香菇优良杂交菌株的初步选育
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摘要
本试验以香菇(Lentinula edodes)4个栽培菌株[武香一号(W1)、L205、秋6、L271]和2个野生菌株(SHL12、SHL44)为试验材料,根据优势互补的原则形成4对杂交组合,采用单-单及单-双两种方式进行杂交,依据吃料速度和漆酶活性测定,筛选获得87个杂交子进行出菇试验,运用灰色关联度分析方法对供试菌种8个主要性状值[菌盖直径、菌盖厚度、菌柄长度、菌柄直径、平均单菇鲜重、单袋平均鲜菇产量、单袋平均菇数、出菇期]进行综合评价。从44个正常出菇的杂交菌株中,初步筛选出综合性状较优良的6个杂交菌株(XZ-3、XZ-8、XZ-40、XZ-74、XZ-81和XZ-87)进入中试。
运用IRAP标记对杂交组合W1×SHL12的39个杂交子进行了遗传分析。采用扩增重复性好、谱带清晰的12对引物,扩增共获得128条带,其中127个多态性条带,多态性比例为99.2%。根据引物扩增图谱,采用软件NTSYS2.10e进行聚类分析并计算菌株间的遗传相似系数。结果显示,所有供试菌株在0.69的水平上聚为两类。第Ⅰ类菌株中,除XZ-27、XZ-28之外,均为单-单杂交获得的杂交子；第Ⅱ类菌株中,除XZ-5、XZ-23之外,均为单-双杂交获得的杂交子。菌株XZ-10与XZ-11出菇很少,个体较大,在0.88相似水平上聚为一类。XZ-3、XZ-8出菇数量适中,个大肉厚,产量远高于两亲本,与XZ-2在0.77水平上聚为一类。XZ-19、XZ-20、XZ-27出菇较少,个体较小,菌袋易吐黄水,在0.83水平上聚为一类。XZ-5、XZ-23两个菌株均未出菇,在0.83水平上聚为一类。XZ-7与XZ-17的菌丝均能长满菌袋,但栽培袋不起瘤状物,亦不转色,在0.86水平上聚为一类。试验结果表明,IRAP分析能较好地反映香菇种质的遗传多样性,依据IRAP指纹图谱进行的聚类分析,可用于香菇优良杂交菌株的初步筛选中,从而进一步提高杂交育种工作的效率。
采用TRAP、ISSR、SSR、RAPD、SRAP分子标记的方法,对杂交组合中6个亲本菌株(武香一号、L205、秋6、L271、SHL12、SHL44)和55个单核体亲本(武香一号单核体16个、L205单核体7个、秋6单核体3个、L271单核体10个、SHL12单核体9个、SHL44单核体10个)进行遗传分析。28对扩增重复性好、谱带清晰的引物,在W1×SHL12组合中共扩增出912个条带,多态性条带数为850个,占总条带数的93.2%；在L205×SHLl2组合中共扩增出924条带,多态性条带数为877条,占总条带数的94.9%；在L205×SHL44组合中共扩增出863条带,多态性条带数为806条,占总条带数的93.4%；在秋6×L271组合中共扩增出780条带,多态性条带数为700条,占总条带数的89.7%。聚类结果很好的反映了各菌株间的亲缘关系,但亲本间的遗传距离与栽培试验中正常出菇的44个杂交子主要农艺性状值及灰色关联度法分析结果间的相关性均较低。用分子标记估算亲本遗传距离与杂种后代综合性状间的相关性存在诸多不确定性,不仅受遗传背景的影响,在一定程度上也受环境因子的影响。
In this study, according to the principle of heterosis, test-strains of four cultivation strains W1, L205, Q6,271 and 2 wild strains SHL12, SHL44 were selected to form four pairs of complementary crossing combination, hybridized by the technique of Mon-Mon crossing and Di-Mon crossing breeding. Then according to the speed of Mycelial growth and laccase activity,87 hybrids were Selected from them. The method of grey correlation degree analysis was applicated to comprehensively analysis the 8 main agronomic characters of test-strains, which were pileus diameter, pileus thinkness, stipe length, stipe diameter, average fresh weight of a single fruitbody, average fresh yield of fruit body per bag, average numbers of fruitbody per bag and fruiting period. Six more excellent strains (XZ-3, XZ-8, XZ-40, XZ-74, XZ-81, XZ-87) selected from 44 normal fruiting of hybrid strains were further studied in the middle test.
IRAP marker was applied to analyze the genetic diversity of 39 hybrid strains from the W1×SHL12 cross combination.12 pairs of IRAP primers were selected and a total of 128 fragments were attained, among which 127 fragments (rate of 99.2%) were polymorphic. Based on amplification map, software NTSYS2.10e was used to calculate the genetic similarity and analyze cluster results between the tested strains. The results show that, all the tested strains were clusted into 2 groups at 0.69 similarity level. the strains of all Mon-Mon crossings except XZ-27, XZ-28 were put into the first group, and all Di-Mon crossings except XZ-5, XZ-23 into the second group. XZ-10 and XZ-11 had few but big fruitbody, so they could be put into one group at 0.88 similarity level. XZ-3 and XZ-8 had big fruitbody, higher yield than their parents, clusted into one group with XZ-2 at 0.77 similarity level. XZ-19, XZ-20, XZ-27 had few and small fruitbody, cultivation bags of which were apt to spit out yellow water, classified into one group at 0.83 similarity level. XZ-5, XZ-23 have no fruitbody, classified into one group at 0.83 similarity level. XZ-7, XZ-17 covered with mycelium in bags completely, which had no tumor and no turn color, were clusted into one group at 0.86 similarity level. The results of IRAP analysis could well reflect the genetic diversity of the Lentinula edodes, and could guid for initial screening excellen hybrid strains of L. edodes strains, which could further improve the efficiency of hybrid breeding.
