中国石蒜属植物叶部微形态特征的研究
|
摘要
本文首先综述了石蒜属植物的研究进展及分类系统学上存在的问题，包括石蒜属种质资源的开发利用、人工栽培和组织培养、生理、形态、染色体、药用和药理、观赏价值以及分子方面的研究。其中，通过形态、染色体、分子方面对石蒜属植物种间分类系统进行了探讨，在石蒜属分类系统方面存在的主要问题有：(1)细胞学与分子系统学上的证据较为一致，而与形态学上的证据存在差异，因此，能否以花冠整齐与否将石蒜属划分为两大亚属值得商榷；(2)石蒜属植物的杂合体较多，这些种的系统位置仍存在分歧；(3)微形态证据探讨石蒜属分类系统方面研究较为缺乏。因此，必须综合形态学、解剖学、细胞学、孢粉学和分子生物学等各方面的性状指标，才能够建立更合理的石蒜属植物分类系统，并绘制出完整的石蒜属分支系统树。本文通过叶表皮的电镜扫描、光镜观察和叶部的石蜡切片对中国石蒜属12种植物叶部微形态特征进行研究，试图为探讨石蒜属植物的系统演化关系提供实验证据和有效性状。研究结果表明：叶表皮的气孔排列方式、长度、长宽比，脊的明显与否，叶表皮上的颗粒纹饰的多少、大小，叶表皮细胞长度、长宽比、垂周壁形状，上部叶栅栏组织细胞层数、叶的维管束数、叶中脉导管组成数目，气腔类型，叶的脊部远轴面的角数，叶缘的形状等这些特征在种间存在差异，同时，在有些种之间又有相似性；因此，这些特征既可以作为区分种的证据，又是研究石蒜属植物系统演化的性状指标，根据这些性状指标，可以将石蒜属植物大致分为两个亚属：一是忽地笑、中国石蒜、短蕊石蒜、安徽石蒜、长筒石蒜，二是江苏石蒜、乳白石蒜、石蒜、玫瑰石蒜、稻草石蒜；在两个亚属中其中有些种如忽地笑、长筒石蒜、中国石蒜的亲缘关系则更近些。换锦花、鹿葱有些性状与一个亚属较近，另一些性状与另一个亚属较近，因此，是过渡类型。
In the first, the research development and problems in the taxonomical systematics of Lycoris were summarized in the article, including the research of germplasma resources and exploitations, artificial cultivation, tissue culture, physiology, morphology, pharmacology, decorative values and molecule. The classification system among species of Lycoris was discussed through morphology, chromosome, molecule. The systematical relationship among 12 species of Lycoris has been discussed. The problems in the taxonomical systematics are as follows: (l)The results of cytology are different from that of molecule, but results of karyotype analycis and that of RAPD are same .So it is seem too hard to divide Lycoris two subgenus according to corolla regularity. (2)The views on systematical tree of those species of Lycoris which have fairly more hybrid body are different . (3)The research of Lycoris on the systematical relationship of micro-morphology and anatomy is absent. It is essential to generalize the character inde
xes of morphology, anatomy, cytology, palynology and molecular physiology. So We can establish classification system of Lycoris and draw a reasonable family tree of Lycoris. Based on the reseaching of Lycoris leaf micro-morphology characters by SEM scanning, light microscope viewing and using paraffin method in the article,we try to provide the proofs of experiment and the efffective characer indexes to discuss the system and evolution of the Lycoris.we find these microscope
characers through reseach are as follows: the arrangement type ,density and the length of stoma; the shape and length of the epidermal cells; the ridge of epidermis; the transverse shape of leaf; the number of the vascular bundle ; the style of vessel arrangement of vascular bundle in the middle vein ; the layers of palisade tissue ; the ratio of the thicken of palisade tissue to that of spongy tissue ; the angle number of the middle leaf; the type of ventilating cavity etc. These characters are different from those of species in Lycoris, meanwhile these characters are similar to those of some species in Lycoris. So these characters can be used as the character indexes for differentiating species and reseaching systematic evolution of Lycoris; We can divide Lycoris into two subgenus by these characteristic indexes: the species of one subgenus are L. aurea (L' Her.) Herb,, chinensis Traub, L. caldweUii Traub, L. anhuiensis Y. Hsu et Q. J. Fan, L. longituba Y. Hsu et Q. J. Fan ; the species of another subgenu
s are L. houdyshelii Traub, L. albiflora Koidz., L. rosea Traub et Moldenke, L. straminea Lindl. Meanwhile, some characters of L. sprengeri Comes ex Baker and L. squamigera Maxim, are close to the genetic relationship among one subgenus and other characters are close to that among the another subgenus . Two species may be transitional taxa.
