間套作功能植物對(duì)茶園生態(tài)系統(tǒng)服務(wù)功能的影響

史凡，黃泓晶，陳燕婷，陳李林*

間套作功能植物對(duì)茶園生態(tài)系統(tǒng)服務(wù)功能的影響

史凡1,2,3，黃泓晶1,2,3，陳燕婷4，陳李林1,2,3*

1.閩臺(tái)作物有害生物生態(tài)防控國家重點(diǎn)實(shí)驗(yàn)室，福建農(nóng)林大學(xué)植物保護(hù)學(xué)院，福建 福州 350002；2.中國白茶研究院，福建 福鼎 355200；3.害蟲綠色防控福建省高校重點(diǎn)實(shí)驗(yàn)室，福建 福州 350002；4.福建省農(nóng)業(yè)科學(xué)院植物保護(hù)研究所，福建 福州 350013

合理間套作功能植物是茶園生境管理的主要措施之一，能夠遮光保溫、保持水土、增肥促長、保持微域環(huán)境穩(wěn)定；還能吸引天敵、驅(qū)避害蟲，減輕茶園病蟲草為害，有助于茶葉提質(zhì)增效。但不合理的間套作會(huì)破壞茶園微域環(huán)境和爭奪養(yǎng)料，進(jìn)而影響茶樹生長。通過綜述近年來在茶園間套作功能植物及其對(duì)茶園生態(tài)系統(tǒng)服務(wù)功能的影響方面的文獻(xiàn)，主要包括茶園常見功能植物的種類、作用方式、功效和常見問題，以及間套作功能植物對(duì)茶樹生長發(fā)育和茶葉產(chǎn)量品質(zhì)的影響及對(duì)病蟲草害的調(diào)控作用；以期為應(yīng)用間套作措施促進(jìn)茶園病蟲草害的綜合調(diào)控和提升茶園生態(tài)系統(tǒng)服務(wù)功能提供指導(dǎo)。

套種；生境管理；生態(tài)調(diào)控；生態(tài)系統(tǒng)服務(wù)；生態(tài)系統(tǒng)功能

茶樹[(L.) Kuntze]是我國重要的經(jīng)濟(jì)作物之一，2020年我國茶園總面積達(dá)316.51萬hm2，國內(nèi)銷售總額為2?888.84億元，出口總額為20.38億美元[1]。目前，我國茶園管理方式朝著生態(tài)調(diào)控的方向發(fā)展[2]。茶園有害生物生態(tài)調(diào)控基于“預(yù)防為主、生態(tài)優(yōu)先、整合治理、精準(zhǔn)施策”的原則，從茶園生態(tài)系統(tǒng)中“茶樹-有害生物-天敵及茶園生態(tài)環(huán)境”相互作用關(guān)系出發(fā)，整合生態(tài)調(diào)控技術(shù)、農(nóng)業(yè)防治、生物防治、理化誘控、現(xiàn)代生物技術(shù)和合理的化學(xué)防治等手段，構(gòu)建“經(jīng)濟(jì)、簡便、有效”的生態(tài)技術(shù)體系，將有害生物控制在生態(tài)經(jīng)濟(jì)閾值水平之下，達(dá)到經(jīng)濟(jì)可實(shí)行、生態(tài)可持續(xù)、社會(huì)可接受的目的[3]。

生境管理是生態(tài)調(diào)控的重要方式之一。生境管理指通過對(duì)生境中各種資源進(jìn)行合理配置，構(gòu)建復(fù)雜生態(tài)系統(tǒng)，提高天敵控害能力的保護(hù)性防治策略[4-5]。生境管理被廣泛應(yīng)用于茶園、果園、菜地和稻田等農(nóng)田生態(tài)系統(tǒng)中。構(gòu)建健康合理的農(nóng)田生態(tài)系統(tǒng)，需要同時(shí)改善生態(tài)環(huán)境和生物類群。改善生態(tài)環(huán)境，主要包括地上部分的溫度、光照、水分；地下部分的保水透氣性、酸堿度及各種營養(yǎng)物質(zhì)等。改善生物類群多樣性，主要包括天敵類群、傳粉者類群、有益微生物及非雜草植物[6]（圖1）。

