王海斌 陳曉婷 趙虎 王裕華 張清旭 汪鵬 葉江華 丁力



摘 ?要??為了分析茶樹根際土壤物質對茶樹根際土壤微生物多樣性的影響,本研究以植茶年限0、3、9、25?a的鐵觀音茶樹根際土壤為材料,采用不同極性樹脂吸附茶樹根際土壤物質并洗脫,探討不同植茶年限茶樹根際土壤物質的自毒潛力及其對土壤微生物多樣性的影響。結果表明,不同極性樹脂吸附洗脫液以ADS-7樹脂洗脫液對受體根長的抑制作用最強。ADS-7樹脂吸附洗脫液處理重新種植的茶樹后,隨著土壤植茶年限的增加,茶樹根際土壤細菌數(shù)量呈現(xiàn)下降趨勢。相關性分析結果表明,與土壤年限呈正相關的細菌T-RFs片段15個,涉及8個綱,31種細菌,按照其功能可分為4類,其中病原菌19種,占比61.29%;負相關細菌T-RFs片段18個,涉及11個綱,31種細菌,按照其功能可分為6類,其中與抑制病原菌、碳素循環(huán)、氮素循環(huán)、硫素循環(huán)、土壤質地改善相關的細菌總占比達到83.87%。綜上表明,ADS-7樹脂洗脫液處理重新種植的茶樹后,隨著土壤植茶年限的增加,茶樹根際土壤病原菌數(shù)量大幅上升,益生菌與土壤養(yǎng)分循環(huán)相關的細菌數(shù)量顯著下降,土壤微生物生態(tài)系統(tǒng)平衡失調(diào)。
關鍵詞 ?茶樹根際土壤;自毒作用;物質;微生物多樣性中圖分類號??S571.1; S154.3??????文獻標識碼??A
The Autotoxicity?ofTea Tree Rhizosphere Soil Chemicals and the Effect of Soil MicrobialDiversity
WANG?Haibin1,2,?CHEN?Xiaoting1,2, ZHAO Hu1,3, WANG Yuhua1, ZHANG Qingxu2, WANG Peng2,YE?Jianghua2,4, DING Li1
1. College of Life Sciences, Longyan University, Longyan, Fujian 364012, China; 2. Fujian Provincial Key Laboratory of Agroecological Processing and Safety Monitoring, Fujian Agriculture and Forestry University, Fuzhou, Fujian?350002, China; 3.?College of Horticulture, Gansu Agricultural University, Lanzhou, Gansu 730070, China; 4. College of Tea and Food Science, Wuyi University, Wuyishan, Fujian?354300, China
Abstract ?In order to analyze the effect?of tea tree rhizosphere soil chemicals on the soil microbial?diversity, the rhizosphere soils from Tieguanyin tea plantations of 0, 3, 9 and 25 years old were extracted and eluted through different polar resins to discuss the autotoxicity?of resins?eluted?and the effect of soil microbial?diversity. The results showed that the inhibitory effect?of ADS-7 resin eluent?on the root length of receptor?was the strongest. After the replanted tea trees was treated by ADS-7 resin eluent, the number of bacteria in the tea trees rhizosphere?soil?decreased with the increase of planting soil age.?Correlation analysis result showed that 15 T-RFs from bacterial?community were significantly?and positively?correlated?with soil?age, which included 31 species of microbes belonging to?8?classes, respectively. Thirty-one microbes?could?be divided into?4 types according to the function, among them, 19 species of pathogenic bacteria accounted for 61.29%.?In addition, 18 T-RFs were significantly and negatively?correlated?with soil age, which included 31 species of bacteria belonging to?11?classes, respectively.?The thirty-one bacteria?could?be divided into?6 types according to the function, and total percentage of the bacteria to inhibit 83.87%. In brief, after the replanted tea trees was treated by ADS-7 resin eluent, the number of pathogenic bacteria?in tea tree?pathogenic bacteria, carbon cycle, nitrogen cycle, sulphur cycle and the bacteria?to?improve the soil quality accounted for?rhizosphere soil?was great enhanced and the number of probiotics and?soil nutrient cycling?bacteria?decreased significantly?as planting soil age increased, which led to the imbalance of soil microbial ecosystem.
