白皮松种子休眠研究进展
|
摘要
种子休眠普遍存在于高等植物中,是植物历经长期的自然选择而保存下来的一种生物学特性。白皮松(Pinus bungeana Zucc.)是中国特有种,观赏价值高,以种子繁殖为主,但种子具有深休眠特性,给苗木繁育带来一定困难。分析关于白皮松种子休眠的研究结果认为,休眠原因主要包括:种子成熟时种胚的发育状况,内、外种皮和胚乳含有种类和含量不尽相同的萌发抑制物且脱落酸类在各部位均存在并有较强的抑制能力,种皮对种子萌发的机械阻力和透水性、透气性障碍以及内源激素在种子休眠中的作用。解除白皮松种子休眠的方法主要有低温层积处理、长时间流水冲洗、植物激素浸种、化学试剂和机械处理以及环境条件处理等。
Seed dormancy is pervasive in advanced plants, and is a biological characteristic that plants possess over long-term natural selection. Pinus bungeana Zucc. is an endemic species in China and has high ornamental value. It reproduces mainly through seeds; however, the seeds with deep dormancy cause difficulties in seedling breeding. In this study, the reasons for dormancy of P. bungeana seeds and the methods of dormancy breaking are summarized. The causes of dormancy mainly include the development of embryo in a mature seed; the presence of different kinds and amounts of germination inhibitors in the seed coat, paper layer, and endosperm; the presence of abscisic acid, which possesses strong inhibitory ability, in these three parts; mechanical resistance and permeability of the seed coat to germination; and endogenous hormones. The common methods of dormancy breaking are cold stratification, long-time flushing, soaking in plant hormones, chemical treatment, mechanical treatment and environmental condition treatment.
Seed dormancy is pervasive in advanced plants, and is a biological characteristic that plants possess over long-term natural selection. Pinus bungeana Zucc. is an endemic species in China and has high ornamental value. It reproduces mainly through seeds; however, the seeds with deep dormancy cause difficulties in seedling breeding. In this study, the reasons for dormancy of P. bungeana seeds and the methods of dormancy breaking are summarized. The causes of dormancy mainly include the development of embryo in a mature seed; the presence of different kinds and amounts of germination inhibitors in the seed coat, paper layer, and endosperm; the presence of abscisic acid, which possesses strong inhibitory ability, in these three parts; mechanical resistance and permeability of the seed coat to germination; and endogenous hormones. The common methods of dormancy breaking are cold stratification, long-time flushing, soaking in plant hormones, chemical treatment, mechanical treatment and environmental condition treatment.
引文
[1] 赵焱,张学忠,王孝安.白皮松天然林地理分布规律研究[J].西北植物学报,1995,15(2):161-166.ZHAO Y,ZHANG X Z,WANG X A.A study on geographical distribution law of Pinus bungeana natural forests in China[J].Acta Botanica Boreali-Occidentalia Sinica,1995,15(2):161-166.
[2] 王小平,王九龄,刘晶岚,等.白皮松分布区的气候区划[J].林业科学,1999,35(4):102-107.DOI:10.3321/j.issn:1001-7488.1999.04.016.WANG X P,WANG J L,LIU J L,et al.Climatic region alization on the distribution area of Pinus bungeana[J].Scientia Silvae Sinicae,1999,35(4):102-107.
[3] 凌宏勤.白皮松扦插繁殖技术及生理学和解剖学研究[D].北京:北京林业大学,2001.LING H Q.Study onpropagation techniques,physiology and anatomy in cutting Pinus bungeana[D].Beijing:Beijing Forestry University,2001.
[4] 郭素娟,凌宏勤,李凤兰.白皮松插穗生根的生理生化基础研究[J].北京林业大学学报,2004,26(2):43-47.DOI:10.13332/j.1000-1522.2004.02.010.GUO S J,LING H Q,LI F L.Physiological and biochemical basis of rooting of Pinus bungeana cuttings[J].Journal of Beijing Forestry University,2004,26 (2):43-47.
[5] 高治中,吝辉,白国章,等.白皮松嫩枝扦插生根试验初报[J].中国园艺文摘,2014(3):45.DOI:10.3969/j.issn.1672-0873.2014.03.019.GAO Z Z,LIN H,BAI G Z,et al.A report on rooting test of Pinus bungeiana cuttings[J].Chinese Horticulture Abstracts,2014 (3):45.
[6] 张昉.植物生长调节剂促进白皮松插穗不定根发生的机理研究[D].北京:北京林业大学,2006.ZHANG F.The studies on the promoting mechanism of plant growth regulators in the initialization of the cuttings' adventitious root of Pinus bungeana[D].Beijing:Beijing Forestry University,2006.
[7] 王霞,刘剑丛,戴小芬.白皮松扦插育苗技术[J].农业科技与信息,2016 (4):123.DOI:10.15979/j.cnki.cn62-1057/s.2016.04.077.WANG X,LIU J C,DAI X F.Technology of cutting seedlings of Pinus bungeana[J].Agricultural Science-Technology and Information,2016 (4):123.
[8] 张全民,辛俊锋.小陇山林区优良乡土树种白皮松繁育推广[J].甘肃科技,2007,23(10):251-252.DOI:10.3969/j.issn.1000-0952.2007.10.101.ZHANG Q M,XIN J F.Breeding of excellent native tree species in the forest region of Xiaolong Mountain[J].Gansu Science and Technology,2007,23 (10):251-252.
[9] 刘教明,孙永兰,王延娜.白皮松嫁接繁育技术[J].河北林业科技,2016 (5):107-108.DOI:10.16449/j.cnki.issn1002-3356.2016.05.036.LIU J M,SUN Y L,WANG Y N.Grafting technology of Pinus bungeana[J].The Journal of Hebei Forestry Science and Technology,2016 (5):107-108.