間套作功能植物是生境管理的重要方式之一，通過將病蟲草害的生態(tài)調(diào)控與提質(zhì)增效技術(shù)相結(jié)合，減緩因生態(tài)系統(tǒng)脆弱對(duì)生物多樣性構(gòu)成的威脅，達(dá)到維持生態(tài)系統(tǒng)平衡和穩(wěn)定的目的[3,7]。合理間套作有利于提高茶園光、水、熱、空間等資源利用率，促進(jìn)茶樹植株健壯生長；驅(qū)避害蟲或誘集天敵，為有益生物類群提供蜜粉資源、棲境、避難所、替代獵物等生態(tài)系統(tǒng)服務(wù)功能，從而提高茶園生物防治效果[8-11]。本文對(duì)近年來在茶園間套作功能植物對(duì)茶園生態(tài)系統(tǒng)影響方面的文獻(xiàn)進(jìn)行了綜述，以期為應(yīng)用間套作措施提升茶園保益控害、提質(zhì)增效等生態(tài)系統(tǒng)服務(wù)功能提供理論依據(jù)和實(shí)踐指導(dǎo)。

1 茶園常見間套作功能植物的選擇及類型

1.1 選擇合適的間套作功能植物

茶園間套作功能植物是為了涵養(yǎng)水土、改善茶園微環(huán)境、增強(qiáng)保益控害效能、茶產(chǎn)品提質(zhì)增效和增加茶旅觀光等生態(tài)系統(tǒng)服務(wù)功能。因此，茶園間套作功能植物的選擇應(yīng)符合種植管理輕簡、適應(yīng)性廣、與茶樹無共同病蟲害、非入侵雜草、對(duì)生境影響具有長效和普適性及作用期與茶樹生長采摘期銜接合適等要求[10-12]。注重景觀生態(tài)格局與過程，空間尺度注意不同植被斑塊的設(shè)計(jì)，時(shí)間尺度注意綜合考慮生物資源及物候期[13]。利用間套作措施對(duì)茶園進(jìn)行生態(tài)調(diào)控成為近年研究熱點(diǎn)，目前常見的適合用于茶園間套作的功能植物見表1。

圖1 間套作功能植物調(diào)節(jié)茶園生態(tài)系統(tǒng)服務(wù)功能

1.2 茶園常見間套作功能植物的功能類型

1.2.1 水土涵養(yǎng)植物（綠肥植物）

水土涵養(yǎng)植物能夠改善土壤肥力、保持水土、維持土壤健康。常見的水土涵養(yǎng)植物包括豆科植物圓葉決明（）和白車軸草（）等，該類植物根系具有固氮根瘤菌，能夠進(jìn)行生物固氮，增加土壤養(yǎng)分含量。彭晚霞等[73]發(fā)現(xiàn)在茶園間作白車軸草能促進(jìn)水分轉(zhuǎn)移至關(guān)鍵土層，提高水分利用率；同時(shí)對(duì)茶園微域溫度具有雙向調(diào)控作用（圖2）。

1.2.2 棲境植物（載體植物、養(yǎng)蟲植物）

棲境植物具有適宜生物類群棲息的物理結(jié)構(gòu)，有助于天敵類群越夏、越冬或繁殖等活動(dòng)；有利于增加有益生物類群的物種數(shù)、種群數(shù)量和群落多樣性，增強(qiáng)天敵控害效果[9,75]。茶園間作功能植物能夠涵養(yǎng)天敵，降低小貫小綠葉蟬（）的發(fā)生[58-59]。Zhang等[76]發(fā)現(xiàn)間作紅豆（）能夠提高天敵東亞小花蝽（）的存活率和產(chǎn)卵量（圖2）。

1.2.3 蜜粉源植物（顯花植物）

蜜粉源植物具有較多的花粉、花蜜或蜜露，可為有益類群（天敵或傳粉者）提供食物，吸引有益生物類群，提高益害比；增強(qiáng)天敵對(duì)有害生物類群的捕食作用[77-78]；還能夠提高寄生性天敵的壽命及寄生能力[79]。張正群等[58]發(fā)現(xiàn)在茶園間作多花植物羅勒（），能為天敵提供生長繁殖所需的花蜜，草蛉、蜘蛛、瓢蟲和寄生蜂的數(shù)量分別提高了9.0、3.3、1.6、1.4倍。取食蜜粉源的昆蟲還可以為植物傳授花粉，具有穩(wěn)定農(nóng)業(yè)生產(chǎn)等作用[80]（圖2）。

表1 茶園常見生境類型及適合的功能植物

圖2 茶園常見功能植物類型（改自肖英方等[74]）

1.2.4 誘集植物（誘蟲植物）

誘蟲植物可以通過物理形態(tài)和揮發(fā)性次生物質(zhì)共同作用形成比主栽作物更強(qiáng)的引誘力[81]。誘蟲植物可以誘捕有害生物類群，減少其對(duì)主栽作物的為害[82-83]。斯里蘭卡茶園管理者通過種植合歡植物（sp.）誘殺白蟻，南印度茶園管理者曾使用千斤拔（）誘集茶枝小蠹成蟲（）[84]。張曉明等[85]發(fā)現(xiàn)在茶園間作黃金菊（）可吸引更多天敵昆蟲取食和停留（圖2）。

