Germination and seedling establishment of two annual grasses on lichen-dominated biological soil crusts

Biological soil crusts dominated by lichens are common components of shrub-steppe ecosystems in northwestern US. We conducted growth chamber experiments to investigate the effects of these crusts on seed germination and initial seedling establishment of two annual grasses; the highly invasive exotic Bromus tectorum L. and the native Vulpia microstachys Nutt. We recorded germination time courses on bare soil and two types of biological soil crusts; one composed predominantly of the lichen Diploschistes muscorum (Scop.) R. Sant. (lichen crust) and the other comprised of an assortment of lichens and mosses (mixed crust). Final germination on the lichen crust for both grass species was about a third of that on the bare soil surface. Mean germination time (MGT) was 3–4 days longer on the lichen crust compared with the bare soil. In contrast, there was no difference in germination percentage or MGT between the mixed crust and bare soil, and results were similar for both grass species. For both species, root penetration of germinating seeds on the lichen crust was lower than on the bare soil or mixed crust surfaces. The combined effects of the lichen crust on germination and root penetration resulted in an overall reduction in seedling establishment of 78% for V. microstachys and 85% for B. tectorum relative to the bare soil treatment. Our results clearly demonstrate that lichen-dominated biological soil crust can inhibit germination and root penetration, but the extent of these effects depends on the composition of the crust. Responsible Editor: Tibor Kalapos     

Initial effects of experimental warming on carbon exchange rates, plant growth and microbial dynamics of a lichen-rich dwarf shrub tundra in Siberia

Biological soil crusts can affect seed germination and seedling establishment. We have investigated the effect of biological soil crusts on seed water status as a potential mechanism affecting seed germination. The seed water potential of two annual grasses, one exotic Bromus tectorum L. and another native Vulpia microstachys Nutt., were analyzed after placing the seeds on bare soil, on a crust that contains various lichens and mosses (mixed crust), or on a crust dominated by the crustose lichen Diploschistes muscorum (Scop.) R. Sant. (Diploschistes crust). Seed water potential and germination were similar on the bare soil and the mixed crust, except for the initial germination of V. microstachys, which was higher on the mixed crust than on the bare soil. For the two grasses studied, seed water potential was significantly higher on the bare soil and mixed crust than on the Diploschistes crust. These differences in water potential correlated with differences in germination, which was much lower on the lichen crust. Experiments were conducted under two watering regimens. Increasing the frequency of watering amplified the differences in seed water potential and germination between the Diploschistes crust and the other two surfaces. For a particular watering regimen, the bare soil, mixed crust, and Diploschistes crust received the same amount of water, but they reached significantly different water potentials. Throughout the experiments, the water potential of the soil and mixed crust remained above −0.6 MPa, while there was a marked decline in the water potential of the Diploschistes surface to about −4 MPa. To ascertain that water was the major factor limiting germination on the Diploschistes crust, we conducted germination tests in an environment with 100% relative humidity. Under these conditions, germination on the Diploschistes crust was similar to that on the bare soil. However, the seeds that germinated on the Diploschistes crust did not penetrate this surface and approximately 60% of their root tips became necrotic. Our results indicate that the presence of D. muscorum can inhibit seedling establishment by two mechanisms: a reduction in seed water absorption and an increase in root tip mortality.     

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.     

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。     

Established native perennial grasses out-compete an invasive annual grass regardless of soil water and nutrient availability

Competition and resource availability influence invasions into native perennial grasslands by non-native annual grasses such as Bromus tectorum. In two greenhouse experiments we examined the influence of competition, water availability, and elevated nitrogen (N) and phosphorus (P) availability on growth and reproduction of the invasive annual grass B. tectorum and two native perennial grasses (Elymus elymoides, Pascopyrum smithii). Bromus tectorum aboveground biomass and seed production were significantly reduced when grown with one or more established native perennial grasses. Conversely, average seed weight and germination were significantly lower in the B. tectorum monoculture than in competition native perennial grasses. Intraspecific competition reduced per-plant production of both established native grasses, whereas interspecific competition from B. tectorum increased production. Established native perennial grasses were highly competitive against B. tectorum, regardless of water, N, or P availability. Bromus tectorum reproductive potential (viable seed production) was not significantly influenced by any experimental manipulation, except for competition with P. smithii. In all cases, B. tectorum per-plant production of viable seeds exceeded parental replacement. Our results show that established plants of Elymus elymoides and Pascopyrum smithii compete successfully against B. tectorum over a wide range of soil resource availability.     