Keywords ?rhizosphere soil of tea tree; autotoxicity; chemical; microbial diversity
DOI10.3969/j.issn.1000-2561.2019.09.025
茶樹[Camellia sinensis(L.)?O.?Kuntze]屬于山茶科、山茶屬灌木或小喬木茶種,為多年生常綠木本植物。鐵觀音茶園的開墾、種植到采摘需要2~3年,正常經(jīng)濟效益旺期在7年左右。21世紀初,安溪鐵觀音進入飛速發(fā)展時期。大型茶葉企業(yè)紛紛在安溪縣及周邊縣市大量開墾新茶園,種植鐵觀音茶樹,并按公司化方式統(tǒng)一管理,安溪縣周圍的山地已形成以茶樹為主要植物種群的單一群落結構。為此,茶園“土壤病”形成,茶園逐漸退化,茶樹病蟲害加劇,茶葉單產(chǎn)水平及品質也逐年下降。茶園退化一方面是茶樹本身自然衰老,另一方面茶樹連年種植后,土壤環(huán)境發(fā)生變化,不利于茶樹生長的因素積累,土壤自毒作用加劇。
王海斌等[1]調(diào)查分析了安溪縣9個鄉(xiāng)鎮(zhèn)茶園土壤的酸化情況發(fā)現(xiàn),調(diào)查的茶園中37.67%的土壤已經(jīng)酸化,10.03%的土壤不適宜種植茶樹,茶樹樹齡與其根際土壤pH呈極顯著負相關,茶葉的產(chǎn)量、品質與茶樹根際土壤pH呈極顯著正相關,該研究認為,茶樹根際土壤酸度隨著茶樹樹齡的增加而加劇,茶葉產(chǎn)量降低、品質呈現(xiàn)下降趨勢。Ye等[2-3]研究也發(fā)現(xiàn),隨著茶樹樹齡的增加,茶樹根際土壤酸度加劇,自毒作用潛力增強,茶葉產(chǎn)量和品質降低,這種現(xiàn)象的形成與土壤中酸類物質的積累增加有關。本課題組前期對不同樹齡茶樹根際土壤物質進行HPLC分析發(fā)現(xiàn),隨著茶樹樹齡的增加,茶樹根際土壤中6種酸類物質含量不斷上升。此外,進一步采用不同極性樹脂吸附土壤物質并進行生物測試發(fā)現(xiàn),以ADS-7樹脂吸附后的洗脫液自毒作用能力最強,GC-MS分析ADS-7樹脂洗脫液的物質成分,發(fā)現(xiàn)13種物質隨著茶樹樹齡的增加呈現(xiàn)上升趨勢,其中9種是酸類物質[4-5]??梢?,退化茶園“土壤病”的形成與茶樹根際土壤物質種類與數(shù)量密切相關。
土壤是一個復雜生態(tài)系統(tǒng),存在著豐富的微生物種類,植物釋放和積累的物質通過土壤載體進行傳播,進而影響土壤中的微生物區(qū)系與種類,土壤生態(tài)系統(tǒng)朝著專一化的趨勢發(fā)展[6-8]。綜上可見,隨著茶樹樹齡的增加,土壤自毒潛力加劇,茶樹根際土壤微生物發(fā)生顯著變化[9-10];而關于這種變化是否與土壤物質有關,土壤物質對微生物多樣性變化是否會產(chǎn)生影響的研究還鮮有報道。據(jù)此,本研究以不同樹齡鐵觀音茶樹根際土壤為材料,采用不同極性樹脂進行吸附并洗脫,土壤洗脫液一方面用于自毒潛力評價,一方面用于處理新種植的茶樹并測定茶樹根際土壤微生物種類、群落結構及其功能的變化,以期為茶園土壤退化的修復提供一定的理論依據(jù)。
1 ?材料與方法
1.1材料
