毛竹(Phyllostachys edulis)是禾本科竹亚科刚竹属的克隆性植物,其强大的竹鞭系统能快速生长并入侵至周围林分进行无性繁殖,由于其强大的空间和光照竞争能力从而排斥其他树种,入侵后不久即可形成纯林[1]。目前本课题组在研究毛竹扩张后土壤生态变化过程中发现有趣的现象:毛竹向周围林地入侵后土壤pH会明显上升,在毛竹入侵初期的混交带就体现出该变化,在完全替代原有林地后表现得更加明显。但由于研究环境不同,各独立研究之间的数据也有差异:毛竹入侵杉木林时土壤pH提升了22.9%[2],但入侵阔叶林时pH反而有下降的趋势[3];毛竹林不同土层深度的pH差异也很大,研究[4]表明竹林10~20 cm的土壤pH为4.67,而40~55 cm的土壤pH却达5.20。为了定量描述毛竹入侵对土壤pH的影响,有必要采用大数据分析以揭示毛竹入侵对土壤pH的影响规律。
土壤pH变化具有级联效应,土壤pH浮动可能会带动土壤养分系列变化,包括但不局限于微生物主导的土壤氮素周转[5]、土壤磷的溶解度[6]、土壤速效钾的含量[7]及土壤有机质[8]等。同时,土壤微生物指标也会因pH的变化而改变,毛竹入侵杉木林后土壤微生物活性和群落结构会发生显著变化[9];入侵亚热带阔叶林会提升微生物的C:N和C:P[10];入侵阔叶林可能还会增加自养细菌的丰富度和活跃度,增加阔叶林CO2的固定能力[11]。因此,有必要同时对毛竹入侵伴随的土壤养分、生物学性质的影响进行分析,并与pH变化关联。
全球土壤酸化是普遍现象,无论是自然界氮硫沉降[12]、酸雨侵蚀[13],还是人为施肥[14]、农作物连作[15],均会使土壤逐渐酸化,由此对森林、农田等生态系统造成诸多消极影响,例如,不利于森林物种生存[16]、降低土壤呼吸、降低根系生物量和微生物生物量碳氮[12]、增加作物对重金属离子的累积[17]等。因此,在全球土壤酸化大背景下,能够使土壤pH提升的自然过程和人为措施均值得关注。基于大数据分析揭示毛竹入侵提升土壤pH的可能机理,对深入了解毛竹入侵的生态过程有重要意义,同时为缓解土壤酸化提供生物防治的全新思路。
Meta分析是一种将不同研究问题的相同研究结果综合分析得出结论的方法,早在1998年就被应用于生态学研究中[18],区别于传统综述文章,这种方法涵盖范围更广、数据更具体,能够更好地评估研究生态学问题。现阶段Meta分析方法仍具有一定的积极意义和生产指导作用[19-20]。本Meta分析除了收集已发表的文献数据外,同时在浙江省采集毛竹入侵带土壤样品,分析土壤pH和养分指标,以增加样本数量,期望解决三个科学问题:(1)毛竹入侵周边林分致土壤pH提高是否具有普遍性?土壤pH变化与毛竹入侵阶段、被入侵林分、毛竹纯林时间、土层深度是否有联系?(2)毛竹入侵如何影响土壤养分和微生物性质?(3)毛竹入侵导致土壤pH变化的可能机制有哪些?