In the first, the research development and problems in the taxonomical systematics of Lycoris were summarized in the article, including the research of germplasma resources and exploitations, artificial cultivation, tissue culture, physiology, morphology, pharmacology, decorative values and molecule. The classification system among species of Lycoris was discussed through morphology, chromosome, molecule. The systematical relationship among 12 species of Lycoris has been discussed. The problems in the taxonomical systematics are as follows: (l)The results of cytology are different from that of molecule, but results of karyotype analycis and that of RAPD are same .So it is seem too hard to divide Lycoris two subgenus according to corolla regularity. (2)The views on systematical tree of those species of Lycoris which have fairly more hybrid body are different . (3)The research of Lycoris on the systematical relationship of micro-morphology and anatomy is absent. It is essential to generalize the character inde
xes of morphology, anatomy, cytology, palynology and molecular physiology. So We can establish classification system of Lycoris and draw a reasonable family tree of Lycoris. Based on the reseaching of Lycoris leaf micro-morphology characters by SEM scanning, light microscope viewing and using paraffin method in the article,we try to provide the proofs of experiment and the efffective characer indexes to discuss the system and evolution of the Lycoris.we find these microscope
characers through reseach are as follows: the arrangement type ,density and the length of stoma; the shape and length of the epidermal cells; the ridge of epidermis; the transverse shape of leaf; the number of the vascular bundle ; the style of vessel arrangement of vascular bundle in the middle vein ; the layers of palisade tissue ; the ratio of the thicken of palisade tissue to that of spongy tissue ; the angle number of the middle leaf; the type of ventilating cavity etc. These characters are different from those of species in Lycoris, meanwhile these characters are similar to those of some species in Lycoris. So these characters can be used as the character indexes for differentiating species and reseaching systematic evolution of Lycoris; We can divide Lycoris into two subgenus by these characteristic indexes: the species of one subgenus are L. aurea (L' Her.) Herb,, chinensis Traub, L. caldweUii Traub, L. anhuiensis Y. Hsu et Q. J. Fan, L. longituba Y. Hsu et Q. J. Fan ; the species of another subgenu