[10] 李茜,张存旭,秦萍.白皮松胚性和非胚性愈伤组织生理生化特性研究[J].西北农林科技大学学报(自然科学版),2008,36(8):151-155.DOI:10.13207/j.cnki.jnwafu.2008.08.026.LI Q,ZHANG C X,QIN P.Physiological and biochemical characteristics of embryogenic callus and non-embryogenic callus in Pinus bungeana[J].Journal of Northwest A & F University(Natural Science Edition),2008,36(8):151-155.
[11] 闫辉,张利,于淑惠,等.秋水仙素诱导白皮松多倍体的研究[J].山东农业大学学报(自然科学版),2012,43(2):184-188.DOI:10.3969/j.issn.1000-2324.2012.02.005.YAN H,ZHANG L,YU S H,et al.Studies on polyploid induction by colchicine in Pinus bungeana[J].Journal of Shandong Agricultural University(Natural Science Edition),2012,43(2):184-188.
[12] 于淑兰.解除白皮松种子休眠和促进发芽的试验研究[J].种子,1983 (1):24-28.DOI:10.16590/j.cnki.1001-4705.1983.01.025.YU S L.Seed dormancy and germination of lacebark pine (Pinus bungeana)[J].Seed,1983 (1):24-28.
[13] 董丽芬,邵崇斌,张宗勤.白皮松种胚的休眠萌发特性研究[J].林业科学,2003,39(6):47-54.DOI:10.3321/j.issn:1001-7488.2003.06.008.DONG L F,SHAO C B,ZHANG Z Q.Characteristics of embryo dormancy and germination of Pinus bungeana[J].Scientia Silvae Sinicae,2003,39(6):47-54.
[14] 董丽芬,肖斌,曹翠萍.白皮松种子休眠原因的初步探讨[J].陕西林业科技,1987 (1):15-17.DONG L F,XIAO B,CAO C P.Investigating seed dormancy in Pinus bungeana[J].Shaanxi Forest Science and Technology,1987 (1):15-17.
[15] 王小平.白皮松生物学及种子生理生态[M].北京:中国环境科学出版社,2002.
[16] 许巧玲.糖浓度、激素、低温对白皮松种子胚萌发的影响[J].山西林业科技,2007 (4):19-20.DOI:10.3969/j.issn.1007-726X.2007.04.008.XU Q L.The effect of sugar concentration,gibberellins and low temperature on embryo germination of Chinese Lace-Bark pine[J].Shanxi Forestry Science and Technology,2007 (4):19-20.
[17] 智信.6种松树种子休眠原因研究[J].西南林学院学报,2008,28(2):5-9.DOI:10.3969/j.issn.2095-1914.2008.02.002.ZHI X.Seed dormancy mechanism of six pine species[J].Journal of Southwest Forestry University,2008,28(2):5-9.
[18] 冷斌.白皮松种子休眠抑制剂的提取、分离、测定的研究[J].中国林副特产,2008 (3):34.DOI:10.13268/j.cnki.fbsic.2008.03.029.LENG B.Studies on extraction,dissociation and determination of inhibitor of Pinus bungeana seed[J].Forest by-Product and Speciality in China,2008 (3):34.
[19] 张利,闫辉,高文强,等.影响白皮松种胚休眠的因素[J].山东林业科技,2011 (5):26-28.DOI:10.3969/j.issn.1002-2724.2011.05.008.ZHANG L,YAN H,GAO W Q,et al.Factors influencing embryos dormancy of Pinus bungeana[J].Journal of Shandong Forestry Science and Technology,2011 (5):26-28.
[20] 杨卫丰,王永超,李皓.引发处理对白皮松种子发芽及生理的影响[J].种子,2013 (5):89-92.DOI:10.16590/j.cnki.1001-4705.2013.05.076.YANG W F,WANG Y C,LI H.Effects of priming on germination and physiology of Pinus bungeana seed[J].Seed,2013 (5):89-92.
[21] 陶章安.白皮松等十三个树种层积催芽的初步观察[J].林业科学,1956 (1):48-54.TAO Z A.Effects of stratification on thirteen tree species seeds including the Pinus bungeana[J].Scientia Silvae Sinicae,1956 (1):48-54.
[22] 王小平,王九龄.白皮松种子内含物的提取、分离及生物测定[J].种子,1998 (5):19-22.DOI:10.16590/j.cnki.1001-4705.1998.05.038.WANG X P,WANG J L.Extraction,separation and biological measurement of endogenous growth regulating substance of seeds of Pinus bungeana[J].Seed,1998 (5):19-22.
[23] 王小平,刘晶岚,王九龄.白皮松种子休眠特性的种源变异[J].应用生态学报,2000,11 (1):9-13.DOI:10.13287/j.1001-9332.2000.0003.WANG X P,LIU J L,WANG J L.Variation of dormancy characteristic of different Pinus bungeana seed sources [J].Chinese Journal of Applied Ecology,2000,11 (1):9-13.
[24] STANLEY R G.Krebs cycle activity of mitochondria from endosperm of sugar pine seed (Pinus lambertiana Dougl.)[J].Plant Physiology,1957,32 (5):409-412.DOI:https://doi.org/10.1104/pp.32.5.409.
[25] REN C,KERMODE A R.Analyses to determine the role of the megagametophyte and other seed tissues in dormancy maintenance of Yellow cedar (Chamaecyparis nootkatensis) seeds:morphological,cellular and physiological changes following moist chilling and during germination[J].Journal of Experimental Botany,1999,50:1403-1419.DOI:10.1093/jexbot/50.337.1403.
[26] DOWNIE B,BEWLEY J D.Dormancy in white spruce (Picea glauca (Moench.) Voss.) seeds is imposed by tissues surrounding the embryo[J].Seed Science Research,1996,6 (1):9-15.DOI:10.1017/S0960258500002944.
[27] BIANCO J,GARELLO G,LE PAGE-DEGIVRY M T.De novo ABA synthesis and expression of seed dormancy in a gymnosperm:Pseudotsuga menziesii[J].Plant Growth Regulation,1997,21 (2):115-119.DOI:10.1023/A:1005707229810.