1 材料与方法 1.1 数据来源与筛选通过中国知网(http://www.cnki.net)和Web of Science搜索了有关毛竹入侵的期刊文章,搜索词为(毛竹林、土壤、扩张、入侵)。本Meta分析包含的研究必须符合以下标准:(1)所研究对象鲜有人为干扰;(2)林分之间地理位置相近,土壤母质一致;(3)可以清楚地获得样本量(n)、平均值(mean)和标准误差(sd)。
本研究共收录了42篇符合要求的文献,101组数据,详情见附表 1。文献中提取的土壤理化数据包含:pH、有效氮、有效磷、速效钾、总有机碳、全氮、硝态氮、铵态氮。并以入侵阶段、毛竹纯林时间、被入侵林分、土层深度将数据分类讨论分析。被入侵林分分为:(1)针叶林(coniferous forest,CF),包含:日本柳杉(Cryptomeria japonica)、杉木(Cunninghamia lanceolata)、红桧(Chamaecyparis formosensis Matsum)、马尾松(Pinus massoniana Lamb)、雪松(Cedrus deodara)、扁柏(Chamaecyparis);(2)阔叶林(broad-leaf forest,BL),包含:苦槠(Castanopsis sclerophylla)、麻栎(Quercus acutissima Carr)、枫香(Liquidambar formosana Hance)、板栗(Castanea mollissima Blume)、青冈(Quercus glauca)、木荷(Schima superba)。此外,不同文献对于土壤深度的划分情况不同,本研究统一将土层分类划分为0~10 cm、0~20 cm、20~40 cm、40 cm以上四种深度以便于统计。如果有按季节采样分析的数据,选择一年内总体的平均值作为代表数据。各养分指标测定方法需是公认的、准确的,参考《土壤农业化学分析方法》[21]。
本研究选择微生物生物量碳、微生物生物量氮、微生物生物量磷、磷脂脂肪酸(PLFA)以及真菌、细菌的香农指数和chao1指数来评估毛竹林入侵后土壤微生物的变化。此外,由于土壤中细菌数量远大于其他微生物,本文还着重分析了五种土壤细菌优势门的相对丰度变化,分别为:变形菌门(Proteobacteria)、酸杆菌门(Acidobacteria)、放线菌门(Actinobacteria)、拟杆菌门(Bacteroidetes)和浮霉菌门(Planctomycetes)。
1.2 Meta分析方法本研究主要摘录的文献数据有平均值(means,m)、标准差(standard deviations,sd)和样本数量(sample sizes,n)。当文献图中未直接标明数据时,通过软件Plot Digitizer(v.2.6.8)来读取图中各点的数据。仅提供标准误差(standard errors,se)的,可以通过下式进行转换:
$ {\text{sd}} = \sqrt {{\text{se}}} $ | (1) |
若无sd与se时,则默认选择均值的百分之十作为sd[22]。根据Hedges等[23]的统计学描述,将效应值(ln RR)作为评价某项研究对实验组和对照组的影响效应:
$ \ln {\text{RR = }}\ln \left( {\frac{{{m_1}}}{{{m_2}}}} \right) = \ln {m_1} - \ln {m_2} $ | (2) |
效应值对应的方差(v)为:
$ v = \frac{{{\text{sd}}_1^2}}{{{n_1}m_1^2}} + \frac{{{\text{sd}}_2^2}}{{{n_2}m_2^2}} $ | (3) |
式中,m1和m2、sd1和sd2、n1和n2分别是实验组和对照组的均值、标准差和样本数量。
特别说明,毛竹入侵是一种变化缓慢、时间跨度大的自然扩展过程,入侵带一般存在毛竹纯林、竹阔(针)混交林、原生纯林三种类型的林分,因此不同分类情况下所要求的对照组与实验组不尽相同。本文选择用毛竹纯林和混交林分别与未被入侵的原生林分作对比,以此来表示毛竹完全入侵、毛竹正在入侵对土壤的影响,具体对照组与实验组如表 1所示。
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表 1 数据分析对象的情况描述 Table 1 Description of the data analysis object |
