4种丛枝菌根真菌对南高丛蓝莓抗旱性的影响

许庆龙， 刘晓敏， 徐小兵， 李晴晴， 张红， 肖家欣

(安徽师范大学生命科学学院，安徽省重要生物资源保护与利用研究重点实验室，安徽 芜湖 241000)



4种丛枝菌根真菌对南高丛蓝莓抗旱性的影响

许庆龙， 刘晓敏， 徐小兵， 李晴晴， 张红， 肖家欣*

(安徽师范大学生命科学学院，安徽省重要生物资源保护与利用研究重点实验室，安徽 芜湖 241000)

采用盆栽试验研究摩西球囊霉(Glomusmosseae)、地表球囊霉(G.versiforme)、根内球囊霉(G.intraradices)和幼套球囊霉(G.etunicatum)4种丛枝菌根(arbuscular mycorrhizal，AM)真菌接种南高丛蓝莓(Vacciniumcorymbosum)品种薄雾对干旱胁迫的生理生化反应。结果表明：干旱胁迫降低了叶片相对含水量与叶绿素含量，增加了可溶性糖含量、超氧化物歧化酶(superoxide dismutase，SOD)活性与丙二醛(malondialdehyde，MDA)含量，但对菌根侵染率的影响较小；在干旱胁迫下，4种AM真菌尤其是摩西球囊霉接种株相对含水量、叶绿素和可溶性糖含量、叶片SOD活性均显著高于未接种株，而接种株叶片MDA含量相对低于未接种株；另外，AM真菌接种处理能够提高蓝莓植株根、茎、叶的磷和钾含量以及根围土壤酸性磷酸酶、脲酶和过氧化氢酶活性，尤以摩西球囊霉处理最为明显。总之，4种AM真菌接种处理提高了蓝莓品种薄雾植株的抗旱性，以摩西球囊霉的效果最好。

球囊霉属； 南高丛蓝莓； 干旱胁迫； 盆栽试验

Summary Blueberry (Vacciniumspp.) is well known for its rich anthocyanins and other bioactive compounds, which helps preventing from cardiovascular disease and other chronic illnesses. Over the past decades, blueberry has been cultured in many areas of China, thus became the fastest-growing species in fruit production, potentially making China one of the largest blueberry-producing countries in the world. However, blueberry growth and production were severely affected by drought events in areas of Yangtze River.Vacciniumplants preferred acidic soils and spontaneously formed mutualistic symbiotic associations mainly with soil fungi of the phylum Ascomycota, called ‘ericoid mycorrhizae’. However, some reports have indicated that the absence of ericoid mycorrhizal fungi inoculum may allowVacciniumplants to associate with arbuscular mycorrhizal (AM) fungi, and AM fungi inoculation significantly enhanced growth ofVacciniumplants. Southern highbush blueberry (Vacciniumcorymbosum) has short or even rare root hairs in field systems, and depends on fungi for optimal growth. Meanwhile, few reports were about the effects of AM fungi on osmotic adjustment and reactive oxygen metabolism of southern highbush blueberry.

The purpose of this study is to evaluate the effects ofGlomusmosseae,G.versiforme,G.intraradicesandG.etunicatumon the southern highbush blueberry seedlings under drought stress conditions, and antioxidant and osmotic adjustment matters, mineral nutrition in blueberry plants and enzymatic activities in rhizosphere soil were investigated.

Two soil water regimes (well-watered [WW] and drought stress [DS]) and five AM fungi inoculations (fourGlomusand non-AM fungi inoculation [CK]) were arranged in a complete randomized block design. Each treatment (one plant per pot) was performed in three replicates. Water treatments began after 134 days (July 31, 2014) adaption of greenhouse conditions, and WW pots were maintained, but DS pots were cut off for 20 days of water supply and resumed on August 20. Meanwhile, the leaves of blueberry cultivar Misty plants were collected for determination of physiological index on 13 d drought, 20 d drought and 2 d rewatering, respectively.

