馬 潔，陳先江，侯扶江

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國(guó)家重點(diǎn)實(shí)驗(yàn)室 蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020)

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草地生物土壤結(jié)皮

馬 潔，陳先江，侯扶江

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國(guó)家重點(diǎn)實(shí)驗(yàn)室 蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020)

摘要：生物土壤結(jié)皮是由隱花植物與土壤中的細(xì)菌、真菌以及土壤顆粒結(jié)合形成的復(fù)雜生物覆蓋體,它們廣泛分布于世界上各草地類組中，在改善生態(tài)環(huán)境和防治草地退化方面起到了積極的作用。多年來(lái)，國(guó)內(nèi)外學(xué)者就相關(guān)問(wèn)題開(kāi)展了大量的研究，但總體來(lái)說(shuō)，主要研究仍集中在對(duì)其功能和作用的認(rèn)知方面，而對(duì)全球變化背景下生物土壤結(jié)皮的演變以及它們維護(hù)生態(tài)系統(tǒng)穩(wěn)定性機(jī)理方面的研究還十分欠缺。本文綜述了國(guó)內(nèi)外該領(lǐng)域的現(xiàn)有研究成果，討論了目前研究的不足以及未來(lái)的研究方向，以期促進(jìn)生物土壤結(jié)皮相關(guān)研究，加深對(duì)草地生態(tài)系統(tǒng)土壤演替過(guò)程的認(rèn)識(shí)。

關(guān)鍵詞：土壤結(jié)皮；組成多樣性；分布；生態(tài)功能；研究展望

草地是面積最大的陸地生態(tài)系統(tǒng)，具有改良土壤、調(diào)節(jié)氣候、凈化環(huán)境、維持物種多樣性等的作用[1]。隨著人類活動(dòng)加強(qiáng)，草原不合理利用，加之全球氣候變化，草地面積減少、生產(chǎn)力下降，制約了農(nóng)牧業(yè)生產(chǎn)和人民生活的提升。草地退化及其生態(tài)恢復(fù)已經(jīng)成為世界性問(wèn)題[2],從生物土壤結(jié)皮(biological soil crusts)的角度，揭示草地退化和恢復(fù)機(jī)制是其中的重要方面[3-4]。

草地生物土壤結(jié)皮是由生活于草地土壤表面的非維管束植物(藍(lán)藻、綠藻、硅藻、地衣和苔蘚)和微生物(土壤細(xì)菌、真菌等)，與土壤顆粒通過(guò)團(tuán)聚和膠結(jié)作用在土壤表層形成的復(fù)合生物體[5]。它能固定大氣中的氮、碳，增加土壤有機(jī)質(zhì)，改良土壤[6]，在荒漠草地中的蓋度甚至超過(guò)生物總蓋度的40%[7]。生物土壤結(jié)皮往往出現(xiàn)在高等植物分布受限的環(huán)境或者植物之間的空地上，生物種類不易辨別，所以長(zhǎng)期受到忽視。20世紀(jì)90年代以后，人們發(fā)現(xiàn)生物土壤結(jié)皮在防止草地退化等方面有重要作用，其研究逐漸引起人們的重視，對(duì)它的保護(hù)已被列入草地生態(tài)系統(tǒng)管理尤其是荒漠草地管理的優(yōu)先等級(jí)[8]。此后，國(guó)內(nèi)外學(xué)者對(duì)生物土壤結(jié)皮的時(shí)空格局、生物組成、形態(tài)結(jié)構(gòu)、生態(tài)功能和管理等開(kāi)展了全面研究[9]。

