白杨年龄与位置效应的生理生化机制研究
摘要
无性繁殖技术在林业生产中应用广泛,具有广阔的发展前景,但无性系生长易受母树年龄和接穗与插条在原株取样部位——成熟效应和位置效应的影响而出现老化的现象。本研究以不同年龄、不同部位繁殖材料嫁接来源的三倍体毛白杨苗木和几种树木不同部位当年生枝条为研究材料,就树木年龄效应和位置效应的相关形态、生长以及生理生化等指标进行研究,期望找到判断树木个体发育阶段的有效标记,为生产中选择适宜繁殖材料作出指导。主要结果如下:
1.三倍体毛白杨不同部位繁殖材料来源的苗木形态和生长指标大多存在显著差异,表现为随繁殖材料在母树上所处高度位置增加,分株的节间长度、叶面积、地径生长量、高生长量、叶柄长度逐渐减小,而且这种差异主要存在于根部与冠部位置材料来源的苗木之间。
2.不同年龄、不同部位繁殖材料来源三倍体毛白杨苗木的光合生理指标和自由基相关指标存在一定的差异,但是这种变化没有一定的规律性,表明在研究三倍体毛白杨年龄与位置效应时,难以通过光合作用以及自由基相关的生理指标进行有效判别。
3.不同年龄、不同部位繁殖材料来源的苗木以及树木不同部位当年生枝条叶片的水溶性总蛋白含量和组分存在一定的差异,但在不同部位、不同年龄以及不同树种中,其蛋白含量或蛋白组分并不存在相同的变化规律,因此难以将蛋白含量或组分与树木不同的发育阶段间建立联系,不能将其用于无性繁殖材料选择的相关标记。
4.单一内源植物激素IAA或ABA的含量变化在不同部位繁殖材料来源的苗木,以及树木不同位置当年生枝条中没有一定的规律可循,但IAA/ABA值则表现出规律性变化,树木基部当年生萌条和根部位置繁殖材料来源苗木的IAA/ABA比值高于中部或冠部冠部位置,表明树木发育阶段转变与内源激素IAA与ABA含量消长相关,因此,可以通过与设置的已知的幼化或老化材料参照系IAA/ABA值的比较,对树木位置效应的相对差异进行判别。
Vegetative propagation technique is applied widely in forestry production and it have wide development prospects, but clones were easily affected by the age of mother tree and the position scions and cuttings in ortet-maturation effect and position effect-appeared ageing phenomenon. In this research, morphological, growth, physiological and biochemical indexes about age and position effect were studied using ramet from different age and different position of triploid Chinese white poplar to find some markers that judge development stage of trees and to guide selecting suitable propagation material in production. The major results are described as follows:
1. The morphological and growth indexes were significant differences among propagation material from different position of triploid Chinese white poplar ortet. The internode length, leaf aera, base diameter growth, height growth and petiole length decrease gradually with the height of propagation material increased in ortet. This differences mainly existed in ramets that from propagation material of root and crown position
2. The photosynthetic physiological indexes and indexes about free radical have some differences between propagation material from different age and different position of triploid Chinese white poplar ortet, but this variation have no discernible regularity. So, it was hard to judge maturation effect and position effect of Chinese white poplar with photosynthetic physiological indexes and indexes about free radical
3. The soluble proteins contents and fractions have some differences between propagation material from different age and position of triploid Chinese white poplar ortet and current-year branch in different position of trees, but this variation have not some regular rule in different positions, ages of trees and different tree species. So, it was hard to construct relationships between soluble proteins contents, proteins fractions and the different development phase of trees, therefore, it was not suitable as a marker in vegetative propagation
4. The single plant hormone contents of IAA and ABA have not same rule neither in ramets from different position propagation materials of triploid Chinese white poplar ortet or in current-year branch at different position of trees. But the IAA/ABA has same rule: the rate of IAA/ABA in current-year branch at basal part of trees and ramets from root is higher than stem and crown propagation materials of triploid Chinese white poplar ortet. This showed that phase change of trees have relationships with contents variation of IAA and ABA. So, the relative differences of position effect of trees can be judged by comparing the IAA/ABA rate to reference materials that already juvenility or aged
1.三倍体毛白杨不同部位繁殖材料来源的苗木形态和生长指标大多存在显著差异,表现为随繁殖材料在母树上所处高度位置增加,分株的节间长度、叶面积、地径生长量、高生长量、叶柄长度逐渐减小,而且这种差异主要存在于根部与冠部位置材料来源的苗木之间。
2.不同年龄、不同部位繁殖材料来源三倍体毛白杨苗木的光合生理指标和自由基相关指标存在一定的差异,但是这种变化没有一定的规律性,表明在研究三倍体毛白杨年龄与位置效应时,难以通过光合作用以及自由基相关的生理指标进行有效判别。