We chosed 6 dicaryotic parents (W1, L205, Q6,271, SHL12, SHL44) and 55 monocaryotic parents (16 monocaryons of W1,7 monocaryons of L205,3 monocaryons of Q6,10 monocaryons of L271,9 monocaryons of SHL12,10 monocaryons of SHL44) as tested strains to analyze their genetic similarity by DNA Fingerprint with TRAP, ISSR, SSR, RAPD, SRAP.28 pairs of primers were selected to amplify the hybrid strains, and a total of 912 fragments were attained from the pair of W1×SHL12, among which 850 fragments were polymorphic, with the rate of 93.2%; 924 fragments from the pair of L205×SHL12, among which 877 fragments were polymorphic, with the rate of 94.9%; 863 fragments from the pair of L205×SHL44, among which 806 fragments were polymorphic, with the rate of 93.4%; 780 fragments from the pair of Q6×L271, among which 700 fragments were polymorphic, with the rate of 89.7%; According to amplification results, amplification products supported the genetic relationship among strains. However, genetic distance and the value of the main agronomic traits and sort results of grey relation degree were not significantly correlated. The method using molecular markers to estimate correlation of genetic distance and traits of hybrids exists many uncertainties, the results of which not only are affected by the genetic background, but also by environmental factors to some extent.
运用IRAP标记对杂交组合W1×SHL12的39个杂交子进行了遗传分析。采用扩增重复性好、谱带清晰的12对引物,扩增共获得128条带,其中127个多态性条带,多态性比例为99.2%。根据引物扩增图谱,采用软件NTSYS2.10e进行聚类分析并计算菌株间的遗传相似系数。结果显示,所有供试菌株在0.69的水平上聚为两类。第Ⅰ类菌株中,除XZ-27、XZ-28之外,均为单-单杂交获得的杂交子；第Ⅱ类菌株中,除XZ-5、XZ-23之外,均为单-双杂交获得的杂交子。菌株XZ-10与XZ-11出菇很少,个体较大,在0.88相似水平上聚为一类。XZ-3、XZ-8出菇数量适中,个大肉厚,产量远高于两亲本,与XZ-2在0.77水平上聚为一类。XZ-19、XZ-20、XZ-27出菇较少,个体较小,菌袋易吐黄水,在0.83水平上聚为一类。XZ-5、XZ-23两个菌株均未出菇,在0.83水平上聚为一类。XZ-7与XZ-17的菌丝均能长满菌袋,但栽培袋不起瘤状物,亦不转色,在0.86水平上聚为一类。试验结果表明,IRAP分析能较好地反映香菇种质的遗传多样性,依据IRAP指纹图谱进行的聚类分析,可用于香菇优良杂交菌株的初步筛选中,从而进一步提高杂交育种工作的效率。
采用TRAP、ISSR、SSR、RAPD、SRAP分子标记的方法,对杂交组合中6个亲本菌株(武香一号、L205、秋6、L271、SHL12、SHL44)和55个单核体亲本(武香一号单核体16个、L205单核体7个、秋6单核体3个、L271单核体10个、SHL12单核体9个、SHL44单核体10个)进行遗传分析。28对扩增重复性好、谱带清晰的引物,在W1×SHL12组合中共扩增出912个条带,多态性条带数为850个,占总条带数的93.2%；在L205×SHLl2组合中共扩增出924条带,多态性条带数为877条,占总条带数的94.9%；在L205×SHL44组合中共扩增出863条带,多态性条带数为806条,占总条带数的93.4%；在秋6×L271组合中共扩增出780条带,多态性条带数为700条,占总条带数的89.7%。聚类结果很好的反映了各菌株间的亲缘关系,但亲本间的遗传距离与栽培试验中正常出菇的44个杂交子主要农艺性状值及灰色关联度法分析结果间的相关性均较低。用分子标记估算亲本遗传距离与杂种后代综合性状间的相关性存在诸多不确定性,不仅受遗传背景的影响,在一定程度上也受环境因子的影响。
In this study, according to the principle of heterosis, test-strains of four cultivation strains W1, L205, Q6,271 and 2 wild strains SHL12, SHL44 were selected to form four pairs of complementary crossing combination, hybridized by the technique of Mon-Mon crossing and Di-Mon crossing breeding. Then according to the speed of Mycelial growth and laccase activity,87 hybrids were Selected from them. The method of grey correlation degree analysis was applicated to comprehensively analysis the 8 main agronomic characters of test-strains, which were pileus diameter, pileus thinkness, stipe length, stipe diameter, average fresh weight of a single fruitbody, average fresh yield of fruit body per bag, average numbers of fruitbody per bag and fruiting period. Six more excellent strains (XZ-3, XZ-8, XZ-40, XZ-74, XZ-81, XZ-87) selected from 44 normal fruiting of hybrid strains were further studied in the middle test.
IRAP marker was applied to analyze the genetic diversity of 39 hybrid strains from the W1×SHL12 cross combination.12 pairs of IRAP primers were selected and a total of 128 fragments were attained, among which 127 fragments (rate of 99.2%) were polymorphic. Based on amplification map, software NTSYS2.10e was used to calculate the genetic similarity and analyze cluster results between the tested strains. The results show that, all the tested strains were clusted into 2 groups at 0.69 similarity level. the strains of all Mon-Mon crossings except XZ-27, XZ-28 were put into the first group, and all Di-Mon crossings except XZ-5, XZ-23 into the second group. XZ-10 and XZ-11 had few but big fruitbody, so they could be put into one group at 0.88 similarity level. XZ-3 and XZ-8 had big fruitbody, higher yield than their parents, clusted into one group with XZ-2 at 0.77 similarity level. XZ-19, XZ-20, XZ-27 had few and small fruitbody, cultivation bags of which were apt to spit out yellow water, classified into one group at 0.83 similarity level. XZ-5, XZ-23 have no fruitbody, classified into one group at 0.83 similarity level. XZ-7, XZ-17 covered with mycelium in bags completely, which had no tumor and no turn color, were clusted into one group at 0.86 similarity level. The results of IRAP analysis could well reflect the genetic diversity of the Lentinula edodes, and could guid for initial screening excellen hybrid strains of L. edodes strains, which could further improve the efficiency of hybrid breeding.