The results showed that the leaf relative water content (RWC) and chlorophyll contents decreased in DS treatments, while the soluble sugar content, superoxide dismutase (SOD) activity and malondialdehyde (MDA) contents increased, but had no significant effects on mycorrhizal colonization. Under the DS conditions, leaf RWC, chlorophyll, soluble sugar contents and SOD activity were significantly higher in AM fungi-inoculated plants, especially inG.mosseae-inoculated plants than in non-AM fungi-inoculated plants, while the leaf MDA content was lower. In addition, AM fungi, especiallyG.mosseae, increased phosphorus (P) and potassium (K) contents in the leaves, stems and roots, as well as acid phosphatase (ACP), catalase (CAT) and urease activities in rhizosphere soil of blueberry plants, in comparison with the non-AM fungi-inoculated treatment (CK).

These results indicate that the drought tolerance of blueberry cultivar Misty is enhanced with AM fungi inoculation through increase of antioxidant enzyme and osmotic adjustment, and the soil environment is improved, accompanied by P and K uptake increases by plants. In addition,G.mosseaeis the most beneficialGlomusamong the four AM fungi to improve the resistance of blueberry cultivar Misty plants against DS.

蓝莓(Vacciniumspp.)属于杜鹃花科越橘属，其浆果内特有的蓝莓花青素具有提高视力、抗衰老和防癌等功效。近年来，我国蓝莓产业发展势头强劲，由北向南已推广到江南、华南等地。蓝莓为浅根性灌木果树，主根不明显，无根毛，不耐旱[1-2]。而我国南方蓝莓主要栽培在丘陵山地，蓄水保水能力差，经常遭遇局部或间歇式干旱的威胁，已成为蓝莓高产优质的一个重要限制因子。

菌根是土壤中的菌根真菌与寄主根系形成的一种互惠共生体。在自然条件下，野生蓝莓菌根真菌资源丰富，其侵染率可达75%；但在人工栽培条件下，蓝莓自身形成菌根比较困难，其菌根真菌侵染率不到3%[3]。已有研究表明，若对蓝莓进行丛枝菌根(arbuscular mycorrhizal，AM)真菌接种处理，其AM真菌侵染率及植株生物量均得到显著提高[4-6]。有关AM真菌提高植物抗逆性尤其是抗旱性的报道较多。GHOLAMHOSEINI等[7]发现，在水分胁迫下，用AM真菌接种处理的向日葵植株干质量、果实产量和出油率均显著提高，并能促进植株对氮(N)、磷(P)的吸收；ABBASPOUR等[8]认为，AM真菌可以提高在干旱胁迫下开心果幼苗的生物量及对矿质元素的吸收，提高其渗透调节能力和抗氧化酶活性；OMIROU等[9]指出，AM真菌可以提高在缺水条件下西瓜对水分的利用率。本研究拟选取4种AM真菌，以南方普遍栽植的南高丛蓝莓品种薄雾为试验材料，研究AM真菌对蓝莓耐旱性的影响，以期筛选出最适合提高蓝莓抗旱性的AM真菌菌种，为生产实际提供理论与实践参考。

1　材料与方法

1.1试验材料

试验基质为V(红壤土)∶V(石英砂)∶V(珍珠岩)=9∶3∶1的混合物，经0.11 MPa、121 ℃高压湿热灭菌2 h后备用。以南高丛蓝莓品系薄雾蓝莓(Vacciniumcorymbosum)为试验材料。供试的AM真菌菌剂为摩西球囊霉(Glomusmosseae，BGC HUN01A)、地表球囊霉(G.versiforme，BGC HUN02B)、根内球囊霉(G.intraradices，BGC AH01)和幼套球囊霉(G.etunicatum，BGC HEB04)，由北京市农林科学院植物营养与资源研究所“中国丛枝菌根真菌种质资源库(Bank of Glomales in China，BGC)”提供。

1.2试验设计

2014年3月19日，将大小相对一致的7月龄扦插蓝莓苗移栽于装有2.2 kg试验基质的塑料盆(盆上口内径22 cm、盆底内径16 cm、盆高18 cm)中，移栽时分别接种摩西球囊霉、地表球囊霉、根内球囊霉和幼套球囊霉菌剂各约200个孢子，以不接种AM真菌作为对照(CK)，每盆栽植1株蓝莓苗，放置于通风和光照条件良好的大棚中培养。设正常供水和自然干旱处理，每个处理3次重复，共30盆。

