生物土壤结皮对荒漠土壤种子库和种子萌发的影响

研究了腾格里沙漠东南缘在不同自然条件（风、温度、水分）下,人工固沙植被区（24龄、41龄、50龄）和相邻天然植被区的两种生物土壤结皮对荒漠土壤种子库和种子萌发的影响。结果表明,荒漠土壤种子库在苔藓结皮上的储量显著高于藻类结皮。随着生物土壤结皮的发育,种子萌发量在苔藓结皮上增加,在藻类结皮上减少。生物土壤结皮层的含水量对种子萌发有显著的影响（p〈0.05）,植物种子在湿润处理的生物土壤结皮上的萌发量高于干燥处理的生物土壤结皮上的种子萌发量。生物土壤结皮表层温度和亚表层温度对荒漠植物种子萌发无显著影响（p〉0.05）,但总体而言,对于苔藓结皮,植物种子在较高温度下的萌发量略高于在较低温度下的萌发量,而对于藻类结皮,植物种子在较低温度下的萌发量略高。     

生物结皮对5种不同形态的荒漠植物种子萌发的影响

生物结皮广泛分布于干旱、半干旱区, 强烈影响着土壤表层理化特性, 进而对种子散布、萌发和定居产生影响。目前关于生物结皮与植物种子萌发关系的研究结论存在争议。该文通过室内人工控制实验, 研究了生物结皮对古尔班通古特沙漠5种具不同种子形态特征的荒漠植物白梭梭(Haloxylon persicum)、蛇麻黄(Ephedra distachya)、角果藜(Ceratocarpus arenaarius)、涩芥(Malcolmia africana)和狭果鹤虱(Lappula semiglabra)的种子萌发的影响。结果表明, 在干燥和湿润两种条件下, 生物结皮对不同形态植物种子萌发均具有不同的作用。在干燥条件下, 生物结皮显著抑制了角果藜和涩芥种子的萌发(p<0.05), 对其它3种植物无显著影响; 而湿润条件下, 生物结皮显著抑制了白梭梭、角果藜和狭果鹤虱种子的萌发(p<0.05), 对蛇麻黄、涩芥则无显著影响。     

Effects of mosaic biological soil crusts on vascular plant establishment in a coastal saline land of the Yellow River Delta,China

黄河三角洲滨海盐碱地镶嵌式分布的生物土壤结皮对维管植物建植的影响 滨海盐碱地生物土壤结皮对维管植物影响研究,鲜见报道。本研究旨在探讨黄河三角洲滨海盐碱 地镶嵌式分布的生物土壤结皮是否以及如何影响两种典型草本植物的种子萌发,以深入理解基质异质性对 植物群落动态的影响。通过室内萌发实验,研究裸斑、芦苇(Phragmites australis)、碱蓬(Suaeda glauca) 和柽柳(Tamarix chinensis)生境下发育的生物土壤结皮和无结皮表土对多年生草本植物芦苇和一年生草本植物碱蓬种子的萌发率和平均萌发时间的影响,并从基质理化性质角度探讨其对种子萌发的影响机制。研究结果发现,与无结皮表土相比,生物土壤结皮提高了含水量、养分积累和大部分盐离子浓度,但降低了 土壤pH值。苔藓生物结皮可直接降低土壤pH值和Mg²⁺浓度,间接提高碱蓬种子的萌发率。较低的土壤pH值也间接降低了芦苇种子在其自身生境中的萌发速度。与有植被生长的生境相比,裸斑生境加剧Cl⁻的积累,间接降低了芦苇种子的萌发速度。结果表明,芦苇生境分布的苔藓结皮为碱蓬种子的萌发提供了一个机会窗口。生物土壤结皮类型和生境类型的共同作用可通过影响维管植物种子的萌发和建植,最终影响植物群落结构。     