We chosed 6 dicaryotic parents (W1, L205, Q6,271, SHL12, SHL44) and 55 monocaryotic parents (16 monocaryons of W1,7 monocaryons of L205,3 monocaryons of Q6,10 monocaryons of L271,9 monocaryons of SHL12,10 monocaryons of SHL44) as tested strains to analyze their genetic similarity by DNA Fingerprint with TRAP, ISSR, SSR, RAPD, SRAP.28 pairs of primers were selected to amplify the hybrid strains, and a total of 912 fragments were attained from the pair of W1×SHL12, among which 850 fragments were polymorphic, with the rate of 93.2%; 924 fragments from the pair of L205×SHL12, among which 877 fragments were polymorphic, with the rate of 94.9%; 863 fragments from the pair of L205×SHL44, among which 806 fragments were polymorphic, with the rate of 93.4%; 780 fragments from the pair of Q6×L271, among which 700 fragments were polymorphic, with the rate of 89.7%; According to amplification results, amplification products supported the genetic relationship among strains. However, genetic distance and the value of the main agronomic traits and sort results of grey relation degree were not significantly correlated. The method using molecular markers to estimate correlation of genetic distance and traits of hybrids exists many uncertainties, the results of which not only are affected by the genetic background, but also by environmental factors to some extent.
引文
1.边银丙,翁曼丽,孙勇,王斌,赵培新,闵家顺.香菇菌丝太空诱变效应研究--Ⅰ.太空环境对香菇菌丝生长特性的影响.食用菌学报,1999,6(4)：11-14
2.蔡衍山,黄秀治.香菇新菌株Cr-20 Cr-62选育报告.中国食用菌,2000(04)：48-50
3.陈五岭,姚胜利.He-Ne激光在香菇速生高产菌株选育中的应用(Ⅰ).光子学报,1997,26(11)：972-976
4.程水明,林范学,徐学锋,李安政,林芳灿.香菇交配型因子偏分离现象的遗传分析.自然科学进展,2005(04)：259-263
5.邓穗生.灰色关联度对腰果新品种的综合评价.热带作物学报,2004,25(2)：10-14
6.方宣钧,吴为人,唐纪良.作物DNA标记辅助育种.北京：科学出版社,2001,2-21
7.黄年来.中国香菇栽培学.上海：上海科学技术文献出版社,1994,83-113
8.贾身茂,秦森.我国白阿魏蘑的驯化与栽培.中国食用菌,2006,(25)：3-7
9.姜海鹰,陈绍江.高油玉米自交系的杂种优势群划分和优势模式分析.作物学报,2005,31(3)：361-367
10.康德灿,李纯华.食用菌细胞融合新品种金凤2-1的生物学特性研究.绵阳经济技术高等专科学校学报,2000,17(3)：10-12
11.李春生.灰色系统理论在农业推广中的应用研究.[硕士学位论文].武汉：华中农业大学图书馆,2006
12.李省印,李孟楼.平菇种内原生质体分离与融合杂交育种技术的研究.生物技术通讯,2004,15(6)：580-583
13.梁文科,张世煌,戚廷香,邱法展,庹洪章,刘永忠,郑用琏,徐尚忠.热带温带玉米群体产量性状遗传力及遗传方差分量的剖析.中国农业科学,2006,39(11)：2178-2185
14.林范学,程水明,李安政,徐学峰,林芳灿.香菇数量性状的相关性分析和主成分分析.菌物学报,2006,25(4)：579-586
15.林芳灿,高国琪,张晓昱,闵家顺.香菇主要数量性状遗传率及相关性研究.华中农业大学学报,1993,12(1)：27-30
16.林芳灿,张树庭.中国香菇栽培菌株不亲和性因子的分析.华中农业大学学报,1995,14(5)：459-466
17.林俊芳,郭丽琼,王杰,赵风云,张凯,草菇基因工程育种研究.中国菌物学会第四届会员代表大会暨全国第七届菌物学学术讨论会论文集,2008,24-33
18.林俊华,邓优锦,谢宝贵.灵芝原生质体单核化杂交子的评价.福建农林大学学报(自然科学版),2010,39(5)：534-537
19.刘春如,易诚.香菇的营养价值与药用价值.60(01)：52-53
20.刘蕾,宁丽,郭立忠,李翠翠,董伟.12个真姬菇菌株拮抗试验及部分同工酶分析.食用菌,2008(01)：23-26
21.刘录祥,孙其信,王士芸.灰色系统理论应用于作物新品种综合评估初探.中国农业科学,1989,22(3)：22-27
22.罗信昌.木耳和毛木耳的极性研究.真菌学报,1988,7：56-61
23.罗信昌,陈大年.食用真菌核荧光染色技术的研究.真菌学报,1990,9(1)：64-68
24.吕作舟.食用菌栽培学.北京：高等教育出版社,2006,343
25.穆平,张洪亮,姜德峰.利用水、旱稻DH系定位产量性状的QTL及其环境互作分析.中国农业科学,2005,38(9)：1725-1733
26.潘慧.杏鲍菇优良杂交子的初步选育.[硕士学位论文],华中农业大学,2010
27.潘迎捷,陈明杰.香菇和虎皮香菇的种间原生质体融合.上海农业学报,1992,8(1)：9-12
28.潘迎捷,陈明杰,汪昭月,马爱民,凌霞芬,贺东梅,冯志勇,伯海英.单核和同核原生质体技术在食用菌遗传育种上的应用.食用菌学报,1994,1(2)：56-62
29.邱贵根,张诚等.金针菇担孢子核荧光染色观察.江西农业学报.1994,6(2)：153-155
30.石红霞,李卫东,袁会卿.秋栽香菇品种比较试验.食用菌,2002,(04)：38-40
31.宋安东,周丽霞,胡渝,谢霞.产木质素降解酶类香菇菌株的筛选.河南农业大学学报,2006,40(3)：307-310
32.宿红艳,王磊,明永飞,等ISSR分子标记技术在金针菇菌株鉴别中的应用[J].生态学杂志,2008,27(10)：1725-1728.