s are L. houdyshelii Traub, L. albiflora Koidz., L. rosea Traub et Moldenke, L. straminea Lindl. Meanwhile, some characters of L. sprengeri Comes ex Baker and L. squamigera Maxim, are close to the genetic relationship among one subgenus and other characters are close to that among the another subgenus . Two species may be transitional taxa.
引文
1．鲍海鸥、陈波红、石蒜属植物资源的开发利用，中国野生植物奖资源，1995，19(5)：31-32
2．陈必义、杜仁华、曾繁典等，二氢石蒜碱的降压作用，中国药理学报，1993，14(1)：45
3．陈耀华、李懋学，四种石蒜属植物的染色体核型研究，园艺学报，1985，12(1)：57-60
4．陈宝梁．叶片解剖特征在木莲属Manglietia BL．分种的应用[J]．中山大学研究生学刊，1989，10(2)：52-55
5．邓传良等，石蒜单染色的显微分离及体外扩增，南京林业大学学报，2002，26(3)：72-74
6．董国良，石蒜外治妙用，中医外治杂志，1995，6：31-32
7．董庆华，石蒜的组织培养，植物生理学通讯，1995(5)：54
8．额特曼(中国科学院植物研究所古植物研究室孢粉组译)，孢粉学手册，北京：科学出版社，1978，88，278-279，339，286-287
9．访台北县淡水镇农会花卉研究班谈，“金花石蒜，台湾花卉，1984，57-59
10．龚新荣等，二氢石蒜碱对大鼠高碳酸性脑缺氧损伤的保护作用，郧阳医学院学报，2002，21(2)：71-72
11．韩锦隆，金花石蒜天然开花期之变温及人工花期调节方法之研究，宜兰农工学报，1991，10：105-121
12．胡庆和等，石蒜内铵盐(AT-1840)综合治疗妇科晚期恶性肿瘤，浙江中西医结合杂志，1997，7，(3)：134-136
13．黄宗春、胡一民，安徽野生石蒜的种质资源及开发利用，中国林副特产，1997．3：46-47
14．柯丽霞、孙叶根、郑艳等，石蒜属三种植物的核型研究，安徽师范大学学报，1998，21(4)：343-348
15．蓝星莲，二氢石蒜碱对家名胜脑血管的选择性作用，中国药理学通报，2002，18(4)：409-411
16．李朝銮：蔷薇科原始属植物叶表皮解剖及其系统学意义．植物分类学报，1989．27(3)：178-183．
17．李荣义．中国水仙．福建科技出版社，1991．
18．李卫东，石蒜的人工栽培成功，厦门晚报，1996，12/21，第四版
19．李建强等．赤爮亚族植物叶片中脉的比较解剖．云南植物研究，1992，14(4)：418-422
20．梁素秋，金花石蒜芽分化及体外培养之探讨，国立台湾大学园艺研究所硕士论文，1992
21．林纯英、马溯轩，金花石蒜之鳞片组织培养毓，中国园艺(台湾)，1987，33(4)：255-264、
22．林定勇，石蒜属球根花卉之分类、形态、生长与开花，中国园艺(台湾)，1993，39(2)：67-72
23．刘顺俊、刘四维，单味石蒜治疗疔疮肿毒，湖北中医杂志，1996，18(4/12
24．刘德仪等．中国木兰属和含笑属导管分子的比较解剖．植物学报，1987，29(1)：22-28
25．刘玉壶等．木兰科分类系统的初步研究[J]．植物分类学报，1984，22(2)：89-109
26．刘琰，徐炳声：石蒜属的核型研究，1989．植物分类学报，27(4)：257-264．
27．鲁雪花、陈扬春，忽地笑胚外植体的培养，云南植物研究，1986，8(4)：467-469
28．吕海亮，吴世安，杨继，饶广远：叶表皮及种皮特征在黄精族系统学研究中的应用，2000．植物
分类学报，38(1)：30-42．
29．吕海亮，吴世安，杨继，饶广远：叶表皮及种皮特征在黄精族系统学研究中的应用，2000植物分类学报，38(1)：30-42．
30．聂剂旺等，安徽产石蒜属植物三种酶同工酶的分析，生物学杂志，2000，17(3)：19-22
31．聂刘旺、张定成等，安徽产石蒜属植物三种酶同工酶的分析，生物学杂志，2000，17(3)：19-22
32．潘龙瑞，二氢石蒜碱对家兔心室乳头肌及血管平滑肌收缩的影响，郧阳医学院学报，2002，21(3)：136-138
33．任秀芳，周守标，郑艳，郭新孤：中国石蒜属植物花粉形态的研究，1995．云南植物研究，17(2)：182-186．
34．任秀芳、周守标、储小慧等，我国石蒜属六种花粉形态的研究，1994，安徽师范大学学报，17(1)：49-52
35．任秀芳、周守标、郑艳等，中国石蒜属植物花粉形态的研究，云南植物石研究，1995，17(2)：182-186
36．邵建章，杨积高，张定成，聂刘旺：二倍体石蒜在安徽的发现，1994．植物分类学报；32(6)：549-552．
37．沈明山、叶志伟等，石蒜的人工栽培，厦门科技，1999，2：21-22
38．孙同兴，胡玉熹，郎楷永：中国兜被兰属植物叶表皮微形态特征的研究，1999，云南植物研究，21(1)：57-62．
39．孙同兴，胡玉熹，郎楷永：中国兜被兰属植物叶表皮微形态特征的研究，1999，云南植物研究，21(1)：57-62．
40．孙叶根、郑艳、张定成，安徽石蒜属四种物核型研究，广西植物，1998，18(4)：363-367
41．唐韬，石蒜治牛瘤胃积食有疗效，贵州畜牧兽医，2002，26(2)：33
42．王兰州等，石蒜叶片微弱电波动信号的测定，西北师范大学学报，2000，36(2)：62-66
43．王意成．中国珍贵球根花卉棗石蒜属植物．中国花卉盆景，1992，6：7．
44．魏岩，周桂玲：十字花科四属植物叶片的表皮特征．云南植物研究，1998．20(4)：407-412．
45．翁素饶等，新抗肿瘤物质—石蒜内铵盐(AT+1840)，科学通报，1976，26(6)：285-287
46．徐炳声，黄少甫，长筒物质的染色体核型分析，植物分类学报，1984，22(1)：43-45
47．徐炳声，黄少甫，红兰石蒜和换锦花的染色体核型分析，南京中山植物园研究论文集，1983，1981：15-19
48．徐炳声，黄少甫，赵治芬，俞志洲，林巾箴：石蒜(Lycoris radiata (L'Her.)Herb., 及其矮小变种var.pumila Gery染色体核型的分析，1984．植物研究，4(1)：112-117．