1.2.5 驅(qū)避植物

驅(qū)避植物含有特定的化學(xué)物質(zhì)，形成較強(qiáng)的化感作用物質(zhì)或刺激性氣味。茶園間套作驅(qū)避植物能夠向環(huán)境分泌芳香物質(zhì)，從而影響生物類群的定位、搜尋、取食和寄生等活動(dòng)[58]。在茶園間作薄荷（）可以驅(qū)避綠盲蝽（）[60]，間作迷迭香（）對(duì)茶尺蠖（）具有干擾和驅(qū)避作用[86]。Cai等[87]在茶園緩釋芳香化合物混合劑，證明芳香植物揮發(fā)物對(duì)茶小綠葉蟬具驅(qū)避作用（圖2）。

1.2.6 殺蟲植物

殺蟲植物可產(chǎn)生有毒物質(zhì)，如生物堿、糖苷類等，直接殺死有害生物。曾維愛[88]開展了15種植物的4種溶劑提取物對(duì)茶尺蠖3齡幼蟲生物測定試驗(yàn)，發(fā)現(xiàn)辣蓼（）中的丁香酚對(duì)茶尺蠖幼蟲具有明顯拒食、胃毒和觸殺作用，抑制幼蟲生長。Kamunya等[89]研究發(fā)現(xiàn)，印加孔雀草（）與茶樹間作后，印加孔雀草分泌的皂苷類物質(zhì)能夠有效毒殺茶園土壤中的根結(jié)線蟲（圖2）。

1.2.7 其他植物

除上述的生態(tài)系統(tǒng)服務(wù)功能外，多花的功能植物還兼具有凈化空氣、散發(fā)香味，以及提供視覺享受的功能。在茶園間套作油菜（）、櫻花（sp.）或杜鵑花（）能夠滿足游客的休閑娛樂需求，具有茶旅融合觀光功能[32-33,90]。部分間套作植物還可作為飼料、菌類栽培料、沼氣池發(fā)酵原料等[27,91]，甚至發(fā)揮“以草治草”功效[61,92]（圖2）。

2 間套作功能植物對(duì)茶園生態(tài)環(huán)境的影響

茶樹為多年生植物，且定植后極少移栽。純茶園和幼齡茶園地表裸露面積較多，土壤保水保肥能力弱；長期施肥和采茶等農(nóng)事操作導(dǎo)致土壤板結(jié)退化，含水量和透氣性降低。功能植物根系的生長能夠促進(jìn)土壤疏松化，增強(qiáng)土壤透氣度，調(diào)節(jié)地表溫度，減少水分蒸發(fā)，降低單位體積土壤容重。同時(shí)，功能植物產(chǎn)生的分泌物和有機(jī)質(zhì)能夠改善土壤微生物群落組成與結(jié)構(gòu)，提高土壤酶活性。

2.1 提高水分利用率

間套作功能植物有助于滯留降雨、減少地表徑流，增加土壤中水分存留；同時(shí)，減緩空氣流動(dòng)，減少土壤水分揮發(fā)和葉片蒸騰，提高水分利用率[17]。在茶園間作白車軸草能促進(jìn)深層土壤中水分向關(guān)鍵土層移動(dòng)[61]。在茶園間作大球蓋菇（）后，冬季茶園土壤含水量提高5.74%～10.81%，且降雨量越小，抗旱保墑能力越強(qiáng)[56]。

2.2 調(diào)節(jié)溫度

茶樹生長的最適氣溫為20～30℃，最適地溫為14～20℃[93]。功能植物對(duì)茶園溫度具有雙向調(diào)節(jié)作用。在氣溫較低的春季，功能植物能夠減緩空氣流動(dòng)，阻礙熱量交換，減少茶叢熱量擴(kuò)散至周圍環(huán)境中，使茶園微域較溫暖，促進(jìn)茶樹提早出芽；在氣溫較高的夏季，功能植物遮陰減少陽光直射，降低茶樹葉面、微域環(huán)境和地表的溫度[73]。在茶行間作大豆（），茶園微域氣溫可降低3.1～3.6℃，間作玉米（）可降低1.7～3.8℃[30-31]。套種杉木茶園有害葉面高溫出現(xiàn)的次數(shù)僅為對(duì)照茶園的20.6%～32.4%，土壤出現(xiàn)的極端高溫比對(duì)照茶園降低約5.0℃[94]。

