吉林省野生果树菌根的调查研究
摘要
果树的菌根，即果树感染有有益的真菌并相互共生的根。果树根的组织与真菌形成双重的有机体，真菌依靠果树供给碳素营养，果树依靠真菌从土壤中吸取果树根系不能吸收的水分和养分。一般果树的根系都和真菌共生。菌根对于增加果树矿质营养，提高果树抗旱、耐盐碱、抗病等抗逆能力，促进果树生长具有重要作用。菌根除显著提高果树叶片的N素、P素营养外，还可显著提高K、Mg、Cu、Zn、Ca、Fe的营养。在抗旱方面，菌根一方面能显著地提高土壤的供水能力，另一方面菌根还能通过帮助植株建立发达的根系来提高树体的吸水能力，使植株避免干旱的威胁。菌根还能提高果树抵抗土传害虫的能力，又可促进果树生长几倍至几十倍。由此可看出，菌根在生产实践中具有广泛应用前景。因此，开发利用菌根真菌，将成为今后果树生产上高产、优质、高效建设的重要新技术。
本实验以吉林省野生果树根系为实验材料，采用石蜡制片、光学显微技术及Phillips和Hayman的染色方法，对其进行结构植物学研究。结果发现，直径＞2mm的木质化根系即根的次生结构中不存在VA菌根。也就是说，VA菌根不能侵染木质化的粗根。显然是由于皮层细胞的木质化阻碍了真菌的侵染。
通过对吉林省18种野生果树菌根的解剖研究，论述了它们在形态结构上的特点，并调查其侵染率。
根据菌物寄生的部位不同，目前的研究将菌根分为三类，即外生菌根、内生菌根和内外生菌根。外生菌根是指菌物的菌丝只寄附于植物根部的皮层外表，并不深入皮层以下的柔膜细胞中；内生菌根是指菌物的菌丝深入到皮层下的柔膜细胞之中，并长成许多分枝状的菌丝，它又可分为VA菌根、杜鹃类菌根和兰科菌根；内外生菌根则兼有外生和内生菌根的特点。本实验调查的吉林省18种野生果树菌根大多属于内生菌根中的VA菌根，即泡囊—丛枝菌根。VA菌根在形态结构上既具相同点，又有各自的独特之处。研究发现，在树莓、杨梅、沙棘菌根中可观察到泡囊。泡囊形成在初生皮层的细胞间和细胞内，形状通常呈卵形、椭圆
形、圆形。沙棘泡囊形状比较特殊，多为纺锤形、球形。在蓝靛果忍冬、山杏中
既可观察到菌根的泡囊，又可观察到菌根的丛枝。丛枝是由菌丝侵入寄生细胞经
过连续的二分叉而形成的一种树枝状结构。在杨梅中可发现带有泡囊的内生菌
丝。内生菌丝无隔，在根的皮层细胞内为纵向或横向伸延，形成大量的分枝。泡
囊、丛枝、内生菌丝均是存在于皮层组织内。不同果树的菌根泡囊形态有差异，
说明它可能被多种V人菌根真菌侵染。不同菌种对不同果树的作用效果不同，因
而菌根形态各异。各种果树菌根的相似之处在于，大多数菌根多存在于皮层细胞
内，形状多为球形、椭圆形、不规则形。菌根数目也不尽相同，且在山梨和山葡
萄中发现有晶体的存在。在山梨皮层薄壁组织中是簇状晶体，山葡萄中是针簇状
晶体，这两种晶体在形态上易区别，以此作为区别植物的解剖特征之一，棒子中
没有发现VA菌根的存在。vA菌根侵染率是菌根化的一个重要指标。本实验对吉
林省野生果树VA菌根自然侵染率进行调查。结果表明:不同果树种类的VA菌根
侵染率相差极大。其中以沙棘、蓝靛果忍冬、软枣桥猴桃的侵染率较高，在60-
700/6左右，沙棘的侵染率达到76.67%;越桔、树毒、山梨、山杏、山萄萄、杨梅、
山喳海棠、黑加仑、稠李、山丁子次之，在30一50%左右;山里红、山樱桃、
五味子较低，只占10一20y0左右。结果还表明:同一品种的果树的不同地点取样，
\’A菌根侵染率也存在差异。山里红、山杏、稠李在靖宇取样的侵染率高于左家，
证实了菌根侵染率除与种类不同而异外，还有地域性的差异。这种差异与立地条
件有关。即使是同一地点同一果树的不同品种，它们侵染率也存在差异。越桔6
个品种中，北空越桔比H。越桔高于20%以上，证实了菌根真菌对不同品种的果树侵
染有所不同。了解果树各种和品种菌根侵染率，以此作为应用菌根可能性的前提。
本文针对各种野生果树菌根的结构特征，进行综合比较研究。研究结果，一
方面为菌根资源的发现提供了依据。另一方面也为果树菌根的分类、发展规律的
研究莫定了基础，对于从事菌根形态功能相关及菌根应用研究具有一定的指导意
义。
Mycorrhizas of fruit-trees, which are the roots that fruit-trees infect some valuable funguses & grow with them symbiotically, Organizations of fruit-trees & funguses can form organism; funguses rely on fruit-trees to provide carbon nutrition; accordingly, fruit-trees rely on funguses to absorb water & nutrition that the roots fail to. Generally speaking, the root family of fruit-trees & funguses co-exist. Funguses can raise nutrition of K, Mg, Cu, Zn, Ca, & Fe as well as that of N element & P element, can gain mineral nutrition of fruit-trees, improve resistance to drought salinization & diseases& promote the growth of fruit-trees. As far as the resistance to drought is concerned, funguses can, on one hand, enlarge providing-water capacity of soil; on the other hand, raise absorbing-water capacity of trees by helping the trees develop perfect root families against threat from drought. Also, funguses can improve the ability of resistance to insects, mean while, promote the growth of fruit-trees several even