7月31日，将每处理试验材料平分成2组，一组仍正常供水(水分充足)，另一组进行自然干旱处理20 d，8月20日对干旱组实施复水处理，浇一遍透水。于干旱13 d(8月13日)、20 d(8月20日)及复水后2 d(8月22日)分别采集各处理材料的叶片，进行相关生理指标的测定。收获植株时进行菌根侵染率、土壤酶活性及矿质元素含量测定。

1.3分析方法

1.4数据处理

采用Excel 2007进行数据处理，利用SPSS 20.0软件的单向方差分析(one-way ANOVA)进行不同处理间的差异显著性检验，采用邓肯法进行多重比较。

2　结果与分析

2.1AM真菌对蓝莓菌根侵染率、相对含水量、叶绿素和可溶性糖含量的影响

摩西球囊霉、地表球囊霉、根内球囊霉和幼套球囊霉接种处理均不同程度地提高了蓝莓菌根侵染率，干旱胁迫对菌根侵染率影响较小，4种AM真菌中以摩西球囊霉和根内球囊霉的侵染率相对较高(图1)。

CK:未接种AM真菌；Gm：接种摩西球囊霉；Gv：接种地表球囊霉；Gi：接种根内球囊霉；Ge：接种幼套球囊霉。WW:水分充足；DS：干旱胁迫。短栅上的不同小写母表示在相同水分条件下不同真菌接种处理间在P<0.05水平差异有统计学意义。CK, Gm, Gv, Gi and Ge represent non-inoculation, inoculation with Glomus mosseae, G. versiforme, G. intraradices and G. etunicatum, respectively. WW and DS represent well-watered and drought stress, respectively. Different lowercase letters above bars represent statistically significant differences at different inoculation treatments under the same water condition at the 0.05 probability level. 图1　AM真菌接种对蓝莓品种薄雾菌根侵染率的影响Fig.1　Effect of AM fungi inoculation on mycorrhizal colonization of blueberry cultivar Misty plants

干旱胁迫降低了蓝莓植株叶片相对含水量与叶绿素含量，增加了叶片可溶性糖含量；干旱13 d时摩西球囊霉接种株及干旱20 d时摩西球囊霉、地表球囊霉和根内球囊霉接种株相对含水量均高于对照；干旱13 d时各接种株叶绿素与可溶性糖含量、干旱20 d时摩西球囊霉与地表球囊霉接种株叶绿素含量以及各接种株可溶性糖含量均显著高于对照，尤以摩西球囊霉接种株明显；复水后2 d，摩西球囊霉与地表球囊霉接种株的相对含水量均显著高于对照，干旱胁迫处理的摩西球囊霉与幼套球囊霉接种株叶绿素含量及各接种株可溶性糖含量高于对照，水分充分处理的各接种株叶绿素含量与对照无显著差异(幼套球囊霉接种株除外)，而摩西球囊霉接种株的可溶性糖含量显著高于对照(图2)。

各处理符号表示的含义详见图1注。13 d、20 d、RW 2 d分别表示停止供水后第13天、第20天和复水后2 d。短栅上的不同小写母表示不同真菌接种处理间在P<0.05水平差异有统计学意义。Please see footnote of Fig.1 for details of each treatment. 13 d, 20 d and RW 2 d represent the cutoff of water supply for 13 d, 20 d and rewatering for 2 d, respectively. Different lowercase letters above bars represent statistically significant differences at different inoculation treatments at the 0.05 probability level. 图2　AM真菌接种对蓝莓品种薄雾叶片相对含水量、叶绿素和可溶性糖含量的影响Fig.2　Effect of AM fungi inoculation on relative water content, chlorophyll and soluble sugar contents in the leaves of blueberry cultivar Misty plants