沙坡头地区生物结皮覆盖区土壤种子库组成及垂直分布特征

采用野外随机取样和室内萌发试验相结合的方法,对沙坡头人工植被区的藓类结皮、地衣结皮和裸沙覆盖区土壤种子库的组成及垂直分布特征进行了研究。结果表明:1)人工植被区土壤种子库的物种组成简单,以1年生草本为主,相较于裸沙,两种生物结皮的存在显著增加了土壤种子库的密度(P0.01),但降低了土壤种子库的物种多样性及其与地上植被的相似性;2)两种生物结皮覆盖区土壤种子库的总密度虽无显著差异,但在0—2 cm土层中,地衣结皮土壤种子库的密度显著高于藓类结皮(P0.05),而在2—5 cm和5—10 cm土层中,藓类结皮土壤种子库的密度高于地衣结皮,但差异不显著;3)不同生物结皮覆盖区土壤种子库的垂直分布特征存在差异,藓类结皮覆盖土壤中有活力的种子主要分布在0—5 cm土层中,而地衣结皮和裸沙覆盖土壤中有活力的种子主要分布在0—2 cm土层中,且随着深度的增加,土壤中有活力的种子急剧减少。该研究表明,相较于裸沙,生物结皮的存在不仅增加了土壤种子库的大小,还改变了土壤种子库的垂直分布格局。生物结皮层下仍有相当比例有活力的小种子,这些种子欠缺萌发条件,对地上植被的贡献作用较小,但它们的存在对荒漠生态系统健康及可持续发展具有重要意义。     

Low allelopathic potential of an invasive forage grass on native grassland plants: a cause for encouragement?

Tall fescue (Festuca arundinacea Schreb.), a highly competitive European grass that invades US grasslands, is reportedly allelopathic to many agronomic plants, but its ability to inhibit the germination or growth of native grassland plants is unknown. In three factorial glasshouse experiments, we tested the potential allelopathic effects of endophyte-infected (E+) and uninfected (E−) tall fescue on native grasses and forbs from Midwestern tallgrass prairies. Relative to a water control, at least one extract made from ground seed, or ground whole plant tissue of E+ or E− tall fescue reduced the germination of 10 of 11 species in petri dishes. In addition, the emergence of two native grasses in potting soil was lower when sown with E+ and E− tall fescue seedlings than when sown with seeds of conspecifics or tall fescue. However, when seeds of 13 prairie species were sown in sterilized, field-collected soil and given water or one of the four tall fescue extracts daily, seedling emergence was lower in one extract relative to water for only one species, and subsequent height growth did not differ among treatments for any species. We conclude that if tall fescue is allelopathic, its inhibitory effects on the germination and seedling growth of native prairie plants are limited, irrespective of endophyte infection. On the other hand, the apparent inability of these plants to detect tall fescue in field soil could hinder prairie restoration efforts if germination near this strong competitor confers fitness consequences. We propose that lack of chemical recognition may be common among resident and recently introduced non-indigenous plants because of temporally limited ecological interactions, and offer a view that challenges the existing allelopathy paradigm. Lastly, we suggest that tall fescue removal will have immediate benefits to the establishment of native grassland plants.     

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.     

