Research progress on soil microorganisms in eucalypt forests

WEI Juxian; WANG Cong; HE Bin; YOU Yeming; HUANG Xueman

doi:10.11833/j.issn.2095-0756.20210701

Volume 39 Issue 5

Sep. 2022

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WEI Juxian, WANG Cong, HE Bin, YOU Yeming, HUANG Xueman. Research progress on soil microorganisms in eucalypt forests[J]. Journal of Zhejiang A&F University, 2022, 39(5): 1144-1154. doi: 10.11833/j.issn.2095-0756.20210701

Citation:

WEI Juxian, WANG Cong, HE Bin, YOU Yeming, HUANG Xueman. Research progress on soil microorganisms in eucalypt forests[J]. Journal of Zhejiang A&F University, 2022, 39(5): 1144-1154. doi: 10.11833/j.issn.2095-0756.20210701

Research progress on soil microorganisms in eucalypt forests

doi: 10.11833/j.issn.2095-0756.20210701

College of Forestry, Guangxi University, Nanning 530004, Guangxi, China

Received Date: 2021-10-19
Accepted Date: 2022-04-22
Rev Recd Date: 2022-04-22

Available Online: 2022-05-18

Publish Date: 2022-10-20

Abstract

As one of the three major fast-growing tree species in the world, eucalypt is featured with great variety, strong resistance to stress and wide adaptability. It is significant to research the diversity and functions of soil microorganisms which affect the growth of trees by participating in the processes of nutrient element cycling and energy flow and play an important role in improving soil fertility and productivity. However, with limitations in knowledge of soil microbial communities and functions due to the complexity of the eucalypt forest ecosystem and research techniques of soil microbiology, researches on the characteristics of soil microbial communities in eucalypt forests so far are still in the primary stage. This study is aimed to conduct a systematic review of the research progress on soil microbial characters in eucalypt forests with different management patterns, stand types and stand ages. Compared with natural forests of eucalypt, there was generally a lower soil microbial abundance in eucalypt plantations whereas there was an increase in the abundance, diversity, and activity of soil microorganisms in eucalypt mixed forests compared with eucalypt plantation pure forests and the abundance of soil microorganisms in eucalypt forests generally increased with the age of the forest, while the diversity of ectomycorrhizal and endomycorrhizal fungi decreased with the age of the forest. In addition, with an analysis of the mechanism of soil microorganisms involved in nutrient element cycling and remediation of heavy metal pollution in eucalypt forests and prospects of future research and analyzing methods of soil microbiome and application of microorganisms, this study will provide scientific guidance for the maintenance of forest soil health and the promotion of green and sustainable forestry development. [Ch, 1 tab. 94 ref.]
- eucalypt,
- soil microorganism,
- soil nutrient cycling,
- management pattern,
- stand type,
- stand age

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沈阳化工大学材料科学与工程学院沈阳 110142

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桉树是桃金娘科Myrtaceae桉属Eucalyptus、杯果木属Angophora和伞房属Corymbia植物的统称，常绿乔木，种类多、适应性广、抗逆性强，与杨树Populus 和松树 Pinus一同被称为世界三大速生树种^[1]。桉树主要原生于澳大利亚(约1.01亿 hm²)，极少数产于印度尼西亚、巴布亚新几内亚和菲律宾^[2]。桉树调节气候、涵养水源和保持水土作用突出，碳汇功能相当强大，研究^[3-4]发现：桉树二氧化碳吸收量达24.3 t·hm⁻²·a⁻¹，是杉木Cunninghamia lanceolata的2.2倍，马尾松 Pinus massoniana的3.0倍，对于实现碳中和具有重要意义。作为优质可再生生物质能源，桉树材是纸浆、人造板、家具的主要原料，还被广泛用于生产各种林副产品，如桉树叶油、桉树多酚、桉树木炭等^[5-6]。目前，全球桉树人工林面积已超过2 257万 hm²(表1)，巴西、中国和印度是桉树人工林面积最大的3个国家，提供约2.5亿 m³·a⁻¹桉树木材，占世界人工林木材年产量的37%^[7-14]。