毛乌素沙地生物土壤结皮对油蒿水分吸收的影响模拟

植物可利用水分是决定沙生灌木生长的主要因子,生物土壤结皮(简称生物结皮)在降雨期影响降水入渗,而在干旱期改变土壤蒸发,从而影响土壤水分分布,最终可能影响灌木水分吸收。然而,关于不同降水条件下生物结皮对灌木水分吸收和水分胁迫的影响机制认识不清。以油蒿为研究对象,基于试验数据和1990—2019年气象数据,采用数学模拟,定量研究了毛乌素沙地不同降水条件下生物结皮对土壤水分分布和油蒿水分吸收的影响,评价干旱期生物结皮对油蒿水分胁迫的影响。结果表明：与无结皮处理相比,生物结皮处理的土壤蒸发降低了5.1%;生物结皮改善了干旱期的土壤水分条件;生物结皮降低了植物水分胁迫的比例,平均降低比例为8.1%;生物结皮提高了植物水分吸收,平均增加比例为12.8%;生物结皮和对照植物水分吸收的比值随季节降水量的增加而降低,均值为1.13。综上,生物结皮的出现并未消极地影响沙生灌木的水分吸收。研究结果有助于理解生物结皮与灌木的共生或竞争关系。     

干旱沙区生物土壤结皮发育对红砂形态及干物质积累的影响

为了探讨干旱沙区生物土壤结皮发育对红砂形态及干物质积累的影响,以巴丹吉林沙漠南缘已发育不同类型生物土壤结皮并有红砂种群成功定居的区域为研究场所,通过野外监测与室内测定的方法对藻类-地衣、地衣、地衣-藓类结皮上的红砂（当年生、幼株、成株）形态特征及生物量进行了调查研究。结果表明：（1）藻类-地衣结皮演替到地衣-藓类结皮的过程中,当年生红砂形态差异不显著,但5 a以上植株基部分枝长、树冠/侧冠投影面积、主根长均显著减小;同时,地衣-藓类结皮的3-5 a植株基部分枝数明显减少,且5 a以上植株明显矮化。（2）生物土壤结皮发育不仅降低了红砂幼株或成株生物量积累能力,还减小了植株根冠比,且降低/减小程度随结皮演替或株龄的增大逐渐增大。（3）红砂形态、生物量指标与物理或藻类结皮面积百分比呈极显著正相关关系,与藓类结皮面积百分比呈极显著的负相关关系。因此认为,生物土壤结皮的演替导致土壤关键生态因子（如土壤水分）发生变化,进而影响红砂植株生长发育能力,从而使得不同发育阶段结皮上的同龄红砂形态特征及生物量存在差异性。     

古尔班通古特沙漠生物结皮不同发育阶段中藻类的变化

通过在古尔班通古特沙漠南缘相同的地貌部位,选择裸沙、藻结皮、地衣结皮和苔藓结皮4种不同演替阶段中的生物结皮,研究了藻类的种类组成、优势种和生物量的变化.结果表明:(1)在结皮的不同演替阶段,藻类种类组成不同,其常见物种有一定的差异,如裸沙中藻类常见种是脆杆藻2(Fragilaria sp.2)、威利颤藻(Oscillatoria willei)和奥克席藻(Phormidium okenii),藻结皮的常见种是小聚球藻(Synechococcus parvus)、颗粒常丝藻 (Tychonema granulatum)、韧氏席藻(Phormidium retzli);同时在不同发育阶段亦存在一些特有种.(2)在裸沙发育到成熟生物结皮的过程中,藻类的优势物种也发生相应的变化.裸沙、藻结皮、地衣结皮和苔藓结皮的优势种分别是脆杆藻1(Fragilaria sp.1)、具鞘微鞘藻(Microcoleus vaginatus)、具鞘微鞘藻、眼点伪枝藻(Scytonema ocellatum)或集球藻(Palmellococcus miniatus).(3)藻类生物量在生物结皮不同演替阶段差异极显著(P<0.01),在裸沙中藻类生物量最低,随着生物结皮的逐渐发育,藻类生物量明显升高,地衣结皮最高,约是裸沙的8.3倍,当发育至苔藓结皮时,藻类生物量又有所下降.(4)在裸沙中基本为松散的沙粒,随着生物结皮的演替,丝状种类占明显的优势,尤其是具鞘微鞘藻,另外真菌菌丝和苔藓假根分别在地衣结皮和苔藓结皮中起着重要作用.     

