黄土高原蓝藻和藓类生物结皮对草本植物多样性及生物量的影响

生物结皮在干旱和半干旱气候区占据了广阔的生态位,深刻影响草本植物的空间格局,但其影响程度和作用途径目前仍存有争议。分别以黄土高原风沙土和黄绵土上的蓝藻结皮和藓结皮为对象,以无结皮为对照,通过样方法调查并测定了生物结皮与无结皮样地中草本植物的多样性和生物量,继而结合灰色关联分析和结构方程模型,对比分析了两类生物结皮对草本植物多样性和生物量的影响程度及作用途径。结果表明：（1）风沙土上,无结皮样地的优势科为禾本科和菊科,蓝藻结皮样地的优势科为藜科,藓结皮样地的优势科为禾本科,而黄绵土上无结皮样地的优势科为菊科、禾本科和豆科,蓝藻结皮样地的优势科与无结皮相似,藓结皮样地的优势科为禾本科和菊科;（2）与无结皮相比,风沙土上蓝藻结皮和藓结皮样地中多年生草本的株数分别增加了30%和16%,而黄绵土上两类生物结皮样地中多年生草本株数则分别增加了44%和减少了46%;（3）与无结皮相比,风沙土上蓝藻结皮和藓结皮样地中草本植物的Patrick和Shannon指数分别降低了1.0、0.2和3.4、0.5;而黄绵土上蓝藻结皮和藓结皮样地中两种多样性指数分别降低了4.8、0.2和4.2、0.4;（4）风沙土上蓝藻结皮和藓结皮样地的草本植物生物量比无结皮减少了111.5 g/m²和145.0 g/m²,而在黄绵土上则分别减少了127.8 g/m²和172.2 g/m²;（5）灰色关联分析显示,土壤穿透阻力对草本植物的多样性影响最大（关联系数分别为0.960和0.996）,而NH₄⁺-N和NO₃^--N含量对草本植物的生物量影响最大（两种土壤上关联系数分别为0.727和0.752）。综上,黄土高原生物结皮显著降低了草本植物的多样性和生物量,其途径为通过增加土壤穿透阻力以降低草本植物的种子萌发几率并限制幼苗根系伸展,同时通过土壤氮竞争限制了草本植物生长。     

生物结皮对古尔班通古特沙漠3种荒漠草本植物生长特性与元素吸收的影响

生物结皮有可能通过物理、水文、养分循环影响与之相邻的维管植物,但二者相互关系尚存在着争议。本文以新疆古尔班通古特沙漠广泛分布的地衣结皮为研究对象,分析了生物结皮对3种荒漠草本植物：尖喙牻牛儿苗（Erodium oxyrrhynchum）、条叶庭芥（Alyssum linifolium）和琉苞菊（Hyalea pulchella）的生长及其对元素吸收的影响。研究结果表明：（1）生物结皮对3种荒漠草本植物生长的影响在生长期不同阶段存在差异。在前期,生物结皮的存在促进了植物生物量的累积;而后期,生物结皮却抑制了植物生长。生物结皮的存在显著影响了荒漠草本植物生物量的累积和冠根比。（2）生物结皮的存在显著增加了3种荒漠草本植物对N和K的吸收,而对P的吸收没有显著影响。生物结皮对3种植物Cu、Ca、Mg、Na、Cl的吸收存在种间差异。本研究结果将为该荒漠生态系统潜在的植被演替方向提供重要的科学根据。     

Vascular Plants and Biocrusts Modulate How Abiotic Factors Affect Wetting and Drying Events in Drylands