3.不同年龄、不同部位繁殖材料来源的苗木以及树木不同部位当年生枝条叶片的水溶性总蛋白含量和组分存在一定的差异,但在不同部位、不同年龄以及不同树种中,其蛋白含量或蛋白组分并不存在相同的变化规律,因此难以将蛋白含量或组分与树木不同的发育阶段间建立联系,不能将其用于无性繁殖材料选择的相关标记。
4.单一内源植物激素IAA或ABA的含量变化在不同部位繁殖材料来源的苗木,以及树木不同位置当年生枝条中没有一定的规律可循,但IAA/ABA值则表现出规律性变化,树木基部当年生萌条和根部位置繁殖材料来源苗木的IAA/ABA比值高于中部或冠部冠部位置,表明树木发育阶段转变与内源激素IAA与ABA含量消长相关,因此,可以通过与设置的已知的幼化或老化材料参照系IAA/ABA值的比较,对树木位置效应的相对差异进行判别。
Vegetative propagation technique is applied widely in forestry production and it have wide development prospects, but clones were easily affected by the age of mother tree and the position scions and cuttings in ortet-maturation effect and position effect-appeared ageing phenomenon. In this research, morphological, growth, physiological and biochemical indexes about age and position effect were studied using ramet from different age and different position of triploid Chinese white poplar to find some markers that judge development stage of trees and to guide selecting suitable propagation material in production. The major results are described as follows:
1. The morphological and growth indexes were significant differences among propagation material from different position of triploid Chinese white poplar ortet. The internode length, leaf aera, base diameter growth, height growth and petiole length decrease gradually with the height of propagation material increased in ortet. This differences mainly existed in ramets that from propagation material of root and crown position
2. The photosynthetic physiological indexes and indexes about free radical have some differences between propagation material from different age and different position of triploid Chinese white poplar ortet, but this variation have no discernible regularity. So, it was hard to judge maturation effect and position effect of Chinese white poplar with photosynthetic physiological indexes and indexes about free radical
3. The soluble proteins contents and fractions have some differences between propagation material from different age and position of triploid Chinese white poplar ortet and current-year branch in different position of trees, but this variation have not some regular rule in different positions, ages of trees and different tree species. So, it was hard to construct relationships between soluble proteins contents, proteins fractions and the different development phase of trees, therefore, it was not suitable as a marker in vegetative propagation
4. The single plant hormone contents of IAA and ABA have not same rule neither in ramets from different position propagation materials of triploid Chinese white poplar ortet or in current-year branch at different position of trees. But the IAA/ABA has same rule: the rate of IAA/ABA in current-year branch at basal part of trees and ramets from root is higher than stem and crown propagation materials of triploid Chinese white poplar ortet. This showed that phase change of trees have relationships with contents variation of IAA and ABA. So, the relative differences of position effect of trees can be judged by comparing the IAA/ABA rate to reference materials that already juvenility or aged
引文
1. 陈大明,沈德绪,李载龙.梨实生树童区和成年区顶端分生组织的细胞学研究.植物生理学通讯,1994,30(5):343-345
2. 陈东攻,赵永波,俎文房,张新忠.桃实生树不同节位叶绿素、可溶性蛋白的动态变化.河北农业大学学报,2005,28(1):16-17
3. 谷瑞升,蒋湘宁.一种省时高效的木本植物蛋白双向电泳分析方法.北京林业大学学报,1999,21 (5):7-10
4. 郝再彬,苍晶,徐仲.植物生理实验.哈尔滨工业大学出版社,2004
5. 金勇丰,李载龙,陈大明.湖北海棠实生树阶段转变过程中多胺!RNA/DNA 和蛋白质含量的变化浙江农业学报,1998,10(5):264-267
6. 李新国,段安安,朱之悌.毛白杨无性系年龄效应及根萌条幼化效果的初步研究.西南林学院学报,1998,18(2):61-67
7. 林晓东.激素调节花芽分化的研究进展.果树学报,1997,14(4):269-274
8. 刘俊,吕波,徐朗莱.植物叶片中过氧化氢含量测定方法的改进.生物化学与生物物理进展,2000,27(5):548-551
9. 刘松泉,黄香等.幼态与老态无性系产量及性状比较的研究.热带作物研究,1985,2:1-4
10. 刘松泉.植物发育阶段理论在提高巴西橡胶产胶力中的应用.热带作物学报,1988,9(2):8-11
11. 龙桂友,沈德绪,林柏年.巨峰葡萄实生苗阶段发育过程中几种同工酶酶谱的变化.果树科学,1991,8(2):91-96
12. 陆定志,傅家瑞,孙松泉.植物衰老及其调控.中国农业出版社,1997
13. 马常耕.世界云杉无性系林业发展现状.世界林业研究,1993,6:24~31.
