黄土丘陵区藓结皮人工培养方法试验研究
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摘要
土壤侵蚀及荒漠化是当今国际社会极为关注的全球性环境问题,生物土壤结皮是干旱、半干旱地区普遍存在的地被物,其主要组成物—藻类、苔藓和地衣是常见的先锋拓殖物种,能够在干旱缺水、营养贫瘠、生境恶劣的环境中生长繁殖,并且能通过其代谢方式改变环境,在防风固沙、防治土壤侵蚀和区域生态环境变化等方面具有重要作用。本文以陕北黄土丘陵区自然发育的藓结皮为研究材料,通过室内培养试验探索了藓结皮人工培养适合的接种方法、接种量,适宜的培养温度和土壤水分条件,葡萄糖以及氮、磷、钾、镁、等几种营养物质对藓结皮形成发育的影响。研究表明,在一定条件下,采用人工接种的方法培养藓结皮切实可行,但接种方式、培养温度、土壤水分及营养物质种类不同,对藓结皮形成发育各项指标影响不尽相同。研究在藓结皮的接种方法、接种量及藓结皮培养条件等方面取得重要进展,可为黄土丘陵区生物土壤结皮的人工修复,及其在土壤侵蚀及沙漠化防治中的应用提供技术支撑。主要结论如下:
1、相同培养条件下,在所采用的混合结皮、茎叶碎片和孢子接种3种方法中,混合结皮法接种有利于藓结皮盖度的形成,不同的接种量对藓结皮形成发育影响显著,茎叶碎片接种萌发成肉眼可辩的绿色配子体最快。在温度20℃,光照5230Lx,光周期12h/d条件下,采用混合结皮法接种,经80d的培养,藓结皮盖度、密度分别达到了76%和59株/cm2。混合结皮法接种量在500-750g/m2时藓结皮的盖度和藓类植物的密度生长较高。茎叶碎片接种萌发成肉眼可辩的绿色配子体最快,约4-5d,混合结皮稍慢需7d左右,孢子用时最长,至少20d。
2、温度和土壤含水量显著影响藓结皮的形成和发育。试验条件下,17℃较11℃、22℃、27℃有利于藓结皮形成发育,单位面积藓类植物的数量及生长态势良好,藓结皮盖度和藓类植物密度生长稳定,并保持持续增长。60%田间持水量是藓结皮萌发的下限,只有土壤含水量>60%时,才有藓结皮形成。当土壤含水量达到超饱和含水量,近地面处空气相对湿度接近饱和时更有利于藓结皮的形成和发育。
3、添加合适浓度的某些营养物质有利于藓结皮的形成和发育,其中以葡萄糖和硫酸镁最为显著。在土壤中添加葡萄糖和硫酸镁可以显著增加藓结皮盖度和藓株密度生长。其中以施用葡萄糖培养基上生长的藓结皮盖度、藓株密度和藓株高三个指标生长均为最高;藓结皮的形成发育与葡萄糖施用量明显正相关。黄土丘陵区藓结皮人工土培较适宜的葡萄糖施用浓度是30g/L;低浓度硝酸铵可以提高单位面积藓类植物的数量及生长态势,高浓度有抑制作用。研究区藓结皮形成发育较适宜的硝酸铵浓度为1.0g/L,当硝酸铵浓度达到8.0g/L时不利于藓结皮形成发育;磷酸二氢钾对藓结皮形成发育没有影响;硫酸镁能够促进藓结皮形成发育,对藓株高生长没有影响,在所设几种浓度处理中,以0.25g/L浓度处理下藓结皮盖度和密度生长最高。
Soil erosion and desertification as the most important environmental problems were considered and studied worldwide. Biological soil crusts, which were consisted mainly by cyanobacteria, algae, mosses and lichens, play important roles in arid and semi-arid region, including soil erosion controlling. Mosses, which were an important organism in biological soil crusts, are a well-known pioneer flora that could survive and reproduce in the drought, poor nutrients and severe habitats. In order to promote the formation and restoration of moss crusts in Hilly Loess Plateau region, the suitable inoculation methods, inoculation amounts, cultivated conditions (temperature and soil moisture) and nutrition conditions (Glucose, ammonium nitrate, potassium, dehydrogenated phosphate, magnesium sulfate) were studied by using a growth chamber under laboratory controlled conditions. In the study, the natural moss crusts from Hilly Loess Plateau region were taken as the propagation materials. The results showed:
1. Among the three inoculation methods, including spores inoculated, moss fragments inoculated and mixed crust, mixed crust inoculated was the most favorable methods for the formation of moss crusts when cultivated at the same conditions. There was a significant effect of different inoculation amount on the formation and development of moss crust. Mosses gametophyte formed firstly when inoculated by moss fragments. The coverage and moss density of crusts were 76% and 59 plant/cm2 after 80 days’cultivated at 20℃under a 12 h/d 5230 lx photoperiod. For mixed moss crusts, the coverage and mosses density were much better for 500-750g/m2 inoculated amount than the others. Compared with mixed crusts and spores inoculation, moss fragments inoculation showed the fastest formation of moss gametophyte, which was approximately 4-5d later; And then was mixed moss crusts, about 7d and the slowest was mosses’spores, which need at least 20d.
2. Temperature and soil moisture showed a significant effect on the formation and development of moss crusts. Under our experiment conditions, compared with 11℃, 22℃ and 27℃,17℃was the favorable temperature for moss crusts formation. The moss density and growth speed could be sustained when cultivated at 17℃. Both formation and development of moss crusts were affected by soil moisture significantly. And the moss crusts could be formed after inoculated only when soil moisture was over 60% field water capacity. The formation and development of moss crusts could be enhanced by supersaturated soil moisture and high air humidity near soil surface.
