土壤氮素内循环对生态覆被变化响应的研究进展*

杨莉琳, 姚琦馥, 梁 琍, 鲁小明

土壤氮素内循环对生态覆被变化响应的研究进展*

杨莉琳, 姚琦馥, 梁 琍, 鲁小明

(铜仁学院 铜仁 554300)

生态覆被; 土地利用变化; 活性氮; 氮转化; 土壤微生物

1 土壤活性氮及其环境效应

活性氮是相对于占地球大气总量79%的非活性氮(N2)而言。由于N2是稳定态氮素, 这个巨大的氮贮库并不能被自然界生物直接利用。只有将N2的分子键破坏, 得到的单个氮原子通过固氮作用或同化作用与其他营养元素如O、H或C相结合, 才能被大多数植物、动物和微生物利用[1]。活性氮直接关系到植被生长、气候变化和生态环境的安全与持续。

工业革命前自然界循环的活性氮总量很少, 人类对自然生态系统干预较少, 生物固氮是植物获得活性氮的主要途径, 植物生产力主要受土壤中氮量的限制。工业革命后, 尤其是近50年来, 工业合成铵和化肥在全球大量施用成为土壤活性氮的主要来源, 土壤因而成了人为活性氮的天然汇。全球人为活性氮在1860年只有15 Tg∙a–1(1 Tg=106t), 到了21世纪初已增加到165 Tg∙a–1, 增长了10倍, 其中来自于粮食生产的活性氮是能源生产的5倍。然而, 粮食生产的活性氮中人类每年只摄取大约12 Tg [每人2 kg(N)∙a–1], 其余近90%的活性氮以硝酸盐、亚硝酸盐或氮氧化物的形态排放到土壤、水体和大气等环境中[2-3]。这些活性氮一旦进入环境, 就会通过各生态系统迅速串联并蔓延开来, 在环境中积累或循环, 并影响着全球生态环境和许多生态过程[2]。如以NO (NO=NO+NO2)形式排放到大气中产生温室效应, 引起全球气候变暖[4]; 生物多样性丧失和亲氮杂草侵入; 生态系统功能发生改变; 之前受活性氮限制的生态系统生产力增加; 森林等系统土壤中氮饱和、水体污染、有毒藻类繁盛、鱼类死亡以及沿海生态系统富营养化; 通过食物链人类易患高铁血红蛋白血症、蓝婴儿综合症、癌症、呼吸系统和心脏疾病等。因此, 活性氮在土壤圈-生物圈-大气圈之间循环的改变与失衡, 对各环节中氮水平维持在适当范围内提出了严格的限制, 也是生态与环境学家、管理工作者和政策制定者的关注重点[5]。

2 生态覆被/土地利用变化对土壤氮库的影响

2.1 不同生态系统的土壤氮库差异分析

自然生态系统中, 不同覆被类型的土壤氮库差异很大。森林不仅影响全球气候的碳汇, 而且还是世界上最大的有机氮库[7]。特别是古森林, 其碎屑生物量和微生物固氮作用比成熟的次生林更能保留氮,是强大的氮汇[8]。尤其是土壤风化程度高的热带林区, 有机质及氮库在土壤功能和森林可持续性方面发挥着重要作用。当森林被清除时, 土壤有机质与有机氮几乎立即开始失去, 并引发一系列土壤退化[9]。

2.2 生态覆被/土地利用变化对土壤氮库的影响

3 土壤氮素内循环对生态覆被/土地利用变化的响应

3.1 人为扰动对自然生态系统土壤氮素内循环产物的影响

天然森林生态系统的净矿化和净硝化均很低, 天然林通常受土壤低氮供应的限制, 氮素处于封闭、积累型循环, 地上部每年的净初级生产力与土壤净氮矿化呈线性正相关[29]。美国北部的天然草地转换为森林35~75 a的土壤氮矿化和微生物活性基本没有变化[30]。因为氮损失与氮净矿化率和枯枝落叶氮通量(植物氮循环指标)密切相关, 尽管原始林和次生林的氮量没有显著差异, 但是原始森林的微生物固氮速度比次生林快, 原始森林的粗木屑和微生物量氮的吸收和转化也比次生林大。美国密西根州半岛西部原始阔叶林总氮矿化大约是次生林的2倍, 而总硝化没有差异[8]。