2.3 保土減流和修復(fù)土壤

降雨或澆水時(shí)產(chǎn)生的徑流會(huì)導(dǎo)致泥土和養(yǎng)分流失，間套作功能植物能夠改善土壤理化性質(zhì)、減少地表徑流和泥土沖刷量，提高養(yǎng)分的生物有效性，有利于茶園水土保持和生態(tài)修復(fù)。在茶園間作白車軸草可減弱雨水沖擊，增加土壤孔隙度和粉砂含量[95-96]。適度遮陰能夠提高茶園土壤養(yǎng)分、酶活性和微生物多樣性，有利于受損生態(tài)系統(tǒng)的恢復(fù)和重建[97]。間套作功能植物能夠提高有效鉬、銅、交換性鈣、鎂和硼的含量；降低土壤中鉛、鎳、錳等對(duì)人體有害重金屬元素的含量[25]。

2.4 平衡酸堿度

茶樹是喜酸性植物，最適合茶樹生長的土壤pH處于4.5～5.5，但我國主要茶區(qū)約72.3%的土壤pH小于4.5，且pH平均每年以0.05的速率下降[98]。土壤過度酸化會(huì)使茶樹生長受到抑制，而種植功能植物能夠緩解土壤的酸化速率，提升土壤pH值。梁麗妮等[32]發(fā)現(xiàn)油菜能夠提高茶樹根圍土的pH值至4.78～5.97，提高根際土壤的pH值至4.34～5.62。

2.5 提高土壤肥力

間套作功能植物對(duì)有機(jī)質(zhì)分解、礦質(zhì)營養(yǎng)循環(huán)和土壤生態(tài)系統(tǒng)的可持續(xù)發(fā)展起著重要的作用[99]。間套作功能植物能提高茶園土壤中礦物質(zhì)和有機(jī)質(zhì)含量[33,58]；提高有效氮、有效磷和有效鉀的含量[100]，促進(jìn)茶樹對(duì)營養(yǎng)元素的吸收。茶園間作的豆科植物會(huì)將氮?dú)夤潭ㄞD(zhuǎn)化為茶樹生長所需的氮素，增加土壤氮肥含量。綠肥植物自然凋落或收割覆蓋，經(jīng)微生物降解后，會(huì)產(chǎn)生大量有機(jī)質(zhì)。

2.6 增加微生物豐度和多樣性

土壤有益微生物具有解磷、釋鉀、固氮，以及增強(qiáng)土壤保濕性等優(yōu)點(diǎn)，隨著茶樹種植年限的增加，會(huì)降低茶園土壤微生物群落多樣性。間套作可誘導(dǎo)有益菌群在土壤中定殖和改善根際微生物群落組成的結(jié)構(gòu)特征[39]。在茶園間作大豆能改變土壤微生物的活性和多樣性，調(diào)節(jié)土壤放線菌、細(xì)菌和真菌的種類及其數(shù)量[101]；間作圓葉決明能夠提高固氮菌和放線菌的數(shù)量[102]。

2.7 提高酶活性

土壤酶催化各種生物化學(xué)反應(yīng)，提高酶活性將促進(jìn)土壤養(yǎng)分供應(yīng)和轉(zhuǎn)化。在茶園間套作功能植物不僅增加植物根系向土壤中釋放的酶量，還增加微生物向土壤中釋放的酶量。楊林等[103]發(fā)現(xiàn)茶行間作光葉紫花苕（）會(huì)提高土壤磷酸酶、蔗糖酶和脲酶的活性。

3 間套作功能植物對(duì)茶園病蟲草害的影響

3.1 間套作功能植物調(diào)控茶園生物種群

昆蟲在生態(tài)系統(tǒng)中有著重要的調(diào)節(jié)作用，具有傳粉、生物控害、傳播種子、分解動(dòng)植物的殘?bào)w和排泄物，以及觀賞等功能，在生態(tài)系統(tǒng)中能夠參與調(diào)節(jié)物質(zhì)循環(huán)、能量流動(dòng)和信息傳遞等過程[104-105]。間套作模式形成適度蔭蔽、溫濕度變幅小的穩(wěn)定微域，有利于天敵類群的定殖繁衍[11,73]。提高生物多樣性，有利于形成復(fù)雜且穩(wěn)定的食物網(wǎng)，但也可能會(huì)增加有害種群的比例[106-107]。不同種植模式可改變茶園生物群落的空間格局和益害比，以及提高生境結(jié)構(gòu)復(fù)雜的茶園中節(jié)肢動(dòng)物群落的物種豐富度和多樣性[35,42]。陳亦根等[52-53]發(fā)現(xiàn)在茶園間作長節(jié)耳草（）提高了物種豐富度和茶園生態(tài)系統(tǒng)穩(wěn)定性。