14. 沈成国等.植物衰老生理与分子生物学.中国农业出版社,2001
15. 师晨娟,刘勇,王春城,司瑞,新张林玉.青海云杉扦插的年龄效应及其生根机理研究.西北农林科技大学学报,2006,34(12):101-104
16. 施季森,何祯祥.林木无性繁殖及其在遗传改良中的地位.世界林业研究,1994,(1):25~30
17. 宋纯鹏.植物衰老生物学.北京大学出版社,1998
18. 苏培玺,张立新,杜明武,毕玉容,赵爱芬,刘新民.胡杨不同叶形光合特征水分利用效率及其对加富 CO2的响应.植物生态学报,2003,27(1):34-40
19. 孙永华,陈学森.杏杂种实生树叶片童性的研究.果树学报,2005,22(4):327-330
20. 田义,张志宏,李春敏,张新忠.木本植物阶段转变分子机制的研究进展.华北农学报,2006,21: 15-18
21. 王爱国,罗广华,邵从本.大豆种子超氧化物歧化酶的研究.植物生理学报,1983,9(1):77-84
22. 王定祥.梨实生树个体发育过程中蛋白质含量、过氧化物酶和多酚氧化酶同工酶的变化.植物生理学报,1986,12(1):40-46
23. 王建华,孙晓梅,王笑山等.母株年龄、激素种类及其浓度对日本落叶松扦插生根的影响.林业科学研究,2006,19(1):102-108
24. 王军辉,张建国,张守攻等.青海云杉硬枝扦插的激素、年龄和位置效应研究.西北农林科技大学学报,2006,34(7):65-71
25. 王明庥.论无性系林业—概念和应用.南京林业大学学报,1993,(1):1~5
26. 王秋玉,赵丽惠,王福来,刘秀琴.红皮云杉扦插繁殖中的年龄效应及其生理机制.植物研究,1997, 17(3):338-343
27. 刑世岩,胡玉芹,苗全盛,张锐.银杏无性繁殖位置效应的研究.武汉植物学研究,1998,16(1):69-76
28. 许智宏,刘春明.植物发育的分子机理.科学出版社,1998
29. 杨江山,常永义,种培芳.樱桃不同节位叶片光合特性与解剖特性比较研究.果树学报,2005,22 (4):323-326
30. 叶金山,季孔庶,王章荣.杂种马褂木无性系插条生根能力的遗传变异.南京林业大学学报,1998, 22(2):71-74
31. 余雪标.论树木阶段发育特性余无性利用问题[J].河北农业大学学报,1994,4:645-651
32. 曾骧.果树生理学.北京农业大学出版社,1992,134-139
33. 张才喜,李载龙,陈大明.湖北海棠阶段转变的生理基础.植物资源与环境学报,2001,10(1):57-59
34. 张才喜,李载龙,陈大明.湖北海棠阶段转变过程中几种同工酶变化研究.植物学通报,2001,18 (1):100-104
35. 张志礼,郑学勤.树木发育阶段理论及其在橡胶树育种中的应用.热带亚热带植物学报,2001,9 (3):262-268
36. 朱广廉,植物生理学试验,北京大学出版社,1990
37. 朱之悌,盛莹萍论树木的老化——幼年性、成年性相互关系及其利用北京林业大学学报1992,14(3):92-104
38. 朱之悌.林木的无性繁殖与无性系育种.林业科学,1986,22(3):280-290.
39. 朱之悌.毛白杨遗传改良.中国林业出版社,2006
40. Aderkas P V, Bonga J M.Influencing micropropagation and somatic embryogenesis in mature trees by manipulation of phase change, stress and culture environment.Tree Physiology,2000,20:921- 928
41. Amo-Marco J B,Vidal N,Vieitez A M.Polypeptide markers differentiating juvenile and adult tissues in chestnut. A Ballester Journal of plant physiology,1993,142(11):117-119
42. Andres H,Fernandez B,Rodriguez R,Rodriguez A.Phytohormone contents in Corylus avellana and their relationship to age and other developmental processes.Plant Cell,Tissue and Organ Culture, 2002,70:173-180
43. Ballester A,San-jose M C,et al.Anatomical and biochemical events during in vitro rooting of microcuttings from juvenile and mature phases of chestnut.Annals of Botany.1999,83:619-629
44. Baurens F C,Nicolleau J,Legavre J L,Monteuuis O.Genomic DNA methylation of juvenile and mature Acacia mangium micropropagated in vitro with reference to leaf morphology as a phase change marker.Tree Physiol,2004,24:401-407
45. Baurens F C,Nicolleau L,LegaverT,et al.Genomic DNA methylation of juvenile and mature Acacia mangium micropropagate dinvitro with reference to leaf morphology as a phase change marker. Tree Physiol,2004,24:401-407.