3. The formation and development of moss crusts could be increased by adding appropriate concentrations of nutrients to soils, such as glucose and magnesium sulfate. There were significant increases in coverage and mosses density when adding glucose and magnesium sulfate to soils. The coverage, mosses’density and mosses’height were the highest for soils that added glucose solution; there was a positive correlation between crusts’formation and glucose concentration, and 30g/l was the favorable concentration for soils from the research region. A little bit ammonium nitrate could increase the amount of mosses and the growth speed, while higher concentrations had an inhibited effect. The suitable concentration of ammonium nitrate in Hilly Loess Plateau region was 1.0 g/l, moss crusts’formation and development were inhibited and when the concentration of ammonium nitrate over 8.0 g/l. Potassium dehydrogenated phosphate had no effect on the mosses formation and development. Magnesium sulfate could promote the mosses formation, while had no effect on the mosses’height. In the concentration treatments, the coverage and mosses density were the highest when the concentration was 0.25g/l.
1、相同培养条件下,在所采用的混合结皮、茎叶碎片和孢子接种3种方法中,混合结皮法接种有利于藓结皮盖度的形成,不同的接种量对藓结皮形成发育影响显著,茎叶碎片接种萌发成肉眼可辩的绿色配子体最快。在温度20℃,光照5230Lx,光周期12h/d条件下,采用混合结皮法接种,经80d的培养,藓结皮盖度、密度分别达到了76%和59株/cm2。混合结皮法接种量在500-750g/m2时藓结皮的盖度和藓类植物的密度生长较高。茎叶碎片接种萌发成肉眼可辩的绿色配子体最快,约4-5d,混合结皮稍慢需7d左右,孢子用时最长,至少20d。
2、温度和土壤含水量显著影响藓结皮的形成和发育。试验条件下,17℃较11℃、22℃、27℃有利于藓结皮形成发育,单位面积藓类植物的数量及生长态势良好,藓结皮盖度和藓类植物密度生长稳定,并保持持续增长。60%田间持水量是藓结皮萌发的下限,只有土壤含水量>60%时,才有藓结皮形成。当土壤含水量达到超饱和含水量,近地面处空气相对湿度接近饱和时更有利于藓结皮的形成和发育。
3、添加合适浓度的某些营养物质有利于藓结皮的形成和发育,其中以葡萄糖和硫酸镁最为显著。在土壤中添加葡萄糖和硫酸镁可以显著增加藓结皮盖度和藓株密度生长。其中以施用葡萄糖培养基上生长的藓结皮盖度、藓株密度和藓株高三个指标生长均为最高;藓结皮的形成发育与葡萄糖施用量明显正相关。黄土丘陵区藓结皮人工土培较适宜的葡萄糖施用浓度是30g/L;低浓度硝酸铵可以提高单位面积藓类植物的数量及生长态势,高浓度有抑制作用。研究区藓结皮形成发育较适宜的硝酸铵浓度为1.0g/L,当硝酸铵浓度达到8.0g/L时不利于藓结皮形成发育;磷酸二氢钾对藓结皮形成发育没有影响;硫酸镁能够促进藓结皮形成发育,对藓株高生长没有影响,在所设几种浓度处理中,以0.25g/L浓度处理下藓结皮盖度和密度生长最高。
Soil erosion and desertification as the most important environmental problems were considered and studied worldwide. Biological soil crusts, which were consisted mainly by cyanobacteria, algae, mosses and lichens, play important roles in arid and semi-arid region, including soil erosion controlling. Mosses, which were an important organism in biological soil crusts, are a well-known pioneer flora that could survive and reproduce in the drought, poor nutrients and severe habitats. In order to promote the formation and restoration of moss crusts in Hilly Loess Plateau region, the suitable inoculation methods, inoculation amounts, cultivated conditions (temperature and soil moisture) and nutrition conditions (Glucose, ammonium nitrate, potassium, dehydrogenated phosphate, magnesium sulfate) were studied by using a growth chamber under laboratory controlled conditions. In the study, the natural moss crusts from Hilly Loess Plateau region were taken as the propagation materials. The results showed:
1. Among the three inoculation methods, including spores inoculated, moss fragments inoculated and mixed crust, mixed crust inoculated was the most favorable methods for the formation of moss crusts when cultivated at the same conditions. There was a significant effect of different inoculation amount on the formation and development of moss crust. Mosses gametophyte formed firstly when inoculated by moss fragments. The coverage and moss density of crusts were 76% and 59 plant/cm2 after 80 days’cultivated at 20℃under a 12 h/d 5230 lx photoperiod. For mixed moss crusts, the coverage and mosses density were much better for 500-750g/m2 inoculated amount than the others. Compared with mixed crusts and spores inoculation, moss fragments inoculation showed the fastest formation of moss gametophyte, which was approximately 4-5d later; And then was mixed moss crusts, about 7d and the slowest was mosses’spores, which need at least 20d.
2. Temperature and soil moisture showed a significant effect on the formation and development of moss crusts. Under our experiment conditions, compared with 11℃, 22℃ and 27℃,17℃was the favorable temperature for moss crusts formation. The moss density and growth speed could be sustained when cultivated at 17℃. Both formation and development of moss crusts were affected by soil moisture significantly. And the moss crusts could be formed after inoculated only when soil moisture was over 60% field water capacity. The formation and development of moss crusts could be enhanced by supersaturated soil moisture and high air humidity near soil surface.