3.1.1 提高天敵類群

在茶園間套作功能植物有助于提高節(jié)肢動(dòng)物群落多樣性和天敵群落多樣性[65,108]。生境結(jié)構(gòu)復(fù)雜的茶園增加蜘蛛物種豐富度和多樣性指數(shù)，增強(qiáng)蜘蛛對(duì)有害生物的跟隨效應(yīng)和控害能力[42,109-110]。與簡單生境茶園相比，復(fù)雜生境茶園中狼蛛科雌蛛、雄蛛和幼蛛個(gè)體數(shù)分別增加了4.36、3.63、7.91倍[111]；但Chen等[64]在茶園間作百喜草（）或圓葉決明，蜘蛛物種豐富度和個(gè)體數(shù)與自然留養(yǎng)雜草對(duì)照茶園的差異不顯著。間套作功能植物能夠提高茶園捕食螨優(yōu)勢類群圓果大赤螨（）的個(gè)體數(shù)，充分發(fā)揮以螨治蟲的生態(tài)控制效應(yīng)[48,65]。Chen等[112]研究還發(fā)現(xiàn)，在間作百喜草或圓葉決明的茶園中，鞘翅目捕食性天敵個(gè)體數(shù)顯著高于自然留養(yǎng)雜草茶園的。間套作功能植物有助于提高寄生蜂物種豐富度、個(gè)體數(shù)和多樣性指數(shù)，提高寄生蜂壽命及寄生率，從而增強(qiáng)其對(duì)茶園主要害蟲茶小綠葉蟬、茶尺蠖的防控效果[48,113]。李慧玲等[69]在茶園間作白車軸草或圓葉決明，纓小蜂蟲口數(shù)與空白對(duì)照相比提高了61.98%。Chen等[49]也發(fā)現(xiàn)間作功能植物茶園能增加纓小蜂類、繭蜂類、姬小蜂類等寄生蜂及捕食性天敵個(gè)體數(shù)，降低茶尺蠖和薊馬個(gè)體數(shù)。

在有害生物流行暴發(fā)期，利用致病力強(qiáng)的蟲生真菌可以較快地降低有害生物種群數(shù)量，對(duì)黑刺粉虱（）等害蟲具有較好的防控效果[114-115]。韓寶瑜等[114,116]調(diào)查了皖南林茶間作茶園，發(fā)現(xiàn)能夠侵染粉虱、蚧殼蟲及鱗翅目害蟲的多種真菌種類。聯(lián)合利用蟲生真菌和天敵昆蟲，可以提高對(duì)茶園有害生物的防控效果，但，目前有關(guān)間套作功能植物強(qiáng)化茶園寄生菌生物防治的研究相對(duì)較少[114]。

3.1.2 降低有害生物類群

茶小綠葉蟬是茶園分布最廣、為害最重的吸汁性害蟲，目前我國各大茶區(qū)小綠葉蟬優(yōu)勢種為小貫小綠葉蟬[117-118]。在茶園間套作功能植物能夠有效驅(qū)避茶小綠葉蟬、減少茶小綠葉蟬著卵量和種群數(shù)量，從而抑制茶小綠葉蟬種群暴發(fā)；同時(shí)，還有研究發(fā)現(xiàn)能夠降低綠盲蝽的種群數(shù)量[58,65,119]。但，陳李林等[57]短期研究結(jié)果發(fā)現(xiàn)，間作功能植物茶園未顯著降低茶小綠葉蟬網(wǎng)捕量和茶梢著卵量。韓寶瑜等[120-121]通過比較發(fā)現(xiàn)，生境結(jié)構(gòu)復(fù)雜的有機(jī)茶園天敵多、害蟲少，植食性螨類和茶尺蠖發(fā)生均較輕。江麗容等[122]研究發(fā)現(xiàn)薄荷和迷迭香等植物對(duì)茶尺蠖具有吸引作用，而薰衣草（）則具有強(qiáng)烈的驅(qū)避效果。張正群等[123-124]發(fā)現(xiàn)了多種具有驅(qū)避氣味的，能夠調(diào)控茶尺蠖行為的植物精油和提取液。葉火香等[40-41]發(fā)現(xiàn)在茶園間套作吊瓜（）或柑桔（）能有效抑制茶蚜（）和粉虱的種群數(shù)量，同時(shí)改變節(jié)肢動(dòng)物垂直空間分布，抑制害蟲為害。