46. Bitonti M B,Cozza R,Chiappetta A,et al.Distinct nuclear organization,DNA methylation pattern and cytokinin distribution mark juvenile-like and adult vegetative apical meristems in peach (Prunuspersica(L.)Batsch).JExpBot,2002,53:1047-1054
47. Bitonti M B,Cozza R,Chiappetta A,Gianniono D,et al.Distrinct nuclear organization,DNA methylation pattern and cytokinin distribution mark juvenile,juvenile-like and vegetative apical meristems in peach (Prunus persica(L.) Batsch).journal of Experimental Botany,2002,53(371): 1047-1054
48. Bon M C ,Monteuuis O.Rejuvenation of a 100-year-old Sequoiadendron giganteum through in vitro meristem culture.II.Biochemical arguments,Physiologia Plantarum,1991,81,116-120
49. Bon M C,Riccardi F,Monteuuis O.Influence of phase change within a 90-year-old Sequoia sempervirens on its in vitro organogenic capacity and protein patterns.Trees,1994,8:283-287
50. Bon M C.J16:An apex protein associated with juvenility of Sequoiadendron giganteum.Tree Physiology,1988,4,381-387
51. Bradford M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein–dye binding.Analytical Biochemistry,1976,72:248-254
52. Cabanne F,et al.Hydroxycinnamic acid amides and ripening to flower of Nicotiana tabacum var. xanthi nc.PlantPhysiol,1981,53(3):445-451
53. Centeno M L,Rodriguez R,Berros B,Rodriguez A.Endogenous hormonal content and somatic embryogenic capacity of Corylus avellana L.cotyledons.Plant Cell,1997,17:139-144
54. Chin T.Y,Meyer M.M,Beevers.Abscisic acid-stimulated rooting of stem cuttings.Planta,1989,88: 192-196
55. Clemens J,Henriod R E,Bailey DG,Jameson P E.Vegetative phase change in Metrosideros: shoot and root restriction.Plant Growth Regulation,1999,28:207-214
56. Elo A,Lemmetyinen J,Turunen M L,et al.Three MADS-box genes similar to APETALA1 and FRUITFULL from silver birch (Betula pendula).Physiologia Plantarum,2001,112:95-103
57. England J R,Attiwill P M.Changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species,Eucalyptus regnans F.Muell.Trees,2006,20:79-90
58. Ferwerda. F P.A possible explanation of the divergence between juvenile type budgrafts and their seeding mother trees in heven.Euphytica,1953,2:15-24
59. Ford YY,Taylor J M,Blake P S,Marks T R.Gibberellin A3 stimulates rooting of cuttings from cherry (Prunus avium).Plant Growth Regulation,2002,37:127-133
60. Fortanier E J,Jonkers H.Juvenility and maturity of plants as influenced by their ontogenetical and physiological ageing.Acta.Hort,1976,56:37-41
61. Fraga M F,Berdasco M,Diego L B,Rodriguez R,et al.Changes in polyamine concentration associated with aging in Pinus radiate and prunus persica.Tree Physiology,2004,24:1221-1226
62. Fraga M F,Canal M J,Rodriguez R.In vitro morphogenic potential of differently aged Pinus radiata trees correlates with polyamines and DNA methylation levels.Plant Cell,Tissue and Organ Culture, 2002,70:139-145
63. Fraga M F,Canal M J,Rodriguez R.Phaes-change related epigenetic and physiological changes in Pinus radiata D.Don.Planta,2002,215:672-678
64. Fraga M F,Rodriguez R,Canal M J.Genomic DNA methlation-demethylation during aging and reinvigoration of Prunusradiata.TreePhysiol,2002,22(11):813-816
65. Fraga M F,Rodriguez R,Canal M J.Phase-change related epigenetic and physiological change in pinus radiata D.Don.Planta,2002,215:672-678
66. Fraga M F,Rodriguez R,Canal M J.Rapid quantification of DNA methylation by high performance capillary electrophoresis.Electrophoresis,2000,54:2990-2994
67. Galoch E.Comparison of the content of growth regulators in juvenile and adult plants of birch.Acta Physiol.Plant.1985,7:205-215
68. Galston A W,Sawhney R K.Polyamines in plant physiology.Plant Physiol,1990,94:406-410
69. Galston A W.Polyamines as modulators of plant development Bioscience,1983,33(6): 382-388
70. Garca J L,Avidan N,Troncoso A,et al.Possible juveni-related proteins in olive tree tissues. SciHort, 2000,85:271-284
71. Greenwood M S,Hopper C A,Hutchison K W.Maturation in Larch 1.effect of age on shoot growth, foliar characteristics,and DNA methylation.Plant physiol.1989,90:406-412
72. Greenwood M S.Phase change in loblolly pine:Shoot development as a function of age.Physiol. Plant,1984,61:518-522
73. Hackett W P.Juvenility,maturation and rejuvenation in woody plants.Hort Rev,1985,7:109-155
74. Hasbun R.Valledor L,Berdasco M,Santamaria E,Canal M J,Rodriguez R,Rios D,Sanchez M.In vitro proliferation and Genome DNA methylation in adult chestnuts.Act Hort,2005,693:333-339
75. Hsu C Y,Liu Y,Luthe D S,Yuceer C.Poplar FT2 Shortens the Juvenile Phase and Promotes Seasonal Flowering.The Plant Cell,2006,18:1846