以福建省泉州市安溪縣龍涓鄉(xiāng)鐵觀音茶園為研究地點(東經(jīng)117°93′、北緯24°97′),收集已種植0、3、9、25年的鐵觀音茶樹根際土壤,用于土壤物質提取及不同極性樹脂的吸附與洗脫后的生物測試。土壤取樣方法:隨機選擇3、9、25?a樹齡的茶樹各100株,去除土壤表層雜質,連根挖出茶樹,收集茶樹根際土壤;以未種植過茶樹的土壤為對照(0?a),首先去除地表植被和雜質,收集15~25?cm范圍的土壤,多點隨機收集;各樣品的取樣量均為約15 kg。取樣點茶園土壤的基本理化指標:有機質、全氮、全磷、全鉀、速效氮、速效磷、速效鉀含量,分別為8.34 g/kg、2.17 g/kg、1.05 g/kg、1.46 g/kg,25.3 mg/kg、56.7 mg/kg、264.6 mg/kg。
1.2方法
1.2.1 ?不同極性樹脂洗脫液對受體萵苣(Lactuca saliva)根長的抑制率分析??土壤樣品置于陰涼處自然風干,研磨過40目篩,稱取0、3、9、25?a茶樹根際土壤各5 kg,加入20 L蒸餾水,360 W超聲提取1 h(每隔10?min均勻攪拌1次),120?r/min振蕩1 h,重復5次,過濾,提取液于45?℃下旋轉蒸發(fā)濃縮至5 L;此時,每毫升浸提液約含有1 g土壤物質[11]。
先將不同極性的樹脂ADS-7、ADS-21、ADS-F8、ADS-17、ADS-8(天津南開合成科技有限公司)采用純乙醇浸泡活化24 h,蒸餾水浸泡清洗至沒有乙醇。取收集的不同樣品提取濃縮液各5 L,分成5組,每組1 L,采用5種不同極性樹脂分別進行靜態(tài)吸附,方法為:每升提取濃縮液加入200 g樹脂,置于搖床上120 r/min振蕩吸附24 h,棄上清液,樹脂中加入600 mL甲醇,置于搖床上120 r/min洗脫12 h,收集甲醇洗脫液過濾并濃縮至200 mL,用于不同極性樹脂洗脫液對受體萵苣根長的抑制率測定,具體參照Wang等[12]的方法進行測定,根長抑制率的計算公式為,相對抑制率=(1-處理值/對照值)×100%。
1.2.2 ?外源添加不同極性樹脂洗脫液處理新種植的茶樹??取不同極性樹脂吸附后的洗脫液各50?mL,于45?℃下旋轉蒸發(fā)濃縮至10 mL,加無菌蒸餾水定容至2 L,-20?℃保存?zhèn)溆谩?/p>
取未種植過茶樹的土壤風干研磨并過40目篩;將土壤裝入盆中,每盆10 kg,選擇1年生、長勢相對一致的鐵觀音茶苗并移栽到盆中,每盆6株,移栽后,恢復生長30 d。適當攪動、松動種植土壤,將配置好的樹脂洗脫液2 L緩慢倒入盆中,盡量使其在土壤中分布均勻,繼續(xù)常規(guī)種植茶樹60 d,收集茶樹根際土壤,用于土壤微生物的T-RFLP分析,每種處理種植3盆,即3個重復。種植土壤的基本理化指標為:有機質、全氮、全磷、全鉀、速效氮、速效磷、速效鉀含量,分別為9.02 g/kg、1.03?g/kg、0.56?g/kg、1.85?g/kg、89.46?mg/kg、15.28?mg/kg、179.62?mg/kg。