1草地生物土壤結(jié)皮的組成及演替

生物土壤結(jié)皮群落主要由自養(yǎng)生物體和異養(yǎng)生物體組成。前者主要包括藻類(algae)、地衣(lichen)和苔蘚(moss)等，其光合作用是結(jié)皮群落生產(chǎn)力的主要來(lái)源；后者主要包括細(xì)菌(bacteria)、微小真菌(microfungi)等，在結(jié)皮中起分解和代謝作用[10]。它們利用菌絲體、假根以及分泌物膠結(jié)微小的土壤顆粒和大氣降塵，團(tuán)聚成厚度為3～10 mm的毯狀結(jié)構(gòu)覆蓋在地表，成為草地生態(tài)系統(tǒng)的“系統(tǒng)工程師”[11]。根據(jù)生物土壤結(jié)皮群落中優(yōu)勢(shì)生物組分，可以劃分為藻類結(jié)皮、地衣結(jié)皮和苔蘚結(jié)皮[12]，各類型結(jié)皮的形態(tài)、功能各異，組成成分具有多樣性。

藻類結(jié)皮是生物土壤結(jié)皮演替的最初階段。它以物理結(jié)皮和微生物結(jié)皮為基礎(chǔ)，隨著時(shí)間的推移，微環(huán)境條件改善，沙粒間隙被大氣降塵所填充，細(xì)菌分泌的胞外多糖黏結(jié)形成表面細(xì)致平滑的無(wú)機(jī)層，藻類出現(xiàn)并迅速繁殖[13-14]。生物土壤結(jié)皮中藍(lán)藻(Cyanobacteria)的種類和數(shù)量比例較高[15]。藻類對(duì)土壤類型的適應(yīng)性因種而異，不同區(qū)域的群落組成差異性較大。譬如黃土高原3個(gè)侵蝕區(qū)藻類的種類組成及優(yōu)勢(shì)種均有差異[16]。藻類雖不易被肉眼觀察到，但有藻類的土壤表面一般會(huì)有相對(duì)應(yīng)的色斑，藍(lán)細(xì)菌占優(yōu)勢(shì)的結(jié)皮往往呈現(xiàn)藍(lán)綠色或黑色，而綠藻會(huì)在土壤濕潤(rùn)時(shí)呈綠色[17]。藻類能夠顯著提高土壤的酶活性和土壤肥力，促進(jìn)藻類結(jié)皮發(fā)育成地衣或苔蘚結(jié)皮[18-19]。

地衣是藻類和菌類的共生體，它具有菌絲和假根等，能夠黏結(jié)土壤，穿透沙土表層并緊密連接葉狀體，形成穩(wěn)定的結(jié)皮形態(tài)，進(jìn)而抵御風(fēng)水侵蝕，促進(jìn)有機(jī)物累積[20]。草地生態(tài)系統(tǒng)中地衣的蓋度與地表環(huán)境的穩(wěn)定性相關(guān)，可以指示環(huán)境的變化[21]。美國(guó)西部、哥倫比亞等地廣泛存在著地衣結(jié)皮[22]。騰格里沙漠、沙坡頭地區(qū)也鑒定出分屬于5目13科16屬的地衣植物22種[23]。地衣結(jié)皮的顏色呈現(xiàn)黑、褐、白、綠色，表面呈凹凸不平的殼狀，可以通過(guò)形態(tài)特征與藻類結(jié)皮相區(qū)別[24]。

苔蘚結(jié)皮是生物土壤結(jié)皮發(fā)育和演替的最高階段，能顯著加快礦物質(zhì)風(fēng)化的速度，積累營(yíng)養(yǎng)元素供植物生長(zhǎng)[25]。古爾班通古特沙漠的壟間低地生存著以刺葉墻蘚(Tortuladesertorum)和真蘚為優(yōu)勢(shì)種的苔蘚植物[26]，沙坡頭有真蘚為優(yōu)勢(shì)種的2科7屬共16種苔蘚植物[27]。苔蘚結(jié)皮的多度和蓋度與土壤粘粒含量呈正比，土表粗糙度的增加有利于種子萌發(fā)及植物的侵入、定居[28-30]。生長(zhǎng)后期，苔蘚結(jié)皮呈現(xiàn)較深的顏色，有效吸收光照，下層土壤在寒冷的天氣下保持較高的溫度，以保證土壤微生物的生長(zhǎng)活動(dòng)[31](表1)。