[28] 王九龄,尹伟伦.中国松属主要树种栽培生理生态与技术[M].北京:高等教育出版社,2001.WANG J L,YIN W L.Cultivation physiology of main species of Pinus in China[M].Beijing:Higher Education Press,2001.
[29] 李正理,张新英.红松后胚离体培养的研究 Ⅱ.具雌配子体与离体后胚培养的比较观察[J].植物学报,1962,10 (3):179-186.LI Z L,ZHANG X Y.Studies on the embryo of Pinus koraiensis grown in vitro Ⅱ.comparative growth of embryos with and without megagametophyte under various culture conditions [J].Bulletin of Botany,1962,10 (3):179-186.
[30] 周健,苏友谊,代松,等.紫荆种子成熟过程中种皮和胚乳超微结构观察[J].南京林业大学学报(自然科学版),2016,40 (6):27-32.DOI:10.3969/j.issn.1000-2006.2016.06.005.ZHOU J,SU Y Y,DAI S,et al.Ultrastructure of seed coat and endosperm in the development of Cercis chinensis seed[J].Journal of Nanjing Forestry University (Natural Sciences Edition),2016,40 (6):27-32.
[31] 杨万霞,洑香香,方升佐.青钱柳种子的种皮构造及其对透水性的影响[J].南京林业大学学报(自然科学版),2005,29 (5):25-28.DOI:10.3969/j.issn.1000-2006.2005.05.006.YANG W X,FU X X,FANG S Z.The seed coat structure of Cyclocarya paliurus and its effects on water permeability[J].Journal of Nanjing Forestry University(Natural Sciences Edition),2005,29 (5):25-28.
[32] GREIPSSON S.Effects of stratification and GA3 on seed germination of a sand stabilising grass Leymus arenarius used in reclamation[J].Seed Science and Technology,2001,29 (1):1-10.
[33] BASKIN J M,BASKIN C C,LI X.Taxonomy,anatomy and evolution of physical dormancy in seeds[J].Plant Species Biology,2000,15 (2):139-152.DOI:10.1046/j.1442-1984.2000.00034.x.
[34] BARNETT J P.Delayed germination of southern pine seeds related to seed coat constraint[J].Canadian Journal of Forest Research,1976,6 (4):504-510.DOI:10.1139/x76-069.
[35] BARON F J.Moisture and temperature in relation to seed structure and germination of sugar pine (Pinus lambertiana Dougl.)[J].American Journal of Botany,1978,65 (7):804-810.DOI:10.2307/2442157.
[36] TILLMAN-SUTELA E,KAUPPI A.The morphological background to imbibition in seeds of Pinus sylvestris L.of different provenances[J].Trees,1995,9 (3):123-133.DOI:10.1007/BF02418201.
[37] TILLMAN-SUTELA E,KAUPPI A.The significance of structure for imbibition in seeds of the Norway spruce,Picea abies L.Karst[J].Trees,1995,9 (5):269-278.DOI:10.1007/BF00202017.
[38] TERSKIKH V V,FEURTADO J A,REN C,et al.Water uptake and oil distribution during imbibition of seeds of western white pine (Pinus monticola Dougl.ex D.Don) monitored in vivo using magnetic resonance imaging[J].Planta,2005,221 (1):17-27.DOI:10.1007/s00425-004-1426-z.
[39] 杨晓玲,张培玉,郭明军,等.山楂种子酚类物质含量与休眠的关系[J].园艺学报,1997,24 (4):393-394.YANG X L,ZHANG P Y,GUO M J,et al.The relationship between the content of phenolic substances and dormancy of hawthorn seeds[J].Acta Horticulturae Sinica,1997,24 (4):393-394.
[40] 黄丹,许岳香,胡海波.植物种子休眠原因与机理的研究进展[J].亚热带植物科学,2010,39 (2):78-83.DOI:10.3969/j.issn.1009-7791.2010.02.023.HUANG D,XU Y X,HU H B.Research progress on causes of seed dormancy and its mechanism[J].Subtropical Plant Science,2010,39 (2):78-83.
[41] BURROWS C J.Germination behaviour of the seeds of the New Zealand species Aristotelia serrata,Coprosma robusta,Cordyline australis,Myrtus obcordata,and Schefflera digitata[J].New Zealand Journal of Botany,1995,33 (2):257-264.DOI:10.1080/0028825X.1995.10410488.
[42] BURROWS C J.Germination behaviour of seeds of the New Zealand woody species Beilschmiedia tawa,Dysoxylum spectabile,Griselinia lucida,and Weinmannia racemosa[J].New Zealand Journal of Botany,1999,37 (1):95-105.DOI:10.1080/0028825X.1999.9512616.
[43] 李正理,张新英.红松后胚离体培养的研究 Ⅰ.不同培养方式下幼苗生长的比较观察[J].植物学报,1962,10 (2):103-112.LI Z L,ZHANG X Y.Studies on the embryo of Pinus koraiensis grown in vitro Ⅰ.growth of seedling under various conditions[J].Bulletin of Botany,1962,10 (2):103-112.
[44] 杨荣超,张海军,王倩,等.植物激素对种子休眠和萌发调控机理的研究进展[J].草地学报,2012,20 (1):1-9.DOI:10.11733/j.issn.1007-0435.2012.01.001.YANG R C,ZHANG H J,WANG Q,et al.Regulatory mechanism of plant hormone on seed dormancy and germination[J].Acta Agrectir Sinica,2012,20 (1):1-9.
[45] FINKELSTEIN R,REEVES W,ARIIZUMI T,et al.Molecular aspects of seed dormancy[J].Annual Review of Plant Biology,2008,59:387-415.DOI:10.1146/annurev.arplant.59.032607.092740.
[46] OGAWA M,HANADA A,YAMAUCHI Y,et al.Gibberellin biosynthesis and response during Arabidopsis seed germination[J].The Plant Cell,2003,15 (7):1591-1604.DOI:10.1105/tpc.011650.ble.