每个观测值的权重(w)为方差(v)的倒数:
$ w = \frac{1}{v} $ | (4) |
同一篇文献中含有多个观测样点,其总数为N。本研究认为这些观测值之间并不相互独立,因此将同一文献中每个观测点重新赋予权重(w′):
$ w' = \frac{w}{N} $ | (5) |
每篇文章中单个样点对应的效应值ln RR′为:
$ \ln {\text{RR'}} = w' \times {\text{RR'}} $ | (6) |
总体平均效应值(RR++)为:
$ {\text{R}}{{\text{R}}_{ + + }} = \frac{{\sum\nolimits_i {\frac{{{w_i}}}{{{N_i}}}\ln {\text{R}}{{\text{R}}_i}} }}{{\sum\nolimits_i {\frac{{{w_i}}}{{{N_i}}}} }} $ | (7) |
式中,wi、Ni、ln RRi表示第i个观测值的权重、观测样点数和效应值。
使用软件METAWIN(v.2.1)利用固定效应模型(fixed effect model,FEM)计算各个平均效应值RR++的95%置信区间,若其95%置信区间的下限大于0,则认为毛竹入侵显著增加了该研究指标;反之则认为毛竹入侵显著降低了该研究指标。为了更好地理解与解释效应关系,本研究将平均效应值(RR++)与其相应的置信区间上下限进行对数转换,用响应百分比(PC,percentage of change)来表示:
$ \text{PC} = ({{\text{e}}^{{\text{R}}{{\text{R}}_{{\text{ + + }}}}}} - 1) \times 100\% $ | (8) |
2021年10月,在浙江省北、中、南部不同地点寻找具有明显“毛竹-混交林-原生林”分布的鲜有人为干扰的毛竹入侵带,沿着毛竹入侵方向在每种林分中设置3个样地(10 m×10 m),共设12个样地。在每个样地内根据五点取样法采集0~20 cm表层土样,混合后装入塑封袋运至实验室。风干后过2 mm筛除去样品中的石块、动植物残体,存入新塑封袋储存,用于后续实验。某些地区的混交林土壤难以采集,则仅选择了毛竹与原生林土壤。本研究共计采集来自6个不同地点、12条入侵带的土壤,总计28个样点、90个样品。分析得到土壤pH、碱解氮(AN)、有效磷(AP)、速效钾(AK)、铵态氮(
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表 2 样地土壤化学性质 Table 2 Soil chemistry properties of sample plots |
有学者将自测的数据与文献数据结合进行Meta分析[24],目的是为了提升分析结果的严谨性和准确性,是对Meta分析的进一步加强和改进。本研究也采用这种结合的方法,将表 2中所得的自测数据按实验组与对照组分类合计为18组,与文献数据结合进行分析,总计得到土壤pH数据119组,铵态氮31组,硝态氮31组,碱解氮39组,土壤有效磷48组,土壤速效钾49组。Meta分析方法如1.2所述。
2 结果 2.1 毛竹入侵对土壤pH的影响基于文献和实测数据(表 2)的119组pH数据的Meta分析结果表明,毛竹入侵致土壤pH升高的样点数占84.9%,入侵阶段、入侵林分、毛竹纯林时间、土层深度等变量均影响土壤pH提高的幅度(图 1)。
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注:四种分类情况分别为:入侵阶段、入侵林分、毛竹纯林时间、土层深度。*表示该亚组总和的效应。未加*且未标注林分对比的组别,其林分对比默认为原生纯林vs.毛竹纯林。括号内数字表示观测样本数量。图中圆点代表平均响应百分比,实心点(●)表示该值显著变化,空心点(○)表示不显著。下同。 