3. The formation and development of moss crusts could be increased by adding appropriate concentrations of nutrients to soils, such as glucose and magnesium sulfate. There were significant increases in coverage and mosses density when adding glucose and magnesium sulfate to soils. The coverage, mosses’density and mosses’height were the highest for soils that added glucose solution; there was a positive correlation between crusts’formation and glucose concentration, and 30g/l was the favorable concentration for soils from the research region. A little bit ammonium nitrate could increase the amount of mosses and the growth speed, while higher concentrations had an inhibited effect. The suitable concentration of ammonium nitrate in Hilly Loess Plateau region was 1.0 g/l, moss crusts’formation and development were inhibited and when the concentration of ammonium nitrate over 8.0 g/l. Potassium dehydrogenated phosphate had no effect on the mosses formation and development. Magnesium sulfate could promote the mosses formation, while had no effect on the mosses’height. In the concentration treatments, the coverage and mosses density were the highest when the concentration was 0.25g/l.
引文
[1]刘拓.中国土地沙漠化经济损失评估[J].中国沙漠,2006,1(26):40-47.
[2]姚德良,李家春,杜岳,等.沙坡头人工植被区陆气耦合模式及生物结皮与植被演变的机理研究[J].生态学报,2002,22 (4):452-460.
[3]肖洪浪,李新荣,段争虎,等.流沙固定过程中土壤——植被系统演变对水环境的影响[J].土壤学报,2003,40(6):810-813.
[4]魏江春.沙漠生物地毯工程——干旱沙漠治理的新途径[J].干旱区研究,2005,22(3):287-288.
[5]吴鹏程.苔藓植物生物学[M] .北京:科学出版社,1998.
[6] Belnap J,Lange O L.Biological Soil Crusts:Structure Function and Management[M].Berlin:Springer Verlag,2001.
[7] Williams JD,Dobrowolski JP,West NE.Microphytic soil crust influence on wind erosion[J].Tran. Amer.Soc.Agric.Eng, 1995,(38):131-137.
[8]吴玉环,高谦,程国栋.生物土壤结皮的生态功能[J].生态学杂志,2002,22(4):41-45.
[9]闫德仁,薛英英,赵春光.沙漠生物结皮国内研究现状[J],内蒙古林业科技.2007,33(3):28-32.
[10]胡春香,张德禄,刘永定.干旱区微小生物结皮中藻类研究的新进展[J].自然科学进展,2003,13(8):791- 795.
[11]张元明,曹同,潘伯荣.新疆古尔班通古特沙漠南缘土壤结皮中苔藓植物的研究[J] .西北植物学报,2002,22(1):8-23.
[12]赵建成,张丙昌,张元明.新疆古尔班通古特沙漠生物结皮绿藻研究[J] .干旱区研究,2006,23(2):189-194.
[13]赵允格,许明祥,王全九,等.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,2006,17(8): 1429-1433.
[14] Eldridge DJ,Greene RSB. Microbiotic crusts:a view of roles in soil and ecological processes in the rangelands of Australia [J]. Australia Journal of Soil Research,1994,32:389-415.
[15] Belnap J.Recovery rates of cryptobiotic crusts:Inculant use and assessment methods [J]. Great Basin Nat,1993,53-89.
[16]聂华丽,吴楠,梁少民,等.不同沙埋深度对刺叶墙藓植株碎片生长的影响[J] .干旱区研究,2006,23(1): 66-70.
[17]武显维,丁朝华,康宁.湿地匍灯藓切茎繁殖研究[J].武汉植物学研究,1996,14(1):82-88.
[18]田桂泉,白学良,徐杰,等.腾格里沙漠固定沙丘藓类植物结皮层的自然恢复及人工培养试验研究[J] .植物生态学报,2005,29(1):164-169.
[19]白学良,王瑶,徐杰,等.沙坡头地区固定沙丘结皮层藓类植物的繁殖和生长特性研究[J] .中国沙漠,2003,23(2):171-176.
[20]肖波,赵允格,邵明安.黄土高原侵蚀区生物结皮的人工培育及其水土保持效应[J] .草地学报,2008,16(1):28-33.
[21] West N E. Structure and function of microphysical soil crusts in wild-land ecosystem of arid to semi-arid regions [J] .Advances in Ecological Research,1990,20:179-223.
[22]李守中,郑怀舟,李守丽,等.沙坡头植被固沙区生物结皮的发育特征[J] .生态学杂志,2008,27(10):1675-1679.
[23] Zaady E,Kuhn U,Wilske B,et al.Patterns of CO2 exchange in biological soil crusts of success ional age[J]. Soil Biology&Biochemistry,2000,32:959-966.
[24]张元明.荒漠地表生物结皮的微结构及其早期发育特征[J].科学通报,2005,50(1):42-47.
[25]胡春香,刘永定,张德禄,等.荒漠藻结皮的胶结机理[J].科学通报,2002,47(12):931-937.
[26] Schulten J A.Soil aggregation by cryptogams of a sand prairie[J].J Bot,1985,72(11):1657-1661.
[27]陈兰周,刘永定,李敦海,等.荒漠藻类及其结皮的研究[J].中国科学基金,2003,(2):90-93.
[28]张丙昌,张元明,赵建成,等.古尔班通古特沙漠生物结皮不同发育阶段中藻类的变化[J] .生态学报,2009,1(29):9-17.
[29] Belnap J,Phillip s S L,MillerM E.Response of desert biological soil crusts to alterations in precip- itation frequency[ J ].Oecologia,2004,141:306-316.
[30] Kidron G J,Barzilay E,Sachs E.Microclimate control upon sand macrobiotic crusts,western Negev desert,Israel[ J ].Geom orphology,2000,36:1-18.
[31] Vies H.Understanding dry land landscape dynamics:Do biological crusts hold the key? [J].Geography Com pass,2008,2 (3) :899-919.
[32] Campbell S E,Seeler J,Golubic S Desert crust formation and soil stabilization [J]. Arid Soil Research and Rehabilitation,1989,(3):217-288.