[47] BEAUDOIN N,SERIZET C,GOSTI F,et al.Interactions between abscisic acid and ethylene signaling cascades[J].2000,12 (7):1103-1115.DOI:10.1105/tpc.12.7.1103.
[48] STEBER C M,MCCOURT P.A role for brassinosteroids in germination in Arabidopsis[J].2001,125 (2):763-769.DOI:10.1104/pp.125.2.763.
[49] BETHKE P C,GUBLER F,JACOBSEN J V,et al.Dormancy of arabidopsis seeds and barley grains can be broken by nitric oxide[J].Planta,2004,219 (5):847-855.DOI:10.1007/s00425-004-1282-x.
[50] MEYER S E,PENDLETON R L.Genetic regulation of seed dormancy in Purshia tridentata (Rosaceae)[J].Annals of Botany,2000,85 (4):521-529.DOI:10.1006/anbo.1999.1099.
[51] LIN C H,LEE L Y,TSENG M J.The effect of stratification and thidiazuron treatment on germination and protein synthesis of Pyrus serotina Rehd cv.Niauli[J].Annals of Botany,1994 (5):515-523.DOI:10.1006/anbo.1994.1064.
[52] 关清霞,刘俊甲.白皮松种子育苗[J].林业实用技术,2003 (4):29-30.DOI:10.13456/j.cnki.lykt.2003.04.019.GUAN Q X,LIU J J.Seedlings fostering of Pinus bungeana[J].Practical Forestry Technology,2003 (4):29-30.
[53] 寇凤仙.白皮松种子高效育苗技术[J].河北林业科技,2009 (3):117.DOI:10.16449/j.cnki.issn1002-3356.2009.03.078.COU F X.Efficient seedling technology of Pinus bungeana[J].The Journal of Hebei Forestry Science and Technology,2009 (3):117.
[54] 韩永军.白皮松生物学特性及育苗技术[J].现代园艺,2015 (7):36-37.DOI:10.14051/j.cnki.xdyy.2015.07.021.HAN Y J.Biological characteristics and seedling fostering of Pinus bungeana[J].Modern Horticulture,2015 (7):36-37.
[55] 余水欢.白皮松的繁殖方法、育苗及管理技术[J].现代农业,2011 (12):77.DOI:10.14070/j.cnki.15-1098.2011.12.101.YU S H.Reproduction,seedlings and management techniques of Pinus bungeana[J].Modern Agriculture,2011 (12):77.
[56] 张瑛春.白皮松播种育苗技术研究[J].甘肃科技,2007,23 (10):249-250.DOI:10.3969/j.issn.1000-0952.2007.10.100.ZHANG Y C.Study on seedling fostering techniques of Pinus bungeana[J].Gansu Science and Technology,2007,23 (10):249-250.
[57] 高澜.白皮松圃地育苗技术[J].山西林业,2015 (1):20-21.DOI:10.3969/j.issn.1005-4707.2015.02.008.GAO L.Seedling fostering techniques of Pinus bungeana[J].Forestry of Shanxi,2015 (1):20-21.
[58] ZHIRI A,JAZIRI M,HOMES J,et al.Factors affecting the in vitro rapid germination of Taxus embryos and the evaluation of taxol content in the plantlets[J].Plant Cell,Tissue and Organ Culture,1994,39 (3):261-263.DOI:10.1007/BF00035979.
[59] BASSEL G W,ZIELINSKA E,MULLEN R T,et al.Down-regulation of DELLA genes is not essential for germination of tomato,soybean,and Arabidopsis seeds[J].Physiologia Plantarum,2004,136 (1):2782-2789.DOI:https://doi.org/10.1104/pp.103.034876.
[60] BEWLEY J D,BLACK M.Seeds:physiology of development and germination[M].New York:Plenum Press,1994.
[61] 高柱,刘腾云,王小玲,等.白木马列桕种子特性及幼苗生长规律[J].北方园艺,2016(8):76-81.DOI:10.11937/bfyy.201608022.GAO Z,LIU T Y,WANG X L,et al.Seed characters and seedlings growth rhythm of Sapium sebiferum[J].Northern Horticulture,2016(8):76-81.
[62] 杨期和,尹小娟,叶万辉.硬实种子休眠的机制和解除方法[J].植物学通报,2006,23 (1):108-118.DOI:10.3969/j.issn.1674-3466.2006.01.014.YANG Q H,YIN X J,YE W H.Dormancy mechanism and breaking methods for hard seeds[J].Chinese Bulletin of Botany,2006,23 (1):108-118.
[63] 周建荣,李仕友.油松种子休眠特性研究[J].阴山学刊(自然科学版),2009,23 (1):37-39.DOI:10.13388/j.cnki.ysajs.2009.01.015.ZHOU J R,LI S Y.Dormancy trait of Piuns tabulaeformis seed research[J].Yinshan Academic Journal(Natural Science Edition),2009,23 (1):37-39.
[64] 尚杰,王文章,赵垦田.白皮松种子萌发过程中有机质代谢的研究[J].东北林业大学学报,1992,20 (2):24-29.DOI:10.13759/j.cnki.dlxb.1992.02.004.SHANG J,WANG W Z,ZHAO K T.Organic substance metabolism during seed germination of Pinus bungeana[J].Journal of Northeast Forestry University,1992,20 (2):24-29.
[65] 李晓洁,徐化成.白皮松种子发芽习性及其种源变异的研究[J].林业科学,1989,25 (2):97-105.LI X J,XU H C.A study on seed germination characteristics of Pinus bungeana Zucc.ex Endl.from different provenances[J].Scientia Silvae Sinicae,1989,25 (2):97-105.
[66] 马建伟,刘林英,王桑,等.卫星搭载华山松、白皮松种子苗木培育试验研究初报[J].甘肃林业科技,2007,32(1):10-13.DOI:10.3969/j.issn.1006-0960.2007.01.003.MA J W,LIU L Y,WANG S,et al.Preliminary report on breeding of Pinus armandii and Pinus bungeana with seed treated in satellitein space[J].Journal of Gansu Forestry Science and Technology,2007,32(1):10-13.