3.1.3 調(diào)控其他生物類群

食腐的蚯蚓和彈尾蟲是農(nóng)業(yè)系統(tǒng)重要組成部分，不僅可以降解枯枝落葉等有機(jī)物，促進(jìn)物質(zhì)循環(huán)，還可以提高土壤排水透氣性。在茶園間作馬唐（）[125]、白車軸草[73]、山毛豆（）[35]等功能植物可增加土壤中蚯蚓數(shù)量和生物量，改善土壤理化性質(zhì)。陳李林等[126]發(fā)現(xiàn)茶園間作百喜草或圓葉決明可提高凋落層彈尾蟲個(gè)體數(shù)。

3.2 間套作功能植物降低病害發(fā)生

間套作促進(jìn)作物對(duì)養(yǎng)分的吸收利用，稀釋、阻擋或驅(qū)避病原菌，提高寄主作物生理生化抗性，抑制帶毒昆蟲種群密度，進(jìn)而抑制病害的發(fā)生發(fā)展[127]。在茶園間套作功能植物能降低茶餅病、赤星病、煤煙病、炭疽病、輪斑病及根結(jié)線蟲的發(fā)病率[19,29,89]。前人研究發(fā)現(xiàn)，間作大豆茶園炭疽病發(fā)病率降低74.34%，茶餅病降低85.12%[29]；間作玉米茶園炭疽病降低49.76%[29]；套種香樟（）茶園輪斑病和赤星病的發(fā)病率分別降低50.47%和60.00%[19]。

3.3 間套作功能植物降低雜草發(fā)生

在茶園間套作功能植物能夠優(yōu)先占據(jù)生態(tài)位，增強(qiáng)雜草種間競爭，利用作物群體優(yōu)勢原理進(jìn)行生態(tài)控草，降低雜草種類和密度；間套作的功能植物刈割后覆蓋于茶行中間能夠阻擋光照和空氣，導(dǎo)致草種不能萌發(fā)[37]。幼齡茶園空白生境比重較大，通過間套作大豆、花生（）等作物，能充分抑制雜草蔓延。羅旭輝等[128]發(fā)現(xiàn)在茶園間作圓葉決明2年后，惡性雜草馬唐密度減少到每平方米7株。張永志等[92]發(fā)現(xiàn)茶園間作鼠茅草（），對(duì)雜草株防效為88.96%，鮮重防效為89.36%。

4 間套作功能植物對(duì)茶樹生長發(fā)育及茶葉產(chǎn)量品質(zhì)的影響

合理間套作功能植物能夠促進(jìn)茶樹對(duì)光的吸收，降低茶葉中酚氨比，提升茶葉產(chǎn)量和品質(zhì)[21,47,61,129]。烏龍茶鮮嫩茶梢的增加，會(huì)吸引茶小綠葉蟬取食，經(jīng)發(fā)酵后可制成濃厚香醇，具有果香的東方美人茶[130]。

4.1 間套作功能植物對(duì)茶樹生長發(fā)育的影響

4.1.1 提高光合速率

茶樹喜漫射光，對(duì)光強(qiáng)需求較低，茶園適度遮陰可調(diào)節(jié)光照強(qiáng)度、有利于改善光質(zhì)，促進(jìn)茶樹春梢生長[131]。間套作能夠減緩茶園光合輻射變化，消除茶樹“午休”現(xiàn)象[20]；降低呼吸速率，提高光合作用[71,132]。功能植物對(duì)光照具有阻擋、吸收、反射等作用，減少因光照過強(qiáng)或直射時(shí)間過長而導(dǎo)致茶樹多花早衰[14]。張明澤等[51]發(fā)現(xiàn)在茶園間作巨菌草（）能促進(jìn)茶葉中葉綠素a的合成，提高葉綠素a/b的比值。間套作功能植物對(duì)茶園的遮光度在30%左右最佳[13]，過度遮陰會(huì)影響茶園通風(fēng)透光性，造成葉部病害加重。

4.1.2 促進(jìn)根系生長

合理間套作功能植物能夠改善土壤環(huán)境，促進(jìn)根系生長。嚴(yán)芳等[63]發(fā)現(xiàn)在茶園間作白車軸草，茶樹根系總長度、表面積、體積和分叉數(shù)分別是單作茶園的2.1、2.6、2.7、4.0倍。余立華等[133]發(fā)現(xiàn)茶園套種栗樹（），能夠提高茶樹根系吸收根比例、比根長和根系活性，其中套種茶園的茶樹吸收根比常規(guī)茶園高19.27%。

4.1.3 增強(qiáng)茶樹長勢

間套作功能植物能夠改變茶園微域氣候，為茶樹提供更適宜的生長條件，提早茶芽萌發(fā)期和全盛期[30-31,56]，增快展葉速度[23-24,92]。與對(duì)照茶園相比，間作花生后茶樹株高增加13.0%，樹幅增大19.5%，茶苗成活率提高6.5%；間作大豆和花生后茶樹枝干直徑增加21.8%[34]。