2.2AM真菌对蓝莓SOD活性和MDA含量的影响

干旱胁迫提高了蓝莓植株叶片SOD活性与MDA含量。干旱13 d时摩西球囊霉接种株及干旱20 d时摩西球囊霉与地表球囊霉接种株叶片SOD活性均显著高于对照；各接种株叶片在干旱胁迫下MDA含量均显著低于对照(干旱13 d时幼套球囊霉接种株除外)；复水2 d后，水分充分与干旱胁迫处理的各接种株叶片MDA含量均显著低于对照，而SOD活性在接种株与对照间差异无统计学意义(P>0.05)(图3)。

各处理符号表示的含义详见图1注。13 d、20 d、RW 2 d分别表示停止供水后第13天、第20天和复水后2 d。短栅上的不同小写母表示不同真菌接种处理间在P<0.05水平差异有统计学意义。Please see footnote of Fig.1 for details of each treatment. 13 d, 20 d and RW 2 d represent the cutoff of water supply for 13 d, 20 d and rewatering for 2 d, respectively. Different lowercase letters above bars represent statistically significant differences at different inoculation treatments at the 0.05 probability level. 图3　AM真菌接种对蓝莓品种薄雾叶片超氧化物歧化酶(SOD)活性和丙二醛(MDA)含量的影响Fig.3　Effect of AM fungi inoculation on SOD activity and MDA content in the leaves of blueberry cultivar Misty plants

2.3AM真菌对蓝莓P、K含量和根围土壤酶活性的影响

在水分充分条件下，各接种株叶片P含量与对照无显著差异，而摩西球囊霉与根内球囊霉接种株根部P含量以及根内球囊霉与幼套球囊霉接种株茎部P含量高于对照，各接种株根部K含量、根内球囊霉接种株茎部K含量以及摩西球囊霉与地表球囊霉接种株叶片K含量均高于对照；在干旱胁迫下，各接种株根部P、K含量(幼套球囊霉接种株K含量除外)以及摩西球囊霉接种株茎、叶K含量均显著高于对照，其他处理的茎、叶K含量以及各接种株茎、叶P含量与对照间差异无统计学意义(P>0.05)(图4)。

无论水分充分还是干旱胁迫，摩西球囊霉与根内球囊霉接种株根围酸性磷酸酶、土壤过氧化氢酶与脲酶活性(根内球囊霉除外)均显著高于对照(图5)。

3　讨论

在本研究中4种AM真菌接种提高了蓝莓的菌根侵染率，其中摩西球囊霉和根内球囊霉的菌根侵染率相对较高，而在对照中未发现菌根侵染。这与已有研究结果[4-6]相一致。说明AM真菌不仅可以侵染蓝莓植株，而且其侵染程度与AM真菌种类及侵染时间长短密切相关。

相对含水量是反映植物水分盈亏程度的最直观指标。在干旱胁迫下，植物叶片相对含水量越高，表明持水力越强，胁迫对细胞膜的伤害程度越小，细胞能更好地维持正常生命活动，抗旱性也就越强[12]。在本研究中，干旱胁迫20 d时摩西球囊霉、地表球囊霉和根内球囊霉接种株尤其是摩西球囊霉接种株相对含水量显著高于对照，复水后可恢复到未胁迫水平。可见，AM真菌尤其是摩西球囊霉接种处理可提高蓝莓品种薄雾在干旱条件下的保水能力。另外，干旱胁迫虽降低了蓝莓叶片叶绿素含量，但AM真菌接种尤其是摩西球囊霉和地表球囊霉接种处理抑制了叶绿素含量的降低。叶绿素是参与光合作用最重要的色素，维持其相对较高含量有利于光合作用的顺利进行。4种AM真菌尤其是摩西球囊霉接种株的可溶性糖含量均显著高于对照。光合产物可溶性糖还是植物体内的一种重要的渗透调节物质，AM真菌接种处理有利于蓝莓在干旱胁迫下积累可溶性糖等渗透调节物质，以维持细胞的渗透势，进而增强细胞的保水能力[13-16]。

各处理符号表示的含义详见图1注。短栅上的不同小写母表示不同真菌接种处理间在P<0.05水平差异有统计学意义。Please see footnote of Fig.1 for details of each treatment. Different lowercase letters above bars represent statistically significant differences at different inoculation treatments at the 0.05 probability level. 图4　AM真菌接种对蓝莓品种薄雾根、茎及叶片中磷钾元素含量的影响Fig.4　Effect of AM fungi inoculation on P and K contents in the roots, stems and leaves of blueberry cultivar Misty plants

各处理符号表示的含义详见图1注。短栅上的不同小写母表示不同真菌接种处理间在P<0.05水平差异有统计学意义。

Please see footnote of Fig.1 for details of each treatment. Different lowercase letters above bars represent statistically significant differences at different inoculation treatments at the 0.05 probability level.