[67] 李洁,梁志英.植物喷施灵对白皮松种子发芽的影响[J].山西林业科技,2010,39 (1):25-26.DOI:10.3969/j.issn.1007-726X.2010.01.009.LI J,LIANG Z Y.Effect of SPF on seed germination of Pinus bungeana[J].Shanxi Forestry and Technology,2010,39(1):25-26.
[68] 袁胜利.激光辐照白皮松种子的效应研究[J].西北农林科技大学学报(自然科学版),2003,31 (5):167-168.DOI:10.13207/j.cnki.jnwafu.2003.05.040.YUAN S L.The irradiating effect of laser on the seeds of Pinus bungeana Zucc.[J].Journal of Northwest A & F University(Natural Science Edition),2003,31(5):167-168.
[69] GHILDIYAL S K,SHARMA C M.Effect of seed size and temperature treatments on germination of various seed sources of Pinus wallichiana and Pinus roxburghii from Garhwal Malaya[J].Indian Forester,2005,131 (1):57-65.
[70] THANOS C A,SKORSILIS A.The effects of light,temperature and osmotic stress on the germination of Pinus halepensis and P.brutia seeds[J].Seed Science & Technology,1987(15):163-174.
[71] DONG L F,ZHANG X Y,MANG H G.Breaking seed dormancy of Pinus bungeana Zucc.with Trichoderma-4030 inoculations[J].New Forests,1987,1(3):245-249.DOI:10.1007/BF00118763.
[2] 王小平,王九龄,刘晶岚,等.白皮松分布区的气候区划[J].林业科学,1999,35(4):102-107.DOI:10.3321/j.issn:1001-7488.1999.04.016.WANG X P,WANG J L,LIU J L,et al.Climatic region alization on the distribution area of Pinus bungeana[J].Scientia Silvae Sinicae,1999,35(4):102-107.
[3] 凌宏勤.白皮松扦插繁殖技术及生理学和解剖学研究[D].北京:北京林业大学,2001.LING H Q.Study onpropagation techniques,physiology and anatomy in cutting Pinus bungeana[D].Beijing:Beijing Forestry University,2001.
[4] 郭素娟,凌宏勤,李凤兰.白皮松插穗生根的生理生化基础研究[J].北京林业大学学报,2004,26(2):43-47.DOI:10.13332/j.1000-1522.2004.02.010.GUO S J,LING H Q,LI F L.Physiological and biochemical basis of rooting of Pinus bungeana cuttings[J].Journal of Beijing Forestry University,2004,26 (2):43-47.
[5] 高治中,吝辉,白国章,等.白皮松嫩枝扦插生根试验初报[J].中国园艺文摘,2014(3):45.DOI:10.3969/j.issn.1672-0873.2014.03.019.GAO Z Z,LIN H,BAI G Z,et al.A report on rooting test of Pinus bungeiana cuttings[J].Chinese Horticulture Abstracts,2014 (3):45.
[6] 张昉.植物生长调节剂促进白皮松插穗不定根发生的机理研究[D].北京:北京林业大学,2006.ZHANG F.The studies on the promoting mechanism of plant growth regulators in the initialization of the cuttings' adventitious root of Pinus bungeana[D].Beijing:Beijing Forestry University,2006.
[7] 王霞,刘剑丛,戴小芬.白皮松扦插育苗技术[J].农业科技与信息,2016 (4):123.DOI:10.15979/j.cnki.cn62-1057/s.2016.04.077.WANG X,LIU J C,DAI X F.Technology of cutting seedlings of Pinus bungeana[J].Agricultural Science-Technology and Information,2016 (4):123.
[8] 张全民,辛俊锋.小陇山林区优良乡土树种白皮松繁育推广[J].甘肃科技,2007,23(10):251-252.DOI:10.3969/j.issn.1000-0952.2007.10.101.ZHANG Q M,XIN J F.Breeding of excellent native tree species in the forest region of Xiaolong Mountain[J].Gansu Science and Technology,2007,23 (10):251-252.
[9] 刘教明,孙永兰,王延娜.白皮松嫁接繁育技术[J].河北林业科技,2016 (5):107-108.DOI:10.16449/j.cnki.issn1002-3356.2016.05.036.LIU J M,SUN Y L,WANG Y N.Grafting technology of Pinus bungeana[J].The Journal of Hebei Forestry Science and Technology,2016 (5):107-108.
[10] 李茜,张存旭,秦萍.白皮松胚性和非胚性愈伤组织生理生化特性研究[J].西北农林科技大学学报(自然科学版),2008,36(8):151-155.DOI:10.13207/j.cnki.jnwafu.2008.08.026.LI Q,ZHANG C X,QIN P.Physiological and biochemical characteristics of embryogenic callus and non-embryogenic callus in Pinus bungeana[J].Journal of Northwest A & F University(Natural Science Edition),2008,36(8):151-155.
[11] 闫辉,张利,于淑惠,等.秋水仙素诱导白皮松多倍体的研究[J].山东农业大学学报(自然科学版),2012,43(2):184-188.DOI:10.3969/j.issn.1000-2324.2012.02.005.YAN H,ZHANG L,YU S H,et al.Studies on polyploid induction by colchicine in Pinus bungeana[J].Journal of Shandong Agricultural University(Natural Science Edition),2012,43(2):184-188.
[12] 于淑兰.解除白皮松种子休眠和促进发芽的试验研究[J].种子,1983 (1):24-28.DOI:10.16590/j.cnki.1001-4705.1983.01.025.YU S L.Seed dormancy and germination of lacebark pine (Pinus bungeana)[J].Seed,1983 (1):24-28.
[13] 董丽芬,邵崇斌,张宗勤.白皮松种胚的休眠萌发特性研究[J].林业科学,2003,39(6):47-54.DOI:10.3321/j.issn:1001-7488.2003.06.008.DONG L F,SHAO C B,ZHANG Z Q.Characteristics of embryo dormancy and germination of Pinus bungeana[J].Scientia Silvae Sinicae,2003,39(6):47-54.