76. Huang L C,Pu S Y,Murashige T,et al.Phase-and age-related differences in protein tyrosine phosphorylation in Sequoia sempervirens.Biologia Plantarum,2003,47(4):601-603
77. Husen A,Pal M.Effect of Serial Bud Grafting and Etiolation on Rejuvenation and Rooting cuttings of Mature Tree of Tectona grandis Linn.f.Silvae Genetica,2003,52(2):84-87
78. Konigshofer H.Polyamine content in needles,shoot-axes,buds and xylem exudates of matures spruce trees(Picea abies (L)Karst).journal of Plant Physiology,1990,136:377-380
79. Lavi U,Lahav E,Degani C,et al.The genetics of the juvenile phase in avocado and its application for breeding.J Amer Soc Hort Sci,1992,117:981-984
80. Liu J H,Moriguchi T,Changes in free polyamines and gene expression during peach flower development.Biologia Plantarum,2007,51(3):530-532
81. Moncalean P,driguez A,Fernandez B.Plant growth regulator as putative physiological markers of developmental stage in Prunus persica .Plant Growth Regulation,2001,00:1-3
82. Monteuuis O,Vallauri D,Poupard C,Chauviere M.Rooting Acacia mangium cuttings of different physiological age with reference to leaf morphology as a phase change marker.Silvae Genetica, 1995,44(2-3):150-154
83. Monteuuis O.In vitro rooting of juvenile and mature Acacia mangium microcuttings with reference to leaf morphology as a phase change marker.Trees,2004,18:77-82
84. Munne-Bosch S,Lalueza P.Age-related changes in oxidative stress markers and abscisic acid levels in a drought-tolerant shrub,Cistus clusii grown under Mediterranean field conditions.Planta,2006, 225(4):1039-1049
85. Nooden L D,Guiametjj.Genetic control of senescence and ageing in plant [A].In: Schneider E L, eds.Handbook of the biology of ageing [C].Orland: Academic Press,1996, 94-118
86. Olesen P O.On cyclophysis and topophysis.Silvae Genetica,1978,27(5):173-178
87. Oliveira C M,Browning G.Gibberellin structure-activity effects on flower initiation in mature trees and on shoot growth in mature and juvenile Prunus avium.Plant Growth Regulation,1993,13:55-63
88. Peer K R,Greenwood M S.Maturation,topophysis and other factors in relation to rooting in Larix. Tree Physiology,2001,21:267-272
89. Poethig R S.Phase Change and the Regulation of Shoot Morphogenesis in Plants.scince,1990,923- 930
90. Poethig R S.Phase Change and the Regulation of Developmental Timing in Plants.science,2003, 301:334-346
91. Power A B,Dodd R S,Libby W J.Cyclophysis and Topophysis in Coast Redwood Stecklings I. Rooting and Nursery Performance.Silvae genetica,1988,37(1):8-15
92. Rey M,Tiburcio A F,Diaz-Sala C,et al.Endogenous polyamine concentration in juvenile,adult and in vitro reinvigorated hazel.Tree Physiology,1994,14:191-200
93. Rottmann W H,Meilan R,Sheppard LA,et al.Diverse effects of overexpression of LEAFY and PTLF,a poplar(Populus) homolog of LEAFY/FLORICAULA,in trans genic poplar and Arabidopsis.The Plant J,2000,22(3):235-245
94. Sanchez M C,Vieitez A M.In vitro morphogenetic competence of basal sprouts and crown branches of mature chestnut.Tree Physiology,1990,8:59-70
95. Scott D B,Jin W,Ledford H K, et al.EAF1 Regulates Vegetative-Phase Change and Flowering Time in Arabidopsis.Plant Physisology,1999,120:675-684
96. Snowball A M,Zeman A M,Tchan Y T,Mullins M G,Goodwin P B.Phase change in Citrus: Immunologically Detectable Differences between Juvenile and Mature Plants.Aust.J.Plant Physiol, 1991,18,385-396
97. Tanimoto E.Regulation of Root Growth by Plant Hormones—Roles for Auxin and Gibberllin. Critical Reviewsin Plant Sciences,2005,24(4):249-265
98. Valledor L,Hasbun R,Meijon M,et al.Involvement of DNA methylation in tree development and micropropagation.Plant Cell Tiss Organ Cell,2007,91:75-86
99. Vega S H,Sauer M,Orkwiszewski J A,Poething R S.The early phase change Gene in Meize.The Plant Cell,2002,14:133-147
100. Zimmerman R H.Juvenility and flowering in fruit trees.AcatHort,1973,34:139-142
101. Zluvova J,Janousek B,Vyskot B.Immunohistochemical study of DNA methylation dynamics during plant development. Journal of Experimental Botany,2001,52(365):2265-2273
2. 陈东攻,赵永波,俎文房,张新忠.桃实生树不同节位叶绿素、可溶性蛋白的动态变化.河北农业大学学报,2005,28(1):16-17
3. 谷瑞升,蒋湘宁.一种省时高效的木本植物蛋白双向电泳分析方法.北京林业大学学报,1999,21 (5):7-10
4. 郝再彬,苍晶,徐仲.植物生理实验.哈尔滨工业大学出版社,2004
5. 金勇丰,李载龙,陈大明.湖北海棠实生树阶段转变过程中多胺!RNA/DNA 和蛋白质含量的变化浙江农业学报,1998,10(5):264-267
6. 李新国,段安安,朱之悌.毛白杨无性系年龄效应及根萌条幼化效果的初步研究.西南林学院学报,1998,18(2):61-67
7. 林晓东.激素调节花芽分化的研究进展.果树学报,1997,14(4):269-274
8. 刘俊,吕波,徐朗莱.植物叶片中过氧化氢含量测定方法的改进.生物化学与生物物理进展,2000,27(5):548-551
9. 刘松泉,黄香等.幼态与老态无性系产量及性状比较的研究.热带作物研究,1985,2:1-4
10. 刘松泉.植物发育阶段理论在提高巴西橡胶产胶力中的应用.热带作物学报,1988,9(2):8-11
11. 龙桂友,沈德绪,林柏年.巨峰葡萄实生苗阶段发育过程中几种同工酶酶谱的变化.果树科学,1991,8(2):91-96
12. 陆定志,傅家瑞,孙松泉.植物衰老及其调控.中国农业出版社,1997
13. 马常耕.世界云杉无性系林业发展现状.世界林业研究,1993,6:24~31.