国家	种植面积/hm²	国家	种植面积/hm²	国家	种植面积/hm²	国家	种植面积/hm²
巴西	4 862 000	摩洛哥	215 000	玻利维亚	41 000	土耳其	20 000
中国	4 500 000	菲律宾	189 000	阿尔及利亚	40 000	卢旺达	17 000
印度	3 942 600	马达加斯加	163 000	尼日利亚	40 000	多哥	17 000
澳大利亚	926 000	印度尼西亚	128 000	孟加拉国	39 000	哥斯达黎加	17 000
乌拉圭	676 024	安哥拉	113 000	肯尼亚	39 000	新西兰	15 000
智利	652 100	埃塞俄比亚	100 000	斯威士兰	33 000	危地马拉	13 000
葡萄牙	647 000	墨西哥	100 000	美国	32 500	津巴布韦	13 000
西班牙	640 000	委内瑞拉	100 000	伊朗	31 000	所罗门群岛	12 000
越南	586 000	厄瓜多尔	81 000	哥伦比亚	27 000	尼加拉瓜	12 000
南非	568 000	意大利	72 000	乌干达	25 000	厄立特里亚	12 000
苏丹	540 400	刚果	68 000	莫桑比克	25 000	赞比亚	12 000
泰国	500 000	塞内加尔	65 000	布隆迪	25 000	尼泊尔	11 000
秘鲁	480 000	伊拉克	60 000	马拉维	25 000	法国	10 000
阿根廷	330 000	突尼斯	56 000	帕劳	21 000
巴基斯坦	245 000	古巴	53 000	巴拉圭	21 000
说明：中国主要分布在广西壮族自治区、广东省、福建省、云南省和其他地区，占比分别为45%、30%、6%、5%和14%；巴西主要分布在　　　　米纳斯吉拉斯州、圣保罗州、巴伊亚州、南马托格罗索州和其他地区，占比分别为29%、23%、14%、13%和21%；印度主要分布　　　　在卡纳塔克邦、西孟加拉邦、马哈拉施特拉邦、北方邦和其他地区，占比分别为40%、13%、10%、6%和31%

Table 1. Distribution of global eucalypt forests ^[7-14]

随着桉树产业的快速发展，不合理经营管理和林木采伐(如长期施肥和缩短桉树轮伐期)问题凸显，林地土壤肥力下降，病虫害频发，土壤生物多样性下降，环境污染加剧，严重危害了土壤健康和林业可持续发展^[15]。土壤微生物是诊断土壤健康的理想指标^[16]，通过自身代谢参与养分循环和污染物降解，驱动元素生物地球化学循环，在增强植物根际免疫力、提高土壤肥力等方面发挥关键作用。本研究系统综述了不同经营方式和林分因子条件下桉树林土壤微生物群落的变化规律，分析土壤微生物和桉树林土壤健康的互作机制，并对今后桉树林土壤微生物研究和分析方法及其应用进行了展望，以期为维持森林土壤健康和促进林业绿色可持续发展提供指导。

