微型月季微繁体系的构建及组培生根机理研究
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摘要
本文在综述国内外有关微型月季（Rosa chinensis. Minima或Rosa.
roelletti）研究进展的基础上，以几个花色具代表性的微型月季品种为试材，
研究了培养基成分及培养条件对微型月季组织培养中无菌系的建立、增殖与壮
苗，瓶内生根与驯化及瓶外生根技术进行了系统的研究，同时还对微型月季的
组培生根机理进行了初步探讨。
试验表明：以开花枝下具完整叶、带腋芽的茎段为外植体（外植体长度/
直径组合为9.0～10.0mm/3.0-3.5mm，茎节上下各带5.0mm亲本组织）有利于
腋芽诱导，将外植体用100mg/L庆大霉素溶液浸泡30分钟可有效防止内生菌
污染。灭菌后接种于100ml三角瓶，每瓶接种4-9个外植体。诱导培养基为
MS+6-BAlmg/L+NAA0.1～0.2mg/L+KT0.5mg/L，接种后先暗处理2天，然后
转入1600—2000Lux的光强下培养，光照16小时。培养温度为20℃。
将腋芽再生芽转入MS+6-BA3～5mg/L+NAA0.1～0.2mg/L+蔗糖（生产上可
用白砂糖）30～35g/L+琼脂6～6.5g/L的培养基上，培养基的pH值调至5.9。
培养温度为22℃，光照800～1600Lux，光照16小时。在以后的继代增殖中以
3芽为一转接单位有利于增殖。
将增殖和壮苗培养后的苗（苗高约20～25mm）转入1/2MS（将Ca~(2+)浓度
提高到1.5倍）+NAA0.2mg/L（或NAA0.1mg/L+IBA0.1mg/L）+蔗糖20～25g/L+
琼脂7g/L+活性炭0.4g/L的生根培养基中，培养基的pH值调至6.0。培养温
度在转接后的第一周为22℃，然后逐渐降到18℃。培养容器以试管最佳，生
产上可用广口瓶。其它培养条件同上。
将培养6～8周、20～25mm高的苗取出，洗净基部琼脂后浸入50g/L的IBA
溶液中5分钟，然后插入由蛭石和粗沙按1：1（v/v）混合而成的基质中，于
自然散射光下培养，培养温度22℃。21天后将驯化成活的苗移入由园土、蛭
石和珍珠岩按！：1：1（V／v／v）混合而成的基质中，其它栽培条件同驯化条件，
并逐渐过渡到自然条件。
在瓶内、瓶外生根苗的比较研究中发现，不同的生根方式对植株生长发育
的影响较大。经两种生根方式处理ZI 天后，瓶外所生的根较长、数量较多、
横截面积较大，其中维管柬所占的比重较大，且维管束发育早，根的韧性好、
白色、有侧根；而瓶内所生的根较细、易碎、黑色、没有侧根。除‘小红’外，
所有瓶外生根的植株其根（横截面积和干重）和茎（高度、横截面积和干重）
的生长量均比瓶内生根的植株多。所有瓶外生根的植株产花量增加，花期提前。
0
通过对微型月季组培苗生根过程中内含物的阶段性测定发现，生根部位的
IAA含量在根原基出现前增加，而根原基出现后下降；iPAs和ZRs在根原基
出现前略有下降，而在根原基出现后的分化过程中急剧上升；可溶性糖和淀粉
在根原基出现前积累，而根原基出现后因消耗含量下降；可溶性蛋白在根原基
发生和分化的过程中增多，而在根原基分化后根的伸长过程中含量下降；POD
活性在根原基发生和分化的过程中增强，而在根原基分化后根的伸长过程中活
性降低：外源IAA处理会加强以上指标的变化强度，从而有利于生根。因此，
在组培苗的生根过程中，根原基的形成为生长素所刺激，而在根原基的分化和
根的伸长过程中需低浓度的细胞分裂素与生长素配合，可溶性糖和淀粉等碳水
化合物为根原基的分化和根的生长提供能源，在根原基的发生和分化过程中还
伴随着一系列的生理生化变化，使蛋白质含量和酶活出现阶段性变化。
By summarizing the research about Miniature Rose (Rosa chinensis.
Minima or Rosa roelletti), this study aimed to select optimum culture
medium and conditions for Miniature Rose with several representative
varieties in flower colour, especially focused on the techniques of the
establishment of aseptic cultures, proliferation, strengthening and the
rooting in vitro and ex vitro of the plantlet. Further more, this article
discussed the mechanism of the rooting during the tissue culture. At last,
the tissue culture system of Miniature Rose was established as follows:
1. Establishment of aseptic cultures: the explant (9.0?0.0mm in
length/3.O?.Smm in diameter, 5.0mm above/under the node) was node
segment with one axillary bud under the intact leaf from the flowering
stem. After being disinfected with 75% alcohol and 0. 1% HgCJ2 , the
explants were dipped in to lOOmg/L gentamicin solution for 30 mm to
eliminate the germs in, then were inoculated in 100m1 vessels, each with
4? explants. The inducing medium was: MS+6桞A lmg/L+NAA 0. 1?. 2mg/L +KT
O.5mg/L. After inoculation, 1600?000Lux light intensity, 16梙s light
period and temperature in 200C was succeeded with treatment of 2梔ay
darkness.