14. 沈成国等.植物衰老生理与分子生物学.中国农业出版社,2001
15. 师晨娟,刘勇,王春城,司瑞,新张林玉.青海云杉扦插的年龄效应及其生根机理研究.西北农林科技大学学报,2006,34(12):101-104
16. 施季森,何祯祥.林木无性繁殖及其在遗传改良中的地位.世界林业研究,1994,(1):25~30
17. 宋纯鹏.植物衰老生物学.北京大学出版社,1998
18. 苏培玺,张立新,杜明武,毕玉容,赵爱芬,刘新民.胡杨不同叶形光合特征水分利用效率及其对加富 CO2的响应.植物生态学报,2003,27(1):34-40
19. 孙永华,陈学森.杏杂种实生树叶片童性的研究.果树学报,2005,22(4):327-330
20. 田义,张志宏,李春敏,张新忠.木本植物阶段转变分子机制的研究进展.华北农学报,2006,21: 15-18
21. 王爱国,罗广华,邵从本.大豆种子超氧化物歧化酶的研究.植物生理学报,1983,9(1):77-84
22. 王定祥.梨实生树个体发育过程中蛋白质含量、过氧化物酶和多酚氧化酶同工酶的变化.植物生理学报,1986,12(1):40-46
23. 王建华,孙晓梅,王笑山等.母株年龄、激素种类及其浓度对日本落叶松扦插生根的影响.林业科学研究,2006,19(1):102-108
24. 王军辉,张建国,张守攻等.青海云杉硬枝扦插的激素、年龄和位置效应研究.西北农林科技大学学报,2006,34(7):65-71
25. 王明庥.论无性系林业—概念和应用.南京林业大学学报,1993,(1):1~5
26. 王秋玉,赵丽惠,王福来,刘秀琴.红皮云杉扦插繁殖中的年龄效应及其生理机制.植物研究,1997, 17(3):338-343
27. 刑世岩,胡玉芹,苗全盛,张锐.银杏无性繁殖位置效应的研究.武汉植物学研究,1998,16(1):69-76
28. 许智宏,刘春明.植物发育的分子机理.科学出版社,1998
29. 杨江山,常永义,种培芳.樱桃不同节位叶片光合特性与解剖特性比较研究.果树学报,2005,22 (4):323-326
30. 叶金山,季孔庶,王章荣.杂种马褂木无性系插条生根能力的遗传变异.南京林业大学学报,1998, 22(2):71-74
31. 余雪标.论树木阶段发育特性余无性利用问题[J].河北农业大学学报,1994,4:645-651
32. 曾骧.果树生理学.北京农业大学出版社,1992,134-139
33. 张才喜,李载龙,陈大明.湖北海棠阶段转变的生理基础.植物资源与环境学报,2001,10(1):57-59
34. 张才喜,李载龙,陈大明.湖北海棠阶段转变过程中几种同工酶变化研究.植物学通报,2001,18 (1):100-104
35. 张志礼,郑学勤.树木发育阶段理论及其在橡胶树育种中的应用.热带亚热带植物学报,2001,9 (3):262-268
36. 朱广廉,植物生理学试验,北京大学出版社,1990
37. 朱之悌,盛莹萍论树木的老化——幼年性、成年性相互关系及其利用北京林业大学学报1992,14(3):92-104
38. 朱之悌.林木的无性繁殖与无性系育种.林业科学,1986,22(3):280-290.
39. 朱之悌.毛白杨遗传改良.中国林业出版社,2006
40. Aderkas P V, Bonga J M.Influencing micropropagation and somatic embryogenesis in mature trees by manipulation of phase change, stress and culture environment.Tree Physiology,2000,20:921- 928
41. Amo-Marco J B,Vidal N,Vieitez A M.Polypeptide markers differentiating juvenile and adult tissues in chestnut. A Ballester Journal of plant physiology,1993,142(11):117-119
42. Andres H,Fernandez B,Rodriguez R,Rodriguez A.Phytohormone contents in Corylus avellana and their relationship to age and other developmental processes.Plant Cell,Tissue and Organ Culture, 2002,70:173-180
43. Ballester A,San-jose M C,et al.Anatomical and biochemical events during in vitro rooting of microcuttings from juvenile and mature phases of chestnut.Annals of Botany.1999,83:619-629
44. Baurens F C,Nicolleau J,Legavre J L,Monteuuis O.Genomic DNA methylation of juvenile and mature Acacia mangium micropropagated in vitro with reference to leaf morphology as a phase change marker.Tree Physiol,2004,24:401-407
45. Baurens F C,Nicolleau L,LegaverT,et al.Genomic DNA methylation of juvenile and mature Acacia mangium micropropagate dinvitro with reference to leaf morphology as a phase change marker. Tree Physiol,2004,24:401-407.