一般情況下，生物土壤結(jié)皮的演替始于大型絲狀藍(lán)細(xì)菌等藻類定殖；隨著地表固定和土壤基質(zhì)的改善，地衣和苔蘚開(kāi)始生長(zhǎng)，結(jié)皮自身的蓄水、抗機(jī)械干擾以及固沙能力依次增強(qiáng)(圖1)。這個(gè)過(guò)程中，組成植物種從低等到高等，種群豐富度和群落多樣性從小到大[42]。但是當(dāng)環(huán)境條件適宜時(shí)，結(jié)皮的演替也可能不經(jīng)過(guò)其中一個(gè)階段而直接發(fā)育到更高級(jí)的階段。在生物結(jié)皮成為典型特征的地域，密集頻繁的放牧、旅游和火燒等造成生物土壤結(jié)皮減少，甚至向上一個(gè)階段退化[43]。

2影響草地結(jié)皮分布的因素

生物土壤結(jié)皮能夠適應(yīng)多種氣候條件和土壤生境，對(duì)外界脅迫具有一定抗性，在世界各類草地生態(tài)系統(tǒng)中多有分布[44]。生物土壤結(jié)皮的組成、發(fā)育以及蓋度、多度等存在地域間甚至地域內(nèi)的差異。

在全球或區(qū)域尺度上，生物土壤結(jié)皮的分布主要受氣候影響。結(jié)皮的酶活性、新陳代謝速率、生理活動(dòng)與溫度和降水密切相關(guān)[45]。降水較多，則適宜形成以苔蘚為優(yōu)勢(shì)種的生物土壤結(jié)皮[46]。溫度變化會(huì)改變地面的蒸散特征，不利于對(duì)溫度敏感的結(jié)皮生物的生長(zhǎng)。溫度升高會(huì)影響結(jié)皮在夜間捕獲凝結(jié)水的能力，從而縮短其生物活性[47]。

在景觀尺度上，土壤的質(zhì)地和化學(xué)特性影響著生物結(jié)皮群落組成，土壤顆粒是主導(dǎo)因素。土壤質(zhì)地較細(xì)能提高藻類、地衣、苔蘚的蓋度和物種豐富度[48]。堿性土壤適宜藍(lán)藻類結(jié)皮的發(fā)育，綠藻則偏好酸性土壤；土壤碳酸鈣含量高促進(jìn)地衣定植[49]。藍(lán)藻的豐富度與鈉、鎘、鋰、銅、鉬、鍶含量呈負(fù)相關(guān)，與鉛和鋅含量呈正相關(guān)[50]。

在生態(tài)系統(tǒng)尺度上，生物結(jié)皮的分布對(duì)地貌部位有強(qiáng)選擇性。在古爾班通古特沙漠，藻結(jié)皮主要分布于沙壟坡部, 從沙坡至丘間低地，逐漸演替為連片分布地衣結(jié)皮和斑塊狀的與地衣鑲嵌而生的苔蘚結(jié)皮，同時(shí)生物結(jié)皮的蓋度、厚度、抗壓性從壟間至壟頂逐漸減小[51]。

表1　各類草地生物土壤結(jié)皮群落組成多樣性Table 1　Community composition diversity of biological soil crusts in several types of grassland

在群落尺度上，同一生境中的高等植物與生物結(jié)皮的形成有密切聯(lián)系。一般蓋度高的植被遮蔽光照和降水，結(jié)皮生物體無(wú)法與其競(jìng)爭(zhēng)資源；植被稀疏的荒漠，高等植物的生長(zhǎng)受水分限制而呈斑塊狀[52]，斑塊之間的裸地為生物土壤結(jié)皮提供了空間、水分和光照，這也是生物土壤結(jié)皮主要集中于干旱、半干旱地區(qū)的原因之一。