荒漠生物结皮中藻类和苔藓植物研究进展

藻类和苔藓植物是荒漠植被演替过程中常见的两类先锋植物, 同时也是生物结皮中生物量最大的2个类群。该文综述了近年来干旱半干旱荒漠地区生物结皮中藻类和苔藓两大类植物区系及其生态作用的研究进展, 重点介绍藻类结皮、苔藓结皮的生态作用以及二者之间存在的生态学关系。在此基础上对荒漠生物结皮中藻类与苔藓植物的研究前景进行了展望, 指出荒漠生物结皮中藻类与苔藓共生机理的探讨是未来的研究重点, 这对进一步探明生物结皮中藻类和苔藓植物之间的相互作用, 揭示它们的生态学关系具有重要的理论意义和实践价值。     

荒漠生物结皮中藻类和苔藓植物研究进展

藻类和苔藓植物是荒漠植被演替过程中常见的两类先锋植物,同时也是生物结皮中生物量最大的2个类群。该文综述了近年来干旱半干旱荒漠地区生物结皮中藻类和苔藓两大类植物区系及其生态作用的研究进展,重点介绍藻类结皮、苔藓结皮的生态作用以及二者之间存在的生态学关系。在此基础上对荒漠生物结皮中藻类与苔藓植物的研究前景进行了展望,指出荒漠生物结皮中藻类与苔藓共生机理的探讨是未来的研究重点,这对进一步探明生物结皮中藻类和苔藓植物之间的相互作用,揭示它们的生态学关系具有重要的理论意义和实践价值。     

黄土丘陵区生物土壤结皮层水稳性

采用改进的土壤水稳性团聚体数量测定方法,研究了黄土丘陵区不同组成和生物量的生物结皮层水稳定性.结果表明：生物结皮层的水稳定性与其生物组成有关,苔藓结皮的水稳定性显著高于藻结皮,震荡390次后,苔藓结皮的厚度和质量损失率仅分别是藻结皮损失率的47.3%和40.1%;生物结皮层水稳定性与生物结皮的生物组成有关,60%以上苔藓覆盖度的生物结皮的稳定性最高,质量和厚度损失率分别是藻结皮（无苔藓覆盖）损失率的28.6%和22.7%;生物结皮层水平方向结构水稳定性显著大于垂直方向,震荡390次后,苔藓结皮的面积损失率仅为厚度损失率的6.4%.试验分析证实,生物结皮层是一种水平方向稳定性极强的层状结构体,这一结构特性增强了其抗风蚀和水蚀的能力.     

生物结皮粗糙特征——以古尔班通古特沙漠为例

摘要：空气动力粗糙度可以反映地表气流与下垫面的相互作用。古尔班通古特沙漠是我国最大的固定、半固定沙漠,其间广泛分布的生物结皮在稳定地表和改善环境方面意义重大。对未经扰动的4种类型生物结皮进行表面微形态观察,并通过风洞实验确定动力粗糙度Z0和摩阻风速u*,结果表明：（1）不同生物结皮类型,其组成和表面微形态等都具有明显差异。藻结皮以表面致密光滑为显著特征,由藻类分泌物和藻丝体粘结细粒物质所形成;地衣结皮表面藻类和真菌形成的叶状体匍匐沙面生长,呈现三维生长方式,形成有明显凹凸的壳状覆被;苔藓结皮以苔藓植物体密集丛生为特点,地上部分出现了茎叶分化,有一定的柔韧性。（2）就动力粗糙度的大小而言,是按地衣结皮>藻类－地衣结皮>苔藓结皮>藻结皮的顺序排列的,Z0平均值依次为（6.5890.850）mm、（4.1790.239）mm、（2.5420.357）mm和（0.3930.220）mm,与定床裸沙面的（0.0420.019）mm相比,生物结皮Z0值提高了10—150倍。随着风速的增大Z0值有所减小,其中以地衣结皮的减小趋势较为明显。（3）由风速廓线对比发现,四类生物结皮对气流阻滞作用的差异主要局限于4 cm以下的高度范围,风速越大这种差异也越大。各类生物结皮摩阻风速u*随风速增大而增大,其中藻结皮的增大速率明显低于其它三类结皮,说明藻结皮随风速增大的阻滞效应较其它三类结皮要差。（4）在净风条件下,地衣结皮具有最好的防风效果,其次为藻类－地衣结皮和苔藓结皮,藻结皮最差。当生物结皮破损后,床面结构和气流性质将发生变化,对空气动力学粗糙度和摩阻风速产生的影响将有待于进行更深入的研究。     