2. Proliferation and Strengthening: The regenerated bud was transferred
to the medium of MS+6桞A3?mg/L+NAA 0. 1?. 2 mg/L +Sucrose 30?5g/L+agar
6?.5g/L. PH5.9, Th22XD, LI (Light intensity) 800?600 Lux, LP(Light
Period>l6hs. The proliferation of Miniature Rose has the quantity
effect Three buds as a transferring unit resulted in the highest
proliferating rate.
3. Rooting in vitro: The strengthened plantlet (height of 20?5mm) was
transferred to the rooting medium of l/2MS(the concentration of Ca2~ was
increased to 1.5 times)+ NAA 0. 2mg/L (or NM 0. lmg/L +IBA
0. lmg/L) +Sucrose 20?5g/L+agar 7g/L+AC 0. 4g/L. after being transferred,
the temperature was 220C in the first week, than decreased to l8~C. Tube
was the best container for rooting. Other culturing conditions were the
same as above.
4. Rooting ex vitro: Cultured 6? week, the plantlet (with the height
of 20?5mm) was taken out from the vessel, after washed off the agar round
the end, the plantlet was dipped into JBA solution(50g/L) for 5mm and
then was inserted to the medium mixed of peat and sand(l :1, V/V), growing
under natural light with T=22 慍 . 2ldays later, transferred the
acclimated plant to the medium mixed of loam, peat and perlite
(l:i:i,V/V/V), slowly developed to the natural condition.
By comparing in vitro and ex vitro rooted plant, we found that
different rooting method affects the plant growth and development.
2ldays after these two rooting treatments, the ex vitro root was white,
longer, larger in quantity and diameter, larger proportion of vascular,
which developed earlier and had better flexible, and had branch root
while the in vitro root was dark, thinner, brittle and without branch
root. Except for the variety of 憀ittle red? the growth of root (diameter
and dry weight) and stem (height, diameter and dry weight) of the ex vitro
rooted plant were larger than those of in vitro rooted plant.
5. Discussion of the mechanism of rooting during the tissue culture: By
determining the endogenous substance periodically during the rooting of
Miniature Rose tissue cultured course, it was found that the content of
IAA in rhizogenic zone increased before the formation of primordium and
decreased after formation. While iPAs and ZRs slightly decreased before
roelletti）研究进展的基础上，以几个花色具代表性的微型月季品种为试材，
研究了培养基成分及培养条件对微型月季组织培养中无菌系的建立、增殖与壮
苗，瓶内生根与驯化及瓶外生根技术进行了系统的研究，同时还对微型月季的
组培生根机理进行了初步探讨。
试验表明：以开花枝下具完整叶、带腋芽的茎段为外植体（外植体长度/
直径组合为9.0～10.0mm/3.0-3.5mm，茎节上下各带5.0mm亲本组织）有利于
腋芽诱导，将外植体用100mg/L庆大霉素溶液浸泡30分钟可有效防止内生菌
污染。灭菌后接种于100ml三角瓶，每瓶接种4-9个外植体。诱导培养基为
MS+6-BAlmg/L+NAA0.1～0.2mg/L+KT0.5mg/L，接种后先暗处理2天，然后
转入1600—2000Lux的光强下培养，光照16小时。培养温度为20℃。
将腋芽再生芽转入MS+6-BA3～5mg/L+NAA0.1～0.2mg/L+蔗糖（生产上可
用白砂糖）30～35g/L+琼脂6～6.5g/L的培养基上，培养基的pH值调至5.9。
培养温度为22℃，光照800～1600Lux，光照16小时。在以后的继代增殖中以
3芽为一转接单位有利于增殖。
将增殖和壮苗培养后的苗（苗高约20～25mm）转入1/2MS（将Ca~(2+)浓度
提高到1.5倍）+NAA0.2mg/L（或NAA0.1mg/L+IBA0.1mg/L）+蔗糖20～25g/L+
琼脂7g/L+活性炭0.4g/L的生根培养基中，培养基的pH值调至6.0。培养温
度在转接后的第一周为22℃，然后逐渐降到18℃。培养容器以试管最佳，生
产上可用广口瓶。其它培养条件同上。
将培养6～8周、20～25mm高的苗取出，洗净基部琼脂后浸入50g/L的IBA
溶液中5分钟，然后插入由蛭石和粗沙按1：1（v/v）混合而成的基质中，于
自然散射光下培养，培养温度22℃。21天后将驯化成活的苗移入由园土、蛭
石和珍珠岩按！：1：1（V／v／v）混合而成的基质中，其它栽培条件同驯化条件，
并逐渐过渡到自然条件。
在瓶内、瓶外生根苗的比较研究中发现，不同的生根方式对植株生长发育
的影响较大。经两种生根方式处理ZI 天后，瓶外所生的根较长、数量较多、
横截面积较大，其中维管柬所占的比重较大，且维管束发育早，根的韧性好、
白色、有侧根；而瓶内所生的根较细、易碎、黑色、没有侧根。除‘小红’外，
所有瓶外生根的植株其根（横截面积和干重）和茎（高度、横截面积和干重）
的生长量均比瓶内生根的植株多。所有瓶外生根的植株产花量增加，花期提前。
0
通过对微型月季组培苗生根过程中内含物的阶段性测定发现，生根部位的
IAA含量在根原基出现前增加，而根原基出现后下降；iPAs和ZRs在根原基
出现前略有下降，而在根原基出现后的分化过程中急剧上升；可溶性糖和淀粉
在根原基出现前积累，而根原基出现后因消耗含量下降；可溶性蛋白在根原基
发生和分化的过程中增多，而在根原基分化后根的伸长过程中含量下降；POD
活性在根原基发生和分化的过程中增强，而在根原基分化后根的伸长过程中活
性降低：外源IAA处理会加强以上指标的变化强度，从而有利于生根。因此，
在组培苗的生根过程中，根原基的形成为生长素所刺激，而在根原基的分化和
根的伸长过程中需低浓度的细胞分裂素与生长素配合，可溶性糖和淀粉等碳水
化合物为根原基的分化和根的生长提供能源，在根原基的发生和分化过程中还
伴随着一系列的生理生化变化，使蛋白质含量和酶活出现阶段性变化。
By summarizing the research about Miniature Rose (Rosa chinensis.