1. 不同经营方式和林分条件下桉树林土壤微生物群落特征

1.1. 不同经营方式下桉树林土壤微生物群落特征

依据经营方式的不同，桉树林分可分为天然林和人工林。受不同林地经营措施如施用肥料和除草剂、连栽等影响，桉树林土壤环境条件存在差异，这会对土壤微生物活性、丰度和群落组成结构产生影响^[17]。桉树天然林土壤细菌的优势群落主要为放线菌门Actinobacteria (15%~42%)和变形菌门Proteobacteria (12%~65%)^[18-22]。施用化肥改变了桉树人工林土壤养分含量和pH，从而影响土壤微生物群落。桉树人工林中土壤细菌的优势群落^[23-26]为绿弯菌门Chloroflexi (10%~44%)、变形菌门(18%~44%)、放线菌门(14%~41%)和酸杆菌门Acidobacteria (9%~18%)，相较天然林，土壤中绿弯菌门和酸杆菌门相对丰度增加，可能是施用化肥导致的^[27]。李超等^[28]采用高通量测序技术对16S rRNA的V3~V4区域进行研究，发现施用复合肥提高了桉树根系的生理活性，改变了土壤细菌群落组成；与不施肥桉树人工林相比，施肥组人工林土壤绿弯菌门和酸杆菌门群落相对丰度更高。有研究^[29]发现：绿弯菌门为兼性厌氧细菌，通过参与氮代谢将土壤内多糖分解成氢和有机酸，加快土壤内有机物的降解^[30]。随着复合肥施用量的增加，施肥组桉树林土壤速效氮和植物根系分泌物增加，绿弯菌门群落相对丰度也随之增加^[31]。酸杆菌门细菌能够在酸性条件下降解木质素与纤维素，参与土壤有机质循环^[32]。随着复合肥施用量的增加，土壤速效氮、有效磷、速效钾含量增加，土壤pH下降，为酸杆菌门细菌提供了良好的生存环境，因而相对丰度增加。子囊菌门Ascomycota和担子菌门Basidiomycota是桉树林土壤真菌的优势群落^[19，33-36]，两者比例在天然林中分别为60%~86%和9%~25%，在人工林中则分别为19%~50%和47%~75%^[37-40]。担子菌门真菌可有效降解木质素。SCHMIDT-DANNERT^[41]研究发现施用复合肥使得桉树人工林土壤中木质素含量升高，为担子菌门真菌提供足够底物，促进其生长和繁殖^[42]，这可能是桉树人工林担子菌门真菌相对丰度增加的原因。

与桉树天然林相比，施用除草剂后桉树人工林林下植被多样性减少，土壤养分含量下降，可能是导致土壤微生物数量减少的原因。SILVA等研究^[43]发现：相比没有施用除草剂的桉树人工林，施用不同剂量和频率的除草剂后，林下植被多样性和根系生理活性降低，土壤微生物生物量碳、氮含量下降。除草剂还会促成植物体内乙烯合成、脱落酸积累和气孔缩小，使二氧化碳吸收量减少，抑制光合作用并影响植物根系的生长发育和活性，引起林下植被多样性降低，间接使土壤养分含量和微生物活性降低，从而影响土壤微生物生物量碳、氮含量^[44-45]。何伟等^[46]采用变性梯度凝胶电泳法研究了桉树人工林表层土壤细菌群落多样性特征，发现施用除草剂的桉树人工林林下植被多样性发生改变，土壤细菌多样性先减少后增加。原因可能是施用除草剂使林下植被多样性下降，土壤凋落物输入的种类与数量减少，影响微生物基质数量和种类，或对某些土壤微生物产生抑制，导致细菌多样性减少。随着时间推移，除草剂被土壤黏粒吸附、微生物降解，随着除草剂毒性减弱，细菌多样性随之增加^[47]。

现存桉树人工林主要为1代植苗林，2代萌芽林，3代萌芽林。魏圣钊等^[48]采用平板菌落计数法发现：随桉树人工林连栽代次增加，土壤细菌、放线菌和真菌数量均呈减少的趋势。ZHU等^[49]研究发现：尾叶桉 Eucalyptus urophylla连续种植后土壤微生物生物量降低，这可能是因为土壤全磷、全氮和含水率随桉树连栽代次增加而逐渐下降，微生物生长所需基质减少，生长受到限制^[50]。可见连栽会造成桉树人工林土壤养分含量下降，影响土壤微生物数量和活性。