49．徐炳声，黄少甫等，安徽石蒜和中国染色体核型的分析，云南植物研究，1984，6(1)：196-198
50．徐炳声，黄少甫等：长筒石蒜的染色体核型分析，1984．植物分类学报，22(1)：43-45．
51．徐垠、胡之壁、黄秀兰等，国产石蒜属的新分类群，植物分类学报，1982，20(2)：196-198
52．徐垠等：中国植物志第十六卷第一分册，北京：科学出版社，1965，152-153．
53．徐垠等：中国植物志第十六卷第一分册，北京：科学出版社，1965，152-153．
54．姚剑虹等，石蒜凝集素基因的克隆，复旦学报，2002，41(1)：102-105
55．叶志伟、沈明山等，石蒜的观赏价值，厦门科技，1999，2：35
56．宰玉良，石蒜糊剂外敷治疗小儿急性肾炎40例疗效观察，江苏临床医学杂志，1998，2(2)：149
57．张定成，中国石蒜核型分析，安徽师范大学学报，1988，2：36-39
58．张定成、孙叶根、郑艳等，三倍体换锦花在安徽发现，植物分类学报，1999，37(1)：35-39
59．张露，石蒜类植物无性繁殖技术，南京林业大学学报，2002，26(4)：1-5
60．张露、曹福亮，石蒜属植物栽培技术研究进展，江西农业大学学报，2001，23(3)：375-378
61．张露等，石蒜属种间亲缘关系RAPD分析，遗传学报，2002，29(10)：915-921
62．张万佛．石蒜的栽培．中国花卉盆景，1988，9：6．
63．赵国举、赵娇玲、胡崇家，力可拉敏与加兰他敏对胆碱反应系统的影响，药学学报，1965，12(1)：362
64．中国科学院中国植物志编辑委员会，中国植物志(第十六卷第一分册)，北京：科学出版社，1989，16-17
65．中国农科院土肥所．土壤肥料分析．农业出版社，1979．
66. Levan, K Fredga and A A Sandberg. Nomenclature for centromeric position on chromosomes, Heredita s, 1964, 52:201-220
67. Adams, P Lycoris-Surprise lilies, Pacific Hort, 1976, 37:23-29
68. Bose S Cytoogical studies in the genus Lycoris, Proc 50th Ind Sci Cong part 3, 1962, 446-447
69. Bose S Phylogeny and karyotype evolution in Lycoris, Proc Ⅸ Int. Bot Congr, 1959, 2:41
70. Bose S. 1963: A new chromosome number and karyotype in L. radiatu. Nature. 197:1229-1230.
71. Bose, S A new chromosome number and karyotype in L radiata, Bature, 1963, 197(4873):1229-1230
72. Bose, S Karyotype alternaion in Lycoris chinensis, Science And Culture, 1966, 32(2):144-145
73. Bose, S. W.S. Flory 1963: Astudy of phyloyeny and of karyotype evolution in Lycoris. The Nucleus. 6(20):141-156.
74. Caldwell, S 1980 Lycoris report, Plant Life, 37:121-128
75. Caldwell, S Amaryllid culture (Lycoris report), Plant Life, 1967, 24:82-92
76. Caldwell, S At long last-seeds on Lycoris squamigera, Plant Life, 1979, 35:43-53
77. Choisk..Studies on the.culture of Lycoris radiata Herb. as a medicinal plant. 1.The effct of bulb at planting on plant growth and bulb yield. Reseach Reports Of The Rural Developmt Administration, Upland Industrial Crops, 1991, 33(2):84-88
78. Creech, J L The genus Lycoris in the mid-Ntlantic states, Natl Hort. Mag, 1952, 31:167-173
79. Dilcher D L.Approaches to the identication of Anhiosperm lemains. Bot Rew, 1974, 40:68130.
80. Erdtman G, Pollen morphology and plant taxonomy V on the occurrence of tetrads and dyads, Svens Bot. Tidakr, 1945, 39.
81. Furuta Y. Nishikawa K. Sagihara M. 1989: Chromosomal evolution in the genus Lycoris. Herbertia. 45, 156-162.
82. Hsu, B. S, Siro Kurita. Synopsis of the genus Lycoris, SIDA, 1994, 16(2):301-331.
83. Hsu, B.S.J.Lin etc, An evaluation of the interspecifice relationships of Lycoris based aon pollen viability and rate of seed set after crossing, Acta. Genet. Sin, 1986, 13:369-376.
84. Hsu, Y. and G.J.Fan, A new species of Lycoris, Acta Phytotax. Sin, 1974, 12:299-300
85. Hunt, W.L. Lycoris sanguinea and L. "cinnabarina", Plant Life, 1963, 19:50-53
86. Inariyama, S, 1931: Cytological studies in the genus Lycoris. Bot May Todyo. 45:11-26.
87． Inariyama, S, 1953: Cytological studies in Lycoris, Rep. Kinara Inst Biol Res 6:5-10．
88． Inariyama, S. Karyotype studies in Amaryllidaceae I, Sci. Rep. Tokyo Bunrika Daigaku, Sect B, 1937,3:95-113
89． Inariyama, S. Origin of Lycoris radiata and Lycoris albiflora, Jap. J. Genet, 1944,20: 87-88
90． Jones, K. Aspects of chromosome evolution in higher plants, Adv. Bot. Res, 1978, 6: 119-194
91． Jones, K. Multiple Robertsonian fusions in the evolution of a plant genus. In: Jones K. Braudham P E. ed, Current Chromosome Research, Amstardam: North-Holl and Pulbishing Company, 1976, 220-221
92． Kihara, H and Koyama, M. Offspring obtained by self-pollination of Lycoris radiata Herb, a triploid species, Jap. J. Genet, 1954, 29: 169-171
93． Kim, M and S. Lee, A taxonomical study on the Korean Lycoris(Amaryllidaceae), Korean J.Pl. Taxon, 1991,21: 123-139
94． Koyama, M. Chromosome pairing in the genus I. Ann. Rep. Doshisha Women's Coll, 1978, 29: 272-282
95． Koyama, M. Chromosome pairing in the genus Lysoris II. Ann. Rep. Doshisha Women's Coll, 1978, 29: 272-282
96． Koyama, M. Meiosis in L. albiflora, Ann. Rep. Doshisha Women's Coll, 1962, 12:9-19
97． Koyama, M. Offsprings of Lycoris radiata obtained by artificial self pollination, Ann. Rep. Doshisha Women's Coll, 1959, 10: 388-394
98． Koyama, M. Somatic chromosomes in the genus Lycoris, Ann. Rep. Doshisha Women's Coll, 1962, 12:9-19
99． Kurita S. 1988: Variation and evolution in the karyotype of Lycoris(Amaryllidaceae). VII. Modes of karyotype alteration within species and probable trend of Karyotype evolution in the genus. Cytologia. 53:323-335．
100． Kurita, S, 1987: Variation and evolution in the Karyotype of Lycoris(Amary llidaceae), IV, Intraspecific variation in karyctype of L. Radiata (L' Her.) Herb. And the origin of this triploid, species, Cytologia. 52:137-140．
101． Kurita, s. Geoclinal change in the pollen ornamentation of Lycoris sanguinea Maxim. Var. sanguinea, J. Jap. Bot, 1985, 60: 275-279．
102． Kurita, S. Lycoris aurea Vs. L. traubii, J.Jap. Bot, 1980, 55: 287-288
103． Kurita, S. Variation and evolution in the karyotype of Lycoris, Amaryllidaceae I, General karyomorphological characteristics of the genus, Cytologia, 1986, 51: 803-815
104． Kurita, S. Variation and evolution in the karyotype of Lycoris, Amaryllidaceae III, intraspecific variation in the karyotype of L. traubii Hayward, Cytologia, 1987, 52: 117-128．
105． Lee, S and Kim, M. Polynological study of some Lycoris species(Amaryllidaceae), Korean J. Pl. Taxon, 1987, 17: 147-154
106． Liu, Y. and B.S.Hsu, Mechanism of sterility of diploid hybrid in genus Lycoris, Acta Agri. Shanghai, 1990, 6: 27-30
107． Melchior. H. A Engler's Syllabus der pflanzenfamilien. Vol. 2． Gebruder Borntae, 1964, Berlin.