dozens of times. In one word, funguses application in production & practice has insignificant prospect. So, to develop & make the best of funguses will be a prominent technique in high output, high quality & high efficiency. From the discovery, we can see that there aren't any mycorrhiza in the woodized root family with diameter more than 2mm or in the secondary structure of roots, that is to say ,VA mycorrhizas can't infect the thick woodized roots, Obviously,it is because the woodization of cortical cells keep funguses from being infected .
The experiment has been done by using the root family of wild fruit-trees in Jilin province as experimental materials; paraffin wax-made slide technique, microscopy, Phillips & Hayman coloration are employed to carry out research into morphological botany.
The thesis explores the formal characteristics of the mycorrhizas in Jilin province & infection rate through dissection research on them.
According to the different parasitic parts of funguses, mycorrhizas can be
III
classified into three parts, namely externally-born mycorrhiza, internally-born mycorrhiza and externally-born internally-born mycorrhiza. The first one refers to the hyphae attached to the cortex; the second one refers to the hyphae stretching into soft mucous membrance of cortex and forming lots of branch-like hyphae which can be divided into VA mycorrhiza. azalea mycorrhiza & orchid family mycorrhiza . The third one has the features of the former two. Most mycorrhizas of seventeen kinds of wild fruit-trees investigated in the experiment in Jilin province are VA mycorrhizas, that is vesicular-arbuscular mycorrhizas, which are similar to or different from each other in structure. From the discovery, vesiculars are found in mycorrhizas ofRubus idaeus, Hippophae rhammoides, which emerge in the cortical cells as well as between the cells with shapes in sphere.oval & ovum forms. Hippophae rhammoides has such particular shapes as spindle and ball forms. Vesiculars &arbusculars are found in Lonicera deulis, Armenica vw/gtfra,Arbusculars are a Kind of branch-like structure with hyphae infecting parasitic cells, which extend in length and breadth in the cortical structure of roots & form many branches Vesicuars, arbusculars & internally-born hyphae are in the cortical organization. The formation varies from one vesicular of mycorrhizas to another, which illustrates the fact that it may be infected by many kinds of VA mycorrhizas. The effect of different funguses on different trees varies,thus, they can't have the same formation. The similarity of mycorrhizas of different fruit-trees is that. Most of them are in cortical cells of initial structure of roots, which are in such shapes as sphere, oval or irregular form. The number of them are different, besides, there exist crystals in pyrus betulaefolia Bge & vitis amurensis Rupr. There exist pile crystals in the cortical organization of Pyrus betulaefolia while pin-pile crystal found in Vitis amurensis.The two kinds of crystals are distinguished Easi
本实验以吉林省野生果树根系为实验材料，采用石蜡制片、光学显微技术及Phillips和Hayman的染色方法，对其进行结构植物学研究。结果发现，直径＞2mm的木质化根系即根的次生结构中不存在VA菌根。也就是说，VA菌根不能侵染木质化的粗根。显然是由于皮层细胞的木质化阻碍了真菌的侵染。
通过对吉林省18种野生果树菌根的解剖研究，论述了它们在形态结构上的特点，并调查其侵染率。