[33] Issa O M,Trichet J,Defarge C,et al. Morphology and microstructure of micro biotic soil crusts on a tiger bush sequence(Niger,Sahel).Catena,1999,37: 175-196.
[34] Fearnehough W,Fullen M A,Mitchen Trueman D J I C,et al.Aeolian deposition and its effect on soil and vegetation changes on stabilized desert dunes in northern China [J].Geomorphology,1998, 23:171-182.
[35] Fullen M A,Mitchell D J.Desertification and reclamation in north-central China[J].AMB IO, 1994,23:131-135.
[36] Malam Issa O. Morphology and microstructure of microbiotic soil crusts on a tiger bush sequence (Niger) Sahel [J] Catena,1999,37:175-196.
[37]吴玉环,高谦,于兴华.生物土壤结皮的分布影响因子及其监测[J].生态学杂志,2003,22(3):38-42.
[38]卢晓杰,张克斌,李瑞.北方农牧交错带生物结皮的主要影响因子探讨[J].水土保持研究,2007,14(6):1-4.
[39] Issa O M,Bissonnais Y L,Defarge C,et al.Role of a cyanobacterial cover on structural stability of sandy soils in the Sahelian part of western Niger[J].Geoderma,2001,101:15-30.
[40] Greene R S B,Tongway D J.The significance of (surface) physical and chemical properties in determining soil surface condition of red-earths in rangelands[J].Australian Journal Soil Research,1989,27:213-225.
[41] Chartres C J. Soil Crusting in Australia[A].Sumner M E,Stewart B A,eds.Soil Crusting Chemical and Physical Processes [C].Florida:Lewis Publishers,1992:339-365.
[42]梁少民,吴楠,王红玲,等.干扰对生物土壤结皮及其理化性质的影响[J].干旱区地理,2005,12(6):818-823.
[43] Metting B.Biological surface features of semiarid lands and deserts[A].Skujins J,eds.Semiarid Lands and Deserts:Soil Resource and Reclamation[C].New York::Marcel Dekker,1991:275-293.
[44]李新荣,张元明,赵允格.生物土壤结皮研究进展前沿与展望[J].地球科学进展,2009,24(1):11-24.
[45]陈宝瑞,辛晓平,朱玉霞,等.内蒙荒漠化年动态变化及与气候因子分析[J] .遥感信息,2007.(6):39-44.
[46]世界防止荒漠化及干旱日,http://www.wmgmw.cn/readnews.asp?newsid=19913,2008.6.17.
[47]肖波,赵允格,邵明安.陕北水蚀风蚀交错区两种生物结皮对土壤理化性质的影响[J].生态学报,2007,27(11):4662-4670.
[48]方海燕,屈建军,俎瑞平,等.防沙工程的结皮效应研究[J].水土保持学报,2005,19(2):17-20.
[49] Gahel A.G.,Shtina E.A.Algae on the sands of arid regions and their role in soil formation [J]. Soviet Soil Sci,1974,6:311-319.
[50]薛英英.鄂尔多斯沙区沙漠生物结皮特征的研究[D].内蒙古:内蒙古农业大学硕士学位论文,2007.
[51] Eldridge D J.Cryptogams,vascular plants,and soil hydrological relations:some preliminary results from the semi-arid woodlands of eastern Australia [J].Great Basin Naturalist,1993,53(1):48-58.
[52]闫德仁,季蒙,薛英英,等.沙漠生物结皮土壤发育特征的研究[J].土壤通报,2006,37(5):990-993.
[53]李新荣,贾玉奎,龙利群,等.干旱半干旱地区土壤生物结皮的生态学意义及若干研究进展[J] .中国沙漠,2001,21(1):4-11.
[54]吴楠,潘伯荣,张元明,等.古尔班通古特沙漠生物结皮中土壤微生物垂直分布特征[J] .应用与环境生物学报,2005,11(3):349-353.
[55]冯金朝,陈荷生.宁夏沙坡头地区人工植被种间水分竞争的初步研究[J].生态学报,1994,14(3):260-265.
[56]徐杰,白学良,杨持,等.固定沙丘结皮层藓类植物多样性及固沙作用研究[J].植物生态学报,2003,27(4):545-551.
[57]田桂泉,白学良,徐杰,等.固定沙丘生物结皮层藓类植物形态结构及其适应性研究[J].中国沙漠,2005,25(2):249-255.
[58]石庆辉,刘家琼.沙坡头铁路两侧人工植被区天然植被动态[M].兰州:甘肃科学技术出版社,1995.
[59]张志山,何明珠,谭会娟,等.沙漠人工植被区生物结皮类土壤的蒸发特性——以沙坡头沙漠研究试验站为例[J].土壤学报,2007,44(3):404-410.
[60] Kidron G J,et al.Two different causes for runoff initiation on microbiotic crusts:hydrophobicity and pore clogging [J].Soil Sci,1999,164(1):18-27.
[61]王雪芹,张元明,张伟民,等.古尔班通古特沙漠生物结皮对地表风蚀作用的影响的风洞实验[J].冰川冻土,2004,26(5):632-637.
[62] Kinnel P.I.A,et a1. The effects of fire on the soil in a degraded semiarid woodland II:susceptibility 0fthesoilto erosion by shallow rain impacted flow[J]. Australia Journal of Soil Research, 1990,28(40):755-777.
[63] Greene RSB,Tonway DJ,The significance of surface physical and chemical properties in determining soil surface condition of red-earths in rangeland[J]. Australia Journal of Soil Research, 1989,27:213-225.
[64]张振国,焦菊英,白文娟.黄土丘陵沟壑区退耕地植被恢复中生物土壤结皮特征[J].水土保持通报,2006,26(4):33-37.
[65] Eldrldge DJ,Greene RSB.Microbiotic soil crusts:A view of the roles in soil and ecological processes in the range lands of Australia [J].Australia Journal of Soil Research,1994,32:389-415.