108． Motomilto, Atsushikawamota, Phylogenetic Relationships Of Amaryllidaceae Based On marK Sequence Data, J. Plant. Res, 1999, 112:207-216
109． Nakai, Lycoris Koreana Nakai, Sp. Nov. Bot. Mag. (Tokyo), 1930,44: 516-517
110． Nishikawa K. Fruta Y. Endo H.1979: Cousideration of the chromosome evolution on the basis of, nuclear DNA content and total chromosome length in Lycoris. Jap J Genet. 54:387-396．
111． Nishiyama, I. Reduction division in Lycoris, Bot. Mag. Tokyo, 1928,42: 509-513
112． N.-S. Lee, Isozyme evidence for the allotriploid origin of Lycoris flavescens( Amaryllidaceae) 2001, Psant Syst. Evol. 227: 227-234．
113． Park, Y.J, E.H.Park, S.O.Yoo, and Y.O.Chung, Oytological study on the genus Lycoris in Korea. I. Epidermal patterns of the leafblades, Abstracts, Annual Conference Of Kor. Soc. Hort. Sci, 1989, 108-109
114． Stebbins, G.L. Chromosomal evolution in higher plants, Edward Arnold Ltd, London, 1971
115． Tae, Kyong-Hwan and Sung-chul Ko. 1995:A taxonomic study on epidermal characters of the genus Lycoris in Korea. Ker J Plant Tax.VOL 25 No 3． 177-193．
116． Tae, Kyoung-Hwan and Sung-Chul Ko, New taxa of the genus Lycoris, Korean J. Pl. Taxon, 1993, 23:233-241
117． Tae, Kyoung-Hwan, Sung-Chul Ko, Kim, Y.S.A Cytotaxonomic study on genus Lycoris in Korea, Kor. J. Plant. Tax, 1987, 17: 135-145
118． Tae. Kyoung-Hwan and Sung-Chul Ko, A taxonomic study on the genus Lycoris, Kor. J. Plant Tax, 1996, 26(1) : 19-35
119． Takemura, E. Morphological and cytological studies on artificial hybrids in the genus Lycoris I. On the F1 hybrid between L.sprengeri Comes and L. straminea Lindl, Bot. Mag. Gokyo, 1961, 74: 514-531．
120． Takenaka, Y. On the chromosomes of Lycoris squamigera Maxim, J. Chosen Nat. Hist. Soc, 1930, 10:54-57
121． Traub, H. P. Lycoris Haywardii, L. houdyshelii, and L. caldwellii, Pl. Life, 1957, 13:42-48
122． Walker, W. Evolution significance of the exine in the pollen of primitive angiosperms, In: I K.Ferguson & J. Muller (eds.) The evolutionary significance of the exine, London: Academic Press, 1976,251-308．
123． Willknson H P The plant surfance (mainly leaf).Anatomy of Dicotyledons (2th edition).Oxford: Clarendon Press, 1979,97117．
124． Worsley, A. Lycoris: A garden review, Gard Chron. Ser. 3 1928, 84:169．
125． Yang, I. S. On the distrbution of the Lycoris, J. Korean, Pl. Taxon, 1976, 7:32．
2．陈必义、杜仁华、曾繁典等，二氢石蒜碱的降压作用，中国药理学报，1993，14(1)：45
3．陈耀华、李懋学，四种石蒜属植物的染色体核型研究，园艺学报，1985，12(1)：57-60
4．陈宝梁．叶片解剖特征在木莲属Manglietia BL．分种的应用[J]．中山大学研究生学刊，1989，10(2)：52-55