1.2.3 ?土壤微生物的T-RFLP分析??采用CTAB-蛋白酶K-液氮凍融法直接抽提不同樣品的土壤微生物總DNA,用于微生物的16S?rDNA的擴增和酶切[13]。16S?rDNA擴增的PCR反應程序為:94?℃?5?min;94?℃?45?s,52?℃?45?s,72?℃?1.5?min,30個循環(huán)后,72?℃?10?min。其中,擴增引物采用帶有FAM熒光標記的細菌通用引物,分別為8F-FAM(5′-AGAGTTTGATCCTGGCTCA G-3′)和926R(5′-CCGTCAATTCCTTTRAGTT T-3′),PCR反應體系的總體積為25?μL,反應體系中各成分含量為:2.5?μL?10×PCR Buffer、正反向引物(10?μmol/L)各0.8?μL、2.0?μL?dNTP(25?μmol/L)、0.2?μL?BSA、17.05?μL?ddH2O、0.15?μL?rTaq、1.5?μL?DNA模板。PCR結束后,電泳檢測,UNIQ-10柱式DNA膠回收試劑盒回收PCR產(chǎn)物中1000?bp左右的片段,用于酶切。
酶切采用HaeIII和MspI 2種內(nèi)切酶進行消化,其中HaeIII酶切體系為:HaeIII?1?μL、H×Buffer?2?μL、ddH2O?7?μL、PCR產(chǎn)物10?μL;MspI酶切體系為:MspI?1?μL、T-Buffer?2?μL、BSA?2?μL、ddH2O?5?μL、PCR產(chǎn)物10?μL。將配置好的2種酶切體系分別置于37?℃水浴酶切5?h,酶切后的產(chǎn)物采用ABI自動測序分析儀(Model 3130 Applied Biosystems)測定。
1.3數(shù)據(jù)處理
土壤微生物測序結果分析采用GeneMarker V1.2軟件,分析參數(shù)參照SoftGenetics Application Note July, 2006。酶切獲得的T-RFs片段分析采用Ribosomal Database Project II數(shù)據(jù)庫比對法,獲取T-RFs雙酶切片段所對應的微生物物種。T-RFs片段豐度計算公式為:T-RFs片段豐度=ni/N×100,式中ni代表可分辨的T-RF的峰面積,N代表所有T-RF峰面積的總和[14]。其余常規(guī)的數(shù)據(jù)分析、方差分析、顯著性分析及變化分析等采用Excel軟件和DPS數(shù)據(jù)處理系統(tǒng)進行處理。
2??結果與分析
2.1不同極性樹脂洗脫液對受體萵苣根長的抑制率分析
不同極性樹脂洗脫液對受體萵苣根長影響的分析結果表明(表1),不同極性樹脂洗脫液對受體根長均存在一定的抑制作用,以25?a茶樹根際土壤的樹脂吸附洗脫液抑制作用最強,表現(xiàn)為ADS-8、ADS-17、ADS-F8、ADS-21、ADS-7樹脂洗脫液對萵苣根長的抑制率分別為15.76%、19.68%、20.57%、29.07%、43.76%。其中以25?a茶樹根際土壤經(jīng)ADS-7樹脂吸附的洗脫液抑制作用最強。
2.2 ?ADS-7樹脂洗脫液處理后茶樹根際土壤細菌多樣性分析
由圖1可見,0、3、9、25?a茶樹根際土壤經(jīng)ADS-7樹脂洗脫液處理后,根際土壤細菌HaeIII和MspI酶切產(chǎn)物的T-RFs片段,隨著土壤年限的增加呈現(xiàn)下降趨勢,表現(xiàn)為HaeIII酶切產(chǎn)物的T-RFs片段數(shù)量從106個下降至58個,而MspI酶切產(chǎn)物的T-RFs片段數(shù)量則從124個下降至68個??梢?,不同種植年限土壤的ADS-7樹脂吸附洗脫液處理后,茶樹根際土壤細菌多樣性發(fā)生顯著的變化。
