生物土壤结皮中微生物群落特征综述

doi:10.7522/j.issn.1000-694X.2021.00137

摘要/Abstract

摘要：

生物土壤结皮（也称生物结皮）是联结荒漠区地表生物成分与非生物成分的纽带，它不仅是荒漠区生物多样性的“热点”，也是退化生态系统修复的“工程师”。生物结皮的拓殖不仅能稳定地表，影响土壤水文过程，对土壤形成起着重要作用，还对荒漠区土壤碳氮循环等产生重要影响。蓝藻、细菌、真菌和古菌等微生物在生物结皮形成和发挥生态功能过程中发挥重要作用。本文系统总结了全球寒漠、冷漠和热漠生物结皮中微生物的组成和群落特征，厘清了主要生物气候区不同演替阶段的生物结皮中主要微生物物种，对相关微生物的研究进展和存在问题进行了分析，最后从微生物相互作用角度为明确关键枢纽微生物种提出了建议，以期对筛选潜在高效固沙菌种、培育人工结皮促进受损生态系统恢复提供参考。

关键词: 生物气候区, 生物土壤结皮, 微生物群落特征, 全球变化

Abstract:

Biological soil crusts （also known as biocrusts） are the link structures between surface biotic components and abiotic components in deserts. Biocrusts are not only the "hot spots" of biodiversity in deserts， but also the "engineers" for the restoration of degraded ecosystems. The colonization of biocrusts not only can stabilize the surface， affect soil hydrological process and play an important role in pedogenesis， but also can affect soil carbon and nitrogen cycles in deserts. As microorganisms （cyanobacteria， bacteria， fungi， archaea， etc.） play an important role in the formation and ecological function of biocrusts， this paper systematically summarized the composition and community assembly of microorganisms in biocrusts in cold deserts， cool deserts and hot deserts， and clarified the main microbial species at different succession stages of biocrusts in different bioclimatic regions at global scale. The existing problems were also discussed. Finally， the authors suggested to focus on microbial interactions and identify the key hub microbial species by co-occurrence ecological network analysis， which will help select potential efficient inoculums for cultivating artificial biocrusts and promoting the restoration of degraded ecosystems.

Key words: bioclimatic regions, biological soil crusts, microbial community assembly, global change

中图分类号:

Q938.1

鲍婧婷, 孙靖尧, 王进. 生物土壤结皮中微生物群落特征综述[J]. 中国沙漠, 2022, 42(6): 33-43.

Jingting Bao, Jingyao Sun, Jin Wang. A review on microbial community assembly in biological soil crusts[J]. Journal of Desert Research, 2022, 42(6): 33-43.