根据菌物寄生的部位不同，目前的研究将菌根分为三类，即外生菌根、内生菌根和内外生菌根。外生菌根是指菌物的菌丝只寄附于植物根部的皮层外表，并不深入皮层以下的柔膜细胞中；内生菌根是指菌物的菌丝深入到皮层下的柔膜细胞之中，并长成许多分枝状的菌丝，它又可分为VA菌根、杜鹃类菌根和兰科菌根；内外生菌根则兼有外生和内生菌根的特点。本实验调查的吉林省18种野生果树菌根大多属于内生菌根中的VA菌根，即泡囊—丛枝菌根。VA菌根在形态结构上既具相同点，又有各自的独特之处。研究发现，在树莓、杨梅、沙棘菌根中可观察到泡囊。泡囊形成在初生皮层的细胞间和细胞内，形状通常呈卵形、椭圆
形、圆形。沙棘泡囊形状比较特殊，多为纺锤形、球形。在蓝靛果忍冬、山杏中
既可观察到菌根的泡囊，又可观察到菌根的丛枝。丛枝是由菌丝侵入寄生细胞经
过连续的二分叉而形成的一种树枝状结构。在杨梅中可发现带有泡囊的内生菌
丝。内生菌丝无隔，在根的皮层细胞内为纵向或横向伸延，形成大量的分枝。泡
囊、丛枝、内生菌丝均是存在于皮层组织内。不同果树的菌根泡囊形态有差异，
说明它可能被多种V人菌根真菌侵染。不同菌种对不同果树的作用效果不同，因
而菌根形态各异。各种果树菌根的相似之处在于，大多数菌根多存在于皮层细胞
内，形状多为球形、椭圆形、不规则形。菌根数目也不尽相同，且在山梨和山葡
萄中发现有晶体的存在。在山梨皮层薄壁组织中是簇状晶体，山葡萄中是针簇状
晶体，这两种晶体在形态上易区别，以此作为区别植物的解剖特征之一，棒子中
没有发现VA菌根的存在。vA菌根侵染率是菌根化的一个重要指标。本实验对吉
林省野生果树VA菌根自然侵染率进行调查。结果表明:不同果树种类的VA菌根
侵染率相差极大。其中以沙棘、蓝靛果忍冬、软枣桥猴桃的侵染率较高，在60-
700/6左右，沙棘的侵染率达到76.67%;越桔、树毒、山梨、山杏、山萄萄、杨梅、
山喳海棠、黑加仑、稠李、山丁子次之，在30一50%左右;山里红、山樱桃、
五味子较低，只占10一20y0左右。结果还表明:同一品种的果树的不同地点取样，
\’A菌根侵染率也存在差异。山里红、山杏、稠李在靖宇取样的侵染率高于左家，
证实了菌根侵染率除与种类不同而异外，还有地域性的差异。这种差异与立地条
件有关。即使是同一地点同一果树的不同品种，它们侵染率也存在差异。越桔6
个品种中，北空越桔比H。越桔高于20%以上，证实了菌根真菌对不同品种的果树侵
染有所不同。了解果树各种和品种菌根侵染率，以此作为应用菌根可能性的前提。
本文针对各种野生果树菌根的结构特征，进行综合比较研究。研究结果，一
方面为菌根资源的发现提供了依据。另一方面也为果树菌根的分类、发展规律的
研究莫定了基础，对于从事菌根形态功能相关及菌根应用研究具有一定的指导意
义。
Mycorrhizas of fruit-trees, which are the roots that fruit-trees infect some valuable funguses & grow with them symbiotically, Organizations of fruit-trees & funguses can form organism; funguses rely on fruit-trees to provide carbon nutrition; accordingly, fruit-trees rely on funguses to absorb water & nutrition that the roots fail to. Generally speaking, the root family of fruit-trees & funguses co-exist. Funguses can raise nutrition of K, Mg, Cu, Zn, Ca, & Fe as well as that of N element & P element, can gain mineral nutrition of fruit-trees, improve resistance to drought salinization & diseases& promote the growth of fruit-trees. As far as the resistance to drought is concerned, funguses can, on one hand, enlarge providing-water capacity of soil; on the other hand, raise absorbing-water capacity of trees by helping the trees develop perfect root families against threat from drought. Also, funguses can improve the ability of resistance to insects, mean while, promote the growth of fruit-trees several even
dozens of times. In one word, funguses application in production & practice has insignificant prospect. So, to develop & make the best of funguses will be a prominent technique in high output, high quality & high efficiency. From the discovery, we can see that there aren't any mycorrhiza in the woodized root family with diameter more than 2mm or in the secondary structure of roots, that is to say ,VA mycorrhizas can't infect the thick woodized roots, Obviously,it is because the woodization of cortical cells keep funguses from being infected .
The experiment has been done by using the root family of wild fruit-trees in Jilin province as experimental materials; paraffin wax-made slide technique, microscopy, Phillips & Hayman coloration are employed to carry out research into morphological botany.