Understanding how organisms control soil water dynamics is a major research goal in dryland ecology. Although previous studies have mostly focused on the role of vascular plants on the hydrological cycle of drylands, recent studies highlight the importance of biological soil crusts formed by lichens, mosses, and cyanobacteria (biocrusts) as a major player in this cycle. We used data from a 6.5-year study to evaluate how multiple abiotic (rainfall characteristics, temperature, and initial soil moisture) and biotic (vascular plants and biocrusts) factors interact to determine wetting and drying processes in a semi-arid grassland from Central Spain. We found that the shrub Retama sphaerocarpa and biocrusts with medium cover (25–75%) enhanced water gain and slowed drying compared with bare ground areas (BSCl). Well-developed biocrusts (>75% cover) gained more water, but lost it faster than BSCl microsites. The grass Stipa tenacissima reduced water gain due to rainfall interception, but increased soil moisture retention compared to BSCl microsites. Biotic modulation of water dynamics was the result of different mechanisms acting in tandem and often in opposite directions. For instance, biocrusts promoted an exponential behavior during the first stage of the drying curve, but reduced the importance of soil characteristics that accentuate drying rates. Biocrust-dominated microsites gained a similar amount of water than vascular plants, although they lost it faster than vascular plants during dry periods. Our results emphasize the importance of biocrusts for water dynamics in drylands, and illustrate the potential mechanisms behind their effects. They will help to further advance theoretical and modeling efforts on the hydrology of drylands and their response to ongoing climate change.     

The Role of Biological Soil Crusts in Nitrogen Cycling and Soil Stabilization in Kangerlussuaq,West Greenland

Ecosystems - Biological soil crusts (biocrusts) naturally coexist with vascular plants in many dryland ecosystems. Although most studies of dryland biocrusts have been conducted in warm deserts,...     

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。     

Effects of biological soil crusts on a Mediterranean perennial grass (Stipa tenacissima L.)

In the last decades, there has been a growing interest on evaluating the effects of biological soil crusts (hereafter, biocrusts) on vascular plants. The purpose of this paper was to investigate the influence of biocrusts on Stipa tenacissima, key species in arid and semi-arid environments of the Mediterranean Basin. In fact, we explored the impact of biocrusts removal underneath S. tenacissima on soil chemical properties and leaf mineral uptake. Indeed, we studied the effect of mixed biocrusts on the seed germination of S. tenacissima under laboratory conditions. The research showed a greater availability of soil resources correlated with enhanced leaf nutrient levels of bioessential elements in the presence of intact biocrusts compared to fragmented biocrusts underneath S. tenacissima tussocks. We suggest that this is related to the loss of soil moisture as a result of biocrusts removal. In addition, we found that mixed biocrusts favored seed germination and viability of S. tenacissima. By improving soil chemical properties and leaf nutrient uptake, biocrusts can provide enhanced growth conditions for Stipa tenacissima. These findings with the positive effect on seed germination suggest that biocrusts can be used in the restoration of S. tenacissima.     

The effect of lichen‐dominated biological soil crusts on growth and physiological characteristics of three plant species in a temperate desert of northwest China

Biocrusts (biological soil crusts) cover open spaces between vascular plants in most arid and semi‐arid areas. Information on effects of biocrusts on seedling growth is controversial, and there is little information on their effects on plant growth and physiology. We examined impacts of biocrusts on growth and physiological characteristics of three habitat‐typical plants, Erodium oxyrhynchum, Alyssum linifolium and Hyalea pulchella, growing in the Gurbantunggut Desert, northwest China. The influence of biocrusts on plant biomass, leaf area, leaf relative water content, photosynthesis, maximum quantum efficiency of PSII (F_v/F_m), chlorophyll, osmotic solutes (soluble sugars, protein, proline) and antioxidant enzymes (superoxide dismutase, catalase, peroxidase) was investigated on sites with or without biocrust cover. Biomass, leaf area, leaf water content, photosynthesis, F_v/F_m and chlorophyll content in crusted soils were higher than in uncrusted soils during early growth and lower later in the growth period. Soluble sugars, proline and antioxidant enzyme activity were always higher in crusted than in uncrusted soils, while soluble protein content was always lower. These findings indicate that biocrusts have different effects on these three ephemeral species during growth in this desert, primarily via effects on soil moisture, and possibly on soil nutrients. The influence of biocrusts changes during plant development: in early plant growth, biocrusts had either positive or no effect on growth and physiological parameters. However, biocrusts tended to negatively influence plants during later growth. Our results provide insights to explain why previous studies have found different effects of biocrusts on vascular plant growth.     