1.2. 不同树种组成下桉树林土壤微生物群落特征

林分可根据树种组成分为纯林和混交林，不同林分的凋落物数量和质量、土壤养分含量、根际土壤化感物质含量和根系活动等均有差异，这会对土壤微生物群落特征产生影响^[51-52]。

桉树人工纯林凋落物难降解化合物(如木质素等)的比例较高，这些结构复杂的化合物降解时会产生大量酚酸类小分子化合物^[53]，对土壤微生物具有抑制作用^[54]。与桉树人工纯林相比，桉树混交林凋落物数量和种类增加，能为更多种类土壤微生物生长提供底物，从而使土壤微生物数量和活性提高。KOUTIKA等^[55]研究发现：桉树-刺槐Robinia pseudoacacia混交林中存在不同种类的凋落物，相比纯林，凋落物碳氮比较低，氮代谢强度提高，酚酸类物质减少，对微生物生长的抑制作用较低，从而使土壤微生物的生物量和多样性增加。PEREIRA等^[56]发现：桉树-马占相思 Acacia mangium混交林中凋落物碳氮比较桉树人工纯林低，土壤细菌多样性增加。原因可能是不同种类和质地的凋落物促进了有机质积累，为更多种类的土壤微生物提供了底物。

与桉树人工纯林相比，不同树种组成的桉树混交林能提高土壤养分，降低有害化感物质，从而提高土壤微生物数量和活性。刘小香等^[57]发现：桉树纯林根系分泌物和凋落物分解产生的化感物质会抑制土壤微生物的生长繁殖。SANTOS等^[58]发现：与桉树人工纯林相比，马占相思-桉树混交林根系分泌物对微生物的抑制作用较低，凋落物分解速率较高，碳代谢活性增强，土壤有机碳含量增加，土壤养分状况得到改善，土壤微生物总数比桉树人工纯林高。马占相思是一种浅根树种，根系较细，吸收根多，与桉树混交后，可提高土壤孔隙度，改善土壤空间结构和养分状况，使土壤微生物总数增加^[59]。李万年等^[60]发现：望天树Parashorea chinensis与桉树混交改善了林地土壤根系分布的空间结构。望天树幼树期根系较浅，与桉树混交后，可有效疏松土壤，改善土壤颗粒结构；混交林土壤全氮、全钾、速效磷含量增加，促进养分的合理利用。

与桉树人工纯林相比，桉树混交林不同树种组成影响根系活动，使真菌群落的生物量降低，土壤细菌和放线菌群落的生物量提高。陈永康等^[61]和黄雪蔓等^[62]采用磷脂脂肪酸法发现：桉树与降香黄檀 Dalbergia odorifera混交可提高根系活性，增加根系分泌物，使土壤细菌和放线菌群落的生物量提高，真菌群落的生物量降低。这可能是由于细菌倾向于分解简单化合物(如根系分泌物中易降解可溶性有机碳等)^[63]，而真菌是有机大分子物质(如木质素等)的主要分解者^[64]。桉树-降香黄檀混交林根系分泌物中易分解活性化合物较桉树人工纯林要多，难降解化合物(如木质素等)则较少，土壤碳氮比降低，导致真菌群落生物量降低，细菌和放线菌群落生物量提高。