[14] 董丽芬,肖斌,曹翠萍.白皮松种子休眠原因的初步探讨[J].陕西林业科技,1987 (1):15-17.DONG L F,XIAO B,CAO C P.Investigating seed dormancy in Pinus bungeana[J].Shaanxi Forest Science and Technology,1987 (1):15-17.
[15] 王小平.白皮松生物学及种子生理生态[M].北京:中国环境科学出版社,2002.
[16] 许巧玲.糖浓度、激素、低温对白皮松种子胚萌发的影响[J].山西林业科技,2007 (4):19-20.DOI:10.3969/j.issn.1007-726X.2007.04.008.XU Q L.The effect of sugar concentration,gibberellins and low temperature on embryo germination of Chinese Lace-Bark pine[J].Shanxi Forestry Science and Technology,2007 (4):19-20.
[17] 智信.6种松树种子休眠原因研究[J].西南林学院学报,2008,28(2):5-9.DOI:10.3969/j.issn.2095-1914.2008.02.002.ZHI X.Seed dormancy mechanism of six pine species[J].Journal of Southwest Forestry University,2008,28(2):5-9.
[18] 冷斌.白皮松种子休眠抑制剂的提取、分离、测定的研究[J].中国林副特产,2008 (3):34.DOI:10.13268/j.cnki.fbsic.2008.03.029.LENG B.Studies on extraction,dissociation and determination of inhibitor of Pinus bungeana seed[J].Forest by-Product and Speciality in China,2008 (3):34.
[19] 张利,闫辉,高文强,等.影响白皮松种胚休眠的因素[J].山东林业科技,2011 (5):26-28.DOI:10.3969/j.issn.1002-2724.2011.05.008.ZHANG L,YAN H,GAO W Q,et al.Factors influencing embryos dormancy of Pinus bungeana[J].Journal of Shandong Forestry Science and Technology,2011 (5):26-28.
[20] 杨卫丰,王永超,李皓.引发处理对白皮松种子发芽及生理的影响[J].种子,2013 (5):89-92.DOI:10.16590/j.cnki.1001-4705.2013.05.076.YANG W F,WANG Y C,LI H.Effects of priming on germination and physiology of Pinus bungeana seed[J].Seed,2013 (5):89-92.
[21] 陶章安.白皮松等十三个树种层积催芽的初步观察[J].林业科学,1956 (1):48-54.TAO Z A.Effects of stratification on thirteen tree species seeds including the Pinus bungeana[J].Scientia Silvae Sinicae,1956 (1):48-54.
[22] 王小平,王九龄.白皮松种子内含物的提取、分离及生物测定[J].种子,1998 (5):19-22.DOI:10.16590/j.cnki.1001-4705.1998.05.038.WANG X P,WANG J L.Extraction,separation and biological measurement of endogenous growth regulating substance of seeds of Pinus bungeana[J].Seed,1998 (5):19-22.
[23] 王小平,刘晶岚,王九龄.白皮松种子休眠特性的种源变异[J].应用生态学报,2000,11 (1):9-13.DOI:10.13287/j.1001-9332.2000.0003.WANG X P,LIU J L,WANG J L.Variation of dormancy characteristic of different Pinus bungeana seed sources [J].Chinese Journal of Applied Ecology,2000,11 (1):9-13.
[24] STANLEY R G.Krebs cycle activity of mitochondria from endosperm of sugar pine seed (Pinus lambertiana Dougl.)[J].Plant Physiology,1957,32 (5):409-412.DOI:https://doi.org/10.1104/pp.32.5.409.
[25] REN C,KERMODE A R.Analyses to determine the role of the megagametophyte and other seed tissues in dormancy maintenance of Yellow cedar (Chamaecyparis nootkatensis) seeds:morphological,cellular and physiological changes following moist chilling and during germination[J].Journal of Experimental Botany,1999,50:1403-1419.DOI:10.1093/jexbot/50.337.1403.
[26] DOWNIE B,BEWLEY J D.Dormancy in white spruce (Picea glauca (Moench.) Voss.) seeds is imposed by tissues surrounding the embryo[J].Seed Science Research,1996,6 (1):9-15.DOI:10.1017/S0960258500002944.
[27] BIANCO J,GARELLO G,LE PAGE-DEGIVRY M T.De novo ABA synthesis and expression of seed dormancy in a gymnosperm:Pseudotsuga menziesii[J].Plant Growth Regulation,1997,21 (2):115-119.DOI:10.1023/A:1005707229810.
[28] 王九龄,尹伟伦.中国松属主要树种栽培生理生态与技术[M].北京:高等教育出版社,2001.WANG J L,YIN W L.Cultivation physiology of main species of Pinus in China[M].Beijing:Higher Education Press,2001.
[29] 李正理,张新英.红松后胚离体培养的研究 Ⅱ.具雌配子体与离体后胚培养的比较观察[J].植物学报,1962,10 (3):179-186.LI Z L,ZHANG X Y.Studies on the embryo of Pinus koraiensis grown in vitro Ⅱ.comparative growth of embryos with and without megagametophyte under various culture conditions [J].Bulletin of Botany,1962,10 (3):179-186.
[30] 周健,苏友谊,代松,等.紫荆种子成熟过程中种皮和胚乳超微结构观察[J].南京林业大学学报(自然科学版),2016,40 (6):27-32.DOI:10.3969/j.issn.1000-2006.2016.06.005.ZHOU J,SU Y Y,DAI S,et al.Ultrastructure of seed coat and endosperm in the development of Cercis chinensis seed[J].Journal of Nanjing Forestry University (Natural Sciences Edition),2016,40 (6):27-32.
[31] 杨万霞,洑香香,方升佐.青钱柳种子的种皮构造及其对透水性的影响[J].南京林业大学学报(自然科学版),2005,29 (5):25-28.DOI:10.3969/j.issn.1000-2006.2005.05.006.YANG W X,FU X X,FANG S Z.The seed coat structure of Cyclocarya paliurus and its effects on water permeability[J].Journal of Nanjing Forestry University(Natural Sciences Edition),2005,29 (5):25-28.