Minima or Rosa roelletti), this study aimed to select optimum culture
medium and conditions for Miniature Rose with several representative
varieties in flower colour, especially focused on the techniques of the
establishment of aseptic cultures, proliferation, strengthening and the
rooting in vitro and ex vitro of the plantlet. Further more, this article
discussed the mechanism of the rooting during the tissue culture. At last,
the tissue culture system of Miniature Rose was established as follows:
1. Establishment of aseptic cultures: the explant (9.0?0.0mm in
length/3.O?.Smm in diameter, 5.0mm above/under the node) was node
segment with one axillary bud under the intact leaf from the flowering
stem. After being disinfected with 75% alcohol and 0. 1% HgCJ2 , the
explants were dipped in to lOOmg/L gentamicin solution for 30 mm to
eliminate the germs in, then were inoculated in 100m1 vessels, each with
4? explants. The inducing medium was: MS+6桞A lmg/L+NAA 0. 1?. 2mg/L +KT
O.5mg/L. After inoculation, 1600?000Lux light intensity, 16梙s light
period and temperature in 200C was succeeded with treatment of 2梔ay
darkness.
2. Proliferation and Strengthening: The regenerated bud was transferred
to the medium of MS+6桞A3?mg/L+NAA 0. 1?. 2 mg/L +Sucrose 30?5g/L+agar
6?.5g/L. PH5.9, Th22XD, LI (Light intensity) 800?600 Lux, LP(Light
Period>l6hs. The proliferation of Miniature Rose has the quantity
effect Three buds as a transferring unit resulted in the highest
proliferating rate.
3. Rooting in vitro: The strengthened plantlet (height of 20?5mm) was
transferred to the rooting medium of l/2MS(the concentration of Ca2~ was
increased to 1.5 times)+ NAA 0. 2mg/L (or NM 0. lmg/L +IBA
0. lmg/L) +Sucrose 20?5g/L+agar 7g/L+AC 0. 4g/L. after being transferred,
the temperature was 220C in the first week, than decreased to l8~C. Tube
was the best container for rooting. Other culturing conditions were the
same as above.
4. Rooting ex vitro: Cultured 6? week, the plantlet (with the height
of 20?5mm) was taken out from the vessel, after washed off the agar round
the end, the plantlet was dipped into JBA solution(50g/L) for 5mm and
then was inserted to the medium mixed of peat and sand(l :1, V/V), growing
under natural light with T=22 慍 . 2ldays later, transferred the
acclimated plant to the medium mixed of loam, peat and perlite
(l:i:i,V/V/V), slowly developed to the natural condition.
By comparing in vitro and ex vitro rooted plant, we found that
different rooting method affects the plant growth and development.
2ldays after these two rooting treatments, the ex vitro root was white,
longer, larger in quantity and diameter, larger proportion of vascular,
which developed earlier and had better flexible, and had branch root
while the in vitro root was dark, thinner, brittle and without branch
root. Except for the variety of 憀ittle red? the growth of root (diameter
and dry weight) and stem (height, diameter and dry weight) of the ex vitro
rooted plant were larger than those of in vitro rooted plant.
5. Discussion of the mechanism of rooting during the tissue culture: By
determining the endogenous substance periodically during the rooting of
Miniature Rose tissue cultured course, it was found that the content of
IAA in rhizogenic zone increased before the formation of primordium and
decreased after formation. While iPAs and ZRs slightly decreased before
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41、Skirrin RM, Chu MC and Young HJ. Rose 1990.In: Ammirato P.V., Sharp W. R. Evans D.A. and Bajaj Y.P.S..Handbook of Plant Cell Culture Vols(Ornamental Species). McGaw-Hill,New York,716-743
42、Elliott RF, Axenic. Culture of Meristem Tips of Rosa Multiflora. Planta, 1970(95): 183-186
43、Zieslin N, Halevy AH. Components of Axillary Bud Inhibition in Rose Plants. 1. The Effect of Different Plant Parts Correlative Inhibition. Bot, Gaz. 1976(137):291-296
44、Hasegawa PM. In vitro Propagation of Rose. Scientia Horticulturae. 1979(14):610-612
45、Davies DR. Papid Propagation of Rose in vitro.Sci Hortic,1980(13):385-389
46、Hasegawa PM. Factors Affecting Shoot and Root Initiation from Cultured Rose Shoot Tip.J.Am. Soc Scientia Horticulturae. 1980(105):216-220
47、Wulster G, Sacalis J. Effects of Auxin and Cytokinnins on Ethylene Evolution and Growth of Rose Callus Tissure in Sealed Vessels. Scientia Horticulturae.