33.谭琦,潘迎捷,黄为一.中国香菇育种的发展历程.食用菌学报,2000,(04)：48-52
34.潭琦,杨建明,陈明杰,贺冬梅,潘迎捷,黄为一.用RAPD技术对香菇非对称杂交后代遗传性状的分析.食用菌学报,2001,8(2)：10-14.
35.唐传红,张劲松,陈明杰,李泰辉,曹晖.利用拮抗试验和RAPD对灵芝属菌株进行分类研究.微生物学通报,2005,32(5)：72-76
36.王波.香菇原生质体再生的单核体形成与遗传生理特性研究.西南农业学报,1995,8(2)：119-122
37.王俊玲.杏鲍菇杂交亲本的筛选及F1代菌株杂种优势的预测.[硕士学位论文],保定：河北农业大学,2004
38.王凯美,陈再鸣,王涛.糙皮侧耳不育株菌丝细胞学研究.食用菌,2000,(2)：6-7.
39.王磊,蔡德华,宿红艳,张丽,徐圣涛.拮抗实验和RADP对鸡腿菇栽培菌株亲缘关系的研究.安徽农业科学,2006,34(16)：3903-3904
40.王卫科,周祖法,袁卫东,闫静,陆娜.秀珍菇菌株比较试验初报.浙江农业科学,2009,1：50-52
41.王秀萍,张国新,鲁雪林,常青.灰色关联度分析法综合评价水稻新品系.中国农学通报,2006,22(08)：557-558
42.王友华.分子标记预测作物杂种优势的研究进展.种子,2005,24(3)：42-44
43.王玉青.香菇种质资源的鉴别及保藏方法的研究.[博士学位论文].福建：华中农业大学图书馆,2009
44.王泽生,廖剑华,李洪荣,王波,陈美元.中国双孢蘑菇野生菌株的生物学特性研究.首届海峡两岸食(药)用菌学术研讨会论文,2005,24：67-70
45.王卓仁,刘启燕.肖扬,吴茜,边银丙.香菇单孢杂交子代群体灰色关联度和ISSR分析.菌物学报,2010,29(2)：267-272
46.吴康云,边银丙.黑木耳种内杂交子的鉴定技术.菌物系统,2002,21(2)：210-214
47.肖炳光,朱军,卢秀萍,李永平等.烤烟主要农艺性状的遗传与相关分析.遗传,2006,28(3)：317-326
48.徐夙侠.采用AFLP分子标记和分子生物学方法对资源植物的评价.[博士学位论文].中国科学院研究生院,2006
49.许益财,焦言英,李艳秋,于文海,薛建臣,庞玉芬,赵国强.香菇单孢杂交新品种“抚香一号”选育.食用菌,2003,(3)：10-11
50.许颖,兰进.真菌漆酶研究进展.食用菌学报,2006,12(1)：57-64
51.闫培生,边银丙,罗信昌.高等真菌基因工程研究进展.食用菌学报,1997,4(2)：47-53
52.杨岱筠,张丕奇,郑美媛,方子若.白色金针菇原生质体单核菌系的建立及其遗传特性分析.食用菌学报,1995,2(4)：18-21
53.杨汉民主编.细胞生物学实验(第二版).北京：高等教育出版社,1997
54.杨新美.中国食用菌栽培学.北京：农业出版社,1988,167-175
55.杨新美.食用菌研究法,中国农业出版社,1998年
56.于淑池,林泽平.香菇的营养保健及免疫调节作用.承德民族师专学报,2003,(02)：40-43
57.袁力行,傅骏骅,刘新芝,彭泽斌,张世煌,李新海,李连城.利用分子标记预测玉米杂种优势的研究.中国农业科学,2000,33(6)：6-12
58.张红,曹晖,潘迎捷,曹娟云.黑木耳交配型的研究.菌物系统,2002,21：559-564
59.张红梅,吕长武,吕杰,曾宪贤,张军.同工酶技术在食用菌育种研究中的应用.种子,2007,26(7)：44-46
60.张金霞,黄晨阳,陈强.食用菌可栽培种类野生种质的评价.植物遗传资源学报,2010,11(2)：127-131.