Synergistic and antagonistic effects of mixing monospecific soils on plant-soil feedbacks

Background and aims

Disturbance affects the ability of organisms to persist on a site, and disturbance history acts as a filter of community composition. This is true for vascular plants and morphological groups of biocrusts, which respond differently to disturbance. Although functioning arid ecosystems include both groups, filtering of morphological groups of biocrusts has not previously been compared simultaneously with the responses of vascular plants.

Methods

Using a chronosequence approach, cover of vascular plants and biocrusts was examined across chronic disturbance gradients related to invasion by exotic species and grazing by livestock, following the acute disturbance of fire using paired burned and unburned plots in Wyoming big sagebrush on 99 plots.

Results

Cover of vascular plants and biocrusts was related to disturbance more so than abiotic factors of precipitation following fire, soil chemistry, percent coarse fragment and heat load index. Over time since fire of 12–23 years, we saw recovery of early successional groups: short mosses, shallow-rooted perennial grasses and annual forbs. Cover of deep and shallow-rooted perennial grasses and annual forbs increased in cover with intermediate levels of disturbance. Perennial forbs lacked a clear relationship with disturbance. Biocrusts decreased in cover with less disturbance when compared with perennial herbaceous plants but differed in sensitivities. Tall mosses were less sensitive to disturbance compared with lichens. Short mosses increased with some disturbance.

Conclusions

Morphological groups of biocrusts and vascular plants are eliminated with increasing variability in the size of gaps between perennials represented by the standard deviation of gaps between perennials. The inclusion of both groups in assessments of ecosystem recovery following disturbance addresses the fact that recovery of either group does not happen in isolation from the other but with interacting contributions to ecosystem functions.

     

黄土丘陵区草本植物覆盖下生物结皮对坡面径流流速的削减作用

生物结皮是干旱半干旱地区的常见地被物,与植物共同影响坡面径流及流速。迄今鲜有研究关注植物和生物结皮共同覆盖对流速的影响,是干旱半干旱地区坡面侵蚀驱动因素研究的薄弱环节。本研究以黄土丘陵区退耕草地为对象,通过野外模拟降雨试验,研究草本植物覆盖下有无生物结皮及不同组成生物结皮(多藻少藓、多藓少藻和藓)对径流流速的影响。结果表明: 植物和植物+生物结皮覆盖显著降低了流速,植物覆盖较裸土降低70.7%,植物+生物结皮覆盖较裸土降低83.1%;植物和生物结皮共同覆盖下,植物和生物结皮对径流流速的削减效益分别为70.7%和12.4%。植物覆盖下生物结皮对流速的影响程度与其组成有关,多藻少藓结皮、多藓少藻结皮和藓结皮对流速的削减效益分别为11.5%、12.4%和19.4%。流速与藓盖度呈显著负相关,与藻盖度呈显著正相关,藓结皮盖度(x)与流速(y)的关系式为:y=-2.081x+0.03(R²=0.469)。当植物盖度一定时(40%±10%),生物结皮组成中藓盖度是影响共同覆盖坡面流速的关键因子。综上,草本植物覆盖下生物结皮有显著减缓流速的作用,且作用程度与其组成有关。表明在研究退耕草地坡面侵蚀动力机制时,生物结皮的作用应予以考虑。     

Biotic communities cannot mitigate the negative effects of grazing on multiple ecosystem functions and services in an arid shrubland

Plant and Soil - Dryland biotic communities (plants and biocrusts) are known to maintain multiple functions (multifunctionality) and services (multiservices) that decline with overgrazing by...     

