楊莉琳, 姚琦馥, 梁 琍, 魯小明

土壤氮素內(nèi)循環(huán)對(duì)生態(tài)覆被變化響應(yīng)的研究進(jìn)展*

楊莉琳, 姚琦馥, 梁 琍, 魯小明

(銅仁學(xué)院 銅仁 554300)

生態(tài)覆被; 土地利用變化; 活性氮; 氮轉(zhuǎn)化; 土壤微生物

1 土壤活性氮及其環(huán)境效應(yīng)

活性氮是相對(duì)于占地球大氣總量79%的非活性氮(N2)而言。由于N2是穩(wěn)定態(tài)氮素, 這個(gè)巨大的氮貯庫(kù)并不能被自然界生物直接利用。只有將N2的分子鍵破壞, 得到的單個(gè)氮原子通過(guò)固氮作用或同化作用與其他營(yíng)養(yǎng)元素如O、H或C相結(jié)合, 才能被大多數(shù)植物、動(dòng)物和微生物利用[1]?；钚缘苯雨P(guān)系到植被生長(zhǎng)、氣候變化和生態(tài)環(huán)境的安全與持續(xù)。

工業(yè)革命前自然界循環(huán)的活性氮總量很少, 人類(lèi)對(duì)自然生態(tài)系統(tǒng)干預(yù)較少, 生物固氮是植物獲得活性氮的主要途徑, 植物生產(chǎn)力主要受土壤中氮量的限制。工業(yè)革命后, 尤其是近50年來(lái), 工業(yè)合成銨和化肥在全球大量施用成為土壤活性氮的主要來(lái)源, 土壤因而成了人為活性氮的天然匯。全球人為活性氮在1860年只有15 Tg?a–1(1 Tg=106t), 到了21世紀(jì)初已增加到165 Tg?a–1, 增長(zhǎng)了10倍, 其中來(lái)自于糧食生產(chǎn)的活性氮是能源生產(chǎn)的5倍。然而, 糧食生產(chǎn)的活性氮中人類(lèi)每年只攝取大約12 Tg [每人2 kg(N)?a–1], 其余近90%的活性氮以硝酸鹽、亞硝酸鹽或氮氧化物的形態(tài)排放到土壤、水體和大氣等環(huán)境中[2-3]。這些活性氮一旦進(jìn)入環(huán)境, 就會(huì)通過(guò)各生態(tài)系統(tǒng)迅速串聯(lián)并蔓延開(kāi)來(lái), 在環(huán)境中積累或循環(huán), 并影響著全球生態(tài)環(huán)境和許多生態(tài)過(guò)程[2]。如以NO (NO=NO+NO2)形式排放到大氣中產(chǎn)生溫室效應(yīng), 引起全球氣候變暖[4]; 生物多樣性喪失和親氮雜草侵入; 生態(tài)系統(tǒng)功能發(fā)生改變; 之前受活性氮限制的生態(tài)系統(tǒng)生產(chǎn)力增加; 森林等系統(tǒng)土壤中氮飽和、水體污染、有毒藻類(lèi)繁盛、魚(yú)類(lèi)死亡以及沿海生態(tài)系統(tǒng)富營(yíng)養(yǎng)化; 通過(guò)食物鏈人類(lèi)易患高鐵血紅蛋白血癥、藍(lán)嬰兒綜合癥、癌癥、呼吸系統(tǒng)和心臟疾病等。因此, 活性氮在土壤圈-生物圈-大氣圈之間循環(huán)的改變與失衡, 對(duì)各環(huán)節(jié)中氮水平維持在適當(dāng)范圍內(nèi)提出了嚴(yán)格的限制, 也是生態(tài)與環(huán)境學(xué)家、管理工作者和政策制定者的關(guān)注重點(diǎn)[5]。