The thesis explores the formal characteristics of the mycorrhizas in Jilin province & infection rate through dissection research on them.
According to the different parasitic parts of funguses, mycorrhizas can be
III
classified into three parts, namely externally-born mycorrhiza, internally-born mycorrhiza and externally-born internally-born mycorrhiza. The first one refers to the hyphae attached to the cortex; the second one refers to the hyphae stretching into soft mucous membrance of cortex and forming lots of branch-like hyphae which can be divided into VA mycorrhiza. azalea mycorrhiza & orchid family mycorrhiza . The third one has the features of the former two. Most mycorrhizas of seventeen kinds of wild fruit-trees investigated in the experiment in Jilin province are VA mycorrhizas, that is vesicular-arbuscular mycorrhizas, which are similar to or different from each other in structure. From the discovery, vesiculars are found in mycorrhizas ofRubus idaeus, Hippophae rhammoides, which emerge in the cortical cells as well as between the cells with shapes in sphere.oval & ovum forms. Hippophae rhammoides has such particular shapes as spindle and ball forms. Vesiculars &arbusculars are found in Lonicera deulis, Armenica vw/gtfra,Arbusculars are a Kind of branch-like structure with hyphae infecting parasitic cells, which extend in length and breadth in the cortical structure of roots & form many branches Vesicuars, arbusculars & internally-born hyphae are in the cortical organization. The formation varies from one vesicular of mycorrhizas to another, which illustrates the fact that it may be infected by many kinds of VA mycorrhizas. The effect of different funguses on different trees varies,thus, they can't have the same formation. The similarity of mycorrhizas of different fruit-trees is that. Most of them are in cortical cells of initial structure of roots, which are in such shapes as sphere, oval or irregular form. The number of them are different, besides, there exist crystals in pyrus betulaefolia Bge & vitis amurensis Rupr. There exist pile crystals in the cortical organization of Pyrus betulaefolia while pin-pile crystal found in Vitis amurensis.The two kinds of crystals are distinguished Easi
引文
[1]鹿金颖，毛永民，申连英，等，果树VA菌根研究进展，河北农业大学学报，1999，2(4)：50～51
[2]胡又厘，中国的果树菌根研究进展，福建农业大学学报，1998，27(1) 52～61
[3]江春祥，常绿果树菌根的研究Ⅰ菌根资源及形态特征[学士论文]，福建农业大学，1995
[4]牛家琪．广东省VA菌根真菌资源调查和应用研究．土壤学报，1994，31(增刊)：54～63
[5]吴铁航，郝文英，林先贵，等，红壤中VA菌根真菌侵染，及接种效应的研究，应用生态学报，1993，4(1)：53～58
[6]苏俐英，天目山自然保护区林木菌根调查，浙江林学院学报，1992，9(3)：263～276
[7]周冲权，吴光林．黄寿波．果树VA菌根侵染率及影响因素的研究，中国果树，1992，(1)：22～24
[8]刘润进：薛炳烨，黄镇，等．山东果树VA菌根调查，山东农业大学学报，1987，18(4)：25～31
[9]秦岭，董清华，郑来友．园艺作物菌根研究进展，北京农学院学报，1995，(2)：101～105
[10]Gnekow M, Marschner H. Role of VA-mycorrhiza in growrh and mineral nutrition of apple (malus pumila var. domestica rootstock cuttings. Plant and Soil.1989, 119(2):285～293