人類活動(dòng)是影響生物土壤結(jié)皮分布的重要因素。放牧、旅游和火燒等高強(qiáng)度的活動(dòng)會(huì)導(dǎo)致結(jié)皮減少甚至消失。放牧家畜的踐踏集中作用于土壤表面的結(jié)皮層，對(duì)微生物的數(shù)量分布及其碳氮代謝產(chǎn)生明顯作用[53]。火燒是常用的生態(tài)系統(tǒng)管理手段，生物土壤結(jié)皮對(duì)輕度火燒具有一定的適應(yīng)性，且輕度火燒可消除地表的凋落物，減少高等植物的生物量，有利于結(jié)皮的再形成和重新分布；但中高強(qiáng)度的火燒顯著降低結(jié)皮蓋度，增加地表侵蝕的風(fēng)險(xiǎn)[54-55]。

生物土壤結(jié)皮廣泛分布于各類型的荒漠生態(tài)系統(tǒng)，以及高山凍原和較濕潤(rùn)地區(qū)植被演替的早期階段[56]，它的出現(xiàn)和發(fā)展特征可以用作評(píng)價(jià)草地生態(tài)環(huán)境健康狀況的指標(biāo)[57](表2)?？傊瑲夂?、海拔、土壤、維管植物群落、家畜和野生動(dòng)物、人類管理以及草地生態(tài)系統(tǒng)的健康狀況等共同影響著生物土壤結(jié)皮的分布。

3草地生物土壤結(jié)皮的生態(tài)功能

3.1生物土壤結(jié)皮對(duì)土壤養(yǎng)分的影響

高等植物是土壤有機(jī)質(zhì)的重要來(lái)源，但植物株叢間隙的裸地上有機(jī)質(zhì)匱乏。生物土壤結(jié)皮在地表繁殖，藍(lán)藻、地衣、綠藻和苔類通過(guò)光合作用固定碳，提高植物間隙的土壤肥力。生物土壤結(jié)皮主要影響0～5 cm土壤有機(jī)質(zhì)，可為淺根系植物種子的萌發(fā)和生長(zhǎng)提供有機(jī)肥，促進(jìn)植物的繁衍與更新[62]。凍原高山草地和冷荒漠草地中以苔蘚、地衣為主的結(jié)皮能固定大量的泥炭，成為陸地生態(tài)系統(tǒng)的重要碳匯[63]。濕度和土壤呼吸等因素限制結(jié)皮，固定有機(jī)碳，生物結(jié)皮減少土壤CO2排放的作用與結(jié)皮厚度正相關(guān)[64]。

圖1　生物土壤結(jié)皮的演替Fig.1　The succession of biological soil crust

注：①降塵填充沙粒間隙，絲狀藻類定殖；②地表固定，外界干擾少；③基質(zhì)穩(wěn)定，水分充足；④人工接種；充足的光照、水分，微地形條件適宜；土壤表面及基質(zhì)穩(wěn)定；⑤火燒，高強(qiáng)度的放牧，高密度的旅游，頻繁的交通；⑥長(zhǎng)期機(jī)械干擾、家畜和人類活動(dòng)；⑦土壤結(jié)構(gòu)遭到破壞；生態(tài)環(huán)境持續(xù)性惡化，水土流失嚴(yán)重，草地退化加劇。

Note： ①The gap among sands full with dustfull, and algae are colonization. ②The process needs stable soil surface and less disturbance. ③Moss needs good soil matrixs and suitable moisture condition. ④Artificial inoculation;enough illumination and moisture and suitable microtopography condition; stable soil surface and matrixs. ⑤Burn,high grazing intensity, excessive tourism, and traffic disturbance frequently. ⑥Long-term mechanical disturbance and activities of livestocks and human. ⑦The soil structure were destroyed; entironment getting worse, soil erosion severly and degenerate seriously.