温带荒漠灌丛和藓类结皮对土壤养分的贡献

灌丛与生物土壤结皮镶嵌分布是温带荒漠常见的地表景观之一,二者的发育均显著影响了地表土壤养分的空间分布特征及循环过程。然而,灌丛和生物土壤结皮对荒漠土壤表层养分的影响存在怎样的差异,二者对养分变化的贡献度如何尚不清楚。因此,选择中国北方典型温带荒漠古尔班通古特沙漠为研究区,以荒漠中的优势灌丛膜果麻黄(Ephedra przewalskii)灌丛和生物土壤结皮发育高级阶段的藓类结皮的结皮层及结皮下层土壤为研究对象,采集不同微生境(裸露地、灌丛下)的裸沙与藓类结皮的土壤样品。为探究不同微生境下不同土层碳、氮、磷和钾养分变化特征,测定了不同土层土壤有机碳(SOC)、全氮(TN)、全磷(TP)、全钾(TK)、速效氮(AN)、速效磷(AP)和速效钾(AK)含量。结果表明:(1)相较于裸露地,灌丛显著提高了藓类结皮不同土层有机碳、全氮和全钾的含量和裸沙全磷的含量,降低了灌丛下藓类结皮土壤全磷的含量。(2)对于速效养分而言,与裸沙相比,裸露地藓类结皮覆盖降低了土壤速效氮含量,增加了速效磷和速效钾含量;而灌丛下藓类结皮覆盖提高了土壤速效氮和速效钾的含量,但降低了速效磷的含量。 (3) 相关性分析显示,在0-2 cm土壤中速效磷与速效氮呈现显著负相关,而在2-6 cm速效磷与速效氮呈现显著正相关(P<0.01)。(4)贡献度分析表明,土壤中灌丛效应对养分的贡献(42.54%)要远大于藓类结皮的贡献(2.43%),但二者交互作用却降低了除速效氮以外的其他土壤养分含量。综上,灌丛、藓类结皮覆盖和土层深度变化对土壤碳、氮、磷和钾养分均具有显著影响(P<0.05),但三者间的交互效应对养分的影响不显著(P>0.05)。相对于裸露地,灌丛与藓类结皮的覆盖均对表层土壤碳、氮、磷和钾养分具有促进作用,且随着土层深度的下降,土壤养分含量呈现显著的下降趋势(P<0.05)。因此,在荒漠生态系统中耐旱灌丛与生物土壤结皮这两个最重要的斑块单元联合调控了微尺度土壤养分的空间异质性变化。     

Allelopathic effect of Aconitum pendulum (Ranunculaceae) on seed germination and seedlings of five native grass species in the Tibetan Plateau

Aconitum pendulum is a dominant weed in bare‐land meadows on the Tibetan plateau and infected areas are very difficult to restore because of the allelopathic effects of this weed. We wanted to select native grasses to restore bare‐land meadows dominated by this toxic weed and tested the allelopathic effect of A. pendulum on seed germination and growth of roots and shoots of five native forage grasses (Elymus nutans, E. sibiricus, Poa crymophila, Festuca sinensis, Bromus inermis). Leaf exudates and high concentrations of root and stem exudates of A. pendulum inhibited seed germination as well as growth of roots and shoots. The allelopathic effect of A. pendulum leaf exudates was stronger than that of roots and stems. Poa crymophila was most resistant to inhibition by A. pendulum, whereas F. sinensis was the most susceptible grass. We conclude that P. crymophila is more suitable than the other four native grasses for sowing in pastures when restoring bare‐land meadows dominated by A. pendulum in the Tibetan plateau.     

Requirement for particular seed-borne fungi for seed germination and seedling growth of <Emphasis Type="Italic">Xyris complanata</Emphasis>, a pioneer monocot in topsoil-lost tropical peatland in Central Kalimantan,Indonesia

Hawaii yellow-eyed grass (Xyris complanata: Xyridaceae) inhabits infertile, acidic peat soil in the rainy tropical zone in Southeast Asia. This monocot plant produces a large number of dormant seeds in order to make a large deposit to seed bank in the soil. Under laboratory conditions, surface-sterilized X. complanata seeds are rarely able to germinate on sterilized peat moss bed; they require inoculation with either seed epiphytic or soil fungi to facilitate active seed germination. In the present study, three different genera of seed epiphytic fungi were isolated, and two common fungal genera, Fusarium sp. (strain R-1) and Penicillium sp. (strain Y-1), were found to promote seed germination of X. complanata. In sterile peat moss beds, the germination-stimulating fungi also showed growth-promoting effects on X. complanata seedlings. These results suggest that the seed germination-promoting fungi likely function as genuine partners for X. complanata in tropical open peat lands.     