Differences in chemical composition ofAlysicarpus vaginalis (L.) DC. growing in saline and non-saline habitats

InAlysicarpus vaginalis (L.) DC. nitrogen, ascorbic acid, proline and epicuticular wax (ECW) contents were higher in the plants growing in the coastal region whereas the protein, soluble sugars and starch contents were lower. The higher contents of proline, nitrogen and ascorbic acid recorded in the plants of the saline habitat are a physiological adaptation to overcome the salt stress. The higher ECW content in the plants of the saline habitat specially in the summer months seems to be an adaptation in these plants to survive in the saline habitat. Communicated by Z. ŠESTáK     

Response of biological soil crusts to raindrop erosivity and underlying influences in the hilly Loess Plateau region,China

Biological soil crusts (biocrusts) are ubiquitous living covers in arid and semiarid regions, playing a critical role in soil erosion control in semiarid regions. So far, research separating the multiple mechanisms of erosion control by biocrusts has been limited. It was problematic to link the influence of biocrusts to existing erosion models. In the present study, the response of biocrusts of different successional stages to raindrop erosivity and underlying influences was investigated. Using single drop simulated rainfall, the erosion controlling capacities of biocrusts were analyzed from an energetic perspective. The results showed that biocrusts caused a dramatic improvement of soil erosion resistance, which depended on species composition and increased considerably with higher succession stages. While the accumulated raindrop kinetic energy sustained by dark cyanobacterial crusts was 0.93 J (~15 times higher than that of bare soil), that of 60 % moss covered crusts reached values up to 20.18 J (~342 times higher than that of bare soil) and for 80 % moss covered crusts even 24.59 J were measured. Besides the composition and successional stages, the resistance of biocrusts to raindrop erosivity was related to the substrate soil moisture, soil texture, slope gradients and seasonal variation. The accumulated raindrop kinetic energy measured for cyanobacterial crusts was highest on silty, followed by loamy and sandy soil. For moss-dominated crusts raindrop kinetic energy was highest on sandy, followed by silty and loamy soil. Dry biocrust samples reached significantly higher accumulated raindrop kinetic energies compared to moist biocrusts, whereas the moisture content within moist crusts did not have a significant influence. Erosion resistance increased significantly with higher slope gradients. The resistance capacities of biocrusts during monsoon and post-monsoon were significantly higher than these of pre-monsoon biocrusts. Our results suggest that the influence of biocrusts can be included into erosion models from an energy point of view. The raindrop kinetic energy resistance capacity provides a potential bridge between biocrust succession and soil erodibility in commonly used erosion models.     

Von Vogelkot zu grünen Teppichen

This study tested whether potash tailing piles can be restored with biological soil crusts using an additive of the company upi. A biocrust community consists of different organisms, such as microalgae, lichens, and mosses. An established biocrust stabilizes the ground and traps rain water, which could in return reduce salt leachate into the environment. This pioneer community promotes formation of a new habitat that can be recolonized by higher plants. For this study microalgae were isolated from biological soil crusts collected at potash tailing pile sites. We characterized their salt tolerance and established first artificial biocrusts on the heap material. Upcoming experiments will focus on the establishment of artificial biocrusts in selected heap areas.     

生物结皮与草本植物共生坡面的产流-入渗过程特征

干旱半干旱地区生物结皮与草本植物互生,发挥了重要的水土保持价值,但二者的耦合作用对坡面土壤产流-入渗过程与机理的影响尚不清楚。本研究利用人工模拟降雨试验,设计裸土、生物结皮、长芒草、长芒草+生物结皮4种坡面处理,研究生物结皮与草本植物的产流-入渗过程与径流水动力学特征。结果表明: 2个有生物结皮处理的坡面产流量变化平稳。4种处理的总产流量大小分别为:生物结皮>长芒草+生物结皮>裸土>长芒草,说明生物结皮具有抑制土壤水分入渗的作用,而长芒草可促进土壤水分入渗。在土壤深度为16和24 cm处,生物结皮处理的累计入渗量均小于长芒草+生物结皮处理,且差异显著,说明随着土壤深度增加,长芒草会减小生物结皮抑制水分入渗的作用。除了裸土坡面的弗劳德数(Fr)大于1以外,其余3种处理的Fr值小于1,属于缓流。生物结皮、长芒草和长芒草+生物结皮处理坡面的径流动能比裸土坡面分别减少83.3%、59.5%和88.1%。综上,水动力学参数的变化说明生物结皮调控径流的作用大于长芒草。     

