刺槐有性生殖过程及交配系统研究
|
摘要
刺槐是一种重要的蜜源、用材、燃料和生态树种。但其人工有性杂交困难,导致刺槐杂交育种进展缓慢。为了更好的开展刺槐杂交育种工作,对刺槐有性生殖过程及交配系统进行系统深入的研究显得尤为迫切和重要。
为此,本研究综合利用人工控制授粉、荧光显微观察、及SSR父本分析对刺槐开花动态、刺槐有性生殖过程中胚珠(种子)成熟及败育规律及交配机制进行了系统深入地研究,在此基础上,给出了在目前刺槐人工杂交座果率低的情况下进行刺槐杂交遗传改良的建议。为刺槐杂交育种工作的开展提供了理论和技术支持。主要研究结果及结论如下：
1)刺槐荚果内胚珠(种子)成熟并不是随机的,而是存在与胚珠位置相关的选择性成熟和败育现象,处于荚果顶端的胚珠更容易发育为成熟种子,而处于荚果基部的胚珠则更容易发生败育。荧光显微观察结果显示,处于荚果顶端的胚珠较基部胚珠优先接受花粉管；SSR父本分析结果显示,荚果顶端区域胚珠异交率高于荚果基部胚珠。而这种胚珠位置效应最终会对植物适合度产生影响。配子体竞争而导致的合子胚获取母本资源能力的差异是造成这一现象的一个重要原因。
2)刺槐在种子成熟期、出苗期和苗期早期生长3个时期存在不同程度的自交衰退。在种子成熟期的自交衰退指数6最高,达到0.5419,在出苗期表现出的自交衰退指数δ为0.3654,苗期早期生长差别不大,在苗期早期生长到第10week和第20week时株高生长方面的自交衰退指数δ分别仅仅为0.0232和0.0873；刺槐自交子代在种子成熟期、出苗期、苗期早期生长3个时期的累积自交衰退指数δ为0.7452。SSR父本分析结果显示,在败育种子期、成熟种子期和子代苗期3个早期生长阶段中,异交率存在显著差异(P=0.042),随着刺槐授粉后早期生长时期的变化,自交子代在逐渐减少,异交率在逐渐升高(败育种子群体异交率为76.25%,成熟种子群体异交率为91.25%,子代苗群体异交率为95.83%)
3)刺槐的繁育系统属于以异交为主的混合交配系统(同花授粉可结实)。刺槐早期的选择作用掩盖了混合交配系统及其近交衰退的事实,这种选择使得后代群体中具有明显的并不显著偏离哈迪温伯格的遗传多样性丰富的异交子代群体。
4)刺槐雌雄蕊空间上的隔离、雌雄蕊异熟、相对稍慢的自交花粉管萌发及其生长率、配子体竞争、近交衰退共同促进了刺槐的异交。
5)配子体竞争和近交衰退在调控刺槐子代遗传组成中扮演着重要角色。
6)在目前刺槐去雄后人工授粉结果率低的情况下,可以尝试利用刺槐的促进异交的机制,通过不去雄利用蜜蜂或人工进行混合竞争授粉获得杂交子代,借助分子生物学父本分析手段进行子代鉴定,进而进行刺槐的杂交遗传改良。
Robinia pseudoacacia L. is a forest legume that is highly valued as a popular and important ecological and honey species, for use as lumber, fuel, bee keeping, and land reclamation. However, artificial hybridization in R. pseudoacacia was proved to be a rather difficult and delicate procedure. In order to advance the artificial breeding of R. pseudoacacia, it is urgent and vital important to study the sexual reproduction and mating system of R. pseudoacacia systematically and thoroughly.
As such, the following researches were conducted by using controlled pollination, fluorescence microscopy observation and SSR paternity analysis. Firstly, the flowering biology was studied to detailed information about the artificial pollination biology of R. pseudoacacia. Secondly, to further understand the ovule maturation patterns, the effect of ovule position on the probability of seed maturation and abortion and their influence on fitness of seedlings in open pollination were detected. Meanwhile, the selection mechanism was investigated. Thirdly, to further detect the breeding system of R. pseudoacacia, pollen donor composition during three early phases of reproduction and gene flow was revealed by SSR paternity analysis and artificial pollination. Finally, the sexual selection and possible mechanism in R. pseudoacacia were discussed. Given these understanding, suggestions on hybridization genetic improvement in R. pseudoacacia were given. It provided theoretical and technical support for us to conduct the hybridization breeding in R. pseudoacacia.