61.张瑞颖.香菇菌株多相鉴定鉴别技术研究.[硕士学位论文].北京：中国农业大学图书馆,2004
62.赵斌清,陈耀武,龚凤萍.秋栽代料香菇品种比较试验.食用菌,2007(05)：19-20
63.朱军.估算遗传方差和协方差的混合模型方法.生物数学学报,1992,7(1)：1-11
64.朱军.遗传模型分析方法.中国农业出版社,1997
65.朱军.数量性状遗传分析的新方法及其在育种中的应用.浙江大学学报(农业与生命科学版),2006,26(1)：1-6
66. Arima T, Morinaga T. Electrophoretic karyotype of Lentinus edodes. Trans Mycol Soc Japan,1993,34:481-485
67. Bernardo R. Relationship between single-cross performance and molecular marker heterozygosity. Theor Appl Genet,1992,83:628-634
68. Boppenmaier J, Melchinger A E, Brunklaus June E. Genetic diversity for RFLPs in European maize in-breeds. I. Relation to performance of flint dent crosses for forage trait. Crop Sci.1992, (32):895-902
69. Boppenmaier J, Melchinger A E, Seitz G. Genetic diversity for RFLPs in European maize inbreds Ⅱ:Performance of crosses within versus between heterosic groups for grain traits. Plant Breeding,1993,111:217-226
70. Charcosset A, M L effortBuson, A Gallais. Relationship between heterosis and heterozygosity at marker loci:atheoretical computation. Theory App Genet.1991, (81):571-575
71. Chen X, Stone M, Schlagnhaufer C. A Fruiting body tissue method for efficient agrobacterium-mediated transformation of Agaricus bisporus. Appl Environ Microb, 2000,4510-4513
72. Diez J, Beguiristain T, Tacon F L, Casacuberta J M, Tagu D. Identification of Tyl-copia retrotransposons in three ectomycorrhizal basidiomycetes:evolutionary relationships and use as molecular markers. Curr Genet,2003,43:34-44
73. Dudley J W, SaghaiMaroofM, A, Rufener G. K. Molecular marker and grouping of parents in maize breeding programs. Crop Science,1991,31:718-723
74. G.F.Liu. Genetic analysis of grain yield conditioned on its component traits in rice. Australian Journal of Agricultural Research,2008,59,139-195
75. Godshalk E. B, Lee M, Lamkey K. R. Analysis of the relationship of restriction fragment length polymorphisms to maize single-cross hybrid performance. Theor Appl Genet,1990,80:273-280
76. Guo D, Zhang H, Luo Z. Genetic relationships of Diospyros kaki Thunb. And related species revealed by IRAP and REMAP analysis. Plant Sci.,2006,170:528-533
77. Hibbett DS, Fukumasa NY, Tsuneda A, Donoghue MJ. Phylogenetic diversity in shiitake inferred from nueear ribosomal DNA sequenees. Mycologia,1995,87: 618-638
78. Kalendar R, Grob T, Regina M, Suoniemi A, Schulman A. IRAP and REMAP:Two new retrotransposon-based DNA fingerprinting techniques. Theor Appl Genet,1999, 98(5):704-711
79. Kangatharalingam N, Ferguson M W. A simple and rapid technique for fluorescence staining of fungal nuclei. Curr microbiol,1984,10:99-104
80. Kunamneni A, Plou F J, Ballesteros A. Laccases and their applications:a patent review. Recent Pat Biotechnol,2008,2(1):10-24
81. Kuo CY, Chou SY, Huang CT. Cloning of glyceraldehyde-3-phosphate dehydrogenase gene and use of the gpd promoter for transformation in Flammulina velutipes. Appl. Microbiol Biotechnol,2004, (65):593-599
82. Larraya LM, Alfonso M, Pisabarro AG. Mapping of genomic re-gions (quantitative trait loci) controlling production and quality in industrial cultures of the edible basidiomycete Pleurotus ostreatus. Appl Environ Microb,2003,69(6):3617-3625.
83. M. Nagai T, Sato H, Watanabe K, Saito-M, Kawata H, Enei. Purification and characterization of an extracellular laccase from the edible mushroom Lentinula edodes and decolorization of chemically different dyes. Appl Microbiol Biotechnol, 2002,60:327-335
84. Manninen O, Kalendar R, Robison J, Schulman A. Application of BARE-1 retrotransposon markers to the mapping of major resistance gene for net blotch in barley. Mol Gen Genet,2000,264:325-334
85. May B, Royse DJ. Application of the electrophoretic methodology to the elucidation of genetic life histories of edible fungi. Mushroom Sci.,1981.11:799-817
86. Mufioz-Rivas A, Specht CA, Drummond BJ. Transformation of the basidiomycete Schizophyllum commune. Mol Genet,1986, (205):103-106
87. Muller J. Enzymatic regulation of temperature dependent truit boby morphogenesis in plearotus ostreatus Abstract of X11 th international congress on the science and cultivation of Edible fungi sept,1987,20-26
88. Murata H, Yamada A. marY1. a member of the gypsy group of long terminal repeat retroelements from the ectomycorrhizal basidiomycete Tricholoma matsutake. Appl Environ Microb,2000,66(8):3642-3645
89. Royse DJ, May B. Identification of Shiitake genotypes by multilocus enzyme electrophoresis:catalog of lines. Biochem Genet,1987,25:705-717
90. Sha T, Zhang H, Ding H, Li Z, Cheng L, Zhao Z. Genetic diversity of Tricholoma matsutake in Yunnan Province. Chinese Sci Bull,2007,52(9):1212-1216
91. Smith O. S, Smith J S. C, Bowen S. L. Similarlties among a goup of elite maize as measured by pedigree:F1 grain yield, grain yield heterosis and RFLPs. Theor Appl Genet,1990,80:833-840
92. Smith, J S C, Chin, E C L, Shu. H, Smith O S, wall S J, Senior M L, Mitchell S E, Kresovich S, Zeigle J. An evaluation the utility of SSR loci as molecular markers in maize (Zea mays L.):comparison with data from RFLPs and pedigree. Theor Appl Genet,1997,95:163-173
93. Terashima K, Matsumoto T, Hasebe K. Genetic diversity and strain-typing in cultivated strains of Lentinula edode (the shiitake mushroom) in Japan by AFLP analysis. Mycol Res,2002,106(1):34-39.