Effects of biological soil crusts on emergence of desert vascular plants in North China

Biological soil crusts are a universal and common feature in arid and semi-arid regions and their appearance profoundly affects soil surface properties which may greatly change the seed dispersal, germination and establishment. To date, only a handful of experiments have exerted to investigate the effects of crusts on vascular plants and the conclusions from these studies are variable. In this study, we investigate the influences of two different crusts universally spreading in southeastern part of the Tengger Desert with four chronosequences (24, 41, 50 years old in artificial vegetation area and natural vegetation crusts) on vascular plants. Crusts were placed at three different sites to simulate different environmental factors (wind velocity and soil crust moisture), we set two soil moisture regimes for crusts to investigate how vascular plants respond under different moisture regimes in crusts. Emergence densities of vascular plants were significantly higher in moss crust than in algae crust. With the development of crusts, seed emergence increased in moss crust while decreased in algae crust. As for effects of moisture, our results showed that soil moisture had a significant effect on seed emergence in both types of crusts at all developing phases. Crusts with higher moisture had more seedlings than those with lower moisture. The above results indicated that the appearance of crusts changed the surface soil properties, which had greatly influenced the entrapment and lodgement of seeds in the study area, thus subsequently influence seed emergence through affecting natural factors.     

Effects of Biological Soil Crusts on Seed Bank, Germination and Establishment of Two Annual Plant Species in the Tengger Desert (N China)

The presence of biological soil crusts can affect the germination and survival of vascular plants, but the reasons are not well investigated. We have conducted a field investigation and greenhouse experiments to test the effect of crusts on two desert annual plants, which occur on the stabilized dunes of the Tengger Desert in N China. The results showed that biological soil crusts negatively influenced the seed bank of Eragrostis poaeoides and Bassia dasyphylla. The important effect of biological soil crusts on seed germination and establishment were performed indirectly through reducing the amount of germinating seeds. Field investigation and experimental results with regard to the seed bank indicated that higher seedling density was found in disturbed crust soil and bare soil surface than in intact crust soils. Greenhouse experiments showed that the effects of biological soil crusts on germination and establishment of the two plants are not obvious in moist condition, while disturbed crusts are more favorable to seed germination in dry treatment. Significant differences in biomass were found between disturbed crust soil and bare soil. Survival and growth of the two annual plants were enhanced in both algal and moss crusts during the season of rainfall or in moist environment, but crusts did not affect seedling survival in the dry period. The small seeded E. poaeoides has higher germination than larger-seeded B. dasyphylla in crust soils, but B. dasyphylla has a relatively higher survival rate than E. poaeoides in crust soils.     

