植被恢复措施影响沙化高寒草地土壤微生物碳氮功能

doi:10.7522/j.issn.1000-694X.2025.00037

摘要/Abstract

摘要：

为探讨植被恢复措施对青藏高原沙化高寒草地与碳氮循环相关的土壤微生物的影响，以分别采用草本和灌丛在沙化草地进行人工植被重建的恢复草地为研究对象，运用宏基因组测序技术对比研究了两种恢复措施对碳氮循环相关土壤微生物功能群结构及其潜在功能的影响。结果表明：植被恢复显著提升了沙化草地中与碳氮循环相关的土壤微生物功能群的多样性、丰富度和均匀度。草地沙化和两种植被恢复措施均显著改变了与碳氮循环相关的土壤微生物功能群结构，进而影响了其潜在生态功能。人工植被恢复草地的绝大多数土壤微生物碳代谢功能与天然草地无显著差异，而大部分氮循环功能仍与天然草地具有明显差异，表明沙化高寒草地土壤微生物碳代谢功能的恢复效果比氮循环功能更佳。碳氮循环相关土壤微生物功能群与植被和土壤环境的改善显著正相关。人工植被重建重塑了沙化高寒草地的土壤微生物功能群结构，但不同植被恢复措施对土壤微生物碳氮循环潜在功能的影响差异不明显。

关键词: 草地退化, 人工植被重建, 碳氮循环, 宏基因组, 微生物功能群

Abstract:

To explore the effects of vegetation to restore the measures on carbon and nitrogen cyclings related soil microorganisms in the desertified Tibetan alpine meadow， this study selected artificially revegetated grasslands using herbs and shrubs to restore the desertified alpine grassland. We compared the structures and potential functions of carbon and nitrogen cyclings related soil microbial communities between the two revegetated grasslands using metagenomic sequencing technology. The results showed that vegetation revegetation significantly improved the Shannon-Wiener diversity， richness and evenness of carbon and nitrogen cyclings related soil microbial functional communities， and thereby affected the potential ecological functions. While there was no significant differences in most soil microbial carbon metabolic functions between revegetated grasslands and the natural desert， many nitrogen cycling related functions showed significant difference， indicating that the restoration of soil microbial carbon metabolic functions in the desertified alpine grassland due to revegetation was better than that of nitrogen cycling related functions. The carbon and nitrogen cyclings related soil microbial functional communities were significantly positively associated with the improvements of vegetation and soil environmental conditions. These results showed that despite artificial revegetation restructed the structures of carbon and nitrogen cycles related soil microbial functional communities in desertified alpine grassland， the effects of different revegetation measures on potential carbon and nitrogen functions of soil microbial communities were limited.

Key words: grassland degradation, artificial revegetation, carbon and nitrogen cycling, metagenomics, microbial functional community

中图分类号:

Q938

刘文静, 胡宜刚, 贺祯子, 张振华, 李以康. 植被恢复措施影响沙化高寒草地土壤微生物碳氮功能[J]. 中国沙漠, 2025, 45(3): 198-209.

Wenjing Liu, Yigang Hu, Zhenzi He, Zhenhua Zhang, Yikang Li. Vegetation restoration measures affect soil microbial carbon and nitrogen functions in desertified alpine grassland[J]. Journal of Desert Research, 2025, 45(3): 198-209.

图/表 9

图1 研究区不同草地类型景观

Fig.1 The views of different grasslands

表1 碳氮循环相关土壤微生物群落α多样性指数

Table 1 The α-diversities of C and N cycles related soil microbial communities in different grasslands

草地类型	碳循环			氮循环
草地类型	丰富度	Shannon-Wiener指数	Simpson指数	丰富度	Shannon-Wiener指数	Simpson指数
天然草地	5 770±292^b	5.82±0.09^a	0.024±0.003^a	661±47^b	5.41±0.07^a	0.011±0.001^a
沙化草地	6 105±227^a	5.12±0.05^b	0.057±0.004^a	746±46^a	4.90±0.05^b	0.039±0.004^a
草本人工草地	6 389±272^a	6.21±0.03^a	0.014±0.001^b	779±31^a	5.74±0.03^a	0.007±0.001^b
灌丛人工草地	6 298±333^a	6.24±0.11^a	0.013±0.002^b	783±73^a	5.73±0.09^a	0.007±0.001^b