94. Thierry N, Labarrere J. Homologous transformation of the edible basidiomycete Agrocybe aegerita with the URAL gene:Characterizition of interative events and of rearranged Plasmids in transformants. Curr Genet.1994,25:432-437
95. Toyomasu T, Zennyozi A. on application of isoenzyme electrophoresis to identification of strains in Lentinus.edodes(Shiitake). Mushroom Sci.,1981,11: 675-684
96. Xiao Y, Dai Y H, Lu Y Y, Liu W, Wang Z R, Bian Y B. Development of IRAP-SCAR markers for strain identification in Lentinula edodes. World Journal of Microbiology and Biotechonology,2010,626-629
97. Yan P S, Jiang J H. Preliminary research of the RAPD molecular marker-assisted breeding of the edible basidiomycete Stropharia rugoso-annulata. World J Microbiol Biotechnol,2005,21:559-563
98. Zhang Z. Q, Zheng Y. L, Wei Y. M. Relationship between Heterosis and Genetic Distance Based on Simple Sequence Repeat Markers in Wheat. Journal of Sichuan Agricultural University,2003,21(1):6-9.
2.蔡衍山,黄秀治.香菇新菌株Cr-20 Cr-62选育报告.中国食用菌,2000(04)：48-50
3.陈五岭,姚胜利.He-Ne激光在香菇速生高产菌株选育中的应用(Ⅰ).光子学报,1997,26(11)：972-976
4.程水明,林范学,徐学锋,李安政,林芳灿.香菇交配型因子偏分离现象的遗传分析.自然科学进展,2005(04)：259-263
5.邓穗生.灰色关联度对腰果新品种的综合评价.热带作物学报,2004,25(2)：10-14
6.方宣钧,吴为人,唐纪良.作物DNA标记辅助育种.北京：科学出版社,2001,2-21
7.黄年来.中国香菇栽培学.上海：上海科学技术文献出版社,1994,83-113
8.贾身茂,秦森.我国白阿魏蘑的驯化与栽培.中国食用菌,2006,(25)：3-7
9.姜海鹰,陈绍江.高油玉米自交系的杂种优势群划分和优势模式分析.作物学报,2005,31(3)：361-367
10.康德灿,李纯华.食用菌细胞融合新品种金凤2-1的生物学特性研究.绵阳经济技术高等专科学校学报,2000,17(3)：10-12
11.李春生.灰色系统理论在农业推广中的应用研究.[硕士学位论文].武汉：华中农业大学图书馆,2006
12.李省印,李孟楼.平菇种内原生质体分离与融合杂交育种技术的研究.生物技术通讯,2004,15(6)：580-583
13.梁文科,张世煌,戚廷香,邱法展,庹洪章,刘永忠,郑用琏,徐尚忠.热带温带玉米群体产量性状遗传力及遗传方差分量的剖析.中国农业科学,2006,39(11)：2178-2185
14.林范学,程水明,李安政,徐学峰,林芳灿.香菇数量性状的相关性分析和主成分分析.菌物学报,2006,25(4)：579-586
15.林芳灿,高国琪,张晓昱,闵家顺.香菇主要数量性状遗传率及相关性研究.华中农业大学学报,1993,12(1)：27-30
16.林芳灿,张树庭.中国香菇栽培菌株不亲和性因子的分析.华中农业大学学报,1995,14(5)：459-466
17.林俊芳,郭丽琼,王杰,赵风云,张凯,草菇基因工程育种研究.中国菌物学会第四届会员代表大会暨全国第七届菌物学学术讨论会论文集,2008,24-33
18.林俊华,邓优锦,谢宝贵.灵芝原生质体单核化杂交子的评价.福建农林大学学报(自然科学版),2010,39(5)：534-537
19.刘春如,易诚.香菇的营养价值与药用价值.60(01)：52-53
20.刘蕾,宁丽,郭立忠,李翠翠,董伟.12个真姬菇菌株拮抗试验及部分同工酶分析.食用菌,2008(01)：23-26
21.刘录祥,孙其信,王士芸.灰色系统理论应用于作物新品种综合评估初探.中国农业科学,1989,22(3)：22-27
22.罗信昌.木耳和毛木耳的极性研究.真菌学报,1988,7：56-61
23.罗信昌,陈大年.食用真菌核荧光染色技术的研究.真菌学报,1990,9(1)：64-68
24.吕作舟.食用菌栽培学.北京：高等教育出版社,2006,343
25.穆平,张洪亮,姜德峰.利用水、旱稻DH系定位产量性状的QTL及其环境互作分析.中国农业科学,2005,38(9)：1725-1733
26.潘慧.杏鲍菇优良杂交子的初步选育.[硕士学位论文],华中农业大学,2010
27.潘迎捷,陈明杰.香菇和虎皮香菇的种间原生质体融合.上海农业学报,1992,8(1)：9-12
28.潘迎捷,陈明杰,汪昭月,马爱民,凌霞芬,贺东梅,冯志勇,伯海英.单核和同核原生质体技术在食用菌遗传育种上的应用.食用菌学报,1994,1(2)：56-62
29.邱贵根,张诚等.金针菇担孢子核荧光染色观察.江西农业学报.1994,6(2)：153-155
30.石红霞,李卫东,袁会卿.秋栽香菇品种比较试验.食用菌,2002,(04)：38-40
31.宋安东,周丽霞,胡渝,谢霞.产木质素降解酶类香菇菌株的筛选.河南农业大学学报,2006,40(3)：307-310
32.宿红艳,王磊,明永飞,等ISSR分子标记技术在金针菇菌株鉴别中的应用[J].生态学杂志,2008,27(10)：1725-1728.
