
西北沙区梭梭根系深色有隔内生真菌等微生物时空分布及对根际土壤环境的响应
赵昕,高慧利,龙俊萌,刘燕霞,李夏,贺学礼
菌物学报 ›› 2021, Vol. 40 ›› Issue (10) : 2716-2734.
西北沙区梭梭根系深色有隔内生真菌等微生物时空分布及对根际土壤环境的响应
Spatial and temporal distribution and the response to rhizosphere soil environment of dark septate endophyte and other microorganisms in roots of Haloxylon ammodendron in sand area of Northwest China
为揭示西北沙区不同时空尺度梭梭Haloxylon ammodendron根系深色有隔内生真菌(dark septate endophyte,DSE)分布规律及其环境驱动机制,分别于2019年7月、9月和12月在安西极旱荒漠国家级自然保护区和民勤连古城国家级自然保护区采集根系和土壤样品,根据真菌形态学特征、组织化学等方法,研究了梭梭根系DSE真菌定殖规律、定殖特征及其与土壤环境因子间的相互关系。结果表明,DSE真菌在自然生境梭梭根系广泛定殖,形成具有时空异质性的典型暗色有隔菌丝和微菌核结构。梭梭根系DSE真菌定殖受季节和空间分布影响,其中总定殖率在生长旺季7月最高(85.6%),且菌丝隔间距显著大于9、12月;安西样地菌丝定殖率(71.5%)和总定殖率(85.6%)显著高于民勤样地(40.7%,60.0%),但民勤样地微菌核直径显著高于安西样地。相关性分析表明,DSE定殖率及形态特征受土壤理化性质影响,温度、有效磷是影响DSE真菌定殖率的主要土壤因子,湿度和碱解氮是影响DSE真菌定殖形态的主要土壤因子。相比空白土壤,梭梭根际革兰氏阳性细菌含量、土壤脲酶活性以及安西样地有效磷含量均更高,预示其对荒漠土壤具有改善作用。本研究对梭梭DSE真菌时空分布和影响因素的系统性调查有助于为荒漠化治理和生态恢复提供依据。
The distribution pattern and driving factors of dark septate endophyte (DSE) in roots of Haloxylon ammodendron across spatial and temporal scales in Northwest China were investigated. Soil and plant root samples were collected from Anxi Extreme-arid Desert National Nature Reserve and Minqin Liangucheng National Nature Reserve in July, September and December in 2019. Based on morphology and histochemistry assay, the relationship between the colonization characteristics of DSE fungi in the roots of H. ammodendron and soil factors were observed. The investigation showed that DSE fungi colonized extensively in the roots of H. ammodendron in natural desert habitats, forming typical dark and septate hyphae and microsclerotia with spatial and temporal heterogeneity. The DSE colonization pattern of H. ammodendron roots was affected by seasonal and spatial variability, and the total colonization rate was the highest in vigorous growth period in July (85.6%), and the hyphal septum interval was significantly distant in July than in September and December. The hyphal colonization rate (71.5%) and total colonization rate (85.6%) in Anxi were significantly higher than that in Minqin (40.7%, 60.0%), but the microsclerotium diameter was significantly increased in Minqin as compared with that in Anxi. The correlation analysis indicated that DSE colonization rate and morphological characteristics were affected by soil physical and chemical factors. Temperature and available phosphorus of soil were the main factors affecting DSE colonization rate, while humidity and alkali-hydrolyzed nitrogen were the main soil factors affecting DSE colonization characteristics. Compared with the blank soil, the content of Gram positive bacteria, the activity of soil urease and the available phosphorus content (in Anxi sample plots) in the rhizosphere soil of H. ammodendron were significantly higher, indicating that H. ammodendron could improve the desert soil. This study provides helpful reference material for desertification control and ecological restoration in dry areas of Northwest China.