The main results and conclusions are as follows:
1) The pattern of seed production in R. pseudoacacia was non-random, but shows a selective ovule (seed) maturation and abortion pattern which depends on the ovule position. Ovules close to the distal positions tend to be maturated more often than those close to basal positions during the ovule development. In contrast, ovules in basal positions were more likely to be aborted in ovule development than ovules in distal positions, although the differences existed between years and populations. Fluorescence microscopy observation results suggested that the ovules that lie close to the distal positions were more likely to be fertilized than those in the most basal positions. SSR paternity analysis results showed that the outcrossing rates were higher in distal position than that in basal position. The above results suggested that seed maturation and abortion might be the results of the gametophyte competition which shows the differences in the ability of developing seeds to gather maternal resources. Hence, ovule position exerts an influence on offspring fitness.
2) We found that progenies resulting from cross-pollination treatments showed significant high fitness compared with progenies resulting from self-pollination, resulting in high levels of selfing depression. Selfing depression was not uniformly manifested, however, over the three stages. Selfing depression was greatest between fertilization and seed maturation stage (δ=0.5419), seedling emergence (0.3654) stage take second place. No significant differences in seedling growth were observed between selfed and corssed progenies. The selfing depression (δ) in10week and20week height growth merely0.0232and0.0873, respectively. Cumulative selfing depression (δ) across all three stages averaged0.7452. SSR paternity analysis results showed that there was an overall decrease in self-fertilized progeny across each life stage (P=0.042). Highest outcrossing rate was observed in seedling stage (95.83%), followed by the mature seed stage (91.25%), finially the abortion stage (76.25%).
3) The breeding system of R. pseudoacacia is self-compatible, and may relay on a mixed-mating system to ensure reproductive success, however, the early viability selection conceals the truth of mixed-mating system and inbreeding depression, resulting in an apparently outcrossed population that does not differ significantly from Hardy-Weinberg expectations.
4) The physical separation of stigmatal and antheral surfaces, protogynous flowering habit, the relatively slow germination and pollen tube growth, gametophyte competition as well as the inbreeding depression encourages outcrossing in R. pseudoacacia.
5) The results suggest that microgametophytic competition and inbreeding depression might contribute to the mechanisms that ultimately determine the production of offspring in R. pseudoacacia.
6) Considering the low fruit set via manual emasculation pollination and features of encouraging outcrossing in R. pseudoacacia, it may be a possible way to obtain hybridprogeny through mixed competition pollination without emasculation, then determined the seedlings by using paternity analysis.