5．邓传良等，石蒜单染色的显微分离及体外扩增，南京林业大学学报，2002，26(3)：72-74
6．董国良，石蒜外治妙用，中医外治杂志，1995，6：31-32
7．董庆华，石蒜的组织培养，植物生理学通讯，1995(5)：54
8．额特曼(中国科学院植物研究所古植物研究室孢粉组译)，孢粉学手册，北京：科学出版社，1978，88，278-279，339，286-287
9．访台北县淡水镇农会花卉研究班谈，“金花石蒜，台湾花卉，1984，57-59
10．龚新荣等，二氢石蒜碱对大鼠高碳酸性脑缺氧损伤的保护作用，郧阳医学院学报，2002，21(2)：71-72
11．韩锦隆，金花石蒜天然开花期之变温及人工花期调节方法之研究，宜兰农工学报，1991，10：105-121
12．胡庆和等，石蒜内铵盐(AT-1840)综合治疗妇科晚期恶性肿瘤，浙江中西医结合杂志，1997，7，(3)：134-136
13．黄宗春、胡一民，安徽野生石蒜的种质资源及开发利用，中国林副特产，1997．3：46-47
14．柯丽霞、孙叶根、郑艳等，石蒜属三种植物的核型研究，安徽师范大学学报，1998，21(4)：343-348
15．蓝星莲，二氢石蒜碱对家名胜脑血管的选择性作用，中国药理学通报，2002，18(4)：409-411
16．李朝銮：蔷薇科原始属植物叶表皮解剖及其系统学意义．植物分类学报，1989．27(3)：178-183．
17．李荣义．中国水仙．福建科技出版社，1991．
18．李卫东，石蒜的人工栽培成功，厦门晚报，1996，12/21，第四版
19．李建强等．赤爮亚族植物叶片中脉的比较解剖．云南植物研究，1992，14(4)：418-422
20．梁素秋，金花石蒜芽分化及体外培养之探讨，国立台湾大学园艺研究所硕士论文，1992
21．林纯英、马溯轩，金花石蒜之鳞片组织培养毓，中国园艺(台湾)，1987，33(4)：255-264、
22．林定勇，石蒜属球根花卉之分类、形态、生长与开花，中国园艺(台湾)，1993，39(2)：67-72
23．刘顺俊、刘四维，单味石蒜治疗疔疮肿毒，湖北中医杂志，1996，18(4/12
24．刘德仪等．中国木兰属和含笑属导管分子的比较解剖．植物学报，1987，29(1)：22-28
25．刘玉壶等．木兰科分类系统的初步研究[J]．植物分类学报，1984，22(2)：89-109
26．刘琰，徐炳声：石蒜属的核型研究，1989．植物分类学报，27(4)：257-264．
27．鲁雪花、陈扬春，忽地笑胚外植体的培养，云南植物研究，1986，8(4)：467-469
28．吕海亮，吴世安，杨继，饶广远：叶表皮及种皮特征在黄精族系统学研究中的应用，2000．植物
分类学报，38(1)：30-42．
29．吕海亮，吴世安，杨继，饶广远：叶表皮及种皮特征在黄精族系统学研究中的应用，2000植物分类学报，38(1)：30-42．
30．聂剂旺等，安徽产石蒜属植物三种酶同工酶的分析，生物学杂志，2000，17(3)：19-22
31．聂刘旺、张定成等，安徽产石蒜属植物三种酶同工酶的分析，生物学杂志，2000，17(3)：19-22
32．潘龙瑞，二氢石蒜碱对家兔心室乳头肌及血管平滑肌收缩的影响，郧阳医学院学报，2002，21(3)：136-138
33．任秀芳，周守标，郑艳，郭新孤：中国石蒜属植物花粉形态的研究，1995．云南植物研究，17(2)：182-186．
34．任秀芳、周守标、储小慧等，我国石蒜属六种花粉形态的研究，1994，安徽师范大学学报，17(1)：49-52
35．任秀芳、周守标、郑艳等，中国石蒜属植物花粉形态的研究，云南植物石研究，1995，17(2)：182-186
36．邵建章，杨积高，张定成，聂刘旺：二倍体石蒜在安徽的发现，1994．植物分类学报；32(6)：549-552．
37．沈明山、叶志伟等，石蒜的人工栽培，厦门科技，1999，2：21-22
38．孙同兴，胡玉熹，郎楷永：中国兜被兰属植物叶表皮微形态特征的研究，1999，云南植物研究，21(1)：57-62．
39．孙同兴，胡玉熹，郎楷永：中国兜被兰属植物叶表皮微形态特征的研究，1999，云南植物研究，21(1)：57-62．
40．孙叶根、郑艳、张定成，安徽石蒜属四种物核型研究，广西植物，1998，18(4)：363-367
41．唐韬，石蒜治牛瘤胃积食有疗效，贵州畜牧兽医，2002，26(2)：33
42．王兰州等，石蒜叶片微弱电波动信号的测定，西北师范大学学报，2000，36(2)：62-66
43．王意成．中国珍贵球根花卉棗石蒜属植物．中国花卉盆景，1992，6：7．
44．魏岩，周桂玲：十字花科四属植物叶片的表皮特征．云南植物研究，1998．20(4)：407-412．
45．翁素饶等，新抗肿瘤物质—石蒜内铵盐(AT+1840)，科学通报，1976，26(6)：285-287
46．徐炳声，黄少甫，长筒物质的染色体核型分析，植物分类学报，1984，22(1)：43-45
47．徐炳声，黄少甫，红兰石蒜和换锦花的染色体核型分析，南京中山植物园研究论文集，1983，1981：15-19
48．徐炳声，黄少甫，赵治芬，俞志洲，林巾箴：石蒜(Lycoris radiata (L'Her.)Herb., 及其矮小变种var.pumila Gery染色体核型的分析，1984．植物研究，4(1)：112-117．
49．徐炳声，黄少甫等，安徽石蒜和中国染色体核型的分析，云南植物研究，1984，6(1)：196-198
50．徐炳声，黄少甫等：长筒石蒜的染色体核型分析，1984．植物分类学报，22(1)：43-45．
51．徐垠、胡之壁、黄秀兰等，国产石蒜属的新分类群，植物分类学报，1982，20(2)：196-198
52．徐垠等：中国植物志第十六卷第一分册，北京：科学出版社，1965，152-153．