The Dynamics of Seed Bank on Shapotou Artificially Stabilized Dunes

There were 46 species belonging to 17 families in the soil seed bank on Shapotou artificially stabilized dunes. Compositae was the family with most species '(11 species) and the others were Chenopodiaceae (7 species), Gramineae (7 species), Leguminosae (4 species), etc. 33.3 % of the species lived in the community also had seeds in soil. The seed production of Artemisia ordosica, Hedysarum scoparium, Caragana korshinskii was 545.57 ± 676.73 grains/m2. 6.67 ±13.74 grains/m2. 0. 032±0. 024 grains/m2, respectively. The seeds of A. ordosica were not destroyed by pests, but 31.9%, 83.29% of the seeds of H. scopariurn. C. korshinskii respectively were destroyed by pests. Under the most suitable burying depth, the cumulative seed germination rate of A. ordosica, H. scoparium, C. korshinskii were 78. 3%, 66.7%. 85.0%, respectively. The rate of germination was in the order of: A. ordosica>H, scoparium>C, korshinskii. The vegetation succession would tend towards community mainly consisted of A. ordosica, Bassia dasyphylla, Eragrostis poaeoides, etc.     

Germination and seed water status of four grasses on moss-dominated biological soil crusts from arid lands

Biological soil crusts dominated by drought-tolerant mosses are commonly found through arid and semiarid steppe communities of the northern Great Basin of North America. We conducted growth chamber experiments to investigate the effects of these crusts on the germination of four grasses: Festuca idahoensis, Festuca ovina, Elymus wawawaiensis and Bromus tectorum. For each of these species, we recorded germination time courses on bare soil and two types of biological soil crusts; one composed predominantly of the tall moss Tortula ruralis and the other dominated by the short moss Bryum argenteum. On the short-moss crust, the final germination percentage was about half of that on bare soil. Also, the mean germination time was 4 days longer on short-mosses than on bare soil. In contrast to the short-moss crust, the tall-moss crust did not reduce the final germination percentage but increased the mean germination time. Similar results were observed in the four grasses studied. To investigate the mechanism by which moss crusts affected germination, we analyzed the water status of seeds on bare soil and moss crusts. Six days after seeding, the water content of seeds on bare soil was approximately twice that of seeds on tall- or short-moss crust. Analysis of the time course of changes in seed weight and water potential in Bromus tectorum revealed that overtime seeds on tall mosses reached higher water content than those on short mosses. The increase in the water content of seeds on tall mosses occurred as the seeds gradually fell through the moss canopy. Taken together, our results indicate that biological soil crusts with distinct structural characteristics can have different effects on seed germination. Furthermore, this study revealed that a biological soil crust dominated by short mosses had a negative effect on seed water status and significantly reduced seed germination.     

Seed characteristics and soil surface patch type interact to affect germination of semi-arid woodland species

Biological soil crusts are common in many arid and semi-arid regions and they can alter microenvironments which are likely to directly and indirectly influence vascular plant establishment. The effect of biological soil crusts on germination is also influenced by the biological characteristics of the seeds themselves but rarely have the effects of both crust type and seed morphology on germination been examined in the same study. In this study, seed of five semi-arid woodland species with contrasting seed morphology were sown on top of patch types that commonly occur in natural woodlands (foliose lichen, short-turf moss, tree leaf litter, disturbed crust) and their emergence was followed. Percent germination varied between patch types and, for the largest-seeded species (Maireana excavata), final germination was significantly lower on the biological soil crust and litter patch types because they strongly acted as a physical barrier to seed penetration into the soil. The remaining four species showed no significant differences in final percent germination with patch type because most seeds either completely or partially penetrated the surface layer. Germination time courses, however, showed that biological soil crusts delayed the timing of germination of these species. Hence, soil crusts might differentially affect the spatial patterning of species in semi-arid woodlands by their subtle influence on seedling emergence that is determined by differences in seed morphology and subsequent positioning within crusts.     