3.2 生态修复对土壤氮库消长的影响

3.2.1 退耕还林还草对土壤氮库积累的效应

植被种类直接影响生态系统的修复速率。在遗弃50 a以上的荒地上种植阔叶林、针叶林、牧场和种植园, 会显著影响土壤氮含量和硝化速率[20]。我国黄土高原退耕还林还草30 a后, 恢复的草原、刺槐()、红松()、油松()以及油松-紫穗槐()混交林等土壤有机质、全氮、有效氮均提高了2倍多, 其中, 刺槐-紫穗槐混合林恢复最迅速, 油松-紫穗槐混交林恢复较慢[45]。晋西黄土丘陵区退耕还林或撂荒地还林还草后, 荒草地土壤全氮表聚效应最强, 0~48 cm土层中以刺槐林地的全氮含量提升最快[46]。黄土高原的退化草地分别栽种油松和柠条()30多年后, 油松地0~20 cm土层有效氮减少, 而柠条灌丛林地的有效氮却增加[47]。在巴西里约热内卢州沿海城镇的退化土地上, 于1991年种植了7种速生的先锋豆科植物用以恢复退化土壤的肥力, 13 a中土壤氮库的年增长率为0.13 Mg×hm-2[9]。豆类植物能增加土壤氮和碳的积累, C3植物和杂草会降低氮和碳的积累率, C4植物提高了土壤C/N比[28]。

尽管种植林草是增加土壤对碳、氮固定的途径, 但生态系统恢复重建绝非易事, 正所谓毁坏容易修复难。明尼苏达州废弃61 a的农田要恢复到之前耕作时95%的水平, 氮库需要180 a, 碳库需要230 a, 且碳的积累速率受氮积累的影响[28]。也有研究结果报道, 在50 a内可以成功恢复严重退化的森林土壤氮的有效性, 但如果在恢复过程中继续收获林下植被和凋落物, 则难以实现这种恢复速率和恢复水平[15]。

3.2.2 恢复重建土壤氮碳库的其他措施

相对于单一植被系统, 农林复合系统对生态的恢复效果一直被广泛推崇。可可()林/农复合系统的土壤碳氮贮量低于天然林, 但高于单一的可可林, 在维持生态系统功能服务方面比单一农业种植更好, 并接近天然森林[48]。尤其是在障碍性土壤区域, 实施农林复合系统是克服土壤障碍, 恢复生态功能的重要途径。我国华北低平原盐碱地区枣()/冬小麦()-夏玉米间作22 a以上的研究表明, 枣/粮间作系统的种植和生态效益仍然比单作农业和单作枣树高[49]。

休耕和轮作也是恢复土壤有机质和氮含量的有效措施[50]。对亚马逊河流域多样化的森林研究表明, 土壤修复除了修复物或施用化肥, 还可以通过延长田间使用时间、缩短休耕期来提升土壤肥力, 提高生物多样性[51]。

3.3 土壤氮素内循环对生态修复的响应

4 土壤微生物对生态覆被/土地利用变化的响应

4.1 生态系统土壤微生物量与氮矿化的关系

土壤微生物是氮素等养分元素循环的引擎, 氨化作用、固氮作用、硝化作用和反硝化作用构成土壤氮循环的主要环节, 且每一个过程都需要相应微生物参与。土壤微生物群落结构控制了不同生态系统中的氮素转化, 进而调节生态系统的功能与稳定。自然森林土壤中有较高的微生物量氮, 净氮矿化相对较少[31], 因此森林系统常受供氮量低的限制。外生菌根等微生物也是森林系统的重要氮源, 澳大利亚国家公园强酸且贫瘠土壤上生长着极度濒危的Wollemi松树, 高度依赖于其根部一个独特的细菌群落[55]。可见不同森林的土壤微生物量差异很大。小兴安岭6种森林类型的土壤微生物碳与微生物氮的大小顺序依次为: 次生白桦()林>人工红松林>择伐林>阔叶红松林>人工落叶松()林>谷地云冷杉()林, 总体表现为阔叶林(次生白桦林、阔叶红松林和择伐林)的土壤微生物量高于针叶林或针叶树占比较高的森林类型[56]。