图2 不同草地类型碳氮循环相关土壤微生物群落Venn图和群落组成

Fig.2 Venn diagrams and C and N cycles related soil microbial structures in different grasslands

图3 KO水平上碳氮循环相关土壤微生物群落主坐标分析

Fig.3 PCoA of C and N cycles related soil microbial communities at the KO level across different grasslands

表2 碳氮循环相关土壤微生物群落结构多元非参数检验

Table 2 Multivariate nonparametric tests （ANOSIM） based on Bray-Curtis distance of C and N cycles related soil microbial communities structures among grasslands

草地类型	碳循环		氮循环
草地类型	r	P	r	P
天然草地-沙化草地	1	0.004	1	0.004
天然草地-草本人工草地	1	0.004	1	0.004
天然草地-灌丛人工草地	1	0.004	0.98	0.004
沙化草地-草本人工草地	1	0.004	1	0.004
沙化草地-灌丛人工草地	1	0.004	1	0.004
草本人工草地-灌丛人工草地	0.292	0.047	0.644	0.014

图4 KEGG注释的Module水平上碳氮循环相关土壤微生物组成差异注：“**”和“*”分别表示在P<0.01和P<0.05水平上差异显著，不同小写字母表示各草地类型间差异显著（P<0.05）

Fig.4 Differences in the relative proportion of C and N cycles related soil microbial communities at the KEGG module level

图5 土壤微生物对碳氮循环主要功能模块的相对贡献度

Fig.5 The relative contributions of soil microbial communities to the key functional modules of C and N cycles

表3 碳氮循环相关微生物KOs结构与环境因子之间的Mantel test分析

Table 3 Mantel test for the correlations of C and N cycles related microbial KOs with vegetation and soil properties

环境因子	碳循环		氮循环
环境因子	r	P	r	P
植被盖度	0.69	0.001	0.65	0.001
植被地上生物量	0.70	0.001	0.63	0.001
植被丰富度	0.67	0.001	0.64	0.002
植被Shannon指数	0.67	0.001	0.64	0.002
土壤pH	0.13	0.107	0.28	0.018
土壤电导率	0.52	0.001	0.67	0.001
土壤容重	0.16	0.135	0.32	0.017
土壤水分	0.62	0.001	0.57	0.001
土壤总有机碳	0.68	0.001	0.62	0.001
土壤全氮	0.34	0.002	0.44	0.001
土壤全磷	-0.11	0.893	-0.04	0.640
土壤铵态氮	0.01	0.405	0.10	0.159
土壤硝态氮	0.62	0.001	0.60	0.001
土壤有效磷	0.35	0.005	0.34	0.006
土壤速效钾	0.23	0.016	0.39	0.002

图6 碳氮循环相关微生物KOs结构与环境因子之间的db-RDA分析注：Richness：植物丰富度；ABs：地上生物量；SM：土壤水分；BD：容重；TP：全磷；AP：有效磷AK：速效钾；Ammonia：铵态氮

Fig.6 db-RDA between C and N cycle related microbial KOs and environmental factors