[11]冯固，杨茂秋，白灯莎，等，VA菌根真菌对石灰性土壤不同形态磷酸盐有效性的影响，植物营养与肥料学报，1997，3(2)：43～48
[12]唐振尧，何首林，菌根促进柑桔吸收难溶性P肥的机理研究．中国柑桔，1991，20(2)：7～10
[13]刘润进，VA菌根对湖北海棠实生苗水分状况的影响．莱阳农学院学报，1989，6(1)：34～39
[14]刘润进，高秋莲，生兆江，等．杜梨菌根苗几种培养方法的试验，莱阳农学院学报，1990，7(4)：298～300
[15]CamprubiA, Calvet C.Isolation and screening ofmycorrhizal fungi from citrus nurseries and orchards and inoculation studies Hort Science, 1996, (31):366～369
[16]郭秀珍，李江山，毕国昌．泡囊丛枝状(VA)菌根真菌对葡萄组培苗的生长效应．园艺学报，1988，15(2)：77～82
[17]Geddeda YI,Trappe JM, Stebbins RL.Effects of vesicular-arbuscular mycorrhizae and phosphorus on apple seedlings. Journal of the American Society for Horticultural Science, 1984, (109):24～27
[18]Covey R P,koch B L,Larsen H J.Influence of vesicular-arbuscular mycorrhizae on growth of apple and cron m low-phosphorous Soil.phytopathology, 1981, 71: 712～715
[19]Koch B L,covey R P.Larsen H J.Response of apple seedlings in fumigated soil tophosphorus and vesicular-arbuscular mycorrhiza Hort Science, 1982,17:232～233
[20]Morin F, Fortin J A. Apple rootstock response to vesicular-arbuscular Mycorrhizal. fungi in a high phosphorus soil. J Amer Soc Hort Sci, 1994, 119(3):578～583
[21]安志强，泡囊一丛枝(VA)菌根(球囊霉属Glomusspp)对八楞海棠(Malus micromalus Mark)实生苗生长和矿质营养的影响：[硕士学位论文]，中国农业大学图书馆，1986
[22]Morin F, Fortin L A, Hamel C, et al. Apple rootstock response to vesicular-arbuscular myeorrhizal fungi in a high phosphorus soil. Journal of the American society for Horticultural Science, 1994(l19):578～583
[23]齐国辉，VA菌根对苹果、葡萄组培苗生理效应的研究：[学位论文]保定：河北农业大学，1996
[24]薛炳烨，罗新书，VAM真菌对再植桃实生苗生长的作用．果树科学，1992，9(2)：106～109
[25]刘润进，罗新书．VA菌根对中国樱桃实生苗生长和养分的影响．莱阳农学院学报，1988，5(1)：6～13
[26]黄万荣，秦岭，刘振亮．碱地上VA菌根苹果苗的生长发现，中国果树，1995(4)：23～2
[27]霍书新，张小红．菌根在果树生产中的应用，北方园艺，1997(4)：58～59
[28]杨兴洪，罗新书，刘润进．利用菌根真菌解决苹果重茬问题．落叶果树，1992(4)：5～7
[29]齐国辉，陈贵林，吕桂云，等．丛枝菌根菌在草莓上的应用试验．河北果树，2001(1)：14～16
[30]姚青，李道高，石井道高，菌根真菌对相桔果汁成分和果皮着色的影响．果树科学，1999，16(1)：38～42
[31]Phillips, J.M. et al. Trans. Br. Mycol. Soc, 55:138～161,1970
[32]毛永民，范培格，贾立清，等，枣树菌根侵染率的田间调查，河北农业大学学报，1999，22(4)：55～57
[33]张梦昌，李悦书，汪矛，等．北细辛、蓝靛果忍冬及软枣猕猴桃VA菌根的调查研究．吉林农业大学学报，1988，10(4)：59～452
[34]Holtett, B.P. et al.:Forest Science, 1976, 22(2):127～130
[35]Nappi P. Effect of methods of soil treatment and seasons on vesicular-arbuscular mycorrhizae of apple Micologia Italiana, 1983,12(12):39～46
[36]李国景，吴光林，黄寿波，等，不同土壤管理方式对金柑VA菌根侵染率的影响．中国柑桔，1992 7 21(2)：5～7
[37]Masuhara G, Kimura S, Katsuya K. Seasonal changes in the mycorrhizae of Bletilla striata(Orchideae).Transaction of the Mycological Society of Japan 1988, 29(1):25～31
[38]Abbott L K, Robson A D. Growth subteranean clover in velation to the farmation of endo-mycorrhizas by introduced and indigeners furgi in a
field soil. New Phytol, 1978, 81:575～587
[39]Sander F E, Tinker P B, Black R L, Palmerley SM. The derelopment of endomycorrhizal root system I spread of infection and growth promoting effect with four spetes of vesicular-arbuscular. New phytol, 1977,78:257～268