Note: The four parts are as follows: stages of invasion, different stands, times of bamboo, and soil depths. Adding * means the effect of the sum of this subgroup. For the group without * and without marked stand contrast, the default stand contrast is original forest vs. bamboo forest. The number in parentheses indicates the number of observed samples. ● means a significant change in the value. ○ means no significant change in the value. The same as below. 图 1 不同维度分析毛竹入侵对土壤pH的影响 Fig. 1 Effects of bamboo invasion on soil pH in a different dimension |
如图 1,基于入侵阶段的分析表明,与原生纯林相比,毛竹纯林土壤pH提升幅度(10.5%)高于混交林(5.5%),两者平均显著提高7.8%(P < 0.05)。基于入侵林分的分析表明,与原生纯林相比,毛竹入侵针叶林后土壤pH的提升幅度(+11.0%)明显高于入侵阔叶林类林分(+1.2%)。基于毛竹纯林时间的分析表明,毛竹纯林时间0~20年区间,土壤pH较原生纯林提升6.0%,低于20~40年提升的13.3%。基于土层深度的分析表明,与原生纯林相比,毛竹入侵对0~10 cm、0~20 cm土层土壤pH提升幅度较大(+8.2%、+11.3%),大于20~40 cm的土层(+4.3%),超过40 cm土层土壤由于文献数据量较小,且变异性较大,故未能得到明确的定量变化结果。
2.2 毛竹入侵对土壤养分的影响基于文献和实测数据(表 2)的Meta分析结果表明,毛竹入侵原生林分后土壤养分发生了明显变化。土壤全氮、铵态氮、硝态氮和有效磷在入侵阶段、入侵林分分类下的分析结果(图 2),表现为共同上升或下降,具有一致规律。土壤全氮和硝态氮表现为显著下降(P < 0.05),平均分别下降15.9%和21.7%,毛竹完全入侵后下降幅度最大,其中毛竹林完全入侵针叶林时土壤全氮下降高达40%,而毛竹完全入侵阔叶林时土壤硝态氮下降达30.4%;毛竹入侵导致土壤铵态氮和有效磷显著(P < 0.05)增加,平均分别提升14.7%和54.9%,当入侵林分为阔叶林时,铵态氮和有效磷的提升幅度分别达15.7%和137.5%。土壤碱解氮和全碳在入侵阶段、入侵林分分类下的分析结果有(图 2)所差异,土壤碱解氮含量除入侵林分为针叶林时显著(P < 0.05)下降(–10.6%)外,其他分类情况下土壤碱解氮含量均表现为显著(P < 0.05)上升。毛竹入侵致土壤总有机碳含量平均下降2%,但入侵林分为阔叶林时显著增加4.2%。毛竹入侵对土壤速效钾的影响较为复杂,将入侵阶段、入侵林分分类情况下一起分析时无显著变化,然而,基于入侵阶段的分析表明,混交林土壤速效钾较原生林显著下降23.6%(P < 0.05)(图 2)。
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注:TN:土壤全氮;TOC:总有机碳。 Note: TN: soil total nitrogen; TOC: soil total organic carbon. 图 2 毛竹入侵的不同阶段和不同林分对土壤养分的影响 Fig. 2 Effects of different stages and different stands of bamboo invasion on soil nutrients |
数据分析表明,毛竹入侵原生林分后土壤微生物群落发生明显变化(图 3)。其中混交林相比原生林土壤真菌香农指数整体增加13.55%(P < 0.05),但毛竹完全入侵形成纯林后有所回落(+5.23%);毛竹入侵后土壤细菌香农指数整体显著增加(P < 0.05),毛竹纯林相比原生林显著增加1.95%(P < 0.05),但混交林与原生林相比并未显著增加。比较毛竹林与原生林土壤优势细菌门的结果发现,毛竹纯林土壤酸杆菌门(-15.5%)、浮霉菌门(-26.7%)、拟杆菌门(-22.58%)的相对丰度显著(P < 0.05)降低,而放线菌门显著增加25.86%(P < 0.05),变形菌门未见显著变化。此外,基于表征不同类别土壤微生物生物量的磷脂肪酸(PLFA)数据分析(图 3c)表明,毛竹纯林土壤除真菌(+14.9%)与放线菌(+7.82%)显著(P < 0.05)高于原生林外,微生物总生物量、细菌、革兰氏阳性菌、革兰氏阴性菌以及丛枝菌根真菌(AMF)等则无显著变化。基于氯仿熏蒸法的土壤微生物生物量碳的分析结果表明,与原生林相比,毛竹入侵显著降低(P < 0.05)土壤微生物生物量碳(-23.6%)和微生物生物量氮(-32.8%),而微生物生物量磷未见显著变化。