53．徐垠等：中国植物志第十六卷第一分册，北京：科学出版社，1965，152-153．
54．姚剑虹等，石蒜凝集素基因的克隆，复旦学报，2002，41(1)：102-105
55．叶志伟、沈明山等，石蒜的观赏价值，厦门科技，1999，2：35
56．宰玉良，石蒜糊剂外敷治疗小儿急性肾炎40例疗效观察，江苏临床医学杂志，1998，2(2)：149
57．张定成，中国石蒜核型分析，安徽师范大学学报，1988，2：36-39
58．张定成、孙叶根、郑艳等，三倍体换锦花在安徽发现，植物分类学报，1999，37(1)：35-39
59．张露，石蒜类植物无性繁殖技术，南京林业大学学报，2002，26(4)：1-5
60．张露、曹福亮，石蒜属植物栽培技术研究进展，江西农业大学学报，2001，23(3)：375-378
61．张露等，石蒜属种间亲缘关系RAPD分析，遗传学报，2002，29(10)：915-921
62．张万佛．石蒜的栽培．中国花卉盆景，1988，9：6．
63．赵国举、赵娇玲、胡崇家，力可拉敏与加兰他敏对胆碱反应系统的影响，药学学报，1965，12(1)：362
64．中国科学院中国植物志编辑委员会，中国植物志(第十六卷第一分册)，北京：科学出版社，1989，16-17
65．中国农科院土肥所．土壤肥料分析．农业出版社，1979．
66. Levan, K Fredga and A A Sandberg. Nomenclature for centromeric position on chromosomes, Heredita s, 1964, 52:201-220
67. Adams, P Lycoris-Surprise lilies, Pacific Hort, 1976, 37:23-29
68. Bose S Cytoogical studies in the genus Lycoris, Proc 50th Ind Sci Cong part 3, 1962, 446-447
69. Bose S Phylogeny and karyotype evolution in Lycoris, Proc Ⅸ Int. Bot Congr, 1959, 2:41
70. Bose S. 1963: A new chromosome number and karyotype in L. radiatu. Nature. 197:1229-1230.
71. Bose, S A new chromosome number and karyotype in L radiata, Bature, 1963, 197(4873):1229-1230
72. Bose, S Karyotype alternaion in Lycoris chinensis, Science And Culture, 1966, 32(2):144-145
73. Bose, S. W.S. Flory 1963: Astudy of phyloyeny and of karyotype evolution in Lycoris. The Nucleus. 6(20):141-156.
74. Caldwell, S 1980 Lycoris report, Plant Life, 37:121-128
75. Caldwell, S Amaryllid culture (Lycoris report), Plant Life, 1967, 24:82-92
76. Caldwell, S At long last-seeds on Lycoris squamigera, Plant Life, 1979, 35:43-53
77. Choisk..Studies on the.culture of Lycoris radiata Herb. as a medicinal plant. 1.The effct of bulb at planting on plant growth and bulb yield. Reseach Reports Of The Rural Developmt Administration, Upland Industrial Crops, 1991, 33(2):84-88
78. Creech, J L The genus Lycoris in the mid-Ntlantic states, Natl Hort. Mag, 1952, 31:167-173
79. Dilcher D L.Approaches to the identication of Anhiosperm lemains. Bot Rew, 1974, 40:68130.
80. Erdtman G, Pollen morphology and plant taxonomy V on the occurrence of tetrads and dyads, Svens Bot. Tidakr, 1945, 39.
81. Furuta Y. Nishikawa K. Sagihara M. 1989: Chromosomal evolution in the genus Lycoris. Herbertia. 45, 156-162.
82. Hsu, B. S, Siro Kurita. Synopsis of the genus Lycoris, SIDA, 1994, 16(2):301-331.
83. Hsu, B.S.J.Lin etc, An evaluation of the interspecifice relationships of Lycoris based aon pollen viability and rate of seed set after crossing, Acta. Genet. Sin, 1986, 13:369-376.