[66]宋阳,严平,张宏,等.荒漠生物结皮研究中的几个问题[J].干旱区研究,2004,21(4):439-443.
[67]吴楠.古尔班通古特沙漠生物中微生物分布的日控差异[J].科学通报,2006,51(z1) :100-107.
[68]赵允格,许明祥,王全九,等.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,2006,l7(8) :1429-1434.
[69]张元明,曹同,潘伯荣.干旱半干旱地区苔藓植物生态学研究综述[J].生态学报,2002,22(7):1129-1134.
[70]徐杰,白学良,田桂泉,等.干旱半干旱地区生物结皮层藓类植物氨基酸和营养物质组成特征及适应性分析[J].生态学报,2005,25(6):1247-1255.
[71]牛玉璐,曹永胜.生物结皮及其在荒漠治理中的作用[J].生物学教学,2005,30(10):5-6.
[72]王世冬,白学良,雍世鹏.沙坡头地区苔藓区系初步研究[J].中国沙漠,2001,9(3):244-249.
[73] Stanley C J,Buxton R. Developing protocols for utilizing moss in revegetation of the Grasberg Mine overburden.http://www.hortresearch.co.nz/product/ncl/reports.
[74] Rochefort L.Sphagnum a keystone genus in habitat restoration [J].The Bryologist,2000,103(3):503-508.
[75]赵建成,黄士良,李敏,等.小扭口藓(Barbula indica)芽孢发育特征的实验研究[J].植物研究,2005,25(2):169-172.
[76]刘晓红.细胞分裂素、氯丙嗪对葫芦藓发育过程的影响[J].南京师范大学学报(自然科学版),1998,23(4):476-480.
[77]高永超,沙伟,张晗.不同植物生长物质对牛角藓愈伤组织诱导的影响[J] .植物生理学通讯,2003,39(1):29-32.
[78]刘世彪,陈军,李菁,等.光照和温度对尖叶拟船叶藓孢子萌发及原丝体发育的影响[J] .西北植物学报,2003,23(1):101-106.
[79]张健,刘国彬,许明祥,等.黄土丘陵区影响生物结皮退化因素的初步研究[J] .中国水土保持科学,2008,12(6):14-20.
[80]黄昌勇.土壤学[M].北京:中国农业出版,2000.
[81]白文娟,焦菊英,张振国.黄土丘陵沟壑区退耕地土壤种子库对植被恢复的影响[J].北京林业大学学报,2008,30(4):65-71.
[82]赵允格,徐冯楠,许明祥.黄土丘陵区藓结皮生物量测定方法及其随发育年限的变化[J].西北植物学报,2008,28(6):1228-1232.
[83]罗应婷,杨钰娟.SPSS统计分析从基础到实践[M].北京:电子工业出版社,2007.
[84]齐红岩,李天来,陈元宏,等.叶面喷施磷酸二氢钾与葡萄糖对番茄光合速率和蔗糖代谢的影响[J].农业工程学报,2005,12:137-142.
[85]刘奎彬,马文荷,安志信.黄瓜对外源糖吸收和运转的研究[J].华北农学,1998,13(4):103-106.
[86]马文苛,刘奎彬,安志信.青椒对外源糖的吸收与分配规律[J].河北农业大学学报,2000,1(23):37-39
[87]浙江农业大学主编.植物营养与肥料[M].北京:中国农业出版社,2004
[88]李延,刘星辉,庄卫民.植物Mg素营养生理的研究进展[J].福建农业大学学报,2000,29(1):74-80.
[89] Glime,Janice M. NUTRIENT RELATIONS:REQUIREMENTS. in Glime,Janice M. Bryophyte Ecology. Volume 1. Physiological Ecology[M]. Michigan Technological University and the International Association of Bryologists,2007.
[90]朱显漠.黄绵土. http://www.chinabaike.com/article/316/327/2007/2007022260712.html.2009.5.7
[91]陈邦杰.中国藓类植物属志上册[M].北京:科学出版社,1963.
[92]刘应迪,朱杰英,陈军,等.3种藓类植物水分含量与光合作用、呼吸作用和水势的关系[J] .武汉植物研究,2001,19(2):135-142.
[93] Martin B.Developmental physiology bryophytes [J].New Mann Bryol,1984,2,2276-2324.
[94]李琴琴,白学良,任向宇.沙漠区生物结皮层中藓类植物繁殖体发育实验研究[J].中国沙漠,2008,3 (28) .289-294.
[95]邢建民,赵德修,李茂寅.水母雪莲悬浮培养细胞生长和黄酮类活性成分合成[J].植物学报1998,40(9):836-841.
[96]高永超,薛红,沙伟.蔗糖对牛角藓愈伤组织悬浮细胞的生理学影响[J].广西植物,2003,23(5):464-469.
[97] Takami S, Yasunaga M, Takio S et al.Establishment of Suspension Cultures of Cells from the Hornwort Anthoceros Punctatusl [M]. J Hattori Bot lab, 1988.
[98]王柳,张福墁,魏秀菊.不同氮肥水平对日光温室黄瓜品质和产量的影响[J].农业工程学报,2007,23(12):225-229.
[99]赵营,同延安,赵护兵.不同供氮水平对夏玉米养分累积、转运及产量的影响[J].植物营养与肥料学报,2006,12(5):622-627.
[100]陈日远,刘厚诚,宋传珍,等.氮素营养对芥蓝生长和品质的影响[J].农业工程学报,2005,12 (21):143-146.
[101]冀瑞萍,张妙娟,温学.水体氮素对藻类植物的影响[J].晋中学院学报,2006,23(6):41-43.
[102]梅赶年.怎样施用磷酸二氢钾[J].新农业,1985,(11):20-26.