1900(15) :736-737
48, Rout GR, Debata BK, Das P. In vitro Mass-scale Propagation of Rosa Hybrida cv.Laudora Curr. Scientia Horticulturae. 1989(58) :876-878
49, Rout GR, Debata BK, Das P. In vitro Clonal Multiplication of Roses . Proc Natl. Acad Sci.(India)1990(60) :311-318
50, Rout GR, Debata BK, Das P. Chloropromazine Induced in vitro Bud Break in Rosa Hybrida cv. Landora Orissa J. Hort. 1992(20) : 8-16
51, Kever C ,Boyer H, Courduroux J, Caspar Th.. The Influence of Ethelene on Proliferation and Growth of Rose Shoot Culture. Plant Cell. Tissue and Organ Culture. 1992(28) : 175-181
52 , Chu CY, Knight SL and Smith AL. Effect of Liquid Culture on the Growth and Development of Miniature Rose (RosaChinensis Jacq 'minima' ) Plant Cell/Tissue and Or-gan Culture. 1993(32) :329-334
53, Lanford PJ, Wainwright H. Effect of Sucrose Concentration on the Photosynthetic Ability of Rose Shoots in Vitro .Ann Bot 1987(60) :633
54, Begin-Sallanon H, Coudret A, Gendraud M. Relations between Intracellular PH.Water Relations and Morphogensis in Rose Plants in Vitro Biol. Plant,1990,32(1) :58-63
55, Hyndman SE, Hasegawa PM, Bressan RA. Stimulation of Root Initiation from Cultured Rose Shoots Through the Use of Reduced Concentration of Mineral Salts. Scientia Horticulturae. 1982(17) :82-83
56, Skirrin RM, Chu MC. In Vitro Propagation of 'Forever Yours 'Rose. Scientia Horticulturae. 1979(14) :608-610
57, Khosh-Khui M, Sink KC. Rooting Enhancrment of Rosa Hybrida for Tissue Cultere Propagation. Scientia Horticulturae. 1982(17) :371-376
58, Khosh-Khui M, Sink KC. Micropropagation of New and Old World Rose Species. J. Scientia Horticulturae. 1982(17) :371-376
59, Sauer A, Walther F, Precl W. Different Suitablity for in Vitro Propagation of Rose Cultivars. Gartenbauwissenschaft, 1985(50) : 133-138
60, Douglas GC, Rutkdge CB, Casey AD, Richardson,DHS. Micropropagtion of Floribunda Ground Cover and Miniature Roses. 1989(19) :55-64
61, Arnold NP, Binns MR, Cloutier CD, Barthakur NN, Pellerin R. Auxin Salt Concentrations and their Interactions during in vitro Rooting of Wiuter-hardy and Hybrid tea Roses. Scientia Horticulturae. 1995(30) : 1436-1440
62, Skirrin RM,Chu MC. The Effect of Light Quality on Root Development on
inVitro Growth Miniature Roses. Scientia Horticulturae.1984(19):575(Abster)
63、Collet GF, Enracinement Ameliore iors de La Production in vitro de Rosiers. Rev. Snisse Vitic Arboric Hortic. 1985(17):259-263
64、Lloyd D, Roberts AV, Short KC. The Induction in Vitro of Adventitious Shoots in Rosa.Euphytica 1988(37):31-36
65、Rout GR. Studies on in vitro Mass Production of Rose Cultivars Ph.D. Thesis Htkal University India 1991
66、Preece JE, Sutter EG. Acclimation of Microprogagated Plants to the Greenhouse and Field.In:DeberghP. C. Zimmer man R.H.(Eds) Micropropagation Kluwer The Nertherlands, 1991
67、Rout GR, Debata BK, Das P. Micropropagation of Rosa Hybrida cv. Queen Elizabeth through in vitro Culture of Axillary buds.Orissa J. Hort. 1989(16): 1-9
68、Scotticampos P, Pais MS. Masspropfgatio of the Dwarf Rose Cultivar'Rosa mini'. Scientia Horticulturae. 1990(43):321-330
69、朱鹿鸣，金韵琴，周娟芳。名贵月季试管苗繁殖和试管苗移栽。园艺学报，1985，12(4)：273-276
70、Dubois LAM, Roggemans J, Soyeurt G, De Vries DP. Comparison of the Growth and Development of Dwarf Rose Cultivars Propagatde in vitro and in Vivo by Soft Cuttings.Scientia Horticulturae. 1988(35):293-299
71、Roberts AV, Walker S, Horan I, Smith EF, Mottley J. The Effects of Growth Retardants,Humidity and Lighting at Stage-3 on Stage-2 of Microprogagation in Chrysanthemum and Rose. Acta Hortic. 1992(319):135-158
72、Horan I, Walker S, Roberts AV, Mottley J. Simpkins I.. Micropropagation of Roses:The Benefits of Pruned Mother-plantlets at Stage-2 and a Greenhouse Environment at Stage-3. J. Scientia Horticulturae. 1995,70(5):799-806
73、Hill GP. Morphogenesis of Shoot Primordia in Cultivated Stem Tissue of a Garden Rose. Nature, 1976(216):596-597
74、Jacobs G,Bornman CH, Allan P. Tissue Culture Studies on Rose:Use of Pith Explants. South Afric J. Agric-sci. 1983(11):673-678
75、Tabaeezadeh Z, Khorh-Khui M. Anther Culture of Rosea Scientia Horticulturae, 1981(15):61-66