2 生態(tài)覆被/土地利用變化對(duì)土壤氮庫(kù)的影響

2.1 不同生態(tài)系統(tǒng)的土壤氮庫(kù)差異分析

自然生態(tài)系統(tǒng)中, 不同覆被類(lèi)型的土壤氮庫(kù)差異很大。森林不僅影響全球氣候的碳匯, 而且還是世界上最大的有機(jī)氮庫(kù)[7]。特別是古森林, 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,是強(qiáng)大的氮匯[8]。尤其是土壤風(fēng)化程度高的熱帶林區(qū), 有機(jī)質(zhì)及氮庫(kù)在土壤功能和森林可持續(xù)性方面發(fā)揮著重要作用。當(dāng)森林被清除時(shí), 土壤有機(jī)質(zhì)與有機(jī)氮幾乎立即開(kāi)始失去, 并引發(fā)一系列土壤退化[9]。

2.2 生態(tài)覆被/土地利用變化對(duì)土壤氮庫(kù)的影響

3 土壤氮素內(nèi)循環(huán)對(duì)生態(tài)覆被/土地利用變化的響應(yīng)

3.1 人為擾動(dòng)對(duì)自然生態(tài)系統(tǒng)土壤氮素內(nèi)循環(huán)產(chǎn)物的影響

天然森林生態(tài)系統(tǒng)的凈礦化和凈硝化均很低, 天然林通常受土壤低氮供應(yīng)的限制, 氮素處于封閉、積累型循環(huán), 地上部每年的凈初級(jí)生產(chǎn)力與土壤凈氮礦化呈線性正相關(guān)[29]。美國(guó)北部的天然草地轉(zhuǎn)換為森林35~75 a的土壤氮礦化和微生物活性基本沒(méi)有變化[30]。因?yàn)榈獡p失與氮凈礦化率和枯枝落葉氮通量(植物氮循環(huán)指標(biāo))密切相關(guān), 盡管原始林和次生林的氮量沒(méi)有顯著差異, 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轉(zhuǎn)化也比次生林大。美國(guó)密西根州半島西部原始闊葉林總氮礦化大約是次生林的2倍, 而總硝化沒(méi)有差異[8]。

3.2 生態(tài)修復(fù)對(duì)土壤氮庫(kù)消長(zhǎng)的影響

3.2.1 退耕還林還草對(duì)土壤氮庫(kù)積累的效應(yīng)

植被種類(lèi)直接影響生態(tài)系統(tǒng)的修復(fù)速率。在遺棄50 a以上的荒地上種植闊葉林、針葉林、牧場(chǎng)和種植園, 會(huì)顯著影響土壤氮含量和硝化速率[20]。我國(guó)黃土高原退耕還林還草30 a后, 恢復(fù)的草原、刺槐()、紅松()、油松()以及油松-紫穗槐()混交林等土壤有機(jī)質(zhì)、全氮、有效氮均提高了2倍多, 其中, 刺槐-紫穗槐混合林恢復(fù)最迅速, 油松-紫穗槐混交林恢復(fù)較慢[45]。晉西黃土丘陵區(qū)退耕還林或撂荒地還林還草后, 荒草地土壤全氮表聚效應(yīng)最強(qiáng), 0~48 cm土層中以刺槐林地的全氮含量提升最快[46]。黃土高原的退化草地分別栽種油松和檸條()30多年后, 油松地0~20 cm土層有效氮減少, 而檸條灌叢林地的有效氮卻增加[47]。在巴西里約熱內(nèi)盧州沿海城鎮(zhèn)的退化土地上, 于1991年種植了7種速生的先鋒豆科植物用以恢復(fù)退化土壤的肥力, 13 a中土壤氮庫(kù)的年增長(zhǎng)率為0.13 Mg×hm-2[9]。豆類(lèi)植物能增加土壤氮和碳的積累, C3植物和雜草會(huì)降低氮和碳的積累率, C4植物提高了土壤C/N比[28]。