图/表 4

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演替阶段	气候区
演替阶段	寒漠	冷漠	热漠
藻结皮	Microcoleus vaginatus和M. steenstrupii (科罗拉多高原的寒漠，犹他州)^[14] Synechococcales (Leptolyngbya antarctica) Oscillatoriales， Nostocales, Gloeobacterales， Chroococcidiopsidales(北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16] Microcoleus， Mastigocladopsis， Wilmottia， Chroococcidiopsis (腾格里沙漠)^[17] Microcoleus， Tolypothrix， Wilmottia， Scytonema (克孜勒库姆沙漠)^[17] Microcoleus， Vaginatus， Chroococcidiopsis， M. steenstrupii， Leptolyngbya， Scytonema， Tolypothrix和 Nostoc (东伊比利亚半岛)^[10]	Microcoleus vaginatus， M. steenstrupii， Leptolyngbya spp.和Phormidium spp.(浅色结皮)； M. vaginatus， M. steenstrupii， Nostoc spp.，和Scytonema spp.(深色结皮) (奇瓦瓦沙漠)^[19]; Chroococcidiopsis sp.， Pseudanbaena spp.， Phormidium spp.， Leptolyngbya spp.， Microcoleus paludosus，和Nostoc spp. (纳米布沙漠)^[20]; Synechococcophycidae， Oscillatoriophycideae， Nostocophycidae (阿塔卡马沙漠)^[22]; Microcoleus like， Leptolyngbya， Chroococcidiopsis， Nostoc， Scytonema， Trichocoleus (内盖夫沙漠)^[21]
地衣结皮	Synechococcales， Oscillatoriales (Phormidium sp. )， Nostocales， Chroococcidiopsidales， Gloeobacterales， Chroococcales (北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16]	Oscillatoriales (Microcoleus， Lyngbya， Phormidium， Oscillatoria)和Nostocales (Nostoc， Scytonema， Tolypothrix) (蒙特沙漠)^[23]
藻-地衣结皮	Synechococcales (Leptolyngbya sp.和Leptolyngbya antarctica)， Oscillatoriales， Nostocales， Gloeobacterales， Chroococcidiopsidales (北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16]	Scytonema和Microcoleus steenstrupii (索诺拉沙漠)^[27]; Chroococcidiopsis， Pseudoanabaena， Phormidium， Leptolyngbya， Microcoleus和Nostoc (非洲的卡鲁沙漠)^[23]
地衣-藓结皮	Pseudanabaenaceae (filamentous species without heterocytes)和Nostoc (heterocytous) (北极苔原土壤)^[28]	Nostocales (Scytonema and Nostoc)和Pseudanabaenaceae (西班牙塔韦纳斯沙漠)^[18]; Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis和Scytonema (古尔班通古特沙漠)^[16]; Microcoleus， Mastigocladopsis， Wilmottia， uncultured_Nostocales， uncultured_Coleofasciculaceae， Nostoc (腾格里沙漠)^[17]; Microcoleus， Mastigocladopsis， Tolypothrix， Wilmottia， Phormidium， uncultured_Nostocales (克孜勒库姆沙漠)^[17]	—
藓结皮	—	Microcoleus， Mastigocladopsis， Wilmottia， uncultured_Coleofasciculaceae， uncultured_Nostocales， Nostoc (腾格里沙漠)^[17] Microcoleus， Wilmottia， Mastigocladopsis， Tolypothrix， uncultured_Nostocales， uncultured_Coleofasciculaceae (克孜勒库姆沙漠)^[17]	—

演替阶段	气候区
演替阶段	寒漠	冷漠	热漠
藻结皮	Microcoleus vaginatus和M. steenstrupii (科罗拉多高原的寒漠，犹他州)^[14] Synechococcales (Leptolyngbya antarctica) Oscillatoriales， Nostocales, Gloeobacterales， Chroococcidiopsidales(北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16] Microcoleus， Mastigocladopsis， Wilmottia， Chroococcidiopsis (腾格里沙漠)^[17] Microcoleus， Tolypothrix， Wilmottia， Scytonema (克孜勒库姆沙漠)^[17] Microcoleus， Vaginatus， Chroococcidiopsis， M. steenstrupii， Leptolyngbya， Scytonema， Tolypothrix和 Nostoc (东伊比利亚半岛)^[10]	Microcoleus vaginatus， M. steenstrupii， Leptolyngbya spp.和Phormidium spp.(浅色结皮)； M. vaginatus， M. steenstrupii， Nostoc spp.，和Scytonema spp.(深色结皮) (奇瓦瓦沙漠)^[19]; Chroococcidiopsis sp.， Pseudanbaena spp.， Phormidium spp.， Leptolyngbya spp.， Microcoleus paludosus，和Nostoc spp. (纳米布沙漠)^[20]; Synechococcophycidae， Oscillatoriophycideae， Nostocophycidae (阿塔卡马沙漠)^[22]; Microcoleus like， Leptolyngbya， Chroococcidiopsis， Nostoc， Scytonema， Trichocoleus (内盖夫沙漠)^[21]
地衣结皮	Synechococcales， Oscillatoriales (Phormidium sp. )， Nostocales， Chroococcidiopsidales， Gloeobacterales， Chroococcales (北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16]	Oscillatoriales (Microcoleus， Lyngbya， Phormidium， Oscillatoria)和Nostocales (Nostoc， Scytonema， Tolypothrix) (蒙特沙漠)^[23]
藻-地衣结皮	Synechococcales (Leptolyngbya sp.和Leptolyngbya antarctica)， Oscillatoriales， Nostocales， Gloeobacterales， Chroococcidiopsidales (北极的Petunia湾)^[15]	Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis， Nostoc， Scytonema (古尔班通古特沙漠)^[16]	Scytonema和Microcoleus steenstrupii (索诺拉沙漠)^[27]; Chroococcidiopsis， Pseudoanabaena， Phormidium， Leptolyngbya， Microcoleus和Nostoc (非洲的卡鲁沙漠)^[23]
地衣-藓结皮	Pseudanabaenaceae (filamentous species without heterocytes)和Nostoc (heterocytous) (北极苔原土壤)^[28]	Nostocales (Scytonema and Nostoc)和Pseudanabaenaceae (西班牙塔韦纳斯沙漠)^[18]; Microcoleus vaginatus， Microcoleus steenstrupii， Chroococcidiopsis和Scytonema (古尔班通古特沙漠)^[16]; Microcoleus， Mastigocladopsis， Wilmottia， uncultured_Nostocales， uncultured_Coleofasciculaceae， Nostoc (腾格里沙漠)^[17]; Microcoleus， Mastigocladopsis， Tolypothrix， Wilmottia， Phormidium， uncultured_Nostocales (克孜勒库姆沙漠)^[17]	—
藓结皮	—	Microcoleus， Mastigocladopsis， Wilmottia， uncultured_Coleofasciculaceae， uncultured_Nostocales， Nostoc (腾格里沙漠)^[17] Microcoleus， Wilmottia， Mastigocladopsis， Tolypothrix， uncultured_Nostocales， uncultured_Coleofasciculaceae (克孜勒库姆沙漠)^[17]	—