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注:AMF:丛枝菌根真菌;PLFA:磷脂脂肪酸法;MBC:微生物生物量碳;MBN:微生物生物量氮;MBP:微生物生物量磷:G+:革兰氏阳性菌;G-:革兰氏阴性菌。 Note: AMF: arbuscular mycorrhizal fungi; PLFA: phospholipid fatty acid; MBC: Microbial biomass carbon; MBN: microbial biomass nitrogen; MBP: microbial biomass phosphorus; G+: gram-positive bacterium; G-: gram-negative bacterium. 图 3 毛竹入侵对微生物指标(a.真菌与细菌的香农指数和Chao1指数,b.五种优势细菌门相对丰度,c.微生物生物量碳、微生物生物量氮、微生物生物量磷与PLFA)的影响 Fig. 3 Effects of bamboo invasion on microbial indices(a. Shannon index and Chao1 index for fungi and bacteria, b. Relative abundances of five dominant bacterial phyla, c. Microbial biomass carbon, microbial biomass nitrogen, microbial biomass phosphorus and PLFA) |
土壤氮素中98%为有机态氮,微生物水解矿化释放出铵态氮[25]。毛竹入侵原生林后全氮下降而碱解氮、铵态氮上升(图 2),表明毛竹具有较强的矿化有机氮作用,致全氮含量下降,而矿化产生的大量氨(NH3),质子化(NH3+H+→
毛竹入侵改变了土壤总有机碳含量(图 2),这可能与土壤碳循环的改变有关。土壤总有机碳积累与否取决于有机碳的输入(植物源碳)和输出(微生物分解释放二氧化碳)平衡。毛竹入侵后有机碳的输入和输出均发生了改变。在亚热带地区常绿阔叶林土壤有机质一般高于其他林分,毛竹入侵阔叶林后土壤总有机碳显著增加,但入侵针叶林则显著下降(图 2)。从有机碳输入端分析,毛竹入侵不同林分生物量的增加量差别不明显,所以导致土壤TOC差异的原因可能是微生物分解的输出端。毛竹入侵后导致微生物生物量碳(MBC)和微生物生物量氮(MBN)总体下降(图 3c),然而,将MBC按照入侵林分(阔叶林、针叶林)分类的Meta分析(图略)发现,入侵阔叶林致土壤MBC显著下降(–26.7%,P<0.05),入侵针叶则以土壤MBC显著增加(+9.74%,P<0.05)。该数据分析很好地解释了入侵阔叶林后土壤微生物分解强度下降、TOC增加,而入侵针叶林则正好相反的结果。混交林土壤总有机碳含量低于原生林的原因可能是毛竹入侵形成混交林时提高土壤微生物活性,从而促进有机质降解。例如:城市森林中混种多果桉(B. polycarpa)与水杉(M. glyptostroboides)可促进微生物对养分的释放作用[32];阔叶林与针叶林凋落物混合后,微生物的分解速率明显提升[33];混合林分重新造林后,相比于单一树种更能提升土壤微生物生物量和土壤基础呼吸速率[34]。
毛竹入侵后土壤有效磷总体显著增加(图 2)。一方面,基于土壤化学过程的分析,在酸性条件下,磷主要被铁铝化合物所吸附固定,较高的土壤pH使OH–与H2PO4–发生交换,提高土壤中磷的有效性。另一方面,基于土壤微生物化学的过程分析,土壤pH提升可能刺激土壤中某些溶磷菌活性[35],加速释放土壤有效磷。入侵阔叶林后土壤有效磷增幅极显著高于入侵针叶林,但从入侵前后两者土壤pH的变化差异无法解释,可能是阔叶林土壤中全磷含量高于针叶林[36],其中的有机磷或无机磷被活化所致。