46. Bitonti M B,Cozza R,Chiappetta A,et al.Distinct nuclear organization,DNA methylation pattern and cytokinin distribution mark juvenile-like and adult vegetative apical meristems in peach (Prunuspersica(L.)Batsch).JExpBot,2002,53:1047-1054
47. Bitonti M B,Cozza R,Chiappetta A,Gianniono D,et al.Distrinct nuclear organization,DNA methylation pattern and cytokinin distribution mark juvenile,juvenile-like and vegetative apical meristems in peach (Prunus persica(L.) Batsch).journal of Experimental Botany,2002,53(371): 1047-1054
48. Bon M C ,Monteuuis O.Rejuvenation of a 100-year-old Sequoiadendron giganteum through in vitro meristem culture.II.Biochemical arguments,Physiologia Plantarum,1991,81,116-120
49. Bon M C,Riccardi F,Monteuuis O.Influence of phase change within a 90-year-old Sequoia sempervirens on its in vitro organogenic capacity and protein patterns.Trees,1994,8:283-287
50. Bon M C.J16:An apex protein associated with juvenility of Sequoiadendron giganteum.Tree Physiology,1988,4,381-387
51. Bradford M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein–dye binding.Analytical Biochemistry,1976,72:248-254
52. Cabanne F,et al.Hydroxycinnamic acid amides and ripening to flower of Nicotiana tabacum var. xanthi nc.PlantPhysiol,1981,53(3):445-451
53. Centeno M L,Rodriguez R,Berros B,Rodriguez A.Endogenous hormonal content and somatic embryogenic capacity of Corylus avellana L.cotyledons.Plant Cell,1997,17:139-144
54. Chin T.Y,Meyer M.M,Beevers.Abscisic acid-stimulated rooting of stem cuttings.Planta,1989,88: 192-196
55. Clemens J,Henriod R E,Bailey DG,Jameson P E.Vegetative phase change in Metrosideros: shoot and root restriction.Plant Growth Regulation,1999,28:207-214
56. Elo A,Lemmetyinen J,Turunen M L,et al.Three MADS-box genes similar to APETALA1 and FRUITFULL from silver birch (Betula pendula).Physiologia Plantarum,2001,112:95-103
57. England J R,Attiwill P M.Changes in leaf morphology and anatomy with tree age and height in the broadleaved evergreen species,Eucalyptus regnans F.Muell.Trees,2006,20:79-90
58. Ferwerda. F P.A possible explanation of the divergence between juvenile type budgrafts and their seeding mother trees in heven.Euphytica,1953,2:15-24
59. Ford YY,Taylor J M,Blake P S,Marks T R.Gibberellin A3 stimulates rooting of cuttings from cherry (Prunus avium).Plant Growth Regulation,2002,37:127-133
60. Fortanier E J,Jonkers H.Juvenility and maturity of plants as influenced by their ontogenetical and physiological ageing.Acta.Hort,1976,56:37-41
61. Fraga M F,Berdasco M,Diego L B,Rodriguez R,et al.Changes in polyamine concentration associated with aging in Pinus radiate and prunus persica.Tree Physiology,2004,24:1221-1226
62. Fraga M F,Canal M J,Rodriguez R.In vitro morphogenic potential of differently aged Pinus radiata trees correlates with polyamines and DNA methylation levels.Plant Cell,Tissue and Organ Culture, 2002,70:139-145
63. Fraga M F,Canal M J,Rodriguez R.Phaes-change related epigenetic and physiological changes in Pinus radiata D.Don.Planta,2002,215:672-678
64. Fraga M F,Rodriguez R,Canal M J.Genomic DNA methlation-demethylation during aging and reinvigoration of Prunusradiata.TreePhysiol,2002,22(11):813-816
65. Fraga M F,Rodriguez R,Canal M J.Phase-change related epigenetic and physiological change in pinus radiata D.Don.Planta,2002,215:672-678
66. Fraga M F,Rodriguez R,Canal M J.Rapid quantification of DNA methylation by high performance capillary electrophoresis.Electrophoresis,2000,54:2990-2994
67. Galoch E.Comparison of the content of growth regulators in juvenile and adult plants of birch.Acta Physiol.Plant.1985,7:205-215
68. Galston A W,Sawhney R K.Polyamines in plant physiology.Plant Physiol,1990,94:406-410
69. Galston A W.Polyamines as modulators of plant development Bioscience,1983,33(6): 382-388
70. Garca J L,Avidan N,Troncoso A,et al.Possible juveni-related proteins in olive tree tissues. SciHort, 2000,85:271-284
71. Greenwood M S,Hopper C A,Hutchison K W.Maturation in Larch 1.effect of age on shoot growth, foliar characteristics,and DNA methylation.Plant physiol.1989,90:406-412
72. Greenwood M S.Phase change in loblolly pine:Shoot development as a function of age.Physiol. Plant,1984,61:518-522
73. Hackett W P.Juvenility,maturation and rejuvenation in woody plants.Hort Rev,1985,7:109-155