为此,本研究综合利用人工控制授粉、荧光显微观察、及SSR父本分析对刺槐开花动态、刺槐有性生殖过程中胚珠(种子)成熟及败育规律及交配机制进行了系统深入地研究,在此基础上,给出了在目前刺槐人工杂交座果率低的情况下进行刺槐杂交遗传改良的建议。为刺槐杂交育种工作的开展提供了理论和技术支持。主要研究结果及结论如下：
1)刺槐荚果内胚珠(种子)成熟并不是随机的,而是存在与胚珠位置相关的选择性成熟和败育现象,处于荚果顶端的胚珠更容易发育为成熟种子,而处于荚果基部的胚珠则更容易发生败育。荧光显微观察结果显示,处于荚果顶端的胚珠较基部胚珠优先接受花粉管；SSR父本分析结果显示,荚果顶端区域胚珠异交率高于荚果基部胚珠。而这种胚珠位置效应最终会对植物适合度产生影响。配子体竞争而导致的合子胚获取母本资源能力的差异是造成这一现象的一个重要原因。
2)刺槐在种子成熟期、出苗期和苗期早期生长3个时期存在不同程度的自交衰退。在种子成熟期的自交衰退指数6最高,达到0.5419,在出苗期表现出的自交衰退指数δ为0.3654,苗期早期生长差别不大,在苗期早期生长到第10week和第20week时株高生长方面的自交衰退指数δ分别仅仅为0.0232和0.0873；刺槐自交子代在种子成熟期、出苗期、苗期早期生长3个时期的累积自交衰退指数δ为0.7452。SSR父本分析结果显示,在败育种子期、成熟种子期和子代苗期3个早期生长阶段中,异交率存在显著差异(P=0.042),随着刺槐授粉后早期生长时期的变化,自交子代在逐渐减少,异交率在逐渐升高(败育种子群体异交率为76.25%,成熟种子群体异交率为91.25%,子代苗群体异交率为95.83%)
3)刺槐的繁育系统属于以异交为主的混合交配系统(同花授粉可结实)。刺槐早期的选择作用掩盖了混合交配系统及其近交衰退的事实,这种选择使得后代群体中具有明显的并不显著偏离哈迪温伯格的遗传多样性丰富的异交子代群体。
4)刺槐雌雄蕊空间上的隔离、雌雄蕊异熟、相对稍慢的自交花粉管萌发及其生长率、配子体竞争、近交衰退共同促进了刺槐的异交。
5)配子体竞争和近交衰退在调控刺槐子代遗传组成中扮演着重要角色。
6)在目前刺槐去雄后人工授粉结果率低的情况下,可以尝试利用刺槐的促进异交的机制,通过不去雄利用蜜蜂或人工进行混合竞争授粉获得杂交子代,借助分子生物学父本分析手段进行子代鉴定,进而进行刺槐的杂交遗传改良。
Robinia pseudoacacia L. is a forest legume that is highly valued as a popular and important ecological and honey species, for use as lumber, fuel, bee keeping, and land reclamation. However, artificial hybridization in R. pseudoacacia was proved to be a rather difficult and delicate procedure. In order to advance the artificial breeding of R. pseudoacacia, it is urgent and vital important to study the sexual reproduction and mating system of R. pseudoacacia systematically and thoroughly.
As such, the following researches were conducted by using controlled pollination, fluorescence microscopy observation and SSR paternity analysis. Firstly, the flowering biology was studied to detailed information about the artificial pollination biology of R. pseudoacacia. Secondly, to further understand the ovule maturation patterns, the effect of ovule position on the probability of seed maturation and abortion and their influence on fitness of seedlings in open pollination were detected. Meanwhile, the selection mechanism was investigated. Thirdly, to further detect the breeding system of R. pseudoacacia, pollen donor composition during three early phases of reproduction and gene flow was revealed by SSR paternity analysis and artificial pollination. Finally, the sexual selection and possible mechanism in R. pseudoacacia were discussed. Given these understanding, suggestions on hybridization genetic improvement in R. pseudoacacia were given. It provided theoretical and technical support for us to conduct the hybridization breeding in R. pseudoacacia.
The main results and conclusions are as follows:
1) The pattern of seed production in R. pseudoacacia was non-random, but shows a selective ovule (seed) maturation and abortion pattern which depends on the ovule position. Ovules close to the distal positions tend to be maturated more often than those close to basal positions during the ovule development. In contrast, ovules in basal positions were more likely to be aborted in ovule development than ovules in distal positions, although the differences existed between years and populations. Fluorescence microscopy observation results suggested that the ovules that lie close to the distal positions were more likely to be fertilized than those in the most basal positions. SSR paternity analysis results showed that the outcrossing rates were higher in distal position than that in basal position. The above results suggested that seed maturation and abortion might be the results of the gametophyte competition which shows the differences in the ability of developing seeds to gather maternal resources. Hence, ovule position exerts an influence on offspring fitness.