76、Zieslin H, Gavish H, Ziv M. Growth Interaction between Calli and Explants of Rose Plants in vitro. Plant Sci., 1987(48):57-62
77、Boxus,1978
78、Dicer and Custer(1988)
79、Burger DW, Liu L, Zary KW, Lee C.I. Organogensis and Plant Regeneration from Immature Embryos of Rosa Hybrida L.Plant Cell Tissue Organ Culture 1990(21):147-152
80、Rout G.R.,Rebata B.K.,Das. R.In vitro Regeneration of Shoots from Callus Cultures of Rosa Hybrida cv.:andora Ind.J. Exp. Biol. 1992(30): 15-18
81、Ishioka N., Tanimoto S..Plant Regeneration from Bulgarian Rose Callus. Plant Cell Tissue Organ Culture 1990(22): 197-199
82、Dubois L.A.M. de Vries D.P. Preliminary Report on the Diret Regeneration of Adventitious Buds on Leaf Explants of in vitro Growth Glass-house Rose Cultivars. Gartenb au Wissenschaft. 1995(60):249-253
83、Rosu A.,Skirrin R.M.,Bein A.,Norton M.A., Kushed M., Otterbacher A.G.. The Development of Putative Adventitious Shoots from a Chimeral Thornlers Rose(Rosa Multiflora Thurb, ex J.Murr) in vitro T. Hort. Sci. 1995(70):901-907
84、Hisa C.N. Korbau S.S. Organ Ogenesis and Somatic Embryogensis in Callus Cultures of Rosa Hybrida and Rosa Chinensis Minima Plant Cell Tissue Organ Culture 1996(44): 1-6
101、Onesto J P, Poupet R and Julien P. Production de fieuries de rosier a partir de plantules obteruces par multiplication in vitro conforme Automne 1983printemps 1984. Hortic, Francaise, 1985:3-10
102、Reist A. Culture in vitro (civ) en pepiniere de rosiers: une alternative au bouturage ou au greffage des varieties. Rev. Suisse vatic Arboric Hortic 1985(17): 361-364
103、谭文澄，戴策刚。观赏植物组织培养技术。中国林业出版社，1991
104、Leifert C, Morris C and Waites W M.Ecology of microbial saprophytes and pathogens in tissue culture and field-grown plants: Reasons for contamination problems in vitro. CRC Crit Rev, Plant Science, 1994(13): 139-183
105、Skirvin RM, chu M C, Young H J. Rose. In: Ammiratio Sharp W R, Evans D A, and Bajaj Y P S(Editors).Handbook of plant cell culture. Vol. 5(Ornamental species) Mc Graw-Hill, New York, 1990:716-743
106、Dodds J R and Roberts L W. Some inhibitory effects of gentamicin on plant tissue cultures. In vitro, 1981(17): 467-470
107、Falkiner F R. The criteria for choosing an antibiotic for control of bacteria in plant tissue culture. Int.assoc Plant Tissue culture. Newsl, 1990(60): 13-23
108、Leifert C, Morris c. and waites W M. Effect of combinations of antibiotics on micropropagation of clematis, Delphinnium, Hosta, Iris and Photinia. Plant cell tissue organ culture, 1992(29): 1353-160
109、Salehi H.In vitro propagation of miniature rose cultivars MS. Thesis. Shiraz university, shiraz Iran, 1995
110、Marcelis-Van, Acker C A m, Scholten Hj. Development of axillary buds on rose in vitro. Sci. Hortic, 1995(63):47-55
111、Horn W A H. Micropropogation of rose. In: Biotechnology in agriculture and foresty. Vol, 20. Springer-verlag, germany, 1992,320-342
112、Bhojwani S S, Mullins K and Cohen D. Micropropagation of Feijoa sellowina Berg. Acta. Hortic, 1987(212):69-76
113、Barre D M, Lyer R S, Kendurkars, Mascarenns A F. An efficient method for mrapid propagation of some budded rose varieties. Ind J. Hort, 1984(41): 1-7
114、Zamski E S, Oshri and Zieslin N.Compartive Morphology and antatomy of axillary buds. Bot Gaz, 1985(146):208-212
115、杨文政，陈树果，王宝档等。月季名贵品种无性系的快速繁殖化生产的万赖俱寂。河南农学院学报，1984(2)：8-14
116、podwyszynska M and Hemple M. the factors influencing acclimation of Rosa hybrida plants multiplied in vitro to greenhouse condition. Acta. Hortic, 1988(226):639-642
117、Alderson P G and Mckinless J. Rooting of cultured rose shoots. Acta. Hortic, 1988(226) 175-182
118、Mcclelland M J, Smith M A and Carothers E B. The effects of in vitro and exvitro root initiation on subsequent microcutting root quality in three woody plants. Plant cell tissue organ culture, 1990(23): 115-123
119、Smith M A L and Mcclelland M T. Gauging the influence of in vitro conditions on in vivo quality and performance of woody plants. In vitro cellular and performance of woody plants. In vitro cellular and developmental bilogy,1991(27): 52-56