梭梭 / 深色有隔内生真菌 / 荒漠 / 时空分布 / 环境因子 {{custom_keyword}} /
Haloxylon ammodendron / dark septate endophyte / desert / spatial and temporal distribution / environmental factors {{custom_keyword}} /
图3 剑孢新赤壳 (HMAS 290889)A-C:自然基物上的子囊壳;D,E:25 ℃培养7 d的菌落形态 (D:PDA;E:SNA);F:子囊壳纵切面结构;G-I:子囊及子囊孢子;J-L:子囊孢子;M,N:分生孢子梗和小型分生孢子;O:小型分生孢子;P-S:大型分生孢子. 标尺:A-C=1 mm;F=50 μm;G-S=10 μm Fig. 3 Neocosmospora protoensiformis (HMAS 290889). A-C: Ascomata on natural substratum; D, E: Colonies after 7 d at 25 °C (D: PDA; E: SNA); F: Median section of an ascoma; G-I: Asci with ascospores; J-L: Ascospores; M, N: Conidiophores and microconidia; O: Microconidia; P-S: Macroconidia. Bars: A-C=1 mm; F=50 μm; G-S=10 μm. |
图5 瘤顶赤壳 (HMAS 290890)A-C:自然基物上的子囊壳;D,E:25 ℃培养14 d的菌落形态 (D:PDA;E:SNA);F:子囊壳纵切面结构;G-K:分生孢子梗和分生孢子;L:厚垣孢子. 标尺:A-C=1 mm;F=50 μm;G-L=10 μm Fig. 5 Tumenectria laetidisca (HMAS 290890). A-C: Ascomata on natural substratum; D, E: Colonies after 14 d at 25 °C (D: PDA; E: SNA); F: Median section of an ascoma; G-K: Conidiophores and conidia; L: Chlamydospores. Bars: A-C=1 mm; F=50 μm; G-L=10 μm. |
图6 基于ITS和LSU序列的MP树粗体显示5个中国新记录种的系统发育位置,MPBP大于50% (左)、BIPP大于90% (右)标注于分支节点上 Fig. 6 Maximum parsimony phylogram reconstructed from the combined sequences of ITS and LSU. the phylogenetic position of the five Nectriaceae species new to China. MPBP above 50% (left) showing and BIPP above 90% (right) are given respectively. |
[1] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[2] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[3] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[4] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[5] |
Native grasses of semi-arid rangelands of the southwestern USA are more extensively colonized by dark septate endophytes (DSE) than by traditional mycorrhizal fungi. Roots of dominant grasses ( Bouteloua sp.) native to arid southwestern USA rangelands were prepared and stained using stains specific for fungi (trypan blue) and for lipids (sudan IV). This revealed extensive internal colonization of physiologically active roots by atypical fungal structures that appear to function as protoplasts, without a distinguishable wall or with very thin hyaline walls that escape detection by methods staining specifically for fungal chitin. These structures were presumed to be active fungal stages that progressed to form stained or melanized septate hyphae and microsclerotia characteristic of DSE fungi within dormant roots. The most conspicuous characteristic of these fungi were the unique associations that formed within sieve elements and the accumulation of massive quantities of lipids. This interface suggests a biologically significant location for carbon transfer between the plant and fungus. The continuous intimate association with all sieve elements, cortical and epidermal cells as well as external extension on the root surface and into the soil indicates that they are systemic and considerably more prevalent than previously thought. A fungal network associated with a mucilaginous complex observed on the root surface and its potential role in root function in dry soil is discussed. It is suggested that those fungi that non-pathogenically and totally colonize plant cells be classed as systemic endophytic fungi (SEF). This would refine the broad designation of DSE fungi. The potential mutualistic benefit of SEF for native plants in arid ecosystems based on the extent of lipid accumulation and its apparent distribution is discussed.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[6] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[7] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[8] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[9] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[10] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[11] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[12] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[13] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[14] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[15] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[16] |
We studied seasonal variation in population attributes of arbuscular mycorrhizal (AM) fungi over 2 years in four sites of temperate grasslands of the Argentinean Flooding Pampas. The sites represent a wide range of soil conditions, hydrologic gradients, and floristic composition. Lotus glaber, a perennial herbaceous legume naturalised in the Flooding Pampas, was dominant at the four plant community sites. Its roots were highly colonised by AM fungi. Temporal variations in spore density, spore type, AM root colonisation, floristic composition and soil chemical characteristics occurred in each site and were different among sites. The duration of flooding had no effect on spore density but depressed AM root colonisation. Eleven different types of spores were recognized and four were identified. Two species dominated at the four sites: Glomus fasciculatum and Glomus intraradices. Spore density was highest in summer (dry season) and lowest in winter (wet season) with intermediate values in autumn and spring. Colonisation of L. glaber roots was highest in summer or spring and lowest in winter or autumn. The relative density of G. fasciculatum and G. intraradices versus Glomus sp. and Acaulospora sp. had distinctive seasonal peaks. These seasonal peaks occurred at all four sites, suggesting differences among AM fungus species with respect to the seasonality of sporulation. Spore density and AM root colonisation when measured at any one time were poorly related to each other. However, spore density was significantly correlated with root colonisation 3 months before, suggesting that high colonisation in one season precedes high sporulation in the next season.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[17] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[18] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[19] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[20] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[21] |