Effects of fungal endophytes on the seed and seedling biology of Lolium perenne and Festuca arundinacea

Summary Many grasses are infected by endophytic fungi that grow intercellularly in leaves, stems, and flowers and are transmitted maternally by hyphal growth into ovules and seeds. The seed biology and seedling growth of endophyte-infected and uninfected perennial ryegrass (Lolium perenne) and tall fescue (Festuca arundinacea) were investigated under controlled environmental conditions. The percentage of filled seeds produced by infected tall fescue was over twice of uninfected tall fescue; infected and uninfected perennial reegrass had similar percentages. Weights of seeds from infected and uninfected plants were similar in both species. Seeds from infected plants of both species exhibited a higher rate of germination than seeds from uninfected plants. Shoot growth in the greenhouse was compared by making three sequential harvests of above-ground plant parts from infected and uninfected plants of both species. Infected perennial ryegrass plants produced significantly more biomass and tillers than uninfected plants after 6 and 10 weeks of growth and significantly more biomass after 14 weeks of growth. Infected tall fescue plants produced significantly more biomass and tillers than uninfected plants after 10 and 14 weeks of growth. The physiological mechanism of enhancement of growth is not known. The results of this study suggest that infected plants may have a selective advantage in populations with uninfected members.     

不同类型生物土壤结皮覆盖下风沙土的入渗特征及模拟

干旱荒漠区广泛分布的生物土壤结皮(BSCs)对土壤水分入渗过程有重要影响。以古尔班通古特沙漠南缘的BSCs为研究对象,基于野外采样与室内模拟实验等方法,探究藓类、地衣和藻等3种类型BSCs覆盖下沙土的入渗特征。结果表明:与无结皮覆盖的风沙土对照,3种类型BSCs均显著降低了沙土初渗速率,藓类结皮、地衣结皮、藻结皮覆盖下初渗速率降低幅度依次为36.10%、46.42%、50.39%;藓类结皮、地衣结皮(P0.05)和藻结皮(P0.05)均明显降低了沙土稳渗速率,降低幅度依次为16.50%、33.98%和35.92%;3种类型BSCs均限制了湿润锋在沙土的推进过程,表现为:藓类结皮、地衣结皮、藻结皮的渗漏时间分别为裸沙对照的2.13、3.04和2.98倍;各类型BSCs均减小了沙土累积入渗量,阻碍了沙土水分入渗,与裸沙对照相比,藓类结皮、地衣结皮、藻结皮的1 h累积入渗量分别降低16.10%、28.56%和26.56%。在实验条件下,Kostiakov模型最适用于模拟不同类型BSCs覆盖下土壤水分入渗过程,Horton模型模拟效果次之。     

古尔班通古特沙漠梭梭冠下裸斑形成土壤学机制

古尔班通古特沙漠梭梭与齿肋赤藓共生区域中,梭梭冠下常形成藓类结皮裸露斑块。为研究梭梭冠下裸斑形成机理,以裸斑内土壤、裸斑外藓类结皮土壤、背景裸地土壤以及背景藓类结皮土壤为研究对象,测定四类土壤营养物质含量和理化因子,并测定土壤代谢组以及分析其之间差异。结果表明,裸斑内土壤养分及生态化学计量比与裸斑外藓类结皮和背景藓类结皮土壤并无显著性差异,土壤养分及生态化学计量并非造成梭梭冠下藓类裸斑的原因。裸斑内土壤Na⁺、K⁺、Mg²⁺、SO^2-₄、CO^2-₃、HCO^-₃含量显著高于裸斑外土壤、背景裸地土壤和背景结皮土壤,总盐含量最高为1.705 g/kg。这个含量不足以对齿肋赤藓正常生长造成影响,因此裸斑内土壤中较高的离子浓度也并非藓类裸斑产生的原因。土壤代谢组结果显示不同土壤代谢物具有极显著差异,油酰胺等酰胺类化合物相对丰度最高,占总代谢物的72.68%,且在裸斑内土壤中相对丰度显著高于裸斑藓类结皮土壤、裸地土壤和背景结皮土壤,因此推测裸斑内土壤中油酰胺等酰胺类化合物可能是抑制齿肋赤藓生长的主要原因,是造成古尔班通古特沙漠梭梭冠下藓类裸斑产生的主要土壤学机制。     

