出不同的变化特征,其中N、P含量总体上呈增加趋势,K含量LAG（除U层外）最高,PLL最低,Ca含量LCQ最高,PLL最低,Mg含量LAG（除U层外）最高,LVR最低;同一恢复阶段N、P（除PLL、LAG外）、K、Ca、Mg含量随着凋落物的分解而下降。3）不同恢复阶段凋落物层主要养分元素的储量依次为：N > Ca > Mg > K > P;凋落物层及各分解层凋落物主要养分元素总储量及各种养分元素的储量总体上随着植被恢复而增加;同一恢复阶段随着凋落物的分解,N、P储量增加,而K、Ca、Mg储量变化不大;随着植被恢复,凋落物层养分元素储存能力和转化归还能力提高,特别是N,养分元素总释放率下降,有利于养分的固持。4）乔木层、灌木层、草本层的植物多样性指数对凋落物层现存量和主要养分元素含量的影响不同,其中乔木层的影响最明显。     

Effects of temperature and litter type on fungal growth and decomposition of leaf litter

The dependence of fungal decomposition of leaf litter on incubation temperature and litter types used as substrata was assessed under pure culture conditions. Isolates of Xylaria sp., a major ligninolytic fungus in cool temperate forests in Japan, were used as the fungal material. Xylaria sp. is mesophilic; maximum growth and decomposition occurred at 25°C. In the temperature test, the decomposition pattern of beech leaf litter by three isolates of Xylaria sp. changed at a threshold at 25°C. Cellulolytic activity increased with temperature from 5 to 25°C, whereas above 25°C ligninolytic activity increased at the expense of cellulolytic activity, leading to suppressed overall decomposition as a result of the higher temperature. The mass loss of leaf litter caused at 20°C by an isolate of Xylaria sp. was variable among 15 litter types and was correlated negatively with acid-unhydrolyzable residue (AUR) content and positively with total carbohydrate content for the 15 litter types. The effects of temperature and litter type on the growth and decomposition of leaf litter by Xylaria sp. may have implications for changes in fungal decomposition of leaf litter that would be predicted in response to future environmental changes.     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

Decomposition of tree leaf litters grown under elevated CO2: Effect of litter quality

Ash (Fraxinus excelsior L.), birch (Betula pubescens Ehrh.), sycamore (Acer pseudoplatanus L.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) leaf litters were monitored for decomposition rates and nutrient release in a laboratory microcosm experiment. Litters were derived from solar domes where plants had been exposed to two different CO₂ regimes: ambient (350 L L^-1 CO₂) and enriched (600 L L^-1 CO₂).Elevated CO₂ significantly affected some of the major litter quality parameters, with lower N, higher lignin concentrations and higher ratios of C/N and lignin/N for litters derived from enriched CO₂. Respiration rates of the deciduous species were significantly decreased for litters grown under elevated CO₂, and reductions in mass loss at the end of the experiment were generally observed in litters derived from the 600 ppm CO₂ treatment. Nutrient mineralization, dissolved organic carbon, and pH in microcosm leachates did not differ significantly between the two CO₂ treatments for any of the species studied. Litter quality parameters were examined for correlations with cumulative respiration and decomposition rates: N concentration, C/N and lignin/N ratios showed the highest correlations, with differences between litter types. The results indicate that higher C storage will occur in soil as a consequence of litter quality changes resulting from higher atmospheric concentrations of CO₂.Abbreviations CHO soluble carbohydrates - DOC dissolved organic carbon - HCel holocellulose - WTREM weight remaining     

Chemical composition of litter affects the growth and enzyme production by the saprotrophic basidiomycete Hypholoma fasciculare

Chemical composition of litter has previously been reported to affect in situ decomposition. To identify its effects on a single species level, the saprotrophic basidiomycete Hypholoma fasciculare was grown on 11 types of litter with variable chemical composition (N content of 3.4–28.9 mg g⁻¹), and the mass loss of litter and lignin, production of extracellular enzymes and fungal biomass were followed. After 12 weeks, mass loss ranged from 16 % to 34 %. During early decomposition stages, litter mass loss, fungal biomass production (estimated by ergosterol content) as well as fungal substrate use efficiency all increased with increasing initial N content of the litter. The initial litter decomposition rate was significantly positively correlated with the activities of arylsulfatase, cellobiohydrolase, endoxylanase and phosphatase. Contrary to expectations, the lignin content did not affect litter mass loss, when covariation with N content was accounted for. The ratio of lignin loss to total mass loss depended on the litter type and did not reflect the activities of ligninolytic enzymes.     

Response of dissolved organic matter in the forest floor to long-term manipulation of litter and throughfall inputs

Dissolved organic matter (DOM) contributes to organic carbon either stored in mineral soil horizons or exported to the hydrosphere. However, the main controls of DOM dynamics are still under debate. We studied fresh leaf litter and more decomposed organic material as the main sources of DOM exported from the forest floor of a mixed beech/oak forest in Germany. In the field we doubled and excluded aboveground litter input and doubled the input of throughfall. From 1999 to 2005 we measured concentrations and fluxes of dissolved organic C and N (DOC, DON) beneath the Oi and Oe/Oa horizon. DOM composition was traced by UV and fluorescence spectroscopy. In selected DOM samples we analyzed the concentrations of phenols, pentoses and hexoses, and lignin-derived phenols by CuO oxidation. DOC and DON concentrations and fluxes almost doubled instantaneously in both horizons of the forest floor by doubling the litter input and DOC concentrations averaged 82 mg C l⁻¹ in the Oe/Oa horizon. Properties of DOM did not suggest a change of the main DOM source towards fresh litter. In turn, increasing ratios of hexoses to pentoses and a larger content of lignin-derived phenols in the Oe/Oa horizon of the Double litter plots in comparison to the Control plots indicated a priming effect: Addition of fresh litter stimulated microbial activity resulting in increased microbial production of DOM from organic material already stored in Oe/Oa horizons. Exclusion of litter input resulted in an immediate decrease in DOC concentrations and fluxes in the thin Oi horizon. In the Oe/Oa horizon DOC concentrations started to decline in the third year and were significantly smaller than those in the Control after 5 years. Properties of DOM indicated an increased proportion of microbially and throughfall derived compounds after exclusion of litter inputs. Dissolved organic N did not decrease upon litter exclusion. We assume a microbial transformation of mineral N from throughfall and N mineralization to DON. Increased amounts of throughfall resulted in almost equivalently increased DOC fluxes in the Oe/Oa horizon. However, long-term additional throughfall inputs resulted in significantly declining DOC concentrations over time. We conclude that DOM leaving the forest floor derives mainly from decomposed organic material stored in Oe/Oa horizons. Leaching of organic matter from fresh litter is of less importance. Observed effects of litter manipulations strongly depend on time and the stocks of organic matter in forest floor horizons. Long-term experiments are particularly necessary in soils/horizons with large stocks of organic matter and in studies focusing on effects of declined substrate availability. The expected increased primary production upon climate change with subsequently enhanced litter input may result in an increased production of DOM from organic soil horizons.     

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

Leaf litter identity alters the timing of lotic nutrient dynamics

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