120、胡祖坤，贾元淑，宋云枝，南晓英，陈惠莉。月季试管苗不定根发生的组织学和细胞学研究。山东农业大学学报，1990(1)：73-77
121、韩碧文，李颖章。植物组织培养中器官建成的生理生化基础。植物学通报，1993，10(2)：1-6
2、余树勋，张本。月季。上海科学出版社，1998
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5、Renate M(?)ller, Arne S. Ansderson, Mangrethe Serek. Ditterences in display life of minatare potted roses. Scientia Horticulturne 1998, 76:59-71
6、北京林业大学园林系花卉教研组。花卉学。中国林业出版社，1998
7、朱键镛。品种、小叶数及季节对玫瑰花单节扦插繁殖之影响。中国园艺，1997，43(1)：29—40
8、Marcelis-van Acker CAM, Leutscher KJ. Effect of Cutting on Heterogeneity and Growth of Rosa Hybrida cv. 'Motrea' and Schefflera arboricola cv.'Compacta', 1993(54): 59-67
9、Stoltz LP, Anderson RG. Rooting of Single Node Cutting of Roses. Acta Horticulturae, 1998(227):230-235
10、Deveries DP and Dubios Lidwien AM.The Effect of BAP and IBA on Sprouting and Adventitious Root Formation of 'Amanda' Rose Single-node Softwood Cuttings. Scientia Horticulturae. 1998(34): 115-121
11、Lidwien AM, Dubis, DP de vries. Variation in Adventitious Root Formation of Wood Cutting of Rosa Chinensis Minima (Sims) voss Cultivars. Scientia Horticulturae. 1991(47):345-349
12、朱键镛，雪伦耐特。由单节扦插苗、芽接扦插苗和微体扦插苗生产迷你玫瑰盆花。中国园艺，1992，38(2)：55—629
13、Moe R.Propagation Growth and Flowering of Potted Roses.Acta Horticulturae,1973(76):40-42
14、朱键镛。玫瑰切花栽培之种苗生产(一)单节扦插苗。台湾花卉园艺，1993(76)：40—42
15、沈再发，林后彦译。花卉之岩棉栽培——玫瑰岩棉在园艺作物栽培上之利用。台北市七星农田水利会研究发展基金会，1992：159-167
16、McCann S. Miniature Roses. Arco Publishing,Inc, NewYork, 1985
17、Sachs RMAM Ko Franck and Hackett WP. Evaluating New Pot Plant Species. Florists Rev. 1976,4116(159):35-36,80-84
18、Chu CY. Budded Cuttings for Propagating Roses. Scientia Horticulturae. 1990(43):163-168
19、Ohkawa K. Cutting-grafts as a Means to Propagate Greenhouse Roses. Scientia Horticulturae. 1980(13): 191-199
20、Van de pol PA, Breukelear A.Stenting of Roses: A Method for Grafting. Scientia Horticulturae. 1982(17): 187-196
21、Van de Pol PA, Joossten MHAJ, Keizer H. Stenting of Rose,Starch Depletion and Accumulation During the Early Development. Acta. Hortic. 1986(189):51-59
22、De Vries DP and. Dubois LAM. Variation for Plant Characters and for Performance of Softwood Cuttingd of 'R.Canina Inermis' Seedling. Euphytica, 1987(36):407-412
23、Dubois LAM, OP de Vries. Varation in an Adventitious Root Formation of Softwood Cutting Rosa Chinensis. Minima (Sims) Voss Cultivars. Scientia Hortic, 1991(47):345-349
24、Hambrick CE, DaviesFT, Pemberton HB. Seasonal Change in Carbonhydrate Nitrogen Levels during Field Rooting of Rosa Multiflora 'Brook56'Hardwood CuRings. Scientia Horticulturae. 1991(46): 137-146
25、Drew RA, Miller RM. Nutritional and Cultural Factors Affecting Rooting of Papaya (Carica Papaya L.) in vitro. Scientia Horticulturae. 1989(64):767-773
26、Kim KY,Hanada T, Takahshi K. Effects of the Magnesium Content of the Culture Solution on the Photosynthetic Activity of Cumcumber Leaves. Horticulturae. And Seri-culture. 1981(23):1-9
27、Mengel DK, Kirkby EK. Principle of Plant Nutrition.4~(th) ed Int. Potash Inst Beru. Switzerland 1987
28、Salidbury FB, Ro-SS CW. Plant Physiology 3~(rd) ed Wadsorth Inc. U.S.A. 1985
29、Kundson L. Flower Production by Orchid Growth Non-symbiotically Bot,Gaz 1922(89): 192
30、黄学林，李筱菊。高等植物组织立体培养的形态建成及其调控。北京：科学技术出版社，1995
31、Routn GR, Samantaray S, Mottley J, Das P. Biotechnology of the Rose: A review of Recent Progress. Scientia Horticulturae. 1990(81):201-228
32、Salehi H and Khosh-Khui M. Effects of Explant Length and Diameter on in vitro Shoot Growth and Proliferation Rate of Miniature Roses. Journal of Scientia Horticulturae. 1997,72(5):673-676
33、Salehi H, Khosh-khui MA. Simple Procedure for Disinfection of 'Baby Masquerade'Miniature Rose Explants. Scientia Horticulturae. 1997(68):145-148
34、郑玉梅，马炜，刘青林。微型月季的离休培养与快速繁殖。植物生理学通讯，2000：96-101
35、Bressan PH, Kim YJ, Hyndman SE, Hasegawa PM,Bressan RA. Factors Affecting in vitro Propagation of Roses. J.Am. Soc. Scientia Horticulturae. 1982(107):97-990
36、Podwyszynka M, Olszewsri J. Influence of Gelling Agents on Shoot Multiplication and the Uptake of Marcoclements by in vitro Culture of Rose. Cordyline and Homalomena Scientia Horticulturae. 1995(64):77-84