2) We found that progenies resulting from cross-pollination treatments showed significant high fitness compared with progenies resulting from self-pollination, resulting in high levels of selfing depression. Selfing depression was not uniformly manifested, however, over the three stages. Selfing depression was greatest between fertilization and seed maturation stage (δ=0.5419), seedling emergence (0.3654) stage take second place. No significant differences in seedling growth were observed between selfed and corssed progenies. The selfing depression (δ) in10week and20week height growth merely0.0232and0.0873, respectively. Cumulative selfing depression (δ) across all three stages averaged0.7452. SSR paternity analysis results showed that there was an overall decrease in self-fertilized progeny across each life stage (P=0.042). Highest outcrossing rate was observed in seedling stage (95.83%), followed by the mature seed stage (91.25%), finially the abortion stage (76.25%).
3) The breeding system of R. pseudoacacia is self-compatible, and may relay on a mixed-mating system to ensure reproductive success, however, the early viability selection conceals the truth of mixed-mating system and inbreeding depression, resulting in an apparently outcrossed population that does not differ significantly from Hardy-Weinberg expectations.
4) The physical separation of stigmatal and antheral surfaces, protogynous flowering habit, the relatively slow germination and pollen tube growth, gametophyte competition as well as the inbreeding depression encourages outcrossing in R. pseudoacacia.
5) The results suggest that microgametophytic competition and inbreeding depression might contribute to the mechanisms that ultimately determine the production of offspring in R. pseudoacacia.
6) Considering the low fruit set via manual emasculation pollination and features of encouraging outcrossing in R. pseudoacacia, it may be a possible way to obtain hybridprogeny through mixed competition pollination without emasculation, then determined the seedlings by using paternity analysis.
引文
戴丽,孙鹏,蒋晋豫,等.刺槐、红花槐、四倍体刺槐花粉体外萌发对比[J].东北林业大学学报,2012,40:1-5.
河南省第二期刺槐良种选育协作组.刺槐速生优质工业用材新无性系选育[J].河南林业科技,1997,17：13-16.
何田华,葛颂.植物种群交配系统、亲本分析以及基因流动研究[J].植物生态学报,2001,25(2):144-154.
解荷锋,于中奎,陈一山,等.种子园刺槐开花结实和控制授粉的初步研究[J].山东林业科技,1994,(4)：4-7.
李继华.刺槐在山东的引种和发展[J].山东林业科技,1983,4:73-75.
刘颖,王国义.刺槐优良无性系的选择[J].宁夏农林科技,1996,(1):19-22.
孙鹏,戴丽,胡瑞阳,等.刺槐开花传粉及交配方式[J].东北林业大学学报,2012,(40):6-11,24.
田砚亭,李云.刺槐新品种介绍[J].中国水土保持科学、2003,1(1):7.
田志和,董健,李继刚.刺槐开花生物学特性的观察初报[J].辽宁林业科技,1981,(02):18-20.
田志和,董健,李继刚.刺槐无性系测定研究报告[J].辽宁林业科技,1982,(1):3-5.
田志和,董健,曲艺.高蜜源刺槐无性系的研究[J].辽宁林业科技,1991,(5):3-6.
王念,王文君,赵蓬晖.温度和时间对储藏刺槐花粉生活力影响的研究[J].河南林业科技,2007,(04)：7-9.
吴全宇,张瑞军,郑宝昌,等.荷刺2号等4个刺槐优良无性系选择研究[J].山东林业科技,2002,(06)：1-6.
吴全宇,郑宝昌,张瑞军,等.刺槐优良无性系荷刺1号选择的研究[J].山东林业科技,1999,(01)：7-10.
荀守华,乔玉玲,张江涛,等.我国刺槐遗传育种现状及发展对策[J].山东林业科技,2009,180(1)：92-96.
原法宪.刺槐开花生物学特性观察[J].山西林业科技,1978,(03):1-10.
张敦伦,张振芬,王方泉.刺槐无性系选种的研究[J].山东林业科技,1990,(2):16-20.
张国君,李云,姜金仲,等.饲料性四倍体刺槐叶粉饲用价值的比较研究[J].草业科学,2007,24(1)：26-31.
周琴宝.刺槐花的形态与解剖结构相关性的观察[J].江苏林业科技,1982,(1):34-39.
朱延林.刺槐优良无性系选育研究[J].河南林业科技,1989,9(1):25-30.