参考文献 46

1	傅伯杰，王晓峰，冯晓明，等.国家生态屏障区生态系统评估［M］.北京：科学出版社，2017.
2	Zhang H， Tang Z， Wang B，et al.A 250 m annual alpine grassland AGB dataset over the Qinghai-Tibet Plateau （2000-2019） in China based on in situ measurements，UAV photos，and MODIS data［J］.Earth System Science Data，2023，15：821-846.
3	Niu Y， Zhu H， Yang S，et al.Overgrazing leads to soil cracking that later triggers the severe degradation of alpine meadows on the Tibetan Plateau［J］.Land Degradation & Development，2019，30（5）：1243-1257.
4	Liu Y， Zhang Z， Tong L，et al.Assessing the effects of climate variation and human activities on grassland degradation and restoration across the globe［J］.Ecological Indicators，2019，106：105504.
5	Yuan Q， Yuan Q Z， Ren P，et al.Coupled effect of climate change and human activities on the restoration/degradation of the Qinghai-Tibet Plateau grassland［J］.Journal of Geographical Sciences，2021，31（8）：1299-1327.
6	Gang C， Zhou W， Chen Y，et al.Quantitative assessment of the contributions of climate change and human activities on global grassland degradation［J］.Environment Earth Science，2014，72（7）：4273-4282.
7	Cai H， Yang X， Xu X，et al.Human-induced grassland degradation/restoration in the central Tibetan Plateau： the effects of ecological protection and restoration projects［J］.Ecological Engineering，2015，83：112-119.
8	Niu Y， Squires V R， Hua L，et al.High-altitude aeolian desertification and sand dunes on the Tibetan Plateau，China［M］.Sand Dunes of the Northern Hemisphere.Boca Raton： CRC Press，2023：179-194.
9	胡宜刚，李睿，辛玉琴，等.青藏铁路植被恢复和黑土型退化草地治理的实践与启示［J］.草业科学，2015，9（9）：1413.
10	张骞，马丽，张中华，等.青藏高寒区退化草地生态恢复：退化现状、恢复措施、效应与展望［J］.生态学报，2019，39（20）：7441-7451.
11	贺金生，卜海燕，胡小文，等.退化高寒草地的近自然恢复：理论基础与技术途径［J］.科学通报，2020，65（34）：3898-3908.
12	晏和飘，李文龙，梁天刚，等.青藏高原退化高寒草地恢复对不同措施响应的Meta分析［J］.草地学报，2021，29（）：190-198.
13	王亚妮，胡宜刚，王增如，等.人工植被重建对沙化高寒草地土壤真菌群落特征的影响［J］.土壤学报，2023，60（1）：280-291.
14	王亚妮，胡宜刚，王增如，等.沙化和人工植被重建对高寒草地土壤细菌群落特征的影响［J］.草业学报，2022，31（5）：26-39.
15	Hartmann M， Six J.Soil structure and microbiome functions in agroecosystems［J］.Nature Reviews Earth & Environment，2023，4：4-18.
16	Jiang J， Song M H.Review of the roles of plants and soil microorganisms in regulating ecosystem nutrient cycling［J］.Plant Ecology，2010，34（8）：979-990.
17	Wilhelm R C， Amsili J P， Kurtz K S M，et al.Ecological insights into soil health according to the genomic traits and environment-wide associations of bacteria in agricultural soils［J］.ISME Communications，2023，3：1.
18	Bahram M， Hildebrand F， Forslund S K，et al.Structure and function of the global topsoil microbiome［J］.Nature，2018，560：233-237.
19	Olsen S R， Sommers L E.Methods of Soil Analysis［M］.Madison，USA：American Society of Agronomy，1982：403-430.
20	鲁如坤.土壤农业化学分析方法［M］.北京：中国农业科技出版社，2000.
21	Hyatt D， Chen G L， LoCascio P F，et al.Prodigal： prokaryotic gene recognition and translation initiation site identification［J］.BMC Bioinformatics，2010，11：119.
22	Hu Y F， Peng J J， Yuan S，et al.Influence of ecological restoration on vegetation and soil microbiological properties in alpine-cold semi-humid desertified land［J］.Ecological Engineering，2016，94：88-94.
23	Ma W W， Li G， Wu J H，et al.Response of soil labile organic carbon fractions and carbon-cycle enzyme activities to vegetation degradation in a wet meadow on the Qinghai-Tibet Plateau［J］.Geoderma，2020，377：114565.
24	Wieder W R， Bonan G B， Allison S D.Global soil carbon projections are improved by modelling microbial processes［J］.Nature Climate Change，2013，3：909-912.
25	Sokol N W， Bradford M A.Microbial formation of stable soil carbon is more efficient from belowground than aboveground input［J］.Nature Geoscience，2019，12：46-53.