4.2 間套作功能植物對(duì)茶葉品質(zhì)的影響

茶葉水浸出物主要包括多酚類、氨基酸和咖啡堿等，決定了茶湯的湯色和滋味；茶多酚與氨基酸的比值決定茶葉鮮爽度，酚氨比低的茶湯鮮爽度較高。施肥過多會(huì)導(dǎo)致茶葉品質(zhì)下降，苦味增加[134]。在茶園間套作功能植物有利于茶葉中氨基酸和蛋白質(zhì)的合成，調(diào)節(jié)可溶性糖、兒茶素、沒食子酸和咖啡堿的含量，降低酚氨比，改善茶葉品質(zhì)[36,45,135]。在茶園套種栗樹會(huì)改變茶樹氨基酸代謝機(jī)制，提高茶葉的鮮爽度和香氣[15-16]。在茶園套種核桃或榿木（）能夠降低茶葉中粗纖維含量，提高鮮嫩度[136]。套種蘋果（）能夠降低茶葉葉片厚度和角質(zhì)層，增加海綿組織和柵欄組織厚度；其中葉片厚度減少0.158%～0.187%，海綿組織增大0.194%～0.462%[44]。間作榆黃菇（），茶水浸出物中茶多酚和花青素含量比單種茶樹分別增加1.04%和0.13%[54]。

4.3 間套作功能植物對(duì)茶葉產(chǎn)量的影響

種植功能植物能夠改變茶園微域氣候，為茶樹提供更適宜生長條件，有利于提高茶樹新梢長度、發(fā)芽密度和百芽重[23-24,92]。與對(duì)照茶園相比，黃槐（）-茶樹-野花生（）的復(fù)合種植模式能夠增加茶樹芽長17.58%、百芽鮮重33.12%、百芽干重39.88%[137]。

5 問題及展望

5.1 常見問題

不科學(xué)合理的間套作措施，將導(dǎo)致間套作植物與茶樹存在養(yǎng)分和光能競爭，會(huì)抑制茶樹生長，甚至降低茶葉品質(zhì)產(chǎn)量，達(dá)不到生境管理的目的。同時(shí)，過度遮陰會(huì)增加水分積累，影響透氣透光，造成葉部病害加重。間套作時(shí)應(yīng)考慮功能植物的高度及種植密度[13]。茶園套種厚樸（）或李樹，明顯加重茶餅病與茶炭疽病發(fā)病[29]。若間套作植物存在較強(qiáng)化感作用或與茶樹產(chǎn)生水肥競爭，則會(huì)抑制茶樹的生長。如在茶園套種巨桉（）或核桃，巨桉根系和葉片中會(huì)分泌大量的化感物質(zhì)[138]，核桃根系會(huì)分泌化感物質(zhì)胡桃醌，從而導(dǎo)致茶樹生長不良和茶葉品質(zhì)下降[139]。

與稻田、菜地等農(nóng)田生態(tài)系統(tǒng)相比，傳統(tǒng)茶園生物群落結(jié)構(gòu)復(fù)雜，可長期保持動(dòng)態(tài)平衡。在通過間套作措施構(gòu)建立體復(fù)合型生態(tài)茶園時(shí)，需要保留一定面積的天然原生植被，可作為本地生物類群的“庫”，以避免造成單一種植功能植物后，茶園的生物多樣性降低[13]。間套作具有誘蟲功能的植物時(shí)，還需要考慮功能植物的種植區(qū)域，防止害蟲向茶園的正向轉(zhuǎn)移[13]。選擇間套作的功能植物時(shí)，需要注意避免與茶樹有共同病蟲害的植物，李祥英[140]發(fā)現(xiàn)在幼齡茶園間作番薯（）后，會(huì)引起茶苗根結(jié)線蟲為害，造成茶樹死亡。

與化學(xué)肥料相比，間套作功能植物提供的養(yǎng)分含量較低、起效慢；如果長期只依靠功能植物為茶樹提供養(yǎng)分，會(huì)造成產(chǎn)量下降、品質(zhì)降低等問題。同時(shí)，受傳統(tǒng)認(rèn)知“草與茶爭肥”的影響，以及種植功能植物需要進(jìn)行開溝、播種和刈割覆蓋等農(nóng)事操作，導(dǎo)致人工成本增加，降低茶農(nóng)在茶園間套作功能植物的積極性。目前在實(shí)際生產(chǎn)過程中，適合茶園種植的新品種選育較為緩慢，種系逐漸退化，可能轉(zhuǎn)變?yōu)殡s草[141]。在茶園間作套種功能植物提高茶葉品質(zhì)產(chǎn)量的效果明顯，但技術(shù)體系尚未完善，生產(chǎn)過程中缺乏相應(yīng)的技術(shù)指導(dǎo)，難以形成大規(guī)模種植。