Aizen MK, Searcy KB, Mulcahy DL. Among-and withinflower comparisons of pollen tube growth following self-and cross-pollinations in Dianthus chinensis (Caryophyltaceae) [J]. American Journal of Botany,1990,77:671-676.
Barrett RP, Mebrahtu T, Hanover JW. Black locust:a multi-purpose tree species for temperate climates Advances in new crops. Janick J, Simon JE, Portland, OR, Timber Press:1990, pp278-283.
Barrett S. Mating strategies in flowering plants:the outcrossing-selfing paradigm and beyond [J]. Philosophical Transactions of the Royal Society of London B,2003,358:991-1004.
Bawa K and Webb C. Flower, fruit, and seed abortion in tropical forest trees:implications for the evolution of paternal and maternal reproductive patterns [J]. American Journal of Botany, 1984, (71):736-751.
Bellusci FG, Pellegrino G, Musacchio A. Different levels of inbreeding depression between outcrossing and selfing Serapias species [J]. Biologia Plantarum,2009,53:175-178.
Bongarten BC. Huber DA, Apsley DK. Environmental and genetic influences on short-rotation biomass production of black locust (Robinia pseudoacacia L.) in the Georgia Piedmont [J]. Forest Ecology and Management,1992,55:315-331.
Brown SK, Iezzoni AF, Fogle HW. et al. (Eds.). Fruit Breeding, vol. I:Tree and Tropical Fruits. John Wiley & Sons, Inc.,NY, pp.1996:213-255.
Bush R and Smouse P. Evidence for the adaptive significance of allozymes in forest trees [J]. New Forests,1992,6:179-196.
Charlesworth D and Charlesworth B. Inbreeding depression and its evolutionary consequences [J]. Annual Review of Ecology and Systematics,1987,18:237-268.
Charnov E. Simultaneous hermaphroditism and sexual selection [J]. Proceedings of the National Academy of Sciences of the United States of America,1979,76:2480-2484.
Dafni A and Firmage D. Pollen viability and longevity:Practical, ecological and evolutionary implications [J]. Plant Systematics and Evolution,2000,222:113-132.
Davis LE, Stephenson AG, Winsor JA. Pollen competition improves performance and reproductive output of the common zucchini squash under field conditions [J]. Journal of the American Society for Horticultural Science,1987,112:712-716.
Diaz A and Macnair M. Pollen tube competition as a mechanism of prezygotic reproductive isolation between Mimulus nasutus and its presumed progenitor M. guttatus [J]. New Phytologist, 1999,144:471-478.
Dini-Papanastasi O and Aravanopoulos FA. Artificial hybridization between Robinia pseudoacacia L. and R pscudoacacia var. monophylla Carr [J]. Forestry,2008,81:91-101.
Franklin T and Franklin F. The different mechanisms of gametophytic self-incompatibility [J]. Philosophical Transactions of the Royal Society of London B,2003,358:1025-1032.
Glaettli M, Pescatore L, Goudet J. Proximity-dependent pollen performance in Silene vulgaris [J]. Annals of Botany,2006,98:431-437.
Goodwillie C, Kalisz S, Eckert G. The evolutionary enigma of mixed mating systems in plants: occurrence, theoretical explanations, and empirical evidence [J]. Annual Review of Ecology, Evolution and Systematics,2005,36:47-79.
Guerra ME, WUnsch A, Margarita LC, Rodrigo J. Flower Emasculation as the Cause for Lack of Fruit Set in Japanese Plum Crosses [J]. Journal of the American Society for Horticultural Science,2010,135:556-562.
Haileslassie T, Mollel M, Skogsmyr I. Effects of nutrient level on maternal choice and siring success in Cucumis sativus (Cucurbitaceae) [J]. Evolution Ecology,2005,19:275-288.
Harder LD and Barrett SCH. Mating cost of large floral displays in hermaphrodite plants [J]. Nature. 1995,373:512-515.
Harlow WM. Harrar ES, White FM. Text book of dendrology. New York. McGraw Hill,1979.