[40]Geddeda Y I, Trappe J M, Stebbins R L. Effect Vesicular-arbuscular mycorrhizae and phosphorus on apple seedlings. Amer Soc Hort Sci, 1984, 109:24～27
[41]Daniels B A,,Menge J A. Eraluation of the commercial potential of the vesicular-arbusular mycorrhizal fungus Glomus epigaeum. New phytol, 1981,87:345～354
[42]Miller D B, Bodmer M, Schuepp H. Spread of endomycorrhizal, colonization and effects on growth of apple seedlings. New phytol, 1989, 111:51～59
[43]Abbott L K, Robson A D.Formation of external hyphae in soil by four species of vesicular-arbuscular mycorrhizal fungi. New phytol, 1985, 99:245～255
[44]刘润进，高秋莲，韩殿一．梨树VA菌根形态结构观察．莱阳农学学报，1989，6(3)：34～38
[45]罗新书，刘润进．苹果砧穗组合习性的研究．园艺，1982，2：6～10
[46]王春梅，谷文祥．VA菌根对苹果苗生长及矿质营养的影响．河北农业技术师范学院学报，1999，13(1)：68～72
[47]张美庆，王幼珊，黄磊．我国北部的八种VA菌根．真菌学报，19927 11(4)：258～267
[48]张美庆，王幼珊．我国北部的七种菌根真菌．真菌学报，1999，10(1)：13～21
[49]Manju G, Lakhanpal TN, Gupta M. Identification of mycorrhizal types mycorrhizosphere mycoflora and vesicular-arbuscular myeorrhizal (VAM) fungi in apple. Annals of Biology Ludhiana 1993, 9(1):87～93
[50]王立军，软枣猕猴桃的解剖学研究．吉林农业大学学报，1990，12(4)：34～38
[51]王立军，长白山区几种野生和栽培越桔菌根的初步研究．东北师范大学学报，1995，12,104～105
[52]景汝勤．中华猕猴桃的解剖学研究．西北农业大学学报，1981(3)：39～49
[53]Metcalfe CR et al. Anatomy of the dicotyledons. Vol I.clarendon press, Oxford. 1957, 191～193
[54]刘润进，李晓林编著．丛枝菌根及其应用．北京：科学出版社，2000
[55]Hoepfner E F, koch B L.Core R P.Enancement of growth and phosphorus concentration in apple seedling by vesicular-arbuscular mycorrhizae. J Amer Soc Hort Sci, 1983, 105:207～209
[56]Miller D D, Domoto P A,Walker. Colonization and efficacy of different endomycorrhizal fungi with apple seedlings at two phosphorus levels. New phytol, 1958, 100:393～402
[57]Johnson C R, Menge J A, Schwab S.Ting I P.Interaction of photoperiod and vesicular-arbuscular mycorrhizae on growth and metabolism of sweet orange. New phytol, 1982, 90:665～667
[58]盖京苹，刘润进．野生植物根围的丛枝菌根真菌I．菌物系统，2000，19(1)：25～29
[59]吴铁航，郝文英，林先贵，等，红壤中VA菌根真菌(球襄霉目)的种类和生态分布．真菌学报，1994，14(2)：81～85
[60]刘润进，沈祟尧，裘维蕃，关于VAM菌与黄萎病菌存在侵染中的竞争作用．第六届全国菌根学术会议(保定)，1993
[61]刘润进，中国盐碱土壤中VAM菌的生态分布，应用生态学报，1999，10(6)：721～724
[62]Michelini, S.,1989,Acta Horticulturae,239:427～430
[63]Rayner, M.C.,1933, Nature, 131:399
[64]Reed, H.S. and Fremont, S.,1934, Phytopathology, 25:634～645
[65]Mller, H.R.A.M, 1936, Landbouw, 12:110
[66]Neill, J.C.,1944, J. Sci.& Techo. A.,25:192～200
[67] Gerdemann, J. W., 1968, Annu. Rer. Phytipathol. 6:397～418
[68] 李亚东，郝瑞，张大鹏．越桔菌根真菌研究进展—文献综述．园艺学报，1(2)：133～138
[69] 郭秀珍，毕国昌，林木菌根及应用技术．中国林业出版社，1989
[70] 邢晓科，李玉．吉林省参地中的10种VA菌根真菌，吉林农业大学学报，2000，22 (2)：41～46
[71] 迟丽华，王立军，刘洪章，等．VA菌根与果树矿质营养(综述)．吉林农业大学学报，2002，24(2)：66～69
[2]胡又厘，中国的果树菌根研究进展，福建农业大学学报，1998，27(1) 52～61
[3]江春祥，常绿果树菌根的研究Ⅰ菌根资源及形态特征[学士论文]，福建农业大学，1995
[4]牛家琪．广东省VA菌根真菌资源调查和应用研究．土壤学报，1994，31(增刊)：54～63
[5]吴铁航，郝文英，林先贵，等，红壤中VA菌根真菌侵染，及接种效应的研究，应用生态学报，1993，4(1)：53～58
[6]苏俐英，天目山自然保护区林木菌根调查，浙江林学院学报，1992，9(3)：263～276
[7]周冲权，吴光林．黄寿波．果树VA菌根侵染率及影响因素的研究，中国果树，1992，(1)：22～24