5.2 展望

間套作功能植物是一種常見的生境管理措施，通過科學(xué)合理地引入功能植物或保護(hù)當(dāng)?shù)刈匀恢脖坏姆绞?，增加有益生物類群的豐度和活性，強(qiáng)化茶園保益控害的生態(tài)系統(tǒng)服務(wù)功能[142-143]。合理的間套作可以通過利用不同植物種間促進(jìn)和互利共生原理，控制茶園病蟲草害的發(fā)生流行，減少化學(xué)肥料和化學(xué)農(nóng)藥施用，提高土地利用率，最終提升茶葉品質(zhì)和產(chǎn)量，保障茶葉安全優(yōu)質(zhì)生產(chǎn)[144-145]。

僅依靠間套作功能植物調(diào)節(jié)茶園生物類群來控制有害生物的發(fā)生是遠(yuǎn)遠(yuǎn)不夠的，還需要結(jié)合其他綠色防控措施進(jìn)行生態(tài)調(diào)控。對(duì)茶園有害生物進(jìn)行生態(tài)調(diào)控應(yīng)從大尺度出發(fā)，拓展至區(qū)域性農(nóng)田景觀的空間范圍。通過景觀設(shè)計(jì)對(duì)茶園植物進(jìn)行合理布局，協(xié)調(diào)“茶樹-病蟲草害-天敵及茶園生態(tài)環(huán)境”相互之間的關(guān)系，提高茶園景觀中有益生物的生物控害服務(wù)功能，將有害生物控制在生態(tài)經(jīng)濟(jì)閾值水平之下，維持茶園生態(tài)系統(tǒng)穩(wěn)定的動(dòng)態(tài)平衡，構(gòu)建長期有效的茶園有害生物生態(tài)調(diào)控體系[3,142]。

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Effects of Intercropping Functional Plants on the Ecosystem Functions and Services in Tea Garden

SHI Fan1,2,3, HUANG Hongjing1,2,3, CHEN Yanting4, CHEN Lilin1,2,3*

1.State Key Laboratory of Ecological Pest Control for Fujian and Taiwan Crops, college of Plant Protection,Fujian Agriculture and Forestry University, Fuzhou 350002, China; 2.Institute of China White Tea, Fuding 355200, China; 3.Key Laboratory of Green Control of Insect Pests, Fuzhou 350002, China; 4.Institute of Plant Protection, Fujian Academy of Agricultural Sciences, Fuzhou 350013, China

As one of the main measures of habitat management, reasonable intercropping of functional plants in tea garden can shade tea bushes and keep them warm, conserve water and soil, increase fertility and promote growth, as well as maintain micro-habitat stability.It also can attract natural enemies, repel pests, reduce the damage caused by diseases, insects, and weeds in tea garden, therefore it is beneficial for improving the quality and efficiency of the tea.However, unreasonable intercropping will destroy the micro-habitat of tea garden, compete for nutrients, thus affecting the growth of tea plants.In this paper, the intercropping of functional plants and their effects on the ecosystem functions and services in tea garden in recent years were reviewed.These effects mainly included the species, management models, functions and common problems with the most widely used functional plants, as well as the regulatory effects of intercropping of functional plants on the growth and development of tea plants, tea quality and yield, and tea pests.The ultimate goal of this paper was to provide guidance for the application of intercropping measures to promote the comprehensive regulation of pests in tea garden, and enhance the ecosystem functions and services of tea garden.

Intercropping, habitat management, ecological control, ecosystem services, ecosystem functions

S571.1

A

1000-369X(2022)02-151-18

2021-07-03

2021-11-02

福建省創(chuàng)新戰(zhàn)略研究計(jì)劃項(xiàng)目（2020R0036）、中國白茶研究院開放課題項(xiàng)目（白茶研院〔2021〕5號(hào)）、女茶師非遺傳習(xí)所科技特派員工作站（安科特派辦〔2021〕3號(hào)）、國家自然科學(xué)基金（31501650）、福建農(nóng)林大學(xué)科技創(chuàng)新專項(xiàng)基金（CXZX2019009G）

史凡，男，碩士研究生，主要從事茶園有害生物生態(tài)調(diào)控。*通信作者：llchen@fafu.edu.cn

（責(zé)任編輯：趙鋒）