37、Scotti Campos P, Salome pais M. Mass Propagation of the Dwarf Rose Cultivar 'Rosa mini' Scientia Horticulturae. 1990(43):321-330
38、朱键镛，雪伦耐特，玛利斯密期。影响迷你玫瑰茎节培养枝稍生产力之因子。中国园艺，1992，38(1)：7-18
39、Gerad C, Doglas,Clare B, Rutedge,AngelsdD, Casey and David HS Richadson. Micropropagation of Floribunda, Ground Cover and Miniature Rose. Plant Cell,Tissue and Culture, 1989,19:55-64
40、Lidwien AM, DubiosJ, Roggemans G, Soywurt and Devries DP. Camparison of the Growth and Development of Dwarf Rose Cultivars Propagated in vitro and in vivo by Softwood Cuttings. Scientia Horticulturae. 1988(35):293-299
41、Skirrin RM, Chu MC and Young HJ. Rose 1990.In: Ammirato P.V., Sharp W. R. Evans D.A. and Bajaj Y.P.S..Handbook of Plant Cell Culture Vols(Ornamental Species). McGaw-Hill,New York,716-743
42、Elliott RF, Axenic. Culture of Meristem Tips of Rosa Multiflora. Planta, 1970(95): 183-186
43、Zieslin N, Halevy AH. Components of Axillary Bud Inhibition in Rose Plants. 1. The Effect of Different Plant Parts Correlative Inhibition. Bot, Gaz. 1976(137):291-296
44、Hasegawa PM. In vitro Propagation of Rose. Scientia Horticulturae. 1979(14):610-612
45、Davies DR. Papid Propagation of Rose in vitro.Sci Hortic,1980(13):385-389
46、Hasegawa PM. Factors Affecting Shoot and Root Initiation from Cultured Rose Shoot Tip.J.Am. Soc Scientia Horticulturae. 1980(105):216-220
47、Wulster G, Sacalis J. Effects of Auxin and Cytokinnins on Ethylene Evolution and Growth of Rose Callus Tissure in Sealed Vessels. Scientia Horticulturae.
1900(15) :736-737
48, Rout GR, Debata BK, Das P. In vitro Mass-scale Propagation of Rosa Hybrida cv.Laudora Curr. Scientia Horticulturae. 1989(58) :876-878
49, Rout GR, Debata BK, Das P. In vitro Clonal Multiplication of Roses . Proc Natl. Acad Sci.(India)1990(60) :311-318
50, Rout GR, Debata BK, Das P. Chloropromazine Induced in vitro Bud Break in Rosa Hybrida cv. Landora Orissa J. Hort. 1992(20) : 8-16
51, Kever C ,Boyer H, Courduroux J, Caspar Th.. The Influence of Ethelene on Proliferation and Growth of Rose Shoot Culture. Plant Cell. Tissue and Organ Culture. 1992(28) : 175-181
52 , Chu CY, Knight SL and Smith AL. Effect of Liquid Culture on the Growth and Development of Miniature Rose (RosaChinensis Jacq 'minima' ) Plant Cell/Tissue and Or-gan Culture. 1993(32) :329-334
53, Lanford PJ, Wainwright H. Effect of Sucrose Concentration on the Photosynthetic Ability of Rose Shoots in Vitro .Ann Bot 1987(60) :633
54, Begin-Sallanon H, Coudret A, Gendraud M. Relations between Intracellular PH.Water Relations and Morphogensis in Rose Plants in Vitro Biol. Plant,1990,32(1) :58-63
55, Hyndman SE, Hasegawa PM, Bressan RA. Stimulation of Root Initiation from Cultured Rose Shoots Through the Use of Reduced Concentration of Mineral Salts. Scientia Horticulturae. 1982(17) :82-83
56, Skirrin RM, Chu MC. In Vitro Propagation of 'Forever Yours 'Rose. Scientia Horticulturae. 1979(14) :608-610
57, Khosh-Khui M, Sink KC. Rooting Enhancrment of Rosa Hybrida for Tissue Cultere Propagation. Scientia Horticulturae. 1982(17) :371-376
58, Khosh-Khui M, Sink KC. Micropropagation of New and Old World Rose Species. J. Scientia Horticulturae. 1982(17) :371-376
59, Sauer A, Walther F, Precl W. Different Suitablity for in Vitro Propagation of Rose Cultivars. Gartenbauwissenschaft, 1985(50) : 133-138
60, Douglas GC, Rutkdge CB, Casey AD, Richardson,DHS. Micropropagtion of Floribunda Ground Cover and Miniature Roses. 1989(19) :55-64
61, Arnold NP, Binns MR, Cloutier CD, Barthakur NN, Pellerin R. Auxin Salt Concentrations and their Interactions during in vitro Rooting of Wiuter-hardy and Hybrid tea Roses. Scientia Horticulturae. 1995(30) : 1436-1440
62, Skirrin RM,Chu MC. The Effect of Light Quality on Root Development on
inVitro Growth Miniature Roses. Scientia Horticulturae.1984(19):575(Abster)
63、Collet GF, Enracinement Ameliore iors de La Production in vitro de Rosiers. Rev. Snisse Vitic Arboric Hortic. 1985(17):259-263
64、Lloyd D, Roberts AV, Short KC. The Induction in Vitro of Adventitious Shoots in Rosa.Euphytica 1988(37):31-36
65、Rout GR. Studies on in vitro Mass Production of Rose Cultivars Ph.D. Thesis Htkal University India 1991