26	Wang X， Zhou M， Yue H，et al.Effects of different artificial vegetation restoration modes on soil microbial community structure in the soil erosion area of southern China［J］.CATENA，2024，237：107803.
27	张志永，樊宝敏，宋超，等.根系构型研究进展：功能、影响因子和研究方法［J］.资源与生态学报，2023，14（1）：15-24.
28	Li Y， Jiang L， Yuan H，et al.The impact of artificial afforestation on the soil microbial community and function in desertified areas of NW China［J］.Forests，2024，15（7）：1140.
29	Kang L， Song Y， Mackelprang R，et al.Metagenomic insights into microbial community structure and metabolism in alpine permafrost on the Tibetan Plateau［J］.Nature Communications，2024，15：5920.
30	孙政国，孙成明，李建龙，等.我国草地生态系统碳循环机制及碳蓄积核算研究回顾与展望［J］.草地科学，2021，28（9）：1611-1616.
31	Celaya-Herrera S， Casados-Vázquez L E， Valdez-Vazquez I，et al.A Cellulolytic Streptomyces Sp.isolated from a highly oligotrophic niche shows potential for hydrolyzing agricultural wastes［J］.Bioenergy Research，2021，14：333-343.
32	Xie G， Zhang Y， Gong Y，et al.Extreme trophic tales： deciphering bacterial diversity and potential functions in oligotrophic and hypereutrophic lakes［J］.BMC Microbiology，2024，24：348.
33	Islam Z F， Cordero P R F， Feng J，et al.Two Chloroflexi classes independently evolved the ability to persist on atmospheric hydrogen and carbon monoxide［J］.ISME Journal，2019，13：1801-1813.
34	Konneke M， Bernhard A E， de la Torre J R，et al.Isolation of an autotrophic ammonia-oxidizing marine archaeon［J］.Nature，2005，437：543-546.
35	Jung M Y， Sedlacek C J， Kits K D，et al.Ammonia-oxidizing archaea possess a wide range of cellular ammonia affinities［J］.ISME，2022，16：272-283.
36	Nikitin D A， Ivanova E A， Semenov M V，et al.Diversity，ecological characteristics and identification of some problematic phytopathogenic fusarium in soil： a review［J］.Diversity，2023，15（1）：49.
37	Bai Y， Cotrufo M F.Grassland soil carbon sequestration： current understanding，challenges，and solutions［J］.Science，2022，377（6606）：603-608.
38	Wang D， Zhou H， Zuo J，et al.Responses of soil microbial metabolic activity and community structure to different degraded and restored grassland gradients of the Tibetan Plateau［J］.Frontiers in Plant Science，2022，13：770315.
39	吴希慧，王蕊，高长青，等.土地利用驱动的土壤性状变化影响微生物群落结构和功能［J］.生态学报，2021，41（20）：7989-8002.
40	Luo Y， Su B， Currie W S，et al.Progressive nitrogen limitation of ecosystem responses to rising atmospheric carbon dioxide［J］.BioScience，2004，54（8）：731-739.
41	李靖宇，张肖冲，陈韵，等.腾格里沙漠东南缘藻结皮与藓结皮放线菌多样性及其潜在代谢功能［J］.生态学报，2020，40（5）：1590-1601.
42	方海富，陈翔，杨红玲，等.草地土壤N₂O排放对人为干扰的响应研究进展［J］.中国沙漠，2025，45（2）：184-190.
43	Wang M， Li D， Frey B，et al.Land use modified impacts of global change factors on soil microbial structure and function：a global hierarchical meta-analysis［J］.Science of The Total Environment，2024，935：173286.
44	Chen J， Li F C， Jia B，et al.Regulation of soil nitrogen cycling by shrubs in grasslands［J］.Soil Biology and Biochemistry，2024，191：109327.
45	Wu D， He X， Jiang L，et al.Root exudates facilitate the regulation of soil microbial community function in the genus Haloxylon ［J］. Frontiers Plant Science，2024，15：1461893.
46	Yang Y， Zheng L， Zhou Y，et al. Changes in soil microbial community structure and function following degradation in a temperate grassland［J］.Journal of Plant Ecology，2021，14（3）：384-397.

[1]	尤全刚, 薛娴, 彭飞, 董斯扬. 高寒草甸草地退化对土壤水热性质的影响及其环境效应[J]. 中国沙漠, 2015, 35(5): 1183-1192.
[2]	戴新刚, 熊喆, K.Kramer, 贾根锁, 赵登海. 羊啃食动力学模拟与内蒙古西部气候变化适应[J]. 中国沙漠, 2012, 32(5): 1442-1450.
[3]	戚登臣;陈文业;郑华平;李广宇;刘正恒;宗文杰. 甘南黄河上游水源补给区“黑土滩”型退化草地现状、成因及综合治理对策[J]. 中国沙漠, 2008, 28(6): 1058-1063.
[4]	李辉霞;刘淑珍. 基于ETM+影像的草地退化评价模型研究——以西藏自治区那曲县为例[J]. 中国沙漠, 2007, 27(3): 412-418.
[5]	王根绪, 程国栋. 江河源区的草地资源特征与草地生态变化[J]. 中国沙漠, 2001, 21(2): 101-107.