实验室条件下两种生物土壤结皮对荒漠植物种子萌发的影响

生物土壤结皮广泛分布在干旱、半干旱区,深刻影响着土壤表层特性,进而对植物种子散布、萌发和定居产生极大的影响。到目前为止,生物土壤结皮与植物关系的研究很少见到,并且这些有限的研究所得出的结论存在着争议。研究了不同年龄的两种生物土壤结皮（苔藓结皮和藻类结皮）对油蒿（Artemisia ordosica）和雾冰藜（Bassia dasyphylla）种子萌发的影响,同时也研究了这两种结皮在失去活性前后对油蒿、雾冰藜和小画眉草（Eragrostis poaeoides）种子萌发的影响。苔藓和藻类结皮的出现对油蒿和雾冰藜种子的萌发均有显著的促进作用,而结皮年龄对植物种子的萌发没有显著的差异。对于不同的植物种,结皮类型和活性对种子的萌发具有不同的作用。雾冰藜在两种结皮上的萌发有显著的差异而油蒿和小画眉草在两种结皮上的萌发没有显著差异。活藻类显著地增加了三种植物的种子萌发,活苔藓仅增加了油蒿和雾冰藜种子萌发量而对小画眉草种子的萌发没有作用,研究表明,生物土壤结皮对一些植物种子萌发具有明显的促进作用。     

Effects of biological soil crusts on seed germination of four endangered herbs in a xeric Florida shrubland during drought

Soil crusts of rosemary scrubs in south-central Florida were examined for effects on seed germination of four herbs that are killed by fire and must recruit from seed: Eryngium cuneifolium (Apiaceae), Hypericum cumulicola (Hypericaceae), Polygonella basiramia (Polygonaceae), and Paronychia chartacea ssp. chartacea (Caryophyllaceae). Biological soil crusts in these sites are dominated by algae, cyanobacteria, fungi, and bacteria. Because crusts can change soil stability, water, and nutrients, they can affect seed germination. A series of greenhouse and field experiments were designed to first examine the effects of crusts in isolation and then to determine their role in the context of other environmental factors – time since fire and distance to the dominant shrub in this system, Ceratiola ericoides. In the greenhouse experiment, germination in autoclaved crusts was dramatically reduced relative to germination in living crusts for all but P. basiramia. In four field experiments where crusts were left intact, disturbed (mechanically or by flaming), or completely removed, the effects of crusts were variable and species-specific, but were significant enough to impact aboveground population sizes. More germination was consistently observed in recently burned sites away from C. ericoides shrubs. Overall rates of germination were generally very low during this study, possibly as a result of seasonal droughts that could have reduced germination, increased seed dormancy, and/or decreased seed viability. The importance of water for germination was confirmed in an experiment with two watering regimes and three crust treatments designed to create a gradient of soil water availability. Germination was significantly greater in the high water treatment and unaffected by soil crust moisture. Dry years are not uncommon in scrub and the results of this study help us to understand how scrub herbs fare during drought and what role biological soil crusts play in germination.     