1.3. 不同林分年龄下桉树林土壤微生物群落特征

桉树林分不同年龄阶段的土壤理化性质^[65]、细根生长状况^[66]、林下植被多样性^[67]、根际土壤化感物质含量的差异等均会导致土壤微生物群落特征发生变化。

不同年龄阶段桉树吸收土壤养分的种类和数量不同，从而影响土壤养分状况，并改变土壤微生物群落特征。张丹桔等^[68]发现：四川丹棱县巨桉E. grandis人工林轮伐期前(4 a)土壤细菌和真菌逐步减少，此后随林龄增加而逐渐增加。这可能是因为轮伐期前(4 a)为桉树人工林速生生长阶段，林木生长需要消耗大量养分，原有有机质的分解大于外界输入，土壤有机碳含量低，微生物生长基质减少，因而活性与数量下降；随着林龄增长，林木生长趋缓，对养分吸收减少；同时林下生物多样性增加，凋落物不断积累，分解产生有机碳输入土壤，使得土壤有机碳质量分数升高，为微生物生长提供足够基质，因而活性与数量均有增加^[69]。XU等^[70]发现：雷州半岛北部尾巨桉 E. urophylla × E. grandis人工林土壤细菌和真菌数量具有林龄效应，土壤细菌和真菌数量表现为随林龄增长而增加的趋势。

桉树人工林细根生长状况一般表现为幼龄林阶段优于成熟林，这可能是导致幼龄林阶段外生菌根真菌群落多样性指数大于成熟林的原因。江瑶等^[71]比较了不同林龄桉树人工林的土壤外生菌根真菌群落多样性，发现桉树栽植2 a时土壤外生菌根真菌物种丰度高于6 a。这可能是2年生桉树处于速生阶段，细根较多，根系活性较高，对营养物质和水分的需求量大^[72]，桉树通过分配大量的养分物质利用外生菌根辅助养分吸收，一定程度上提升了外生菌根真菌多样性。随着林龄增长，桉树生长速度减缓，养分吸收速率降低^[73]，对外生菌根真菌的依赖性降低，细根活性下降，外生菌根真菌群落多样性减少。

土壤磷含量依赖于土壤本身的母质特征^[74]。幼龄林阶段桉树人工林土壤养分循环缓慢，土壤磷元素来源少，磷含量一般小于成熟林，这可能导致了幼龄林阶段内生菌根真菌群落多样性指数大于成熟林。李佳雨等^[75]采用湿筛倾析法研究比较不同林龄桉树人工林的土壤内生菌根真菌群落多样性的变化后发现：栽植2 a的桉树土壤内生菌根真菌香农指数和辛普森指数均最高，5 a时均为最低，但组间多样性指数差异不显著。随着人工营林措施(如施磷肥)的进行，桉树人工林土壤磷含量增加，植物对内生菌根真菌的依赖性降低，分配给内生菌根真菌的养分减少，造成内生菌根真菌多样性减少^[76]。随着桉树林龄增加，内生菌根真菌群落多样性指数逐渐减少。

2. 微生物对桉树林土壤养分循环和重金属污染修复的影响

2.1. 土壤微生物和元素循环

土壤微生物通过自身代谢参与碳、氮、磷、铁和硫等元素循环^[77]来改善土壤养分状况，提升土壤健康。目前对桉树碳、氮和磷元素循环与土壤微生物群落间关系的研究较多^[78]。

李永双等^[79]通过盆栽实验对云南省建水地区植被根际土壤中产碳酸酐酶的微生物进行分离培养，获得1株具有较强溶蚀效果的沙雷氏菌属 Serratia细菌。SALEEM等^[80]研究认为沙雷氏菌菌液分泌物对碳酸钙类岩石的溶蚀具有促进作用，根际微生物降解有机残体产生的二氧化碳可增大难溶性碳酸盐的溶解性^[81]。文晓萍等^[82]将回接相思树Acacia根瘤作用于非豆科Leguminosea植物巨尾桉苗木的生长，发现接种根瘤菌处理土壤含氮量显著提高。可能是根瘤菌可以将植物不能吸收的氮、土壤固定态磷、钾以及微量元素变成可吸收利用的状态^[83]，接种根瘤菌后产生的胞外多糖为土壤微生物提供了更多的碳源^[84]，促进了土壤可培养微生物(细菌、真菌和放线菌)的生长^[85]，有利于土壤微生物固定无机态氮，因此有效地提高巨尾桉土壤氮素的积累。张辉等^[86]研究发现：接种促生菌(固氮菌、解钾菌、解磷菌)后，巨尾桉土壤有效磷质量分数提高。可能是因为这些联合固氮菌可以分泌有机酸，而有机酸可使土壤中不溶性磷素转变为可溶性磷素，使磷有效地溶解和被植物吸收^[87]。