[103]李晓鸣.矿质镁对水稻产量及品质影响的研究[J].植物营养与肥料学报,2002,8(1):125-126.
[104]陈圆圆,郭水良,曹同.藓类植物的无性繁殖及其应用[J] .生态学杂志,2008,27(6):993-998.
[2]姚德良,李家春,杜岳,等.沙坡头人工植被区陆气耦合模式及生物结皮与植被演变的机理研究[J].生态学报,2002,22 (4):452-460.
[3]肖洪浪,李新荣,段争虎,等.流沙固定过程中土壤——植被系统演变对水环境的影响[J].土壤学报,2003,40(6):810-813.
[4]魏江春.沙漠生物地毯工程——干旱沙漠治理的新途径[J].干旱区研究,2005,22(3):287-288.
[5]吴鹏程.苔藓植物生物学[M] .北京:科学出版社,1998.
[6] Belnap J,Lange O L.Biological Soil Crusts:Structure Function and Management[M].Berlin:Springer Verlag,2001.
[7] Williams JD,Dobrowolski JP,West NE.Microphytic soil crust influence on wind erosion[J].Tran. Amer.Soc.Agric.Eng, 1995,(38):131-137.
[8]吴玉环,高谦,程国栋.生物土壤结皮的生态功能[J].生态学杂志,2002,22(4):41-45.
[9]闫德仁,薛英英,赵春光.沙漠生物结皮国内研究现状[J],内蒙古林业科技.2007,33(3):28-32.
[10]胡春香,张德禄,刘永定.干旱区微小生物结皮中藻类研究的新进展[J].自然科学进展,2003,13(8):791- 795.
[11]张元明,曹同,潘伯荣.新疆古尔班通古特沙漠南缘土壤结皮中苔藓植物的研究[J] .西北植物学报,2002,22(1):8-23.
[12]赵建成,张丙昌,张元明.新疆古尔班通古特沙漠生物结皮绿藻研究[J] .干旱区研究,2006,23(2):189-194.
[13]赵允格,许明祥,王全九,等.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,2006,17(8): 1429-1433.
[14] Eldridge DJ,Greene RSB. Microbiotic crusts:a view of roles in soil and ecological processes in the rangelands of Australia [J]. Australia Journal of Soil Research,1994,32:389-415.
[15] Belnap J.Recovery rates of cryptobiotic crusts:Inculant use and assessment methods [J]. Great Basin Nat,1993,53-89.
[16]聂华丽,吴楠,梁少民,等.不同沙埋深度对刺叶墙藓植株碎片生长的影响[J] .干旱区研究,2006,23(1): 66-70.
[17]武显维,丁朝华,康宁.湿地匍灯藓切茎繁殖研究[J].武汉植物学研究,1996,14(1):82-88.
[18]田桂泉,白学良,徐杰,等.腾格里沙漠固定沙丘藓类植物结皮层的自然恢复及人工培养试验研究[J] .植物生态学报,2005,29(1):164-169.
[19]白学良,王瑶,徐杰,等.沙坡头地区固定沙丘结皮层藓类植物的繁殖和生长特性研究[J] .中国沙漠,2003,23(2):171-176.
[20]肖波,赵允格,邵明安.黄土高原侵蚀区生物结皮的人工培育及其水土保持效应[J] .草地学报,2008,16(1):28-33.
[21] West N E. Structure and function of microphysical soil crusts in wild-land ecosystem of arid to semi-arid regions [J] .Advances in Ecological Research,1990,20:179-223.
[22]李守中,郑怀舟,李守丽,等.沙坡头植被固沙区生物结皮的发育特征[J] .生态学杂志,2008,27(10):1675-1679.
[23] Zaady E,Kuhn U,Wilske B,et al.Patterns of CO2 exchange in biological soil crusts of success ional age[J]. Soil Biology&Biochemistry,2000,32:959-966.
[24]张元明.荒漠地表生物结皮的微结构及其早期发育特征[J].科学通报,2005,50(1):42-47.
[25]胡春香,刘永定,张德禄,等.荒漠藻结皮的胶结机理[J].科学通报,2002,47(12):931-937.
[26] Schulten J A.Soil aggregation by cryptogams of a sand prairie[J].J Bot,1985,72(11):1657-1661.
[27]陈兰周,刘永定,李敦海,等.荒漠藻类及其结皮的研究[J].中国科学基金,2003,(2):90-93.
[28]张丙昌,张元明,赵建成,等.古尔班通古特沙漠生物结皮不同发育阶段中藻类的变化[J] .生态学报,2009,1(29):9-17.
[29] Belnap J,Phillip s S L,MillerM E.Response of desert biological soil crusts to alterations in precip- itation frequency[ J ].Oecologia,2004,141:306-316.
[30] Kidron G J,Barzilay E,Sachs E.Microclimate control upon sand macrobiotic crusts,western Negev desert,Israel[ J ].Geom orphology,2000,36:1-18.
[31] Vies H.Understanding dry land landscape dynamics:Do biological crusts hold the key? [J].Geography Com pass,2008,2 (3) :899-919.
[32] Campbell S E,Seeler J,Golubic S Desert crust formation and soil stabilization [J]. Arid Soil Research and Rehabilitation,1989,(3):217-288.
[33] Issa O M,Trichet J,Defarge C,et al. Morphology and microstructure of micro biotic soil crusts on a tiger bush sequence(Niger,Sahel).Catena,1999,37: 175-196.
[34] Fearnehough W,Fullen M A,Mitchen Trueman D J I C,et al.Aeolian deposition and its effect on soil and vegetation changes on stabilized desert dunes in northern China [J].Geomorphology,1998, 23:171-182.