盡管種植林草是增加土壤對(duì)碳、氮固定的途徑, 但生態(tài)系統(tǒng)恢復(fù)重建絕非易事, 正所謂毀壞容易修復(fù)難。明尼蘇達(dá)州廢棄61 a的農(nóng)田要恢復(fù)到之前耕作時(shí)95%的水平, 氮庫(kù)需要180 a, 碳庫(kù)需要230 a, 且碳的積累速率受氮積累的影響[28]。也有研究結(jié)果報(bào)道, 在50 a內(nèi)可以成功恢復(fù)嚴(yán)重退化的森林土壤氮的有效性, 但如果在恢復(fù)過(guò)程中繼續(xù)收獲林下植被和凋落物, 則難以實(shí)現(xiàn)這種恢復(fù)速率和恢復(fù)水平[15]。

3.2.2 恢復(fù)重建土壤氮碳庫(kù)的其他措施

相對(duì)于單一植被系統(tǒng), 農(nóng)林復(fù)合系統(tǒng)對(duì)生態(tài)的恢復(fù)效果一直被廣泛推崇?？煽?)林/農(nóng)復(fù)合系統(tǒng)的土壤碳氮貯量低于天然林, 但高于單一的可可林, 在維持生態(tài)系統(tǒng)功能服務(wù)方面比單一農(nóng)業(yè)種植更好, 并接近天然森林[48]。尤其是在障礙性土壤區(qū)域, 實(shí)施農(nóng)林復(fù)合系統(tǒng)是克服土壤障礙, 恢復(fù)生態(tài)功能的重要途徑。我國(guó)華北低平原鹽堿地區(qū)棗()/冬小麥()-夏玉米間作22 a以上的研究表明, 棗/糧間作系統(tǒng)的種植和生態(tài)效益仍然比單作農(nóng)業(yè)和單作棗樹(shù)高[49]。

休耕和輪作也是恢復(fù)土壤有機(jī)質(zhì)和氮含量的有效措施[50]。對(duì)亞馬遜河流域多樣化的森林研究表明, 土壤修復(fù)除了修復(fù)物或施用化肥, 還可以通過(guò)延長(zhǎng)田間使用時(shí)間、縮短休耕期來(lái)提升土壤肥力, 提高生物多樣性[51]。

3.3 土壤氮素內(nèi)循環(huán)對(duì)生態(tài)修復(fù)的響應(yīng)

4 土壤微生物對(duì)生態(tài)覆被/土地利用變化的響應(yīng)

4.1 生態(tài)系統(tǒng)土壤微生物量與氮礦化的關(guān)系

土壤微生物是氮素等養(yǎng)分元素循環(huán)的引擎, 氨化作用、固氮作用、硝化作用和反硝化作用構(gòu)成土壤氮循環(huán)的主要環(huán)節(jié), 且每一個(gè)過(guò)程都需要相應(yīng)微生物參與。土壤微生物群落結(jié)構(gòu)控制了不同生態(tài)系統(tǒng)中的氮素轉(zhuǎn)化, 進(jìn)而調(diào)節(jié)生態(tài)系統(tǒng)的功能與穩(wěn)定。自然森林土壤中有較高的微生物量氮, 凈氮礦化相對(duì)較少[31], 因此森林系統(tǒng)常受供氮量低的限制。外生菌根等微生物也是森林系統(tǒng)的重要氮源, 澳大利亞國(guó)家公園強(qiáng)酸且貧瘠土壤上生長(zhǎng)著極度瀕危的Wollemi松樹(shù), 高度依賴(lài)于其根部一個(gè)獨(dú)特的細(xì)菌群落[55]。可見(jiàn)不同森林的土壤微生物量差異很大。小興安嶺6種森林類(lèi)型的土壤微生物碳與微生物氮的大小順序依次為: 次生白樺()林>人工紅松林>擇伐林>闊葉紅松林>人工落葉松()林>谷地云冷杉()林, 總體表現(xiàn)為闊葉林(次生白樺林、闊葉紅松林和擇伐林)的土壤微生物量高于針葉林或針葉樹(shù)占比較高的森林類(lèi)型[56]。