3.2 毛竹入侵影响微生物群落的可能机制诸多研究证明土壤pH可直接和间接影响土壤微生物[37]。本研究数据分析表明,放线菌门相对丰度(高通量结果)以及放线菌门PLFA含量显著增加,而酸杆菌门、浮霉菌门、拟杆菌门的丰度显著下降(图 3)。放线菌门细菌偏好碱性环境,毛竹入侵致土壤pH升高有利于放线菌门细菌繁殖。此外,作为最优势细菌门之一,偏好酸性土壤[37]的酸杆菌门,在毛竹入侵提升土壤pH后其生存空间被抑制,也为放线菌门留出更大的生存空间。严淑娴等[38]进一步分析放线菌门的优势物种后发现毛竹林土壤中Frankiales(属放线菌门)的相对丰度显著(P < 0.05)高于阔叶林与杉木林,而Frankiales是被证实具有强烈固氮能力的菌种,能与大多数乔木和灌木形成共生固氮系统[39],毛竹林土壤的这种固氮能力也可能是其土壤铵态氮高的原因之一。此外,本研究结果还表明毛竹入侵后细菌的香农多样性指数有显著提升(图 3a),这可能与毛竹入侵导致土壤pH提升有关,因为大量研究证明,土壤细菌香农多样性指数随着土壤pH升高呈线性增加[40]。
真菌喜酸性环境,土壤真菌的多样性与生物量一般随着pH的降低而增加[41]。但本研究毛竹入侵前后土壤真菌变化的数据分析结果表明,毛竹入侵后土壤pH提升并未致土壤真菌的多样性及PLFA下降,反而混交林土壤真菌多样性显著高于原生林(图 3a);从有机输入的数量和质量对土壤真菌的影响分析,毛竹庞大的地下鞭根系统为真菌提供了大量材料,激发土壤真菌的大量繁殖,土壤真菌的多样性增加,如在中纬度地区,植物入侵增加土壤真菌的周转率从而提高真菌β多样性[42]。至于真菌生物量增加(图 3c)的原因可从以下几个方面解释,一是土壤中某些特定真菌门有响应pH升高的能力,有研究发现毛竹土壤pH较高并且担子菌门丰度显著高于阔叶和杉木[38];Tedersoo等[43]发现担子菌门的丰度随着土壤pH的提升而增加;改变土地利用情况后,土壤pH提升时子囊菌门的丰度也相应提升[44];全球尺度的统计分析表明,在pH3~5范围内,子囊菌门相对丰度随着土壤pH升高而增加,pH5~7则下降,而壶菌门的相对丰度则是pH3~7范围内直线增加[45]。二是真菌的重要分支AMF(丛枝菌根真菌)对pH的积极响应,虽然毛竹入侵后原始植物物种减少,导致仅定殖竹根的AMF保留,基于乘客假设(passenger hypothesis)的原理,毛竹依然会刺激根鞭中AMF生物量的增加[46];类似研究证明,在提升土壤pH后AMF的定殖量和生物量显著增加,并提高土壤磷的有效性[47]。本研究数据分析表明,毛竹入侵后土壤AMF的PLFA变化不显著(图 3c),可能是因为样本数不足导致。
3.3 土壤pH升高可能的机制毛竹入侵原生林分显著提升土壤pH的普遍性得到证实(图 1),并且此现象具有随着时间推移而加剧的正效应。毛竹根系主要分布在0~20 cm土层[48],因而对此区间的土壤pH提升幅度最大。此外,本研究发现毛竹入侵针叶林后土壤pH的提升幅度明显高于入侵阔叶林类林分(图 1),可能是原生针叶林土壤平均pH 4.31低于阔叶林pH 4.40导致。虽然土壤pH升高的现象占总体比值为84.9%,但仍然有15.1%未呈现上升结果,推测是因为这些点位毛竹林存在时间较短,毛竹提升土壤pH累加效应尚不明显。
植物改变土壤pH的案例很多,植物吸收阳离子[49]时会泌出氢离子而使根际土壤酸化;若植物残体中有机阴离子与质子的比值较大,土壤pH随该比值线性增加[50]。氮素循环过程也会影响土壤pH,微生物的氨化作用消耗质子致土壤pH上升,而其硝化作用释放质子致土壤pH降低[51]。因此,推测毛竹入侵后土壤pH的变化可能是通过改变土壤特定离子浓度所致。基于本研究Meta分析中几种氮素的变化结果(图 2),土壤中氮相关的特定离子浓度发生变化,在所有研究情况下土壤硝态氮显著下降,而铵态氮显著上升,因此推测:(1)毛竹入侵可能增强NH3+H+→
本研究数据分析结果证明毛竹林入侵周围原生林分提高土壤pH的科学性和普遍性,并且随着入侵阶段、时间的增加而加剧,并对根系分布层(0~20 cm)影响最大。毛竹入侵总体上使土壤总有机碳、硝态氮含量下降,但铵态氮和有效磷含量提升。毛竹入侵提高了土壤细菌和真菌多样性,表明对土壤微生物的正反馈作用。
附表1
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