[32] GREIPSSON S.Effects of stratification and GA3 on seed germination of a sand stabilising grass Leymus arenarius used in reclamation[J].Seed Science and Technology,2001,29 (1):1-10.
[33] BASKIN J M,BASKIN C C,LI X.Taxonomy,anatomy and evolution of physical dormancy in seeds[J].Plant Species Biology,2000,15 (2):139-152.DOI:10.1046/j.1442-1984.2000.00034.x.
[34] BARNETT J P.Delayed germination of southern pine seeds related to seed coat constraint[J].Canadian Journal of Forest Research,1976,6 (4):504-510.DOI:10.1139/x76-069.
[35] BARON F J.Moisture and temperature in relation to seed structure and germination of sugar pine (Pinus lambertiana Dougl.)[J].American Journal of Botany,1978,65 (7):804-810.DOI:10.2307/2442157.
[36] TILLMAN-SUTELA E,KAUPPI A.The morphological background to imbibition in seeds of Pinus sylvestris L.of different provenances[J].Trees,1995,9 (3):123-133.DOI:10.1007/BF02418201.
[37] TILLMAN-SUTELA E,KAUPPI A.The significance of structure for imbibition in seeds of the Norway spruce,Picea abies L.Karst[J].Trees,1995,9 (5):269-278.DOI:10.1007/BF00202017.
[38] TERSKIKH V V,FEURTADO J A,REN C,et al.Water uptake and oil distribution during imbibition of seeds of western white pine (Pinus monticola Dougl.ex D.Don) monitored in vivo using magnetic resonance imaging[J].Planta,2005,221 (1):17-27.DOI:10.1007/s00425-004-1426-z.
[39] 杨晓玲,张培玉,郭明军,等.山楂种子酚类物质含量与休眠的关系[J].园艺学报,1997,24 (4):393-394.YANG X L,ZHANG P Y,GUO M J,et al.The relationship between the content of phenolic substances and dormancy of hawthorn seeds[J].Acta Horticulturae Sinica,1997,24 (4):393-394.
[40] 黄丹,许岳香,胡海波.植物种子休眠原因与机理的研究进展[J].亚热带植物科学,2010,39 (2):78-83.DOI:10.3969/j.issn.1009-7791.2010.02.023.HUANG D,XU Y X,HU H B.Research progress on causes of seed dormancy and its mechanism[J].Subtropical Plant Science,2010,39 (2):78-83.
[41] BURROWS C J.Germination behaviour of the seeds of the New Zealand species Aristotelia serrata,Coprosma robusta,Cordyline australis,Myrtus obcordata,and Schefflera digitata[J].New Zealand Journal of Botany,1995,33 (2):257-264.DOI:10.1080/0028825X.1995.10410488.
[42] BURROWS C J.Germination behaviour of seeds of the New Zealand woody species Beilschmiedia tawa,Dysoxylum spectabile,Griselinia lucida,and Weinmannia racemosa[J].New Zealand Journal of Botany,1999,37 (1):95-105.DOI:10.1080/0028825X.1999.9512616.
[43] 李正理,张新英.红松后胚离体培养的研究 Ⅰ.不同培养方式下幼苗生长的比较观察[J].植物学报,1962,10 (2):103-112.LI Z L,ZHANG X Y.Studies on the embryo of Pinus koraiensis grown in vitro Ⅰ.growth of seedling under various conditions[J].Bulletin of Botany,1962,10 (2):103-112.
[44] 杨荣超,张海军,王倩,等.植物激素对种子休眠和萌发调控机理的研究进展[J].草地学报,2012,20 (1):1-9.DOI:10.11733/j.issn.1007-0435.2012.01.001.YANG R C,ZHANG H J,WANG Q,et al.Regulatory mechanism of plant hormone on seed dormancy and germination[J].Acta Agrectir Sinica,2012,20 (1):1-9.
[45] FINKELSTEIN R,REEVES W,ARIIZUMI T,et al.Molecular aspects of seed dormancy[J].Annual Review of Plant Biology,2008,59:387-415.DOI:10.1146/annurev.arplant.59.032607.092740.
[46] OGAWA M,HANADA A,YAMAUCHI Y,et al.Gibberellin biosynthesis and response during Arabidopsis seed germination[J].The Plant Cell,2003,15 (7):1591-1604.DOI:10.1105/tpc.011650.ble.
[47] BEAUDOIN N,SERIZET C,GOSTI F,et al.Interactions between abscisic acid and ethylene signaling cascades[J].2000,12 (7):1103-1115.DOI:10.1105/tpc.12.7.1103.
[48] STEBER C M,MCCOURT P.A role for brassinosteroids in germination in Arabidopsis[J].2001,125 (2):763-769.DOI:10.1104/pp.125.2.763.
[49] BETHKE P C,GUBLER F,JACOBSEN J V,et al.Dormancy of arabidopsis seeds and barley grains can be broken by nitric oxide[J].Planta,2004,219 (5):847-855.DOI:10.1007/s00425-004-1282-x.
[50] MEYER S E,PENDLETON R L.Genetic regulation of seed dormancy in Purshia tridentata (Rosaceae)[J].Annals of Botany,2000,85 (4):521-529.DOI:10.1006/anbo.1999.1099.
[51] LIN C H,LEE L Y,TSENG M J.The effect of stratification and thidiazuron treatment on germination and protein synthesis of Pyrus serotina Rehd cv.Niauli[J].Annals of Botany,1994 (5):515-523.DOI:10.1006/anbo.1994.1064.
[52] 关清霞,刘俊甲.白皮松种子育苗[J].林业实用技术,2003 (4):29-30.DOI:10.13456/j.cnki.lykt.2003.04.019.GUAN Q X,LIU J J.Seedlings fostering of Pinus bungeana[J].Practical Forestry Technology,2003 (4):29-30.