Harmon-Threatt AN, Burns JH, Shemyakina LA, Knight TM. Breeding system and pollination ecology of introduced plants compared to their native relatives [J]. American Journal of Botany.2009,96:1544-1550.
Hedhly A, Hormaza JI, Herrero, M. Flower emasculation accelerates ovule degeneration and reduces fruit set in sweet cherry [J]. Science Hortic-Amsterdam,2009,119: 455-457.
Hedrick P and Kalinowski S. Inbreeding depression in conservation biology [J]. Annual Review of Ecology Evolution and Systematics,2000,31:139-162.
Hufford KM and Hamrick JL. Viability selection at three early life stages of the tropical tree, Platypodium elegans (Fabaceae, Papilionoideae) [J]. Evolution 2003,57:518-526.
Husband BC and Schemske DW. Magnitude and timing of inbreeding depression in a diploid population of Epilobium angustifolium (Onagraceae) [J]. Heredity,1995,75:206-215.
Husband BC and Schemske DW. Evolution of the magnitude and timing of inbreeding depression in plants [J]. Evolution; international journal of organic evolution,1996,50:54-70.
Ishida KH, Yoshimaru H, Ito H. Effects of geitonogamy on the seed set of Magnolia obovata Thumb. (Magnoliaceae) [J]. International Journal of Plant Sciences,2003,164:729-735.
Jarne P and Charlesworth D. The evolution of the selfing rate in functionally hermaphrodite plants and animals [J]. Annual Review of Ecology and Systematics,1993,24:441-466.
Johnston MO and Schoen DJ. On the measurement of inbreeding depression [J]. Evolution,1994,48: 1735-1741.
Keresztesi B. Breeding and cultivation of black locust, Robinia pseudoacacia, in Hungary [J]. Forest Ecology and Management,1983,6:217-244.
Kittelson PM and Maron JL. Outcrossing rate and inbreeding depression in the perennial yellow bush lupine, Lupinus arboreus (Fabaceae) [J]. American Journal of Botany,2000,87(5): 652-660.
Kittelson PM and Maron JL. Outcrossing rate and inbreeding depression in the perennial yellow bush lupine. Lupinus arboreus (Fabaceae) [J]. American Journal of Botany,2000,87: 652-660.
Korbecka G, Klinkhamer PGL, Vrieling K. Selective embryo abortion hypothesis revisited-a molecular approach [J]. Plant biology,2002,4:298-310.
Kruszewski L and Galloway L. Explaining outcrossing rate in Campanulastrum americanum (Campanulaceae):geitonogamy and cryptic self-incompatibility [J]. International Journal of Plant Science,2006,167:455-461.
Lande R, Schemske DW. The evolution of self-fertilization and inbreeding depression in plants. I: Genetic models [J]. Evolution,1985,39:24-40.
Lee CB, Page LE, McClure BA, Holtsford. Post-pollination hybridization barriers in Nicotiana section Alatae [J]. Sexual Plant Reproduction,2008,21:183-195.
Lee T. Patterns of fruit and seed production. In:Lovett Doust J, Lovett Doust L, eds. Plant reproductive ecology:patterns and strategies. New York:Oxford University Press:1988, 179-202.
Lee T and Bazzaz F. Regulation of fruit and seed production in an annual legume, Cassia fasciculata [J]. Ecology,1982a,63:1363-1373.
Lee T and Bazzaz F. Regulation of fruit maturation pattern in an annual legume, Cassia fasciculata [J]. Ecology,1982b,63:1374-1388.
Lian C and Hogetsu T. Development of microsatellite markers in black locust (Robinia pseudoacacia) using a dual-suppression-PCR technique [J]. Molecular Ecology Notes.2002,2:211-213.
Lian C, Oishi R, Miyashita, Hogetsu T. High somatic instability of a microsatellite locus in a clonal tree, Robinia pseudoacacia [J]. Theoretical and Applied Genetics,2004,108:836-841.
Lloyd D. Self-and cross-fertilization in plants.Ⅱ. The selection of self-fertilization [J]. International Journal of Plant Science,1992,153:370-380.