表2　生物土壤結(jié)皮與草地生態(tài)系統(tǒng)健康狀況Table 2　The relationship between biological soil crusts and grassland ecosystem healthy condition

荒漠中的氮主要來(lái)自于大氣沉降和生物固氮，其中土壤結(jié)皮中的藍(lán)細(xì)菌可以固定大氣中的氮，提供荒漠植物和土壤生物重要營(yíng)養(yǎng)源[65]。北極地區(qū)，與苔蘚結(jié)皮共生的藻青菌所固定的氮占凍原高山草地氮年固定總量的80%[45]。美國(guó)猶他州膠衣屬地衣的氮輸入量為13 kg·(hm2·a)-1，相當(dāng)于無(wú)結(jié)皮土壤表層氮輸入量[1.4 kg·(hm2·a)-1]的10倍[66]。苔蘚結(jié)皮下土壤氮素轉(zhuǎn)化速率及其對(duì)溫度的敏感性均高于無(wú)結(jié)皮土壤，從而提高土壤的供氮能力，促進(jìn)氮素轉(zhuǎn)化與循環(huán)，加快草地土壤的修復(fù)[67]。

結(jié)皮形成過(guò)程中，藻類首先在植物根部出現(xiàn)，然后具有固氮功能的地衣開(kāi)始定殖，改善了土壤的結(jié)構(gòu)和持水力，為苔蘚植物的生長(zhǎng)創(chuàng)造了條件[68]。苔蘚和地衣植物富集又促進(jìn)表土的腐殖質(zhì)累積，增加鉀、鈣、鎂、磷等元素，從而提高草地生產(chǎn)力。

圖2　生物土壤結(jié)皮的主要生態(tài)功能Fig.2　The main ecological function of biological soil crusts

注：①促進(jìn)植物生長(zhǎng)，捕獲種子；②降低風(fēng)速，為結(jié)皮發(fā)育提供穩(wěn)定的陰蔽環(huán)境；③穩(wěn)定地表，累積營(yíng)養(yǎng)，參與水循環(huán)；④結(jié)皮生長(zhǎng)的載體,影響其演替過(guò)程；⑤增加土壤溫度、濕度,加快微生物活動(dòng)；⑥結(jié)皮形成和生長(zhǎng)的參與者；⑦為節(jié)肢動(dòng)物提供食物，為昆蟲(chóng)提供適宜的生境；⑧適度放牧促進(jìn)結(jié)皮發(fā)育。

Note： ①Promote plants growth and capture their seeds. ②Reduce wind speed, and provide stable environment for crusts. ③Fix soil surface, accumulate nutriention and attend the water cycle. ④Carrier of crusts growth. and effect it’s evolution process. ⑤Elevated soil moisture and temperature to quicken it’s activities. ⑥Participant of crusts formation. and development. ⑦Provide foods for arthropods, and suitable habitat for insects. ⑧Moderate grazing would beneficial to crusts’ growth.

3.2生物土壤結(jié)皮與微生物

微生物既是土壤結(jié)皮形成的參與者，反過(guò)來(lái)又受到結(jié)皮發(fā)育程度的影響[69]。通過(guò)調(diào)節(jié)土壤的溫度和濕度，生物土壤結(jié)皮顯著地增加了細(xì)菌、放線菌和真菌等的分布，它的發(fā)育也為微生物的生長(zhǎng)繁殖創(chuàng)造了良好的條件，研究表明，生物土壤結(jié)皮類型與土壤微生物生物量密切相關(guān)[70-71],其微生物活性隨著結(jié)皮發(fā)育而持續(xù)增加[72]；苔蘚結(jié)皮下土壤微生物量碳氮顯著高于其它類型結(jié)皮[73]。