[53] 寇凤仙.白皮松种子高效育苗技术[J].河北林业科技,2009 (3):117.DOI:10.16449/j.cnki.issn1002-3356.2009.03.078.COU F X.Efficient seedling technology of Pinus bungeana[J].The Journal of Hebei Forestry Science and Technology,2009 (3):117.
[54] 韩永军.白皮松生物学特性及育苗技术[J].现代园艺,2015 (7):36-37.DOI:10.14051/j.cnki.xdyy.2015.07.021.HAN Y J.Biological characteristics and seedling fostering of Pinus bungeana[J].Modern Horticulture,2015 (7):36-37.
[55] 余水欢.白皮松的繁殖方法、育苗及管理技术[J].现代农业,2011 (12):77.DOI:10.14070/j.cnki.15-1098.2011.12.101.YU S H.Reproduction,seedlings and management techniques of Pinus bungeana[J].Modern Agriculture,2011 (12):77.
[56] 张瑛春.白皮松播种育苗技术研究[J].甘肃科技,2007,23 (10):249-250.DOI:10.3969/j.issn.1000-0952.2007.10.100.ZHANG Y C.Study on seedling fostering techniques of Pinus bungeana[J].Gansu Science and Technology,2007,23 (10):249-250.
[57] 高澜.白皮松圃地育苗技术[J].山西林业,2015 (1):20-21.DOI:10.3969/j.issn.1005-4707.2015.02.008.GAO L.Seedling fostering techniques of Pinus bungeana[J].Forestry of Shanxi,2015 (1):20-21.
[58] ZHIRI A,JAZIRI M,HOMES J,et al.Factors affecting the in vitro rapid germination of Taxus embryos and the evaluation of taxol content in the plantlets[J].Plant Cell,Tissue and Organ Culture,1994,39 (3):261-263.DOI:10.1007/BF00035979.
[59] BASSEL G W,ZIELINSKA E,MULLEN R T,et al.Down-regulation of DELLA genes is not essential for germination of tomato,soybean,and Arabidopsis seeds[J].Physiologia Plantarum,2004,136 (1):2782-2789.DOI:https://doi.org/10.1104/pp.103.034876.
[60] BEWLEY J D,BLACK M.Seeds:physiology of development and germination[M].New York:Plenum Press,1994.
[61] 高柱,刘腾云,王小玲,等.白木马列桕种子特性及幼苗生长规律[J].北方园艺,2016(8):76-81.DOI:10.11937/bfyy.201608022.GAO Z,LIU T Y,WANG X L,et al.Seed characters and seedlings growth rhythm of Sapium sebiferum[J].Northern Horticulture,2016(8):76-81.
[62] 杨期和,尹小娟,叶万辉.硬实种子休眠的机制和解除方法[J].植物学通报,2006,23 (1):108-118.DOI:10.3969/j.issn.1674-3466.2006.01.014.YANG Q H,YIN X J,YE W H.Dormancy mechanism and breaking methods for hard seeds[J].Chinese Bulletin of Botany,2006,23 (1):108-118.
[63] 周建荣,李仕友.油松种子休眠特性研究[J].阴山学刊(自然科学版),2009,23 (1):37-39.DOI:10.13388/j.cnki.ysajs.2009.01.015.ZHOU J R,LI S Y.Dormancy trait of Piuns tabulaeformis seed research[J].Yinshan Academic Journal(Natural Science Edition),2009,23 (1):37-39.
[64] 尚杰,王文章,赵垦田.白皮松种子萌发过程中有机质代谢的研究[J].东北林业大学学报,1992,20 (2):24-29.DOI:10.13759/j.cnki.dlxb.1992.02.004.SHANG J,WANG W Z,ZHAO K T.Organic substance metabolism during seed germination of Pinus bungeana[J].Journal of Northeast Forestry University,1992,20 (2):24-29.
[65] 李晓洁,徐化成.白皮松种子发芽习性及其种源变异的研究[J].林业科学,1989,25 (2):97-105.LI X J,XU H C.A study on seed germination characteristics of Pinus bungeana Zucc.ex Endl.from different provenances[J].Scientia Silvae Sinicae,1989,25 (2):97-105.
[66] 马建伟,刘林英,王桑,等.卫星搭载华山松、白皮松种子苗木培育试验研究初报[J].甘肃林业科技,2007,32(1):10-13.DOI:10.3969/j.issn.1006-0960.2007.01.003.MA J W,LIU L Y,WANG S,et al.Preliminary report on breeding of Pinus armandii and Pinus bungeana with seed treated in satellitein space[J].Journal of Gansu Forestry Science and Technology,2007,32(1):10-13.
[67] 李洁,梁志英.植物喷施灵对白皮松种子发芽的影响[J].山西林业科技,2010,39 (1):25-26.DOI:10.3969/j.issn.1007-726X.2010.01.009.LI J,LIANG Z Y.Effect of SPF on seed germination of Pinus bungeana[J].Shanxi Forestry and Technology,2010,39(1):25-26.
[68] 袁胜利.激光辐照白皮松种子的效应研究[J].西北农林科技大学学报(自然科学版),2003,31 (5):167-168.DOI:10.13207/j.cnki.jnwafu.2003.05.040.YUAN S L.The irradiating effect of laser on the seeds of Pinus bungeana Zucc.[J].Journal of Northwest A & F University(Natural Science Edition),2003,31(5):167-168.
[69] GHILDIYAL S K,SHARMA C M.Effect of seed size and temperature treatments on germination of various seed sources of Pinus wallichiana and Pinus roxburghii from Garhwal Malaya[J].Indian Forester,2005,131 (1):57-65.
[70] THANOS C A,SKORSILIS A.The effects of light,temperature and osmotic stress on the germination of Pinus halepensis and P.brutia seeds[J].Seed Science & Technology,1987(15):163-174.
[71] DONG L F,ZHANG X Y,MANG H G.Breaking seed dormancy of Pinus bungeana Zucc.with Trichoderma-4030 inoculations[J].New Forests,1987,1(3):245-249.DOI:10.1007/BF00118763.