[35] Fullen M A,Mitchell D J.Desertification and reclamation in north-central China[J].AMB IO, 1994,23:131-135.
[36] Malam Issa O. Morphology and microstructure of microbiotic soil crusts on a tiger bush sequence (Niger) Sahel [J] Catena,1999,37:175-196.
[37]吴玉环,高谦,于兴华.生物土壤结皮的分布影响因子及其监测[J].生态学杂志,2003,22(3):38-42.
[38]卢晓杰,张克斌,李瑞.北方农牧交错带生物结皮的主要影响因子探讨[J].水土保持研究,2007,14(6):1-4.
[39] Issa O M,Bissonnais Y L,Defarge C,et al.Role of a cyanobacterial cover on structural stability of sandy soils in the Sahelian part of western Niger[J].Geoderma,2001,101:15-30.
[40] Greene R S B,Tongway D J.The significance of (surface) physical and chemical properties in determining soil surface condition of red-earths in rangelands[J].Australian Journal Soil Research,1989,27:213-225.
[41] Chartres C J. Soil Crusting in Australia[A].Sumner M E,Stewart B A,eds.Soil Crusting Chemical and Physical Processes [C].Florida:Lewis Publishers,1992:339-365.
[42]梁少民,吴楠,王红玲,等.干扰对生物土壤结皮及其理化性质的影响[J].干旱区地理,2005,12(6):818-823.
[43] Metting B.Biological surface features of semiarid lands and deserts[A].Skujins J,eds.Semiarid Lands and Deserts:Soil Resource and Reclamation[C].New York::Marcel Dekker,1991:275-293.
[44]李新荣,张元明,赵允格.生物土壤结皮研究进展前沿与展望[J].地球科学进展,2009,24(1):11-24.
[45]陈宝瑞,辛晓平,朱玉霞,等.内蒙荒漠化年动态变化及与气候因子分析[J] .遥感信息,2007.(6):39-44.
[46]世界防止荒漠化及干旱日,http://www.wmgmw.cn/readnews.asp?newsid=19913,2008.6.17.
[47]肖波,赵允格,邵明安.陕北水蚀风蚀交错区两种生物结皮对土壤理化性质的影响[J].生态学报,2007,27(11):4662-4670.
[48]方海燕,屈建军,俎瑞平,等.防沙工程的结皮效应研究[J].水土保持学报,2005,19(2):17-20.
[49] Gahel A.G.,Shtina E.A.Algae on the sands of arid regions and their role in soil formation [J]. Soviet Soil Sci,1974,6:311-319.
[50]薛英英.鄂尔多斯沙区沙漠生物结皮特征的研究[D].内蒙古:内蒙古农业大学硕士学位论文,2007.
[51] Eldridge D J.Cryptogams,vascular plants,and soil hydrological relations:some preliminary results from the semi-arid woodlands of eastern Australia [J].Great Basin Naturalist,1993,53(1):48-58.
[52]闫德仁,季蒙,薛英英,等.沙漠生物结皮土壤发育特征的研究[J].土壤通报,2006,37(5):990-993.
[53]李新荣,贾玉奎,龙利群,等.干旱半干旱地区土壤生物结皮的生态学意义及若干研究进展[J] .中国沙漠,2001,21(1):4-11.
[54]吴楠,潘伯荣,张元明,等.古尔班通古特沙漠生物结皮中土壤微生物垂直分布特征[J] .应用与环境生物学报,2005,11(3):349-353.
[55]冯金朝,陈荷生.宁夏沙坡头地区人工植被种间水分竞争的初步研究[J].生态学报,1994,14(3):260-265.
[56]徐杰,白学良,杨持,等.固定沙丘结皮层藓类植物多样性及固沙作用研究[J].植物生态学报,2003,27(4):545-551.
[57]田桂泉,白学良,徐杰,等.固定沙丘生物结皮层藓类植物形态结构及其适应性研究[J].中国沙漠,2005,25(2):249-255.
[58]石庆辉,刘家琼.沙坡头铁路两侧人工植被区天然植被动态[M].兰州:甘肃科学技术出版社,1995.
[59]张志山,何明珠,谭会娟,等.沙漠人工植被区生物结皮类土壤的蒸发特性——以沙坡头沙漠研究试验站为例[J].土壤学报,2007,44(3):404-410.
[60] Kidron G J,et al.Two different causes for runoff initiation on microbiotic crusts:hydrophobicity and pore clogging [J].Soil Sci,1999,164(1):18-27.
[61]王雪芹,张元明,张伟民,等.古尔班通古特沙漠生物结皮对地表风蚀作用的影响的风洞实验[J].冰川冻土,2004,26(5):632-637.
[62] Kinnel P.I.A,et a1. The effects of fire on the soil in a degraded semiarid woodland II:susceptibility 0fthesoilto erosion by shallow rain impacted flow[J]. Australia Journal of Soil Research, 1990,28(40):755-777.
[63] Greene RSB,Tonway DJ,The significance of surface physical and chemical properties in determining soil surface condition of red-earths in rangeland[J]. Australia Journal of Soil Research, 1989,27:213-225.
[64]张振国,焦菊英,白文娟.黄土丘陵沟壑区退耕地植被恢复中生物土壤结皮特征[J].水土保持通报,2006,26(4):33-37.
[65] Eldrldge DJ,Greene RSB.Microbiotic soil crusts:A view of the roles in soil and ecological processes in the range lands of Australia [J].Australia Journal of Soil Research,1994,32:389-415.
[66]宋阳,严平,张宏,等.荒漠生物结皮研究中的几个问题[J].干旱区研究,2004,21(4):439-443.
[67]吴楠.古尔班通古特沙漠生物中微生物分布的日控差异[J].科学通报,2006,51(z1) :100-107.