[8]刘润进：薛炳烨，黄镇，等．山东果树VA菌根调查，山东农业大学学报，1987，18(4)：25～31
[9]秦岭，董清华，郑来友．园艺作物菌根研究进展，北京农学院学报，1995，(2)：101～105
[10]Gnekow M, Marschner H. Role of VA-mycorrhiza in growrh and mineral nutrition of apple (malus pumila var. domestica rootstock cuttings. Plant and Soil.1989, 119(2):285～293
[11]冯固，杨茂秋，白灯莎，等，VA菌根真菌对石灰性土壤不同形态磷酸盐有效性的影响，植物营养与肥料学报，1997，3(2)：43～48
[12]唐振尧，何首林，菌根促进柑桔吸收难溶性P肥的机理研究．中国柑桔，1991，20(2)：7～10
[13]刘润进，VA菌根对湖北海棠实生苗水分状况的影响．莱阳农学院学报，1989，6(1)：34～39
[14]刘润进，高秋莲，生兆江，等．杜梨菌根苗几种培养方法的试验，莱阳农学院学报，1990，7(4)：298～300
[15]CamprubiA, Calvet C.Isolation and screening ofmycorrhizal fungi from citrus nurseries and orchards and inoculation studies Hort Science, 1996, (31):366～369
[16]郭秀珍，李江山，毕国昌．泡囊丛枝状(VA)菌根真菌对葡萄组培苗的生长效应．园艺学报，1988，15(2)：77～82
[17]Geddeda YI,Trappe JM, Stebbins RL.Effects of vesicular-arbuscular mycorrhizae and phosphorus on apple seedlings. Journal of the American Society for Horticultural Science, 1984, (109):24～27
[18]Covey R P,koch B L,Larsen H J.Influence of vesicular-arbuscular mycorrhizae on growth of apple and cron m low-phosphorous Soil.phytopathology, 1981, 71: 712～715
[19]Koch B L,covey R P.Larsen H J.Response of apple seedlings in fumigated soil tophosphorus and vesicular-arbuscular mycorrhiza Hort Science, 1982,17:232～233
[20]Morin F, Fortin J A. Apple rootstock response to vesicular-arbuscular Mycorrhizal. fungi in a high phosphorus soil. J Amer Soc Hort Sci, 1994, 119(3):578～583
[21]安志强，泡囊一丛枝(VA)菌根(球囊霉属Glomusspp)对八楞海棠(Malus micromalus Mark)实生苗生长和矿质营养的影响：[硕士学位论文]，中国农业大学图书馆，1986
[22]Morin F, Fortin L A, Hamel C, et al. Apple rootstock response to vesicular-arbuscular myeorrhizal fungi in a high phosphorus soil. Journal of the American society for Horticultural Science, 1994(l19):578～583
[23]齐国辉，VA菌根对苹果、葡萄组培苗生理效应的研究：[学位论文]保定：河北农业大学，1996
[24]薛炳烨，罗新书，VAM真菌对再植桃实生苗生长的作用．果树科学，1992，9(2)：106～109
[25]刘润进，罗新书．VA菌根对中国樱桃实生苗生长和养分的影响．莱阳农学院学报，1988，5(1)：6～13
[26]黄万荣，秦岭，刘振亮．碱地上VA菌根苹果苗的生长发现，中国果树，1995(4)：23～2
[27]霍书新，张小红．菌根在果树生产中的应用，北方园艺，1997(4)：58～59
[28]杨兴洪，罗新书，刘润进．利用菌根真菌解决苹果重茬问题．落叶果树，1992(4)：5～7
[29]齐国辉，陈贵林，吕桂云，等．丛枝菌根菌在草莓上的应用试验．河北果树，2001(1)：14～16
[30]姚青，李道高，石井道高，菌根真菌对相桔果汁成分和果皮着色的影响．果树科学，1999，16(1)：38～42
[31]Phillips, J.M. et al. Trans. Br. Mycol. Soc, 55:138～161,1970
[32]毛永民，范培格，贾立清，等，枣树菌根侵染率的田间调查，河北农业大学学报，1999，22(4)：55～57
[33]张梦昌，李悦书，汪矛，等．北细辛、蓝靛果忍冬及软枣猕猴桃VA菌根的调查研究．吉林农业大学学报，1988，10(4)：59～452
[34]Holtett, B.P. et al.:Forest Science, 1976, 22(2):127～130
[35]Nappi P. Effect of methods of soil treatment and seasons on vesicular-arbuscular mycorrhizae of apple Micologia Italiana, 1983,12(12):39～46
[36]李国景，吴光林，黄寿波，等，不同土壤管理方式对金柑VA菌根侵染率的影响．中国柑桔，1992 7 21(2)：5～7
[37]Masuhara G, Kimura S, Katsuya K. Seasonal changes in the mycorrhizae of Bletilla striata(Orchideae).Transaction of the Mycological Society of Japan 1988, 29(1):25～31
[38]Abbott L K, Robson A D. Growth subteranean clover in velation to the farmation of endo-mycorrhizas by introduced and indigeners furgi in a