Lyons EE, Waser NM, Price MV, Antonovics J, Motten AF. Sources of variation in plant reproductive success and implications for concepts of sexual selection [J]. American Naturalist,1989,134:409-433.
Marshall D and Diggle P. Mechanisms of differential pollen donor performance in wild radish, Raphanus sativus (Brassicaceae) [J]. American Journal of Botany.2001,88:242-257.
Marshall D and Folsom M. Mate choice in plants:an anatomical to population perspective [J]. Annual Review of Ecology and Systematics,1991,22:37-63.
Marshall D, Shaner MGM, Oliva JP. Effects of pollen load size on seed paternity in wild radish:the roles of pollen competition and mate choice [J]. Evolution,2007,61:1925-1937.
Melser C and Klinkhamer P. Selective seed abortion increases offspring survival in Cynoglossum officinale (Boraginaceae) [J]. American Journal of Botany,2001,88:1033-1040.
Menaali J and Rocha O. Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata [J]. Annals of Botany,2005,95:449-455.
Mishima K, Hirao T, Urano S, Watanabe. Isolation and characterization of microsatellite markers from Robinia pseudoacacia L [J]. Theoretical and Applied Genetics,2008,108:836-841.
Mulcahy D. The rise of the angiosperms:a genecological factor [J]. Science,1974,206:20-23.
Mulcahy D. Rise of the angiosperms:a genecological factor [J]. Science,1979,206:20-23.
Mulugeta D. Maxwell BD, Fay PK, Dyer WE. Kochia (Kochia scoparia) pollen dispersion, viability and germination [J]. Weed Science,1994,42:548-552.
Murawski D and Hamrick J. The effect of the density of flowering individuals on the mating systems of nine tropical tree species [J]. Heredity,1991,67:167-174.
Murawski DA, Hamrick JL, Hubbell SP, Foster RB. Mating systems of two Bombacaceous trees of a neotropical moist forest [J]. Oecologia,1990,82:501-506.
Nettancourt Dde. Incompatibility in angiosperms [J]. Sexual Plant Reproduction,1997,10:185-199.
Obaysshi K, Tsumura Y, Iharaujino T, Niiyama K, Tanouchi H, Suyama Y, Washitani I, Lee CT, Lee SL, Norwati M. Genetic diversity and outcrossing rate between undisturbed and selectively logged forests of Shorea curtisii (Dipterocarpaceae)using microsattellite DNA analysis [J]. International Journal of Plant Science,2002,163:151-158.
Okie WR, Weinberger JH. Plums. In:Janick, J, Moore, JN (Eds) Fruit Breeding, vol. I: Tree and Tropical Fruits. John Wiley & Sons. Inc., NY,1996, pp 213-255.
Queller D. Sexual selection in a hermaphroditic plant [J]. Nature,1983,305:706-707.
Real D. Rizza MD, Quesenberry, Echenique M. Reproductive and molecular evidence for allogamy in Lotononis bainesii Baker [J]. Crop Science,2004,44:394-400.
Redei K. Black Locust (Robinia pseudoacacia L.) Growing in Hungary. Budapest, Publications of the Hungarian Forest Research Institute,1998.
Richards AJ. Plant breeding systems. London, UK, George Allen & Unwin,1986.
Rocha O and Stephenson A. Effects of ovule position on seed production, seed weight, and progeny performance in Phaseolus coccineus L [J]. American Journal of Botany,1990,77: 1320-1329.
Rocha O and Stephenson A. Effects of nonrandom seed abortion on progeny performance in Phaseolus coccineus L [J]. Evolution,1991,45:1198-1208.
Rocha O and Stephenson A. Order of fertilization within the ovary in Phaseolus coccineus L. (Leguminosae) [J]. Sexual Plant Reproduction,1991.9:273-278.
Schemske DW and Lande R. The evolution of self-fertilization and inbreeding depression in plants. Ⅱ. Empirical observations [J]. Evolution,1985,39:41-52.
Seavey S and Bawa K. Late-acting self-incompatibility in angiosperms [J]. The Botanical Review, 1986,52:195-219.
Skogsmyr I and Lankinen A. Sexual selection:an evolutionary force in plants? [J]. Biological Reviews,2002,77:537-562.