74. Hasbun R.Valledor L,Berdasco M,Santamaria E,Canal M J,Rodriguez R,Rios D,Sanchez M.In vitro proliferation and Genome DNA methylation in adult chestnuts.Act Hort,2005,693:333-339
75. Hsu C Y,Liu Y,Luthe D S,Yuceer C.Poplar FT2 Shortens the Juvenile Phase and Promotes Seasonal Flowering.The Plant Cell,2006,18:1846
76. Huang L C,Pu S Y,Murashige T,et al.Phase-and age-related differences in protein tyrosine phosphorylation in Sequoia sempervirens.Biologia Plantarum,2003,47(4):601-603
77. Husen A,Pal M.Effect of Serial Bud Grafting and Etiolation on Rejuvenation and Rooting cuttings of Mature Tree of Tectona grandis Linn.f.Silvae Genetica,2003,52(2):84-87
78. Konigshofer H.Polyamine content in needles,shoot-axes,buds and xylem exudates of matures spruce trees(Picea abies (L)Karst).journal of Plant Physiology,1990,136:377-380
79. Lavi U,Lahav E,Degani C,et al.The genetics of the juvenile phase in avocado and its application for breeding.J Amer Soc Hort Sci,1992,117:981-984
80. Liu J H,Moriguchi T,Changes in free polyamines and gene expression during peach flower development.Biologia Plantarum,2007,51(3):530-532
81. Moncalean P,driguez A,Fernandez B.Plant growth regulator as putative physiological markers of developmental stage in Prunus persica .Plant Growth Regulation,2001,00:1-3
82. Monteuuis O,Vallauri D,Poupard C,Chauviere M.Rooting Acacia mangium cuttings of different physiological age with reference to leaf morphology as a phase change marker.Silvae Genetica, 1995,44(2-3):150-154
83. Monteuuis O.In vitro rooting of juvenile and mature Acacia mangium microcuttings with reference to leaf morphology as a phase change marker.Trees,2004,18:77-82
84. Munne-Bosch S,Lalueza P.Age-related changes in oxidative stress markers and abscisic acid levels in a drought-tolerant shrub,Cistus clusii grown under Mediterranean field conditions.Planta,2006, 225(4):1039-1049
85. Nooden L D,Guiametjj.Genetic control of senescence and ageing in plant [A].In: Schneider E L, eds.Handbook of the biology of ageing [C].Orland: Academic Press,1996, 94-118
86. Olesen P O.On cyclophysis and topophysis.Silvae Genetica,1978,27(5):173-178
87. Oliveira C M,Browning G.Gibberellin structure-activity effects on flower initiation in mature trees and on shoot growth in mature and juvenile Prunus avium.Plant Growth Regulation,1993,13:55-63
88. Peer K R,Greenwood M S.Maturation,topophysis and other factors in relation to rooting in Larix. Tree Physiology,2001,21:267-272
89. Poethig R S.Phase Change and the Regulation of Shoot Morphogenesis in Plants.scince,1990,923- 930
90. Poethig R S.Phase Change and the Regulation of Developmental Timing in Plants.science,2003, 301:334-346
91. Power A B,Dodd R S,Libby W J.Cyclophysis and Topophysis in Coast Redwood Stecklings I. Rooting and Nursery Performance.Silvae genetica,1988,37(1):8-15
92. Rey M,Tiburcio A F,Diaz-Sala C,et al.Endogenous polyamine concentration in juvenile,adult and in vitro reinvigorated hazel.Tree Physiology,1994,14:191-200
93. Rottmann W H,Meilan R,Sheppard LA,et al.Diverse effects of overexpression of LEAFY and PTLF,a poplar(Populus) homolog of LEAFY/FLORICAULA,in trans genic poplar and Arabidopsis.The Plant J,2000,22(3):235-245
94. Sanchez M C,Vieitez A M.In vitro morphogenetic competence of basal sprouts and crown branches of mature chestnut.Tree Physiology,1990,8:59-70
95. Scott D B,Jin W,Ledford H K, et al.EAF1 Regulates Vegetative-Phase Change and Flowering Time in Arabidopsis.Plant Physisology,1999,120:675-684
96. Snowball A M,Zeman A M,Tchan Y T,Mullins M G,Goodwin P B.Phase change in Citrus: Immunologically Detectable Differences between Juvenile and Mature Plants.Aust.J.Plant Physiol, 1991,18,385-396
97. Tanimoto E.Regulation of Root Growth by Plant Hormones—Roles for Auxin and Gibberllin. Critical Reviewsin Plant Sciences,2005,24(4):249-265
98. Valledor L,Hasbun R,Meijon M,et al.Involvement of DNA methylation in tree development and micropropagation.Plant Cell Tiss Organ Cell,2007,91:75-86
99. Vega S H,Sauer M,Orkwiszewski J A,Poething R S.The early phase change Gene in Meize.The Plant Cell,2002,14:133-147
100. Zimmerman R H.Juvenility and flowering in fruit trees.AcatHort,1973,34:139-142
101. Zluvova J,Janousek B,Vyskot B.Immunohistochemical study of DNA methylation dynamics during plant development. Journal of Experimental Botany,2001,52(365):2265-2273