图5AM真菌接种对蓝莓品种薄雾根围土壤酸性磷酸酶、过氧化氢酶和脲酶活性的影响

Fig.5Effect of AM fungi inoculation on acid phosphatase (ACP), catalase (CAT) and urease activities in rhizosphere soils of blueberry cultivar Misty plants

在干旱胁迫下植物会积累活性氧，而SOD的主要功能是清除生物体内超氧阴离子，防御活性氧对细胞膜的伤害。MDA是膜脂过氧化的终产物，其含量高低可以反映膜脂过氧化的程度[17-19]。本研究结果显示，干旱胁迫处理的摩西球囊霉和地表球囊霉接种株的SOD活性显著高于对照，而各AM接种株MDA含量均显著低于对照，复水后各接种株MDA含量也均低于对照。说明在干旱胁迫下接种AM真菌能够增强蓝莓品种薄雾对活性氧的清除能力，减少活性氧的伤害，从而增强蓝莓植株对干旱胁迫的耐受性。

研究表明，在干旱胁迫下AM真菌能够促进植物对P、K等矿质元素的吸收，改善植物的营养环境[20-23]。本研究结果显示，在干旱胁迫下AM真菌接种株P、K含量相对高于对照。说明AM真菌尤其是摩西球囊霉接种提高了蓝莓对P、K的吸收。P是核酸和磷脂的组分，直接参与细胞内能量代谢和糖的运输，并能维持细胞的渗透势[24]；K是影响细胞渗透势的主要因素，它能维持细胞水势、气孔开张和光合作用等一系列生理过程，同时还是许多酶的辅酶[14]。在干旱胁迫条件下，AM真菌接种后蓝莓体内P、K水平的提高，有利于进一步提高植株体内的渗透调节能力，进而增强抗旱性。当然，P、K营养的改善，与土壤环境的改良是分不开的。

本研究显示，无论是水分充足还是干旱胁迫，摩西球囊霉和根内球囊霉接种株根围土壤的酸性磷酸酶、过氧化氢酶及脲酶活性均显著高于其他处理组和未接种株。土壤酸性磷酸酶能促进有机磷向无机磷转化，有利于植物体对P的吸收；脲酶能促进尿素水解，使之转化为可利用的有效态，提高氮素利用率[25]；过氧化氢酶的作用是水解土壤中的过氧化氢，减少对生物体的毒害作用。可见，摩西球囊霉和根内球囊霉接种株根围土壤酶活性的提高与上述AM接种株P、K含量相对较高的结果相吻合.说明AM真菌接种处理能够改善寄主根际土壤环境，提高根际土壤酶活性[4,8,20,26]，进而促进寄主对养分与水分的吸收。

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XU Qinglong, LIU Xiaomin, XU Xiaobing, LI Qingqing, ZHANG Hong, XIAO Jiaxin*

(KeyLaboratoryfortheConservationandUtilizationofImportantBiologicalResourcesinAnhuiProvince,CollegeofLifeSciences,AnhuiNormalUniversity,Wuhu241000,Anhui,China)

Glomus;Vacciniumcorymbosum; drought stress; pot experiment

国家自然科学基金(31372014)；安徽省高等学校自然科学研究项目(KJ2016SD24).

Corresponding author)：肖家欣(http://orcid.org/0000-0002-8427-6551)，E-mail:xjx0930@163.com

联系方式：许庆龙(http://orcid.org/0000-0003-3912-0299)，E-mail:hebau2010@sina.com

2015-09-06；接受日期(Accepted)：2015-11-27；网络出版日期(Published online)：2016-07-19

S 663； Q 945

A

URL:http://www.cnki.net/kcms/detail/33.1247.S.20160719.1836.004.html