33.谭琦,潘迎捷,黄为一.中国香菇育种的发展历程.食用菌学报,2000,(04)：48-52
34.潭琦,杨建明,陈明杰,贺冬梅,潘迎捷,黄为一.用RAPD技术对香菇非对称杂交后代遗传性状的分析.食用菌学报,2001,8(2)：10-14.
35.唐传红,张劲松,陈明杰,李泰辉,曹晖.利用拮抗试验和RAPD对灵芝属菌株进行分类研究.微生物学通报,2005,32(5)：72-76
36.王波.香菇原生质体再生的单核体形成与遗传生理特性研究.西南农业学报,1995,8(2)：119-122
37.王俊玲.杏鲍菇杂交亲本的筛选及F1代菌株杂种优势的预测.[硕士学位论文],保定：河北农业大学,2004
38.王凯美,陈再鸣,王涛.糙皮侧耳不育株菌丝细胞学研究.食用菌,2000,(2)：6-7.
39.王磊,蔡德华,宿红艳,张丽,徐圣涛.拮抗实验和RADP对鸡腿菇栽培菌株亲缘关系的研究.安徽农业科学,2006,34(16)：3903-3904
40.王卫科,周祖法,袁卫东,闫静,陆娜.秀珍菇菌株比较试验初报.浙江农业科学,2009,1：50-52
41.王秀萍,张国新,鲁雪林,常青.灰色关联度分析法综合评价水稻新品系.中国农学通报,2006,22(08)：557-558
42.王友华.分子标记预测作物杂种优势的研究进展.种子,2005,24(3)：42-44
43.王玉青.香菇种质资源的鉴别及保藏方法的研究.[博士学位论文].福建：华中农业大学图书馆,2009
44.王泽生,廖剑华,李洪荣,王波,陈美元.中国双孢蘑菇野生菌株的生物学特性研究.首届海峡两岸食(药)用菌学术研讨会论文,2005,24：67-70
45.王卓仁,刘启燕.肖扬,吴茜,边银丙.香菇单孢杂交子代群体灰色关联度和ISSR分析.菌物学报,2010,29(2)：267-272
46.吴康云,边银丙.黑木耳种内杂交子的鉴定技术.菌物系统,2002,21(2)：210-214
47.肖炳光,朱军,卢秀萍,李永平等.烤烟主要农艺性状的遗传与相关分析.遗传,2006,28(3)：317-326
48.徐夙侠.采用AFLP分子标记和分子生物学方法对资源植物的评价.[博士学位论文].中国科学院研究生院,2006
49.许益财,焦言英,李艳秋,于文海,薛建臣,庞玉芬,赵国强.香菇单孢杂交新品种“抚香一号”选育.食用菌,2003,(3)：10-11
50.许颖,兰进.真菌漆酶研究进展.食用菌学报,2006,12(1)：57-64
51.闫培生,边银丙,罗信昌.高等真菌基因工程研究进展.食用菌学报,1997,4(2)：47-53
52.杨岱筠,张丕奇,郑美媛,方子若.白色金针菇原生质体单核菌系的建立及其遗传特性分析.食用菌学报,1995,2(4)：18-21
53.杨汉民主编.细胞生物学实验(第二版).北京：高等教育出版社,1997
54.杨新美.中国食用菌栽培学.北京：农业出版社,1988,167-175
55.杨新美.食用菌研究法,中国农业出版社,1998年
56.于淑池,林泽平.香菇的营养保健及免疫调节作用.承德民族师专学报,2003,(02)：40-43
57.袁力行,傅骏骅,刘新芝,彭泽斌,张世煌,李新海,李连城.利用分子标记预测玉米杂种优势的研究.中国农业科学,2000,33(6)：6-12
58.张红,曹晖,潘迎捷,曹娟云.黑木耳交配型的研究.菌物系统,2002,21：559-564
59.张红梅,吕长武,吕杰,曾宪贤,张军.同工酶技术在食用菌育种研究中的应用.种子,2007,26(7)：44-46
60.张金霞,黄晨阳,陈强.食用菌可栽培种类野生种质的评价.植物遗传资源学报,2010,11(2)：127-131.
61.张瑞颖.香菇菌株多相鉴定鉴别技术研究.[硕士学位论文].北京：中国农业大学图书馆,2004
62.赵斌清,陈耀武,龚凤萍.秋栽代料香菇品种比较试验.食用菌,2007(05)：19-20
63.朱军.估算遗传方差和协方差的混合模型方法.生物数学学报,1992,7(1)：1-11
64.朱军.遗传模型分析方法.中国农业出版社,1997
65.朱军.数量性状遗传分析的新方法及其在育种中的应用.浙江大学学报(农业与生命科学版),2006,26(1)：1-6
66. Arima T, Morinaga T. Electrophoretic karyotype of Lentinus edodes. Trans Mycol Soc Japan,1993,34:481-485
67. Bernardo R. Relationship between single-cross performance and molecular marker heterozygosity. Theor Appl Genet,1992,83:628-634
68. Boppenmaier J, Melchinger A E, Brunklaus June E. Genetic diversity for RFLPs in European maize in-breeds. I. Relation to performance of flint dent crosses for forage trait. Crop Sci.1992, (32):895-902
69. Boppenmaier J, Melchinger A E, Seitz G. Genetic diversity for RFLPs in European maize inbreds Ⅱ:Performance of crosses within versus between heterosic groups for grain traits. Plant Breeding,1993,111:217-226
70. Charcosset A, M L effortBuson, A Gallais. Relationship between heterosis and heterozygosity at marker loci:atheoretical computation. Theory App Genet.1991, (81):571-575
71. Chen X, Stone M, Schlagnhaufer C. A Fruiting body tissue method for efficient agrobacterium-mediated transformation of Agaricus bisporus. Appl Environ Microb, 2000,4510-4513
72. Diez J, Beguiristain T, Tacon F L, Casacuberta J M, Tagu D. Identification of Tyl-copia retrotransposons in three ectomycorrhizal basidiomycetes:evolutionary relationships and use as molecular markers. Curr Genet,2003,43:34-44
73. Dudley J W, SaghaiMaroofM, A, Rufener G. K. Molecular marker and grouping of parents in maize breeding programs. Crop Science,1991,31:718-723
74. G.F.Liu. Genetic analysis of grain yield conditioned on its component traits in rice. Australian Journal of Agricultural Research,2008,59,139-195
75. Godshalk E. B, Lee M, Lamkey K. R. Analysis of the relationship of restriction fragment length polymorphisms to maize single-cross hybrid performance. Theor Appl Genet,1990,80:273-280
76. Guo D, Zhang H, Luo Z. Genetic relationships of Diospyros kaki Thunb. And related species revealed by IRAP and REMAP analysis. Plant Sci.,2006,170:528-533
77. Hibbett DS, Fukumasa NY, Tsuneda A, Donoghue MJ. Phylogenetic diversity in shiitake inferred from nueear ribosomal DNA sequenees. Mycologia,1995,87: 618-638