生物结皮影响下沙漠土壤表面凝结水的形成与变化特征

在水资源匮乏的沙漠生境,凝结水是植物、生物结皮、无脊椎和脊椎小动物的重要水分来源之一.采用微渗计法对比研究3种生物结皮类型(藻结皮、地衣结皮和苔藓结皮)和自然裸沙对地表凝结水量及凝结水蒸散过程的影响.微渗计的规格为内径6 cm,高3.5 cm的PVC管.研究结果表明:不同类型地表的总凝结水量之间存在极显著的差异(P < 0.01),总凝结水量随生物结皮发育水平呈显著增加的趋势,依次为:裸沙 < 藻结皮 < 地衣结皮 < 苔藓结皮,即生物结皮的存在有利于沙漠地表凝结水的形成.不同类型地表凝结水量的日均值有所差异.对于同一地表类型,凝结水量的最大值为最小值的数倍.黎明时,苔藓结皮的凝结水量最大,而裸沙的凝结水量最小,地衣结皮和藻结皮居中.凝结现象自20:00～22:00,次日8:00～9:00结束.大多数日出后凝结现象仍继续发生.不同类型地表的凝结及蒸散过程经历2个阶段:日出前凝结水量呈缓慢增加的趋势,日出后随温度的升高凝结水量快速减少,其中以苔藓结皮凝结水量下降最为迅速.凝结水量主要受温度、大气湿度、凝结面类型、气象条件和生境等方面因素的影响.     

荒漠结皮层藓类植物死亡对表层土壤水分蒸发和入渗的影响

水分是荒漠植物生长最主要的限制因子,藓类结皮作为荒漠土壤表层重要覆被物,对土壤水分蒸发入渗具有重要影响。研究表明,在全球气候变化背景下,不确定的降水格局变化导致结皮层藓类植物出现集群死亡现象,但这一过程对荒漠地表土壤水分蒸发与入渗过程的影响及其机理尚不清楚。以古尔班通古特沙漠齿肋赤藓结皮为研究对象,利用便携式渗透计和蒸发仪,研究了结皮层藓类植物死亡对土壤水分蒸发与入渗的影响。结果表明,与裸沙相比,藓类结皮的存在显著抑制了水分入渗,而藓类植物死亡的结皮层抑制作用最大,其初渗速率、稳渗速率和累积入渗量分别是活藓类结皮的39.89%、85.91%及64.48%,仅为裸沙的5.96%、13.13%及20.42%。在水分蒸发初期,裸沙的水分蒸发速率明显高于活藓类结皮和藓类植物死亡的结皮层,但藓类植物死亡的结皮层维持相对稳定的蒸发速率的时间长于裸沙和活藓类结皮,这也导致最终累计蒸发量以藓类植物死亡的结皮层最高、裸沙最低。可见,荒漠生物土壤结皮中藓类植物死亡会明显减少土壤水分入渗、增大水分蒸发,进一步影响荒漠表层土壤水分格局,从而影响生物土壤结皮与维管植物的水分利用关系。     

荒漠地表生物土壤结皮形成与演替特征概述

土壤表面结皮是世界范围内干旱沙漠地区土壤表面广泛存在的自然现象,包括物理结皮和生物土壤结皮两大类型。其中,生物土壤结皮作为干旱沙漠地区特殊环境的产物,是由细菌、真菌、蓝绿藻、地衣和苔藓植物与土壤形成的有机复合体。它的形成使土壤表面在物理、化学和生物学特性上均明显不同于松散沙土,具有较强的抗风蚀功能和重要的生态效应,成为干旱沙漠地区植被演替的重要基础。随着形成生物土壤结皮的物种更替,维持结皮结构的主要胶结方式亦随之发生变化,即由胞外多糖的粘结作用逐渐转变为蓝藻和荒漠藻的藻丝体、地衣菌丝体以及苔藓植物假根的缠绕和捆绑作用,物种更替是结皮微结构和胶结方式转化的生物基础。生物土壤结皮的形成通常可以分为以下几个阶段:生物土壤结皮的早期阶段(土壤酶和土壤微生物),藻结皮阶段、地衣结皮阶段和苔藓结皮阶段。即随着土壤微生物在沙土表面的生长,随后出现丝状蓝藻和荒漠藻类植物,形成以藻类植物为主体的荒漠藻结皮;当土壤表面得到一定固定后,便开始出现地衣和苔藓植物,形成以地衣和苔藓植物为优势的生物结皮类型。其中,前一阶段的完成又为后一阶段的开始提供良好的环境条件。当环境条件适宜时,生物土壤结皮也可以不经历其中某个阶段而直接发育到更高级的阶段。