3.3生物土壤結(jié)皮與植物的相互作用

由于研究區(qū)域、植物種類、結(jié)皮組成、所受干擾程度和土壤基本情況的復(fù)雜性，生物結(jié)皮和種子植物之間的關(guān)系尚無(wú)定論。一些研究認(rèn)為，生物結(jié)皮累積了大量植物生長(zhǎng)所必需的元素，促進(jìn)了植物吸收營(yíng)養(yǎng)和生長(zhǎng)，而植被能降低風(fēng)速、形成陰敝而穩(wěn)定的環(huán)境，從而加快結(jié)皮的發(fā)育[74]；但也有研究表明，苔蘚植物迅速吸收養(yǎng)分，與維管植物競(jìng)爭(zhēng)資源，植被與苔蘚結(jié)皮的蓋度呈負(fù)相關(guān)[75]；此外有觀點(diǎn)認(rèn)為兩者占據(jù)著不同的生態(tài)位，沒(méi)有明顯的互惠或競(jìng)爭(zhēng)關(guān)系[76]。

除了與植株的直接關(guān)系，生物結(jié)皮的粗糙度也直接影響到土壤對(duì)種子的捕獲能力[77]。荒漠土壤種子庫(kù)在苔蘚結(jié)皮上的儲(chǔ)量顯著高于表面光滑藻類結(jié)皮[78]，而種子的萌發(fā)與結(jié)皮層的含水量存在正相關(guān)關(guān)系[79]。

3.4生物土壤結(jié)皮在水文過(guò)程中的作用

草地生物結(jié)皮參與的水文過(guò)程主要包括降水入滲、凝結(jié)水捕獲以及蒸發(fā)。起伏型結(jié)皮可以增加入滲和土壤的含水量，而光滑型結(jié)皮容易造成地表徑流[80]，去除生物結(jié)皮明顯提高水分的入滲率[81]。生物土壤結(jié)皮可以對(duì)水分再分配，增加土壤條件的多樣性[82]。

凝結(jié)水是荒漠生態(tài)系統(tǒng)持續(xù)而穩(wěn)定的水源，生物土壤結(jié)皮捕獲凝結(jié)水能夠促進(jìn)有效水分的淺層化和生物量積累，促進(jìn)結(jié)皮固氮、光合作用和生長(zhǎng)，結(jié)皮葉綠素含量與其凝結(jié)水捕獲能力正相關(guān)[83]。藻類結(jié)皮對(duì)土壤水分蒸發(fā)的影響與土壤含水量相關(guān)；土壤含水量較高時(shí)，藻類結(jié)皮下土壤的蒸發(fā)明顯高于無(wú)結(jié)皮土壤；土壤變干時(shí)，結(jié)皮有效減小蒸發(fā)量，將水分保持在土壤中[84]。

3.5其它

生物土壤結(jié)皮對(duì)于草地生態(tài)系統(tǒng)環(huán)境的穩(wěn)定有重要作用。生物土壤結(jié)皮顯著降低風(fēng)和水等對(duì)土壤的侵蝕，結(jié)皮穩(wěn)定地表的能力隨著其發(fā)育而增強(qiáng)[85-86]。顏色較深的苔蘚和地衣結(jié)皮顯著降低地表反射率，吸收更多能量，提高土壤溫度，加速微生物活動(dòng)和種子萌發(fā)[87]。生物土壤結(jié)皮可以維護(hù)草地生態(tài)系統(tǒng)的土壤食物網(wǎng)，其發(fā)育過(guò)程中累積的養(yǎng)分和生物量直接被食草動(dòng)物、原生動(dòng)物(變形蟲(chóng)、纖毛蟲(chóng)和鞭毛蟲(chóng)等)、線蟲(chóng)以及節(jié)肢動(dòng)物(螨等)等利用，調(diào)控土壤養(yǎng)分分配和分解[88]。