84. Hsu, Y. and G.J.Fan, A new species of Lycoris, Acta Phytotax. Sin, 1974, 12:299-300
85. Hunt, W.L. Lycoris sanguinea and L. "cinnabarina", Plant Life, 1963, 19:50-53
86. Inariyama, S, 1931: Cytological studies in the genus Lycoris. Bot May Todyo. 45:11-26.
87． Inariyama, S, 1953: Cytological studies in Lycoris, Rep. Kinara Inst Biol Res 6:5-10．
88． Inariyama, S. Karyotype studies in Amaryllidaceae I, Sci. Rep. Tokyo Bunrika Daigaku, Sect B, 1937,3:95-113
89． Inariyama, S. Origin of Lycoris radiata and Lycoris albiflora, Jap. J. Genet, 1944,20: 87-88
90． Jones, K. Aspects of chromosome evolution in higher plants, Adv. Bot. Res, 1978, 6: 119-194
91． Jones, K. Multiple Robertsonian fusions in the evolution of a plant genus. In: Jones K. Braudham P E. ed, Current Chromosome Research, Amstardam: North-Holl and Pulbishing Company, 1976, 220-221
92． Kihara, H and Koyama, M. Offspring obtained by self-pollination of Lycoris radiata Herb, a triploid species, Jap. J. Genet, 1954, 29: 169-171
93． Kim, M and S. Lee, A taxonomical study on the Korean Lycoris(Amaryllidaceae), Korean J.Pl. Taxon, 1991,21: 123-139
94． Koyama, M. Chromosome pairing in the genus I. Ann. Rep. Doshisha Women's Coll, 1978, 29: 272-282
95． Koyama, M. Chromosome pairing in the genus Lysoris II. Ann. Rep. Doshisha Women's Coll, 1978, 29: 272-282
96． Koyama, M. Meiosis in L. albiflora, Ann. Rep. Doshisha Women's Coll, 1962, 12:9-19
97． Koyama, M. Offsprings of Lycoris radiata obtained by artificial self pollination, Ann. Rep. Doshisha Women's Coll, 1959, 10: 388-394
98． Koyama, M. Somatic chromosomes in the genus Lycoris, Ann. Rep. Doshisha Women's Coll, 1962, 12:9-19
99． Kurita S. 1988: Variation and evolution in the karyotype of Lycoris(Amaryllidaceae). VII. Modes of karyotype alteration within species and probable trend of Karyotype evolution in the genus. Cytologia. 53:323-335．
100． Kurita, S, 1987: Variation and evolution in the Karyotype of Lycoris(Amary llidaceae), IV, Intraspecific variation in karyctype of L. Radiata (L' Her.) Herb. And the origin of this triploid, species, Cytologia. 52:137-140．
101． Kurita, s. Geoclinal change in the pollen ornamentation of Lycoris sanguinea Maxim. Var. sanguinea, J. Jap. Bot, 1985, 60: 275-279．
102． Kurita, S. Lycoris aurea Vs. L. traubii, J.Jap. Bot, 1980, 55: 287-288
103． Kurita, S. Variation and evolution in the karyotype of Lycoris, Amaryllidaceae I, General karyomorphological characteristics of the genus, Cytologia, 1986, 51: 803-815
104． Kurita, S. Variation and evolution in the karyotype of Lycoris, Amaryllidaceae III, intraspecific variation in the karyotype of L. traubii Hayward, Cytologia, 1987, 52: 117-128．
105． Lee, S and Kim, M. Polynological study of some Lycoris species(Amaryllidaceae), Korean J. Pl. Taxon, 1987, 17: 147-154
106． Liu, Y. and B.S.Hsu, Mechanism of sterility of diploid hybrid in genus Lycoris, Acta Agri. Shanghai, 1990, 6: 27-30
107． Melchior. H. A Engler's Syllabus der pflanzenfamilien. Vol. 2． Gebruder Borntae, 1964, Berlin.
108． Motomilto, Atsushikawamota, Phylogenetic Relationships Of Amaryllidaceae Based On marK Sequence Data, J. Plant. Res, 1999, 112:207-216
109． Nakai, Lycoris Koreana Nakai, Sp. Nov. Bot. Mag. (Tokyo), 1930,44: 516-517
110． Nishikawa K. Fruta Y. Endo H.1979: Cousideration of the chromosome evolution on the basis of, nuclear DNA content and total chromosome length in Lycoris. Jap J Genet. 54:387-396．
111． Nishiyama, I. Reduction division in Lycoris, Bot. Mag. Tokyo, 1928,42: 509-513
112． N.-S. Lee, Isozyme evidence for the allotriploid origin of Lycoris flavescens( Amaryllidaceae) 2001, Psant Syst. Evol. 227: 227-234．
113． Park, Y.J, E.H.Park, S.O.Yoo, and Y.O.Chung, Oytological study on the genus Lycoris in Korea. I. Epidermal patterns of the leafblades, Abstracts, Annual Conference Of Kor. Soc. Hort. Sci, 1989, 108-109
114． Stebbins, G.L. Chromosomal evolution in higher plants, Edward Arnold Ltd, London, 1971
115． Tae, Kyong-Hwan and Sung-chul Ko. 1995:A taxonomic study on epidermal characters of the genus Lycoris in Korea. Ker J Plant Tax.VOL 25 No 3． 177-193．
116． Tae, Kyoung-Hwan and Sung-Chul Ko, New taxa of the genus Lycoris, Korean J. Pl. Taxon, 1993, 23:233-241
117． Tae, Kyoung-Hwan, Sung-Chul Ko, Kim, Y.S.A Cytotaxonomic study on genus Lycoris in Korea, Kor. J. Plant. Tax, 1987, 17: 135-145
118． Tae. Kyoung-Hwan and Sung-Chul Ko, A taxonomic study on the genus Lycoris, Kor. J. Plant Tax, 1996, 26(1) : 19-35
119． Takemura, E. Morphological and cytological studies on artificial hybrids in the genus Lycoris I. On the F1 hybrid between L.sprengeri Comes and L. straminea Lindl, Bot. Mag. Gokyo, 1961, 74: 514-531．
120． Takenaka, Y. On the chromosomes of Lycoris squamigera Maxim, J. Chosen Nat. Hist. Soc, 1930, 10:54-57
121． Traub, H. P. Lycoris Haywardii, L. houdyshelii, and L. caldwellii, Pl. Life, 1957, 13:42-48
122． Walker, W. Evolution significance of the exine in the pollen of primitive angiosperms, In: I K.Ferguson & J. Muller (eds.) The evolutionary significance of the exine, London: Academic Press, 1976,251-308．
123． Willknson H P The plant surfance (mainly leaf).Anatomy of Dicotyledons (2th edition).Oxford: Clarendon Press, 1979,97117．
124． Worsley, A. Lycoris: A garden review, Gard Chron. Ser. 3 1928, 84:169．
125． Yang, I. S. On the distrbution of the Lycoris, J. Korean, Pl. Taxon, 1976, 7:32．