演替阶段	气候区
演替阶段	寒漠	冷漠	热漠
藻结藻结皮	Actinobacteria， β-Proteobacteria， Bacteriodetes， Low-GC Gram-positives (Bacilli)， α-Proteobacteria， Acidobacteria和Thermomicrobiales (科罗拉多高原的寒漠，犹他州)^[14]	Actinobacteria，Proteobacteria， Firmicutes， Chloroflexi， Acidobacteria，Cyanobacteria， Gemmatimonadetes， Bacteroidetes (腾格里沙漠)^[45]; Cyanobacteria， Bacteroidetes， Proteobacteria， unclassified bacteria， Actinobacteria (古尔班通古特沙漠)^[46]; Leptolyngbya (Cyanobacteria)， Rubrobacter， Solirubrobacter， Geodermatophilus (Actinomycetes) (塔韦纳斯沙漠)^[39]	Rhizobiales (phylum Proteobacteria)， Chroococcales， Nostocophycideae (Cyanobacteria) (西澳)^[47]; Cyanobacteria， Alphaproteobacteria， Bacteroidetes， Chloroflexi， Actinobacteria， Acidobacteria (阿曼)^[29]; Cyanobacteria， Proteobacteria， Bacteroidetes， Acidobacteria (莫哈韦沙漠)^[48]
地衣结皮	Actinobacteria， Proteobacteria, Acidobacteria， Firmicutes， Armatimonadetes Cyanobacteria， Bacteroidetes和Planctomycetes（南极洲大陆维多利亚岛)^[49]	Proteobacteria， Actinobacteria， Chloroflexi， Cyanobacteria， Bacteroidetes， Planctomycetes， Acidobacteria， Armatimonadetes (腾格里沙漠)^[50]; Cyanobacteria， Proteobacteria， Bacteroidetes， unclassified bacteria， Acidobacteria， Actinobacteria， Verrucomicrobia (古尔班通古特沙漠)^[46]; Proteobacteria， Actinobacteria， Bacteroidetes， Acidobacteria， Cyanobacteria and Verrucomicrobia (西班牙塔韦纳斯沙漠)^[31]	Acidobacteriales (phylum Acidobacteria)， Rhodospirillales (Proteobacteria)， Actinomycetales (Actinobacteria)和unclassified Proteobacteria (西澳)^[47]; Cyanobacteria， Bacteroidetes， Acidobacteria， Alphaproteobacteria， Actinobacteria， Chloroflexi (阿曼)^[29]
藻-地衣结皮	—	Nostocales， Chroococcidiopsaceae， Coleofasciculaceae (Cyanobacteria) (塔韦纳斯沙漠)^[39]	Cyanobacteria (Scytonema and Microcoleus steenstrupii)， Proteobacteria (β-Proteobacteria)， Actinobacteria， Bacteriodetes，和Chloroflexi (索诺拉沙漠)^[27]; Cyanobacteria， Proteobacteria， Bacteroidetes， Chloroflexi， Armatimonadetes (莫哈韦沙漠)^[48]; Bacteroidetes， Proteobacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Chloroflexi， Verrucomicrobia和Planctomycetes (非洲的卡鲁沙漠)^[32]
地衣-藓结皮	Cyanobacteria， Bacteroidetes， Proteobacteria， Actinobacteria， Chloroflexi， Acidobacteria (美国峡谷国家公园)^[51];Actinobacteria， Bacteroidetes， Proteobacteria， Cyanobacteria， Firmicutes， Verrucomicrobia， Acidobacteria， Chloroflexi (美国西部山间寒漠)^[52]; Cyanobacteria， Actinobacteria， Bacteriodetes， Chloroflexi和Proteobacteria (科罗拉多高原的寒漠)^[53]	Actinobacteria， Proteobacteria， Chloroflexi， Cyanobacteria， Acidobacteria， Firmicutes， Bacteroidetes (腾格里沙漠)^[45]	Cyanobacteria， Proteobacteria， Bacteroidetes， Actinobacteria和Acidobacteria (奇瓦瓦沙漠的Actopan和Atexcac地区)^[54]
藓结皮	—	Proteobacteria， Bacteroidetes， unclassified bacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Verrucomicrobia (古尔班通古特沙漠)^[46]; Bacteroidetes， Acidobacteria和Proteobacteria (新墨西哥桑迪亚山脉)^[34]; Acidobacteria， Proteobacteria， Chloroflexi， Actinobacteria (黄土高原)^[33]	Proteobacteria， Actinobacteria， Acidobacteria， Chloroflexi， Bacteroidetes， Cyanobacteria， Deinococcus-Thermus， Planctomycetes (莫哈韦沙漠)^[48]; Bacteroidetes， Proteobacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Chloroflexi， Verrucomicrobia和Planctomycetes (非洲卡鲁沙漠)^[32]