field soil. New Phytol, 1978, 81:575～587
[39]Sander F E, Tinker P B, Black R L, Palmerley SM. The derelopment of endomycorrhizal root system I spread of infection and growth promoting effect with four spetes of vesicular-arbuscular. New phytol, 1977,78:257～268
[40]Geddeda Y I, Trappe J M, Stebbins R L. Effect Vesicular-arbuscular mycorrhizae and phosphorus on apple seedlings. Amer Soc Hort Sci, 1984, 109:24～27
[41]Daniels B A,,Menge J A. Eraluation of the commercial potential of the vesicular-arbusular mycorrhizal fungus Glomus epigaeum. New phytol, 1981,87:345～354
[42]Miller D B, Bodmer M, Schuepp H. Spread of endomycorrhizal, colonization and effects on growth of apple seedlings. New phytol, 1989, 111:51～59
[43]Abbott L K, Robson A D.Formation of external hyphae in soil by four species of vesicular-arbuscular mycorrhizal fungi. New phytol, 1985, 99:245～255
[44]刘润进，高秋莲，韩殿一．梨树VA菌根形态结构观察．莱阳农学学报，1989，6(3)：34～38
[45]罗新书，刘润进．苹果砧穗组合习性的研究．园艺，1982，2：6～10
[46]王春梅，谷文祥．VA菌根对苹果苗生长及矿质营养的影响．河北农业技术师范学院学报，1999，13(1)：68～72
[47]张美庆，王幼珊，黄磊．我国北部的八种VA菌根．真菌学报，19927 11(4)：258～267
[48]张美庆，王幼珊．我国北部的七种菌根真菌．真菌学报，1999，10(1)：13～21
[49]Manju G, Lakhanpal TN, Gupta M. Identification of mycorrhizal types mycorrhizosphere mycoflora and vesicular-arbuscular myeorrhizal (VAM) fungi in apple. Annals of Biology Ludhiana 1993, 9(1):87～93
[50]王立军，软枣猕猴桃的解剖学研究．吉林农业大学学报，1990，12(4)：34～38
[51]王立军，长白山区几种野生和栽培越桔菌根的初步研究．东北师范大学学报，1995，12,104～105
[52]景汝勤．中华猕猴桃的解剖学研究．西北农业大学学报，1981(3)：39～49
[53]Metcalfe CR et al. Anatomy of the dicotyledons. Vol I.clarendon press, Oxford. 1957, 191～193
[54]刘润进，李晓林编著．丛枝菌根及其应用．北京：科学出版社，2000
[55]Hoepfner E F, koch B L.Core R P.Enancement of growth and phosphorus concentration in apple seedling by vesicular-arbuscular mycorrhizae. J Amer Soc Hort Sci, 1983, 105:207～209
[56]Miller D D, Domoto P A,Walker. Colonization and efficacy of different endomycorrhizal fungi with apple seedlings at two phosphorus levels. New phytol, 1958, 100:393～402
[57]Johnson C R, Menge J A, Schwab S.Ting I P.Interaction of photoperiod and vesicular-arbuscular mycorrhizae on growth and metabolism of sweet orange. New phytol, 1982, 90:665～667
[58]盖京苹，刘润进．野生植物根围的丛枝菌根真菌I．菌物系统，2000，19(1)：25～29
[59]吴铁航，郝文英，林先贵，等，红壤中VA菌根真菌(球襄霉目)的种类和生态分布．真菌学报，1994，14(2)：81～85
[60]刘润进，沈祟尧，裘维蕃，关于VAM菌与黄萎病菌存在侵染中的竞争作用．第六届全国菌根学术会议(保定)，1993
[61]刘润进，中国盐碱土壤中VAM菌的生态分布，应用生态学报，1999，10(6)：721～724
[62]Michelini, S.,1989,Acta Horticulturae,239:427～430
[63]Rayner, M.C.,1933, Nature, 131:399
[64]Reed, H.S. and Fremont, S.,1934, Phytopathology, 25:634～645
[65]Mller, H.R.A.M, 1936, Landbouw, 12:110
[66]Neill, J.C.,1944, J. Sci.& Techo. A.,25:192～200
[67] Gerdemann, J. W., 1968, Annu. Rer. Phytipathol. 6:397～418
[68] 李亚东，郝瑞，张大鹏．越桔菌根真菌研究进展—文献综述．园艺学报，1(2)：133～138
[69] 郭秀珍，毕国昌，林木菌根及应用技术．中国林业出版社，1989
[70] 邢晓科，李玉．吉林省参地中的10种VA菌根真菌，吉林农业大学学报，2000，22 (2)：41～46
[71] 迟丽华，王立军，刘洪章，等．VA菌根与果树矿质营养(综述)．吉林农业大学学报，2002，24(2)：66～69