66、Preece JE, Sutter EG. Acclimation of Microprogagated Plants to the Greenhouse and Field.In:DeberghP. C. Zimmer man R.H.(Eds) Micropropagation Kluwer The Nertherlands, 1991
67、Rout GR, Debata BK, Das P. Micropropagation of Rosa Hybrida cv. Queen Elizabeth through in vitro Culture of Axillary buds.Orissa J. Hort. 1989(16): 1-9
68、Scotticampos P, Pais MS. Masspropfgatio of the Dwarf Rose Cultivar'Rosa mini'. Scientia Horticulturae. 1990(43):321-330
69、朱鹿鸣，金韵琴，周娟芳。名贵月季试管苗繁殖和试管苗移栽。园艺学报，1985，12(4)：273-276
70、Dubois LAM, Roggemans J, Soyeurt G, De Vries DP. Comparison of the Growth and Development of Dwarf Rose Cultivars Propagatde in vitro and in Vivo by Soft Cuttings.Scientia Horticulturae. 1988(35):293-299
71、Roberts AV, Walker S, Horan I, Smith EF, Mottley J. The Effects of Growth Retardants,Humidity and Lighting at Stage-3 on Stage-2 of Microprogagation in Chrysanthemum and Rose. Acta Hortic. 1992(319):135-158
72、Horan I, Walker S, Roberts AV, Mottley J. Simpkins I.. Micropropagation of Roses:The Benefits of Pruned Mother-plantlets at Stage-2 and a Greenhouse Environment at Stage-3. J. Scientia Horticulturae. 1995,70(5):799-806
73、Hill GP. Morphogenesis of Shoot Primordia in Cultivated Stem Tissue of a Garden Rose. Nature, 1976(216):596-597
74、Jacobs G,Bornman CH, Allan P. Tissue Culture Studies on Rose:Use of Pith Explants. South Afric J. Agric-sci. 1983(11):673-678
75、Tabaeezadeh Z, Khorh-Khui M. Anther Culture of Rosea Scientia Horticulturae, 1981(15):61-66
76、Zieslin H, Gavish H, Ziv M. Growth Interaction between Calli and Explants of Rose Plants in vitro. Plant Sci., 1987(48):57-62
77、Boxus,1978
78、Dicer and Custer(1988)
79、Burger DW, Liu L, Zary KW, Lee C.I. Organogensis and Plant Regeneration from Immature Embryos of Rosa Hybrida L.Plant Cell Tissue Organ Culture 1990(21):147-152
80、Rout G.R.,Rebata B.K.,Das. R.In vitro Regeneration of Shoots from Callus Cultures of Rosa Hybrida cv.:andora Ind.J. Exp. Biol. 1992(30): 15-18
81、Ishioka N., Tanimoto S..Plant Regeneration from Bulgarian Rose Callus. Plant Cell Tissue Organ Culture 1990(22): 197-199
82、Dubois L.A.M. de Vries D.P. Preliminary Report on the Diret Regeneration of Adventitious Buds on Leaf Explants of in vitro Growth Glass-house Rose Cultivars. Gartenb au Wissenschaft. 1995(60):249-253
83、Rosu A.,Skirrin R.M.,Bein A.,Norton M.A., Kushed M., Otterbacher A.G.. The Development of Putative Adventitious Shoots from a Chimeral Thornlers Rose(Rosa Multiflora Thurb, ex J.Murr) in vitro T. Hort. Sci. 1995(70):901-907
84、Hisa C.N. Korbau S.S. Organ Ogenesis and Somatic Embryogensis in Callus Cultures of Rosa Hybrida and Rosa Chinensis Minima Plant Cell Tissue Organ Culture 1996(44): 1-6
101、Onesto J P, Poupet R and Julien P. Production de fieuries de rosier a partir de plantules obteruces par multiplication in vitro conforme Automne 1983printemps 1984. Hortic, Francaise, 1985:3-10
102、Reist A. Culture in vitro (civ) en pepiniere de rosiers: une alternative au bouturage ou au greffage des varieties. Rev. Suisse vatic Arboric Hortic 1985(17): 361-364
103、谭文澄，戴策刚。观赏植物组织培养技术。中国林业出版社，1991
104、Leifert C, Morris C and Waites W M.Ecology of microbial saprophytes and pathogens in tissue culture and field-grown plants: Reasons for contamination problems in vitro. CRC Crit Rev, Plant Science, 1994(13): 139-183
105、Skirvin RM, chu M C, Young H J. Rose. In: Ammiratio Sharp W R, Evans D A, and Bajaj Y P S(Editors).Handbook of plant cell culture. Vol. 5(Ornamental species) Mc Graw-Hill, New York, 1990:716-743
106、Dodds J R and Roberts L W. Some inhibitory effects of gentamicin on plant tissue cultures. In vitro, 1981(17): 467-470
107、Falkiner F R. The criteria for choosing an antibiotic for control of bacteria in plant tissue culture. Int.assoc Plant Tissue culture. Newsl, 1990(60): 13-23
108、Leifert C, Morris c. and waites W M. Effect of combinations of antibiotics on micropropagation of clematis, Delphinnium, Hosta, Iris and Photinia. Plant cell tissue organ culture, 1992(29): 1353-160
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