78. Kalendar R, Grob T, Regina M, Suoniemi A, Schulman A. IRAP and REMAP:Two new retrotransposon-based DNA fingerprinting techniques. Theor Appl Genet,1999, 98(5):704-711
79. Kangatharalingam N, Ferguson M W. A simple and rapid technique for fluorescence staining of fungal nuclei. Curr microbiol,1984,10:99-104
80. Kunamneni A, Plou F J, Ballesteros A. Laccases and their applications:a patent review. Recent Pat Biotechnol,2008,2(1):10-24
81. Kuo CY, Chou SY, Huang CT. Cloning of glyceraldehyde-3-phosphate dehydrogenase gene and use of the gpd promoter for transformation in Flammulina velutipes. Appl. Microbiol Biotechnol,2004, (65):593-599
82. Larraya LM, Alfonso M, Pisabarro AG. Mapping of genomic re-gions (quantitative trait loci) controlling production and quality in industrial cultures of the edible basidiomycete Pleurotus ostreatus. Appl Environ Microb,2003,69(6):3617-3625.
83. M. Nagai T, Sato H, Watanabe K, Saito-M, Kawata H, Enei. Purification and characterization of an extracellular laccase from the edible mushroom Lentinula edodes and decolorization of chemically different dyes. Appl Microbiol Biotechnol, 2002,60:327-335
84. Manninen O, Kalendar R, Robison J, Schulman A. Application of BARE-1 retrotransposon markers to the mapping of major resistance gene for net blotch in barley. Mol Gen Genet,2000,264:325-334
85. May B, Royse DJ. Application of the electrophoretic methodology to the elucidation of genetic life histories of edible fungi. Mushroom Sci.,1981.11:799-817
86. Mufioz-Rivas A, Specht CA, Drummond BJ. Transformation of the basidiomycete Schizophyllum commune. Mol Genet,1986, (205):103-106
87. Muller J. Enzymatic regulation of temperature dependent truit boby morphogenesis in plearotus ostreatus Abstract of X11 th international congress on the science and cultivation of Edible fungi sept,1987,20-26
88. Murata H, Yamada A. marY1. a member of the gypsy group of long terminal repeat retroelements from the ectomycorrhizal basidiomycete Tricholoma matsutake. Appl Environ Microb,2000,66(8):3642-3645
89. Royse DJ, May B. Identification of Shiitake genotypes by multilocus enzyme electrophoresis:catalog of lines. Biochem Genet,1987,25:705-717
90. Sha T, Zhang H, Ding H, Li Z, Cheng L, Zhao Z. Genetic diversity of Tricholoma matsutake in Yunnan Province. Chinese Sci Bull,2007,52(9):1212-1216
91. Smith O. S, Smith J S. C, Bowen S. L. Similarlties among a goup of elite maize as measured by pedigree:F1 grain yield, grain yield heterosis and RFLPs. Theor Appl Genet,1990,80:833-840
92. Smith, J S C, Chin, E C L, Shu. H, Smith O S, wall S J, Senior M L, Mitchell S E, Kresovich S, Zeigle J. An evaluation the utility of SSR loci as molecular markers in maize (Zea mays L.):comparison with data from RFLPs and pedigree. Theor Appl Genet,1997,95:163-173
93. Terashima K, Matsumoto T, Hasebe K. Genetic diversity and strain-typing in cultivated strains of Lentinula edode (the shiitake mushroom) in Japan by AFLP analysis. Mycol Res,2002,106(1):34-39.
94. Thierry N, Labarrere J. Homologous transformation of the edible basidiomycete Agrocybe aegerita with the URAL gene:Characterizition of interative events and of rearranged Plasmids in transformants. Curr Genet.1994,25:432-437
95. Toyomasu T, Zennyozi A. on application of isoenzyme electrophoresis to identification of strains in Lentinus.edodes(Shiitake). Mushroom Sci.,1981,11: 675-684
96. Xiao Y, Dai Y H, Lu Y Y, Liu W, Wang Z R, Bian Y B. Development of IRAP-SCAR markers for strain identification in Lentinula edodes. World Journal of Microbiology and Biotechonology,2010,626-629
97. Yan P S, Jiang J H. Preliminary research of the RAPD molecular marker-assisted breeding of the edible basidiomycete Stropharia rugoso-annulata. World J Microbiol Biotechnol,2005,21:559-563
98. Zhang Z. Q, Zheng Y. L, Wei Y. M. Relationship between Heterosis and Genetic Distance Based on Simple Sequence Repeat Markers in Wheat. Journal of Sichuan Agricultural University,2003,21(1):6-9.