4展望

作為草地生態(tài)系統(tǒng)的結(jié)構(gòu)和功能組分，生物土壤結(jié)皮能夠改良土壤，提高生態(tài)系統(tǒng)抵御風(fēng)水侵蝕的能力，為維管植物的定居和繁育提供基礎(chǔ)條件。20世紀(jì)后期以來(lái)，國(guó)內(nèi)開(kāi)展了多種草地生態(tài)系統(tǒng)生物結(jié)皮的研究，揭示了結(jié)皮在生態(tài)系統(tǒng)的作用和獨(dú)特的生物多樣性，對(duì)于認(rèn)識(shí)景觀的功能、結(jié)構(gòu)和反饋機(jī)制具有重要作用。以往的研究重點(diǎn)集中在結(jié)皮功能和作用的認(rèn)知上，對(duì)全球變化背景下生物土壤結(jié)皮的演變，及其對(duì)生態(tài)系統(tǒng)管理的長(zhǎng)期反饋等研究較少，而這些知識(shí)是判別草地退化發(fā)生和發(fā)展及其機(jī)理的基礎(chǔ)。人類活動(dòng)對(duì)生物土壤結(jié)皮的破壞短期內(nèi)難以通過(guò)自然恢復(fù)，適度利用閾值和人工輔助修復(fù)土壤結(jié)皮的技術(shù)對(duì)于退化草地治理具有實(shí)踐意義。生物土壤結(jié)皮在區(qū)域草地生態(tài)系統(tǒng)能量和物質(zhì)平衡中有重要作用，利用遙感監(jiān)測(cè)，建立生物土壤結(jié)皮的生態(tài)系統(tǒng)管理體系，預(yù)測(cè)結(jié)皮在區(qū)域尺度上的時(shí)空變化，也亟待研究。

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(責(zé)任編輯王芳)

DOI:10.11829/j.issn.1001-0629.2015-0414

*收稿日期：2015-07-23接受日期：2015-10-09

基金項(xiàng)目：國(guó)家自然科學(xué)基金(31172249)；長(zhǎng)江學(xué)者和創(chuàng)新團(tuán)隊(duì)發(fā)展計(jì)劃(IRT13019)；國(guó)家科技支撐計(jì)劃課題(2011BAD17B02-03，2012BAD13B05)

通信作者：侯扶江(1971-)，男，河南扶溝人，教授，博導(dǎo)，博士，主要從事草地-家畜互作研究。 E-mail:cyhoufj@lzu.edu.cn

中圖分類號(hào)：S812.2

文獻(xiàn)標(biāo)識(shí)碼：A

文章編號(hào)：1001-0629(2016)7-1243-10*

Corresponding author:Hou Fu-jiangE-mail:cyhoufj@lzu.edu.cn

A review on biological soil crusts of grassland

Ma Jie, Chen Xian-jiang, Hou Fu-jiang

(Key Laboratory of Grassland Farming Systems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China)

Abstract:Biological soil crusts are a complex covering integrities developing by combining cryptigams with bacteria,fungi and soil particles, and are widely distributed in each grassland type in the world, and play an active role in improving the ecological environment and prevention of grassland degradation. In recently years, many studies throughout the world have been conducted in this field, and these studies focus on the cognition of its function and effect. However, how biological soil crusts develop under global change and how they maintain the stability of the ecosystem are still not well documented. This paper reviewed the research achievements in this filed, and discussed the current shortage and future directions of the research, and these would encourage the biological soil crusts to further advance and helpful understand the soil evolution in grassland ecosystem.

Key words：biological soil crusts; composition diversity; distribution; ecological functions; prospect

馬潔，陳先江，侯扶江.草地生物土壤結(jié)皮.草業(yè)科學(xué),2016,33(7)：1243-1252.

Ma J,Chen X J,Hou F J.A review on biological soil crusts of grassland.Pratacultural Science，2016,33(7)：1243-1252.

前植物生產(chǎn)層

第一作者：馬潔(1992-)，女，甘肅定西人，在讀碩士生，主要從事生物土壤結(jié)皮研究。E-mail:majie14@lzu.edu.cn