生物土壤结皮演替对土壤生态化学计量特征的影响

生物土壤结皮(生物结皮)是干旱半干旱地区重要的地表覆被物,能够固定碳氮,影响养分循环,从而可能引起土壤化学计量特征的变化。以黄土丘陵区不同演替阶段生物结皮为研究对象,研究该区生态恢复初期生物结皮演替对土壤碳(C)、氮(N)、磷(P)含量及其化学计量特征的影响。结果表明:(1)生物结皮显著增加了生物结皮层C、N、P含量,后期藓结皮较初期藻结皮C、N、P含量分别增加了161%、127%和9%,均显著高于其下0—10 cm土壤;(2)土壤C/N随着生物结皮演替变化较小,范围为10.0—11.8,C/P和N/P在演替后期分别是初期的2.4倍和2.1倍,均显著高于其下0—10 cm土壤;(3)生物结皮土壤C/N、C/P与N/P受坡向影响较大,并与藓生物量显著正相关,与土壤容重显著负相关;(4)生物结皮显著影响土壤C、N、P含量及其化学计量特征,使生物结皮层显著不同于其下层土壤及裸地;(5)生物结皮演替加速表层土壤养分恢复,影响程度可至其下2 cm土层。研究初次从土壤化学计量特征的角度揭示生物结皮对土壤养分循环的贡献,对干旱半干旱地区生态修复和管理具有指导意义。     

黄土高原丘陵区生物结皮土壤的斥水性

采用滴水穿透时间法和酒精溶液入渗法,研究了黄土高原丘陵区浅色藻结皮、深色藻结皮、藻+少量藓结皮、藓+少量藻结皮、藓结皮5种不同发育阶段的原状生物结皮土壤的斥水性及其与土壤含水量的关系.结果表明: 生物结皮增加了土壤的斥水性,其斥水强度和持久性均显著增加.生物结皮土壤的斥水性随生物结皮的演替逐渐降低,当生物结皮中藓类植物盖度在20%以上时,斥水持久性显著低于藻结皮.生物结皮土壤的斥水性与土壤含水量及优势种密切相关,藓类生物结皮土壤的斥水性随着含水量的降低逐渐增加,藻类生物结皮土壤的斥水性随含水量的变化呈双峰曲线.     

生物土壤结皮对雨滴动能的响应及削减作用

在野外调查采样的基础上,采用单滴雨滴法研究了黄土高原水蚀风蚀交错区不同类型生物结皮对雨滴动能的响应及削减作用.结果表明:生物结皮对雨滴动能的响应与其生物组成密切相关,击穿1 cm厚浅色藻结皮和80％盖度藓结皮的累积雨滴动能分别为0.99 J和75.56 J;相同组成的生物结皮对雨滴动能的响应与其生物量有关,生物量越大,击穿生物结皮所需累积雨滴动能越大;当藻结皮叶绿素a含量(表征藻结皮生物量)从3.32 μg·g-1增至3.73 μg·g-1时,对应的雨滴动能则由0.99 J增至2.17 J;当藓结皮的生物量从2.03 g·dm-2升至4.73 g·dm-2时,其对应的雨滴动能由6.08 J增至75.56 J;生物结皮演替过程中对雨滴动能的响应呈“S”形曲线变化.不同生物量的藻结皮对雨滴动能耐受能力的差异不显著;随单位面积苔藓生物量的增加,藓结皮对雨滴动能的耐受能力显著增加.藓结皮对雨滴动能的耐受能力在生物量2.03～4.73 g·dm-2之间呈线性增加;当藓结皮平均生物量达3.70 g·dm-2时,可以抗击62.03 J的雨滴动能.生物结皮对雨滴动能有显著的削弱作用,且这种削弱作用随着生物量的增加而增加.     