演替阶段	气候区
演替阶段	寒漠	冷漠	热漠
藻结藻结皮	Actinobacteria， β-Proteobacteria， Bacteriodetes， Low-GC Gram-positives (Bacilli)， α-Proteobacteria， Acidobacteria和Thermomicrobiales (科罗拉多高原的寒漠，犹他州)^[14]	Actinobacteria，Proteobacteria， Firmicutes， Chloroflexi， Acidobacteria，Cyanobacteria， Gemmatimonadetes， Bacteroidetes (腾格里沙漠)^[45]; Cyanobacteria， Bacteroidetes， Proteobacteria， unclassified bacteria， Actinobacteria (古尔班通古特沙漠)^[46]; Leptolyngbya (Cyanobacteria)， Rubrobacter， Solirubrobacter， Geodermatophilus (Actinomycetes) (塔韦纳斯沙漠)^[39]	Rhizobiales (phylum Proteobacteria)， Chroococcales， Nostocophycideae (Cyanobacteria) (西澳)^[47]; Cyanobacteria， Alphaproteobacteria， Bacteroidetes， Chloroflexi， Actinobacteria， Acidobacteria (阿曼)^[29]; Cyanobacteria， Proteobacteria， Bacteroidetes， Acidobacteria (莫哈韦沙漠)^[48]
地衣结皮	Actinobacteria， Proteobacteria, Acidobacteria， Firmicutes， Armatimonadetes Cyanobacteria， Bacteroidetes和Planctomycetes（南极洲大陆维多利亚岛)^[49]	Proteobacteria， Actinobacteria， Chloroflexi， Cyanobacteria， Bacteroidetes， Planctomycetes， Acidobacteria， Armatimonadetes (腾格里沙漠)^[50]; Cyanobacteria， Proteobacteria， Bacteroidetes， unclassified bacteria， Acidobacteria， Actinobacteria， Verrucomicrobia (古尔班通古特沙漠)^[46]; Proteobacteria， Actinobacteria， Bacteroidetes， Acidobacteria， Cyanobacteria and Verrucomicrobia (西班牙塔韦纳斯沙漠)^[31]	Acidobacteriales (phylum Acidobacteria)， Rhodospirillales (Proteobacteria)， Actinomycetales (Actinobacteria)和unclassified Proteobacteria (西澳)^[47]; Cyanobacteria， Bacteroidetes， Acidobacteria， Alphaproteobacteria， Actinobacteria， Chloroflexi (阿曼)^[29]
藻-地衣结皮	—	Nostocales， Chroococcidiopsaceae， Coleofasciculaceae (Cyanobacteria) (塔韦纳斯沙漠)^[39]	Cyanobacteria (Scytonema and Microcoleus steenstrupii)， Proteobacteria (β-Proteobacteria)， Actinobacteria， Bacteriodetes，和Chloroflexi (索诺拉沙漠)^[27]; Cyanobacteria， Proteobacteria， Bacteroidetes， Chloroflexi， Armatimonadetes (莫哈韦沙漠)^[48]; Bacteroidetes， Proteobacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Chloroflexi， Verrucomicrobia和Planctomycetes (非洲的卡鲁沙漠)^[32]
地衣-藓结皮	Cyanobacteria， Bacteroidetes， Proteobacteria， Actinobacteria， Chloroflexi， Acidobacteria (美国峡谷国家公园)^[51];Actinobacteria， Bacteroidetes， Proteobacteria， Cyanobacteria， Firmicutes， Verrucomicrobia， Acidobacteria， Chloroflexi (美国西部山间寒漠)^[52]; Cyanobacteria， Actinobacteria， Bacteriodetes， Chloroflexi和Proteobacteria (科罗拉多高原的寒漠)^[53]	Actinobacteria， Proteobacteria， Chloroflexi， Cyanobacteria， Acidobacteria， Firmicutes， Bacteroidetes (腾格里沙漠)^[45]	Cyanobacteria， Proteobacteria， Bacteroidetes， Actinobacteria和Acidobacteria (奇瓦瓦沙漠的Actopan和Atexcac地区)^[54]
藓结皮	—	Proteobacteria， Bacteroidetes， unclassified bacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Verrucomicrobia (古尔班通古特沙漠)^[46]; Bacteroidetes， Acidobacteria和Proteobacteria (新墨西哥桑迪亚山脉)^[34]; Acidobacteria， Proteobacteria， Chloroflexi， Actinobacteria (黄土高原)^[33]	Proteobacteria， Actinobacteria， Acidobacteria， Chloroflexi， Bacteroidetes， Cyanobacteria， Deinococcus-Thermus， Planctomycetes (莫哈韦沙漠)^[48]; Bacteroidetes， Proteobacteria， Actinobacteria， Cyanobacteria， Acidobacteria， Chloroflexi， Verrucomicrobia和Planctomycetes (非洲卡鲁沙漠)^[32]