Dark septate root endophytes (DSE) are conidial or sterile fungi (Deuteromycotina, Fungi Imperfecti) likely to be ascomycetous and colonizing plant roots. They have been reported for nearly 600 plant species representing about 320 genera and 100 families. DSE fungi occur from the tropics to arctic and alpine habitats and comprise a heterogeneous group that functionally and ecologically overlaps with soil fungi, saprotrophic rhizoplane-inhabiting fungi, obligately and facultatively pathogenic fungi and mycorrhizal fungi. Numerous species of undescribed sterile and anamorphic taxa may also await discovery. Although DSE are abundant in washed root and soil samples from various habitats, and are easily isolated from surface-sterilized roots of ecto-, ectendo-, endo- and non-mycorrhizal host species, their ecological functions are little understood. Studies of DSE thus far have yielded inconsistent results and only poorly illustrate the role of DSE in their natural habitats. These inconsistencies are largely due to the uncertain taxonomic affinities of the strains of DSE used. In addition, because different strains of a single anamorph taxon seem to vary greatly in function, no clear generalizations on their ecological role have been drawn. This paper reviews the current literature on DSE and the ecology and discusses the need for and direction of future research.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[22] |
Arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungi are ubiquitous in grass roots, but their colonizations may vary according to latitudinal gradient and site conditions. We investigated how vegetation zone (boreal vs. subarctic), humus thickness, and site openness affect root fungal colonizations of the grass Avenella flexuosa. More precisely, we hypothesized that AM and DSE fungal colonizations would have different responses to environmental conditions such that AM fungi could be more common in boreal zone, whereas we expected DSE fungi to be more affected by the amount of humus. We found site openness to affect AM and DSE fungi in a contrasting manner, in interaction with the vegetation zone. AM colonization was high at open coastal dunes, whereas DSE fungi were more common at forested sites, in the boreal zone. Humus thickness affected AM fungi negatively and DSE fungi positively. To conclude, the observed AM and DSE fungal colonization patterns were largely contrasting. AM fungi were favored in seashore conditions characterized by thin humus layer, whereas DSE fungi were favored in conditions of higher humus availability.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[23] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[24] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[25] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[26] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[27] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[28] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[29] |
The ecology of weed plants includes their interactions with soil microorganisms, such as mutualistic partners that may contribute to their adaptation and competitive success in the agricultural fields. Despite the importance of microorganisms to plant growth, knowledge on weed-symbiont associations is still incipient compared to crops. Thus, a survey for the presence of arbuscular mycorrhiza (AM) and dark septate endophyte (DSE) associations in the roots of 50 weed species was done in three distinct areas during the dry and rainy seasons. We found that 41 and 29 out of the 50 species were associated with AM fungi and DSE, respectively, and 27 species presented both associations. All the plant species not forming AM belong to families thought to be nonmycorrhizal, such as Amaranthaceae, Commelinaceae, Brassicaceae, and Cyperaceae. The most common morphotype of AM observed was the Arum-type. No significant differences were found in root length colonization between the areas or seasons. For 19 species surveyed, this is the first report on their mycorrhizal status.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[30] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[31] |