地上部植被和土壤環(huán)境均對(duì)土壤微生物生物量氮和氮礦化有影響[37]。幼齡草甸土壤微生物生物量氮較低, 隨著年限增加, 微生物氮與全氮比(Nmic/TN)增加, 土壤總氮礦化率下降, 氮對(duì)植物的有效性降低, 植物產(chǎn)量下降[57]。

4.2 土壤微生物對(duì)生態(tài)覆被/土地利用變化的響應(yīng)

城市化過(guò)程對(duì)土壤氮循環(huán)的關(guān)鍵微生物影響很大。城市草坪土壤中的氨氧化古菌(AOA)豐度高于郊區(qū)和農(nóng)村的農(nóng)田土壤, AOA對(duì)城市草坪土壤硝化起關(guān)鍵作用。城市草坪土壤中的根瘤菌、變形桿菌()和綠彎菌()也比農(nóng)田土壤豐富, 但城市草坪土壤AOB和反硝化細(xì)菌nirS, nosZ)的相對(duì)豐度低于郊區(qū)草坪和農(nóng)田[27]。

4.3 土壤氮素內(nèi)循環(huán)對(duì)生態(tài)覆被/土地利用變化響應(yīng)的研究方法展望

由于土壤環(huán)境的多樣性和土壤微生物的復(fù)雜性,導(dǎo)致生態(tài)覆被/土地利用變化與土壤氮循環(huán)過(guò)程的效應(yīng)至今難有確定結(jié)論。在生態(tài)覆被/土地利用加劇和全球氣候變暖趨勢(shì)下, 對(duì)這一科學(xué)問(wèn)題的探索仍是研究熱點(diǎn)。其瓶頸仍然在于土壤微生物, 因?yàn)橥寥乐卸鄶?shù)微生物在休眠狀態(tài)下長(zhǎng)時(shí)間存活, 休眠期間細(xì)胞活性很低甚至沒(méi)有活性。傳統(tǒng)的平板培養(yǎng)法無(wú)法將它們分離出來(lái)。目前常用的熏蒸法測(cè)定土壤微生物碳和微生物氮誤差非常大, 結(jié)果重現(xiàn)性差。運(yùn)用分子生物學(xué)技術(shù)研究土壤微生物與土壤氮素循環(huán)之間的關(guān)系是目前的方向, 但是, 微生物群落的數(shù)量(豐度)以及多樣性指數(shù)(如Shannon和Simpson指數(shù))難以揭示復(fù)雜的微生物群落結(jié)構(gòu)與功能微生物之間量化關(guān)系[63-65]。特別是找出微生物群落中控制生態(tài)功能的關(guān)鍵物種是本方向研究中的一個(gè)難點(diǎn)。近年來(lái), 利用高通量基因芯片數(shù)據(jù)和微生物群落的生態(tài)網(wǎng)絡(luò)分析方法是提升土壤微生物群落結(jié)構(gòu)研究定量化和可視化新方向。

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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

楊莉琳, 姚琦馥, 梁琍, 魯小明. 土壤氮素內(nèi)循環(huán)對(duì)生態(tài)覆被變化響應(yīng)的研究進(jìn)展[J]. 中國(guó)生態(tài)農(nóng)業(yè)學(xué)報(bào)(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

* 國(guó)家自然科學(xué)基金項(xiàng)目(31270521)、貴州省教育廳創(chuàng)新群體重大研究項(xiàng)目(黔教合KY字[2016]053號(hào))、貴州省科技計(jì)劃項(xiàng)目(黔科合基礎(chǔ)[2019]1312)、貴州省創(chuàng)新人才團(tuán)隊(duì)(黔教合人才團(tuán)隊(duì)字[2015]67號(hào))、銅仁學(xué)院博士基金項(xiàng)目(trxyDH1525)和農(nóng)業(yè)生態(tài)創(chuàng)新團(tuán)隊(duì)(CXTD[2020-10])資助

楊莉琳, 主要研究方向?yàn)橥寥鲤B(yǎng)分循環(huán)與環(huán)境生態(tài)。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