演替阶段	气候区
演替阶段	寒漠	冷漠	热漠
藻结皮	Ascomycota， Sordariomycetes， Pezizomycetes和Basidiomycota (科罗拉多高原的寒漠，犹他州)^[61-62]	Ascomycota， unclassified_Fungi， Basidiomycota Chytridiomycota和Zygomycota (腾格里沙漠)^[45]; Dothideomycetes， Eurotiomycetes， Lecanoromycetes (古尔班通古特沙漠)^[37]	Ascomycota (Dothideomycetes， Eurotiomycetes)， Basidiomycota和Chytridiomycota (阿曼)^[29，68]; Ascomycota (莫哈韦沙漠)^[48]
地衣结皮	Ascomycota， Pezizomycetes， Basidiomycota， Sordariomycetes， Tremellomycetes和Leotiomycetes (科罗拉多高原的寒漠，犹他州)^[61]	Ascomycota， no_rank_Fungi， unclassified_Fungi， Basidiomycota， Chytridiomycota和Zygomycota (腾格里沙漠)^[50]; Eurotiomycetes， Lecanoromycetes和unidentified Ascomycota (古尔班通古特沙漠)^[37]	Ascomycota (Dothideomycetes， Eurotiomycetes)， Basidiomycota和Chytridiomycota (阿曼)^[29，68]; Ascomycota， Pezizomycetes， Basidiomycota和Eurotiomycetes (奇瓦瓦沙漠)^[61]; Sordariomycetes， Ascomycota， Lichinomycetes， Pezizomycetes和Basidiomycota (索诺拉沙漠)^[61]
藻-地衣结皮	—	—	Ascomycota， Sordariomycetes， Pezizomycetes和Eurotiomycetes (奇瓦瓦沙漠)^[61]
藓结皮	—	Ascomycota， Basidiomycota， unclassified_Fungi， Chytridiomycota和Zygomycota (腾格里沙漠)^[45]; Penicillium和Alternaria(可培养真菌)， Unidentified Ascomycota， Eurotiomycetes和Dothideomycetes (不可培养真菌) (古尔班通古特沙漠)^{[37， 69]}; Ascomycota和Basidiomycota (黄土高原)^[33]	Ascomycota (莫哈韦沙漠)^[48]