2.2. 土壤微生物和土壤重金属污染修复

当前，中国林地土壤重金属污染和土壤农药污染等问题^[88]严重，土壤健康受到威胁。目前研究主要是关于土壤微生物群落对土壤重金属污染的影响^[89]。

黄佳玉等^[90]通过盆栽实验发现：接种摩西球囊霉菌Glomus mosseae且土壤中外加铜和锌质量分数分别为25和200 mg·kg⁻¹处理，桉树苗中铜质量分数由26 mg·kg⁻¹降低为13 mg·kg⁻¹，较未接种处理差异显著(P＜0.05)。这可能是由于菌根真菌的结构阻碍金属元素向地上部分运输，降低重金属元素对植物的毒性，从而保护桉树的生长^[91]。廖妤婕等^[92]采用差速离心法和化学试剂逐步提取法研究了桉树对铅的耐受机制，从铅在桉树不同部位中的亚细胞分布和化学形态来看，高质量分数(200和400 mg·kg⁻¹) 铅处理下，接种丛枝菌根真菌(AMF)的桉树根部细胞壁组分和叶片可溶组分的分配比例增大，细胞壁滞留作用和液泡区隔化作用增强，可能是AMF作用下桉树耐铅的主要机制。细胞壁和以液泡为主的可溶组分是桉树储存铅的2个主要场所，桉树根部细胞壁对铅的滞留作用最强，叶片中可溶组分对铅的区隔化作用最强。随着铅胁迫的增加，接种AMF的桉树根部对铅的固持作用及细胞壁组分的滞留作用增强，同时，滞留在桉树根部的铅从活性较强的乙醇提取态向活性较弱的醋酸提取态转化，减弱了铅的活性，也减少了铅向地上部分茎和叶的迁移；此外，叶片可溶组分对铅的区隔化增强，也减轻了迁移到地上部分的铅对桉树的毒害作用。

3. 展望

微生物分子生态技术的快速发展和应用，为拓宽微生物群落结构和多样性研究提供了新的途径和方法，推动了碳、氮、磷等元素生物地球化学循环机制和土壤污染的深入探索。加强对土壤微生物组成和功能的研究，强化其在桉树林生态系统中的作用，对减少化肥和农药的施用、维持森林土壤健康，进而促进桉树林绿色可持续发展具有重要意义。

当前对桉树林土壤微生物群落特征的研究大多处于初级阶段，多数通过采集少量次数的土壤样本，基于平板菌落计数法、磷脂脂肪酸法、16S rRNA高通量测序等方法进行。与高通量测序法相比，平板菌落计数法和磷脂脂肪酸法等方法灵敏度和准确度低、检测微生物种类少、数据量少^[93]，基于非功能基因的扩增子测序等方法无法准确评估微生物群落功能特征。因此，目前的研究尚未充分了解桉树林土壤微生物活性、数量、群落组成结构和功能特征，难以准确评价桉树林土壤健康状况。为实现桉树林土壤健康与可持续发展，建议从3个方面进一步开展相关研究：①运用多种组学方法(如宏基因组学、宏转录组学和宏代谢组学等)研究桉树林土壤微生物群落特征，探究土壤微生物的功能并使其有益功能最大化，从而提高桉树林生态系统中林木的抗病性和资源的有效利用。②集成桉树林土壤微生物和土壤健康大数据自动化分析系统，为土壤微生物作为土壤健康评估的重要指标提供有力支撑。③微生物移植工程已经成功应用于改变植物微生物菌群组成^[94]，发挥了改善土壤健康和促进林业可持续发展的巨大潜力。今后可深入探究土壤微生物参与特定生物过程的功能，并将其广泛应用于治疗性微生物组工程。

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