The broad distribution and high colonization rates of plant roots by a variety of endophytic fungi suggest that these symbionts have an important role in the function of ecosystems. Semiarid and arid lands cover more than one-third of the terrestrial ecosystems on Earth. However, a limited number of studies have been conducted to characterize root-associated fungal communities in semiarid grasslands. We conducted a study of the fungal community associated with the roots of a dominant grass, Bouteloua gracilis, at the Sevilleta National Wildlife Refuge in New Mexico. Internal transcribed spacer ribosomal DNA sequences from roots collected in May 2005, October 2005, and January 2006 were amplified using fungal-specific primers, and a total of 630 sequences were obtained, 69% of which were novel (less than 97% similarity with respect to sequences in the NCBI database). B. gracilis roots were colonized by at least 10 different orders, including endophytic, coprophilous, mycorrhizal, saprophytic, and plant pathogenic fungi. A total of 51 operational taxonomic units (OTUs) were found, and diversity estimators did not show saturation. Despite the high diversity found within B. gracilis roots, the root-associated fungal community is dominated by a novel group of dark septate fungi (DSF) within the order Pleosporales. Microscopic analysis confirmed that B. gracilis roots are highly colonized by DSF. Other common orders colonizing the roots included Sordariales, Xylariales, and Agaricales. By contributing to drought tolerance and nutrient acquisition, DSF may be integral to the function of arid ecosystems.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[32] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[33] |
Arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungal colonization of Alchemilla glomerulans, Carex vaginata, Ranunculus acris ssp. pumilus and Trollius europaeus growing in low-alpine meadows in the Finnish subarctic were studied at different times during the growing season. Fungal colonization was correlated to soil soluble phosphorus (P) concentration. The influence of flower bud removal on fungal colonization was investigated in A. glomerulans, C. vaginata and R. acris and the correlation between AM and DSE colonization was studied. The fungal colonization patterns were found to be species-specific. R. acris maintained a relatively high rate of fungal colonization throughout the summer, while the rates of colonization of T. europaeus were lower and decreased towards the end of the season. A. glomerulans had constant arbuscular and vesicular colonization throughout the summer, but hyphal and DSE colonization declined towards the end of the season. C. vaginata did not form arbuscular mycorrhiza, but was colonized by DSE fungi and hyaline septate hyphae throughout the season. The soil soluble P concentration showed some seasonal variation, but was also highly variable between the study sites. Bud removal decreased arbuscular colonization of R. acris, but no unique effects were seen in any other parameters or the other species studied. The root fungal parameters correlated with soil P in some species at some sites, but no consistent trend was found. DSE colonization was positively correlated with root vesicular and hyphal colonization in some cases. The differences in fungal colonization parameters may be related to species-specific phenologies.
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[34] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[35] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[36] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[37] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[38] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[39] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[40] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[41] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[42] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[43] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[44] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[45] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[46] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[47] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[48] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[49] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[50] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[51] |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[52] |
毕银丽, 解琳琳, 2021. 丛枝菌根真菌与深色有隔内生真菌生态修复功能与作用. 微生物学报, 61(1):58-67
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[53] |
曹艳峰, 李彦, 李晨华, 吕光辉, 2016. 荒漠灌木梭梭(Haloxylon ammodendron)周围土壤微生物的空间分布. 生态学报, 36(6):1628-1635
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[54] |
贺超, 陈晓玉, 王文全, 侯俊玲, 2020. 西北旱区甘草深色有隔内生真菌物种多样性及空间分布. 菌物学报, 39(8):1487-1501
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[55] |
李丽娟, 谢婷婷, 张松林, 袁中勋, 刘明辉, 李昌晓, 2020. 三峡库区消落带4种适生植物根际与非根际土壤养分与酶活性特征研究. 生态学报, 40(21):7611-7620
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[56] |
徐高兴, 徐先英, 王立, 付贵全, 赵鹏, 丁爱强, 2019. 梭梭不同密度与配置固沙效果风洞模拟试验. 干旱区资源与环境, 33(9):189-195
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[57] |
姚丹, 牛舒琪, 赵祺, 曹靖, 韩庆庆, 李慧萍, 缑晶毅, 张金林, 2020. 梭梭根际枯草芽孢杆菌WM13-24调控黑麦草耐盐性的研究. 生态学报, 40(20):1-17
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[58] |
张海涵, 2011. 黄土高原枸杞根际微生态特征及其共生真菌调控宿主生长与耐旱响应机制. 西北农林科技大学博士论文,西安. 87-101
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[59] |
张雪, 郭亚楠, 李烨东, 王文彬, 郑盈盈, 赵丽莉, 贺学礼, 2019. 西北荒漠带细枝岩黄耆AM真菌与DSE真菌时空分布研究. 菌物学报, 38(11):2030-2042
{{custom_citation.content}}
{{custom_citation.annotation}}
|
[60] |
赵业婷, 李志鹏, 常庆瑞, 2013. 关中盆地县域农田土壤碱解氮空间分异及变化研究. 自然资源学报, 28(6):1030-1038
{{custom_citation.content}}
{{custom_citation.annotation}}
|
{{custom_ref.label}} |
{{custom_citation.content}}
{{custom_citation.annotation}}
|
/
〈 |
|
〉 |