[68]赵允格,许明祥,王全九,等.黄土丘陵区退耕地生物结皮理化性状初报[J].应用生态学报,2006,l7(8) :1429-1434.
[69]张元明,曹同,潘伯荣.干旱半干旱地区苔藓植物生态学研究综述[J].生态学报,2002,22(7):1129-1134.
[70]徐杰,白学良,田桂泉,等.干旱半干旱地区生物结皮层藓类植物氨基酸和营养物质组成特征及适应性分析[J].生态学报,2005,25(6):1247-1255.
[71]牛玉璐,曹永胜.生物结皮及其在荒漠治理中的作用[J].生物学教学,2005,30(10):5-6.
[72]王世冬,白学良,雍世鹏.沙坡头地区苔藓区系初步研究[J].中国沙漠,2001,9(3):244-249.
[73] Stanley C J,Buxton R. Developing protocols for utilizing moss in revegetation of the Grasberg Mine overburden.http://www.hortresearch.co.nz/product/ncl/reports.
[74] Rochefort L.Sphagnum a keystone genus in habitat restoration [J].The Bryologist,2000,103(3):503-508.
[75]赵建成,黄士良,李敏,等.小扭口藓(Barbula indica)芽孢发育特征的实验研究[J].植物研究,2005,25(2):169-172.
[76]刘晓红.细胞分裂素、氯丙嗪对葫芦藓发育过程的影响[J].南京师范大学学报(自然科学版),1998,23(4):476-480.
[77]高永超,沙伟,张晗.不同植物生长物质对牛角藓愈伤组织诱导的影响[J] .植物生理学通讯,2003,39(1):29-32.
[78]刘世彪,陈军,李菁,等.光照和温度对尖叶拟船叶藓孢子萌发及原丝体发育的影响[J] .西北植物学报,2003,23(1):101-106.
[79]张健,刘国彬,许明祥,等.黄土丘陵区影响生物结皮退化因素的初步研究[J] .中国水土保持科学,2008,12(6):14-20.
[80]黄昌勇.土壤学[M].北京:中国农业出版,2000.
[81]白文娟,焦菊英,张振国.黄土丘陵沟壑区退耕地土壤种子库对植被恢复的影响[J].北京林业大学学报,2008,30(4):65-71.
[82]赵允格,徐冯楠,许明祥.黄土丘陵区藓结皮生物量测定方法及其随发育年限的变化[J].西北植物学报,2008,28(6):1228-1232.
[83]罗应婷,杨钰娟.SPSS统计分析从基础到实践[M].北京:电子工业出版社,2007.
[84]齐红岩,李天来,陈元宏,等.叶面喷施磷酸二氢钾与葡萄糖对番茄光合速率和蔗糖代谢的影响[J].农业工程学报,2005,12:137-142.
[85]刘奎彬,马文荷,安志信.黄瓜对外源糖吸收和运转的研究[J].华北农学,1998,13(4):103-106.
[86]马文苛,刘奎彬,安志信.青椒对外源糖的吸收与分配规律[J].河北农业大学学报,2000,1(23):37-39
[87]浙江农业大学主编.植物营养与肥料[M].北京:中国农业出版社,2004
[88]李延,刘星辉,庄卫民.植物Mg素营养生理的研究进展[J].福建农业大学学报,2000,29(1):74-80.
[89] Glime,Janice M. NUTRIENT RELATIONS:REQUIREMENTS. in Glime,Janice M. Bryophyte Ecology. Volume 1. Physiological Ecology[M]. Michigan Technological University and the International Association of Bryologists,2007.
[90]朱显漠.黄绵土. http://www.chinabaike.com/article/316/327/2007/2007022260712.html.2009.5.7
[91]陈邦杰.中国藓类植物属志上册[M].北京:科学出版社,1963.
[92]刘应迪,朱杰英,陈军,等.3种藓类植物水分含量与光合作用、呼吸作用和水势的关系[J] .武汉植物研究,2001,19(2):135-142.
[93] Martin B.Developmental physiology bryophytes [J].New Mann Bryol,1984,2,2276-2324.
[94]李琴琴,白学良,任向宇.沙漠区生物结皮层中藓类植物繁殖体发育实验研究[J].中国沙漠,2008,3 (28) .289-294.
[95]邢建民,赵德修,李茂寅.水母雪莲悬浮培养细胞生长和黄酮类活性成分合成[J].植物学报1998,40(9):836-841.
[96]高永超,薛红,沙伟.蔗糖对牛角藓愈伤组织悬浮细胞的生理学影响[J].广西植物,2003,23(5):464-469.
[97] Takami S, Yasunaga M, Takio S et al.Establishment of Suspension Cultures of Cells from the Hornwort Anthoceros Punctatusl [M]. J Hattori Bot lab, 1988.
[98]王柳,张福墁,魏秀菊.不同氮肥水平对日光温室黄瓜品质和产量的影响[J].农业工程学报,2007,23(12):225-229.
[99]赵营,同延安,赵护兵.不同供氮水平对夏玉米养分累积、转运及产量的影响[J].植物营养与肥料学报,2006,12(5):622-627.
[100]陈日远,刘厚诚,宋传珍,等.氮素营养对芥蓝生长和品质的影响[J].农业工程学报,2005,12 (21):143-146.
[101]冀瑞萍,张妙娟,温学.水体氮素对藻类植物的影响[J].晋中学院学报,2006,23(6):41-43.
[102]梅赶年.怎样施用磷酸二氢钾[J].新农业,1985,(11):20-26.
[103]李晓鸣.矿质镁对水稻产量及品质影响的研究[J].植物营养与肥料学报,2002,8(1):125-126.
[104]陈圆圆,郭水良,曹同.藓类植物的无性繁殖及其应用[J] .生态学杂志,2008,27(6):993-998.