地上部植被和土壤环境均对土壤微生物生物量氮和氮矿化有影响[37]。幼龄草甸土壤微生物生物量氮较低, 随着年限增加, 微生物氮与全氮比(Nmic/TN)增加, 土壤总氮矿化率下降, 氮对植物的有效性降低, 植物产量下降[57]。

4.2 土壤微生物对生态覆被/土地利用变化的响应

城市化过程对土壤氮循环的关键微生物影响很大。城市草坪土壤中的氨氧化古菌(AOA)丰度高于郊区和农村的农田土壤, AOA对城市草坪土壤硝化起关键作用。城市草坪土壤中的根瘤菌、变形杆菌()和绿弯菌()也比农田土壤丰富, 但城市草坪土壤AOB和反硝化细菌nirS, nosZ)的相对丰度低于郊区草坪和农田[27]。

4.3 土壤氮素内循环对生态覆被/土地利用变化响应的研究方法展望

由于土壤环境的多样性和土壤微生物的复杂性,导致生态覆被/土地利用变化与土壤氮循环过程的效应至今难有确定结论。在生态覆被/土地利用加剧和全球气候变暖趋势下, 对这一科学问题的探索仍是研究热点。其瓶颈仍然在于土壤微生物, 因为土壤中多数微生物在休眠状态下长时间存活, 休眠期间细胞活性很低甚至没有活性。传统的平板培养法无法将它们分离出来。目前常用的熏蒸法测定土壤微生物碳和微生物氮误差非常大, 结果重现性差。运用分子生物学技术研究土壤微生物与土壤氮素循环之间的关系是目前的方向, 但是, 微生物群落的数量(丰度)以及多样性指数(如Shannon和Simpson指数)难以揭示复杂的微生物群落结构与功能微生物之间量化关系[63-65]。特别是找出微生物群落中控制生态功能的关键物种是本方向研究中的一个难点。近年来, 利用高通量基因芯片数据和微生物群落的生态网络分析方法是提升土壤微生物群落结构研究定量化和可视化新方向。

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Research progress on soil nitrogen internal cycling response to ecological cover change*

YANG Lilin, YAO Qifu, LIANG Li, LU Xiaoming

(Tongren University, Tongren 554300, China)

Ecosystem cover; Land use change; Active N; Nitrogen cycling; Soil microbe

S153

10.13930/j.cnki.cjea.190908

杨莉琳, 姚琦馥, 梁琍, 鲁小明. 土壤氮素内循环对生态覆被变化响应的研究进展[J]. 中国生态农业学报(中英文), 2020, 28(10): 1543-1550

YANG L L, YAO Q F, LIANG L, LU X M. Research progress on soil nitrogen internal cycling response to ecological cover change[J]. Chinese Journal of Eco-Agriculture, 2020, 28(10): 1543-1550

* 国家自然科学基金项目(31270521)、贵州省教育厅创新群体重大研究项目(黔教合KY字[2016]053号)、贵州省科技计划项目(黔科合基础[2019]1312)、贵州省创新人才团队(黔教合人才团队字[2015]67号)、铜仁学院博士基金项目(trxyDH1525)和农业生态创新团队(CXTD[2020-10])资助

杨莉琳, 主要研究方向为土壤养分循环与环境生态。E-mail: yangllin@sjziam.ac.cn

2019-12-24

2020-04-09

* This study was supported by the National Natural Sciences Foundation of China (31270521), the Major Research Project of Innovation Group for Guizhou Education Department (Qian Education NO. [2016] 053th), the Science and Technology Plan Project for Guizhou Province (Qian Science NO.[2019] 1312), the Guizhou Innovation Talent Group (Qian Education NO. [2015] 67th), the Doctoral Fund Project for Tongren University (trxyDH1525), and Agro-ecological Innovation Research Group (CXTD[2020-10]).

, YANG Lilin, E-mail: yangllin@sjziam.ac.cn

Dec. 24, 2019;

Apr. 9, 2020