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。     

斑块面积对干燥-复水条件下藓类植物生理活性的影响

干扰会导致生物结皮斑块破碎并退化。为明确生物结皮斑块破碎诱发生物结皮退化机理,以黄土丘陵区土生对齿藓(Didymodon vinealis)结皮为研究对象,研究了干燥-复水条件下,单株和直径1 cm、2 cm、3 cm、4 cm、5 cm的藓结皮斑块内土生对齿藓的干燥速率、渗透调节物质、丙二醛和光合色素含量等的变化,以期揭示干扰后生物结皮退化的生物学机理。结果表明(1)除单株外,干燥速率随斑块面积减小而增加,直径1 cm斑块内藓的干燥速率是直径5 cm的2倍。(2)反复干燥-复水25天后,直径小于5 cm的斑块内藓的可溶性糖、可溶性蛋白和叶绿素含量低于直径5 cm斑块内藓的含量,丙二醛含量随斑块面积变化无明显规律。(3)干燥速率与斑块面积、可溶性糖、可溶性蛋白及叶绿素含量均呈极显著负相关。以上结果表明,藓结皮斑块面积通过影响斑块内藓类植物干燥速率进而影响其渗透调节和光合作用能力。藓结皮斑块面积减小,藓类植物干燥速率增大,生理活性降低,是藓结皮斑块破碎诱发其退化的原因。研究从藓类植物生理的角度,阐明了干扰后藓类植物衰亡的生理学原因,为生物结皮的保护和管理提供了科学依据。     

Nitrogen Fixation and Leaching of Biological Soil Crust Communities in Mesic Temperate Soils

Biological soil crust is composed of lichens, cyanobacteria, green algae, mosses, and fungi. Although crusts are a dominant source of nitrogen (N) in arid ecosystems, this study is among the first to demonstrate their contribution to N availability in xeric temperate habitats. The study site is located in Lucas County of Northwest Ohio. Using an acetylene reduction technique, we demonstrated potential N fixation for these crusts covering sandy, acidic, low N soil. Similar fixation rates were observed for crust whether dominated by moss, lichen, or bare soil. N inputs from biological crusts in northwestern Ohio are comparable to those in arid regions, but contribute substantially less N than by atmospheric deposition. Nitrate and ammonium leaching from the crust layer were quantified using ion exchange resin bags inserted within intact soil cores at 4 cm depth. Leaching of ammonium was greater and nitrate less in lichen than moss crusts or bare soil, and was less than that deposited from atmospheric sources. Therefore, biological crusts in these mesic, temperate soils may be immobilizing excess ammonium and nitrate that would otherwise be leached through the sandy soil. Moreover, automated monitoring of microclimate in the surface 7 cm of soil suggests that moisture and temperature fluctuations in soil are moderated under crust compared to bare soil without crust. We conclude that biological crusts in northwestern Ohio contribute potential N fixation, reduce N leaching, and moderate soil microclimate.     

黄土高原水蚀风蚀交错区藓结皮对土壤酶活性的影响

以黄土高原水蚀风蚀交错区黄绵土和风沙土上发育30年的典型藓结皮为对象,分6层采集0～12 cm土层土样,研究了藓结皮对不同土层4种水解酶活性的影响及其与土壤养分的关系.结果表明: 与无结皮相比,黄绵土藓结皮的脲酶、碱性磷酸酶、蔗糖酶和蛋白酶活性分别提升了2.4、7.6、20.7、2.4倍,而风沙土藓结皮的这4种水解酶活性分别提升了3.5、22.2、22.3、2.0倍;黄绵土和风沙土藓结皮的体积含水量分别降低了6.5%和0.8%,温度分别降低了0.8和2.5 ℃;黄绵土藓结皮的有机质、碱解氮和速效磷含量分别提升了2.5、2.9和3.6倍,风沙土分别提升了3.6、3.0和6.6倍.4种水解酶的活性与土壤养分含量呈显著正相关,与土壤含水量呈显著负相关;而土壤温度与脲酶活性呈显著正相关,与蛋白酶活性呈显著负相关.黄土高原水蚀风蚀交错区2种土壤藓结皮均能显著提升水解酶活性,这是藓结皮加速土壤养分周转的重要因素;通过改变土壤含水量和温度等酶促反应条件间接影响酶活性,是藓结皮促进土壤酶活性的重要途径.