2.3 ?ADS-7樹脂洗脫液處理后茶樹根際土壤正、負相關細菌分析
以不同茶樹根際土壤細菌HaeIII酶切后的T-RFs片段豐度進行相關性分析發(fā)現(xiàn)(圖2),不同年限茶樹根際土壤的ADS-7樹脂吸附洗脫液處理后,與土壤種植年限呈顯著或極顯著正相關的細菌T-RFs片段共15個,呈顯著或極顯著負相關的有18個。
進一步將T-RFs片段與Ribosomal Database Project II數(shù)據(jù)庫進行比對,結果表明(圖3),正相關細菌T-RFs片段15個,涉及31種細菌,由8個綱組成,占比分別為β-變形菌綱12.90%、γ-變形菌綱16.13%、δ-變形菌綱6.45%、芽孢桿菌綱38.71%、放線菌亞綱9.68%、鞘脂桿菌綱9.68%、梭菌綱3.23%、螺旋體綱3.23%;負相關細菌T-RFs片段18個,涉及31種細菌,由11個綱組成,占比分別為α-變形菌綱25.81%、β-變形菌綱6.45%、γ-變形菌綱12.90%、放線菌亞綱19.35%、梭菌綱12.90%、芽孢桿菌綱6.45%、螺旋體綱3.23%、柔膜菌綱3.23%、鞘脂桿菌綱3.23%、紅色桿菌亞綱3.23%、梭桿菌綱3.23%。可見,不同種植年限土壤的ADS-7樹脂吸附洗脫液處理后,茶樹根際土壤細菌種類發(fā)生顯著變化。
2.4 ?ADS-7樹脂洗脫液處理后茶樹根際土壤正相關細菌功能分析
由表2可見,0、3、9、25年茶樹根際土壤經(jīng)ADS-7樹脂洗脫液處理后,茶樹根際土壤正相關細菌有31種,按照其功能可分為4大類:病原菌19種,占比61.29%;抑制病原菌相關細菌3種,占比9.68%;碳素循環(huán)相關細菌4種,占比12.90%;改善土壤質地相關細菌5種,占比16.13%??梢?,不同種植年限土壤的ADS-7樹脂吸附洗脫液處理后,茶樹根際土壤不同功能類別的細菌數(shù)量發(fā)生顯著變化,特別是病原菌數(shù)量顯著增加,表現(xiàn)為正相關細菌中病原菌占比最大。
2.5 ?ADS-7樹脂洗脫液處理后茶樹根際土壤負相關細菌功能分析
由表3可見,0、3、9、25 a茶樹根際土壤經(jīng)ADS-7樹脂洗脫液處理后,茶樹根際土壤負相關細菌31種,按照其功能可分為6大類:病原菌5種,占比16.13%;抑制病原菌相關細菌4種,占比12.90%;碳素循環(huán)相關細菌10種,占比32.26%;氮素循環(huán)相關細菌8種,占比25.81%;改善土壤質地相關細菌3種,占比9.68%;硫素循環(huán)細菌1種,占比3.23%??梢?,年限土壤經(jīng)ADS-7樹脂吸附洗脫液處理后,茶樹根際土壤不同功能類別的細菌數(shù)量發(fā)生顯著變化,特別是與抑制病原菌、碳素循環(huán)、氮素循環(huán)、土壤質地改善、硫素循環(huán)相關的細菌數(shù)量顯著下降,總占比達到83.87%。
3??討論