Response and resilience of soil biocrust bacterial communities to chronic physical disturbance in arid shrublands

The impact of 10 years of annual foot trampling on soil biocrusts was examined in replicated field experiments at three cold desert sites of the Colorado Plateau, USA. Trampling detrimentally impacted lichens and mosses, and the keystone cyanobacterium, Microcoleus vaginatus, resulting in increased soil erosion and reduced C and N concentrations in surface soils. Trampled biocrusts contained approximately half as much extractable DNA and 20–52% less chlorophyll a when compared with intact biocrusts at each site. Two of the three sites also showed a decline in scytonemin-containing, diazotrophic cyanobacteria in trampled biocrusts. 16S rRNA gene sequence and terminal restriction fragment length polymorphism (T-RFLP) analyses of soil bacteria from untrampled and trampled biocrusts demonstrated a reduced proportion (23–65% reduction) of M. vaginatus and other Cyanobacteria in trampled plots. In parallel, other soil bacterial species that are natural residents of biocrusts, specifically members of the Actinobacteria, Chloroflexi and Bacteroidetes, became more readily detected in trampled than in untrampled biocrusts. Replicate 16S rRNA T-RFLP profiles from trampled biocrusts at all three sites contained significantly more fragments (n=17) than those of untrampled biocrusts (n⩽6) and exhibited much higher variability among field replicates, indicating transition to an unstable disturbed state. Despite the dramatic negative impacts of trampling on biocrust physical structure and composition, M. vaginatus could still be detected in surface soils after 10 years of annual trampling, suggesting the potential for biocrust re-formation over time. Physical damage of biocrusts, in concert with changing temperature and precipitation patterns, has potential to alter performance of dryland ecosystems for decades.     

沙埋对干旱沙区真藓结皮层细菌群落结构和多样性的影响

作为干旱沙区常见干扰之一的沙埋显著影响着生物结皮的结构和功能,但其内在的生物学机理还不清楚。利用高通量测序技术,通过对0(对照)、0.5(浅层)、2和10mm(深层)沙埋处理后的腾格里沙漠东南缘沙坡头地区真藓(Bryum argenteum)结皮层细菌群落物种组成与丰度的测定,研究了沙埋对真藓结皮层细菌群落结构和多样性的影响。结果表明:(1)共检测到沙坡头地区真藓结皮层细菌38门106纲181属,以放线菌、变形菌、蓝藻、浮霉菌、拟杆菌和酸杆菌等为主(占细菌群落的78.4%—83.0%);(2)PCA分析表明沙埋导致该地区真藓结皮层细菌群落结构组成发生明显改变。无沙埋时,真藓结皮层细菌群落中相对丰度最高的是蓝藻(18.6%),随着沙埋厚度的增加,依次变为变形菌(21.5%,沙埋厚度0.5mm)、浮霉菌(21.5%,沙埋厚度2mm)和放线菌(23.3%,沙埋厚度10mm);浅层沙埋显著增加了真藓结皮层细菌群落中光合菌、固氮菌和产菌丝体细菌等关键功能菌的丰度,但深层沙埋降低了它们的丰度;(3)沙埋显著增加了真藓结皮层细菌群落多样性(P0.05)和物种丰富度(P0.05),0.5mm沙埋后的细菌群落丰富度指数最高,2mm沙埋后的结皮层细菌群落多样性指数最高。揭示了沙埋对干旱沙区真藓结皮层细菌群落结构与多样性的影响,为深入理解沙埋对沙区生物结皮结构和生态功能影响的生物学机制提供了一定的理论依据。     

黄土丘陵区不同类型生物结皮下的土壤生态化学计量特征

生物结皮在土壤养分累积和循环中起着重要作用.本研究以黄土丘陵区浅色藻结皮、深色藻结皮、藻藓混合结皮、藓结皮、地衣结皮和普通念珠藻结皮6类典型生物结皮为对象,分析不同类型生物结皮土壤碳(C)、氮(N)、磷(P)含量及其化学计量学特征,研究不同类型生物结皮对土壤养分的影响.结果表明:不同类型生物结皮土壤C、N、P、C/N、C/P、N/P差异显著;生物结皮层C、N、P、C/N、C/P、N/P均显著高于0~10 cm土层土壤.6类生物结皮土壤C、N含量均随土层加深而下降,P含量受土层深度影响较小.对于生物结皮层,藓结皮C、N、P含量分别为27.07、2.42、0.67 g·kg^-1,显著高于其他类型生物结皮.念珠藻结皮的0~2 cm土层土壤C、N、P、C/N、C/P、N/P显著高于其他类型生物结皮.