植物土壤生態(tài)系統(tǒng),主要涉及植物、土壤、微生物三者,三者之間相互協(xié)調(diào),影響著植物的生長,土壤的質量,微生物的生存[63]。茶樹長期種植后,根系分泌物在土壤中大量積累,使土壤微生物在選擇性壓力影響下發(fā)生了顯著的變化,這種變化可能朝著對茶樹生長有利或有弊的方向發(fā)展。本研究結果表明,隨著茶樹樹齡的增加,茶樹根際土壤不同極性樹脂吸附洗脫液對受體萵苣根長存在一定的抑制作用,以ADS-7樹脂洗脫液的抑制作用最強。其次,不同種植年限土壤的ADS-7樹脂吸附洗脫液處理重新種植的茶樹后發(fā)現(xiàn),隨著土壤種植年限的增加,茶樹根際土壤細菌數(shù)量呈現(xiàn)下降趨勢。眾多學者在研究不同作物——太子參、山銀花、茶樹等連續(xù)種植后,土壤微生物數(shù)量變化時也發(fā)現(xiàn)類似的趨勢[64-66]??梢?,ADS-7樹脂洗脫液處理對茶樹根際土壤微生物數(shù)量產(chǎn)生影響,這種影響與不同年限茶樹原位種植現(xiàn)象類似。
作物長期種植,土壤物質積累后會對微生物數(shù)量與種類產(chǎn)生影響。Wang等[12]研究發(fā)現(xiàn),百香果連續(xù)種植后,產(chǎn)量和品質呈現(xiàn)下降趨勢;郝慧榮等[67]研究發(fā)現(xiàn),牛膝連年種植后,反而有利于促進其生長和品質提高??梢姡钊敕治鲎魑镞B續(xù)種植后土壤微生物種類及功能變化對于明晰“土壤病”的形成具有重要的意義。本研究結果表明,不同種植年限土壤的ADS-7樹脂吸附洗脫液處理重新種植的茶樹后,隨著土壤年限的增加,與其正相關細菌T-RFs片段15個,涉及8個綱、31種細菌,其中病原菌19種,占比61.29%??梢?,ADS-7樹脂吸附洗脫液處理后,茶樹根際土壤的病原菌數(shù)量增多,進而影響茶樹的生長。此外,進一步分析發(fā)現(xiàn),隨著土壤年限的增加,與其負相關細菌T-RFs片段18個,涉及11個綱、31種細菌,其中與抑制病原菌、碳素循環(huán)、氮素循環(huán)、土壤質地改善、硫素循環(huán)相關的細菌數(shù)量顯著下降,總占比達到83.87%??梢姡珹DS-7樹脂吸附洗脫液處理后,茶樹根際土壤的益生菌與土壤養(yǎng)分循環(huán)相關的細菌數(shù)量下降,茶樹根際土壤質地變劣,養(yǎng)分循環(huán)受阻。
土壤微生物是土壤生態(tài)系統(tǒng)的重要組成部分,其數(shù)量及多樣性水平高低對于生態(tài)系統(tǒng)穩(wěn)定具有重要的作用,豐富的土壤微生物有利于降低“土壤病”的發(fā)生機率,反之則提高[68-69]。可見,不同年限土壤的ADS-7樹脂吸附洗脫液處理重新種植的茶樹后,茶樹根際土壤細菌多樣性及群落結構失去平衡,進而可能導致茶樹生長受阻。
綜上,本研究探討了茶樹根際土壤物質的自毒潛力及其對土壤微生物多樣性的影響,結果表明,ADS-7樹脂吸附洗脫液對受體萵苣根長的抑制作用最強,ADS-7樹脂洗脫液處理重新種植的茶樹后,隨著土壤年限的增加,茶樹根際土壤病原菌數(shù)量大幅上升,益生菌與土壤養(yǎng)分循環(huán)相關的細菌數(shù)量顯著下降,土壤微生物生態(tài)系統(tǒng)平衡失調(diào),“土壤病”形成,進而可能導致茶樹生長和品質受阻。然而,對于茶樹—土壤—微生物,三者之間是如何實現(xiàn)相互調(diào)控與影響,還需進一步深入研究。
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