Snow A. Postpollination selection and male fitness in plants [J]. American Naturalist,1994,144: 69-83.
Souto CP, Aizen MA, Premoli AC. Effects of crossing distance and genetic relatedness on pollen performance in Alstroemeria aurea (Alstroemeriaceae) [J]. American Journal of Botany, 2002,89:427-432.
Stephenson AG. Flower and fruit abortion:proximate causes and ultimate functions [J]. Annual Review of Ecology and Systematics,1981,12:253-279.
Surles S, Hamrick JL, Bongarten, BC. Mating systems in open-pollinated families of black locust (Robinia pseudoacacia L.) [J]. Silvae Genet,1990,39:35-40.
Susko DJ. Effect of ovule position on patterns of seed maturation and abortion in Robinia pseudoacacia (Fabaceae) [J]. Canadian Journal of Botany,2006,84:1259-1265.
Torres C, Eynard MC, Aizen MA, Galetto L. Selective fruit maturation and seedling performance in Acacia caven (Fabaceae) [J]. International Journal of Plant Sciences,2002,163:809-813.
Travers S and Mazer S. The absence of cryptic self-incompatibility in Clarkia unguiculata (Onagraceae) [J]. American Journal of Botany,2000,87:191-196.
Waser N and Price M. Crossing distance effects on prezygotic performance in plants:an argument for female choice [J]. Oikos,1993,68:303-308.
Weir B. Genetic data analysis Ⅱ. Sinauer Associates. Sunderland, MA,1996.
Wiens D, Calvin CL, Wilson CA, Davern CI, Frank D, Seavy SR. Reproductive success, spontaneous embryo abortion, and genetic load in flowering plants [J]. Oecologia,1987,71: 501-509.
Wilcock C and Neiland R. Pollination failure in plants:why it happens and when it matters [J]. Trends in Plant Science,2002,7:270-277.
Williams E and Rouse J. Relationships of pollen size, pistil length and pollen tube growth rates in Rhododendron and their influence on hybridization [J]. Sexual Plant Reproduction,1990.3: 7-17.
Willson M. Sexual selection in plants and animals [J]. Trends in Ecology Evolution,1979,5: 210-214.
Willson M. Sexual selection and dicliny in angiosperms [J]. American Naturalist,1982,119: 579-583.
Winsor JA, Peretz S, Stephenson AG. Pollen competition in a natural population of Cucurbita foetidissima (Cucurbitaceae) [J]. American Journal of Botany,2000,87:527-532.
Wright S. Systems of mating. Ⅱ. The effects of inbreeding on the genetic composition of a population [J]. Genetics,1921,6:124-143.
河南省第二期刺槐良种选育协作组.刺槐速生优质工业用材新无性系选育[J].河南林业科技,1997,17：13-16.
何田华,葛颂.植物种群交配系统、亲本分析以及基因流动研究[J].植物生态学报,2001,25(2):144-154.
解荷锋,于中奎,陈一山,等.种子园刺槐开花结实和控制授粉的初步研究[J].山东林业科技,1994,(4)：4-7.
李继华.刺槐在山东的引种和发展[J].山东林业科技,1983,4:73-75.
刘颖,王国义.刺槐优良无性系的选择[J].宁夏农林科技,1996,(1):19-22.
孙鹏,戴丽,胡瑞阳,等.刺槐开花传粉及交配方式[J].东北林业大学学报,2012,(40):6-11,24.
田砚亭,李云.刺槐新品种介绍[J].中国水土保持科学、2003,1(1):7.
田志和,董健,李继刚.刺槐开花生物学特性的观察初报[J].辽宁林业科技,1981,(02):18-20.
田志和,董健,李继刚.刺槐无性系测定研究报告[J].辽宁林业科技,1982,(1):3-5.
田志和,董健,曲艺.高蜜源刺槐无性系的研究[J].辽宁林业科技,1991,(5):3-6.
王念,王文君,赵蓬晖.温度和时间对储藏刺槐花粉生活力影响的研究[J].河南林业科技,2007,(04)：7-9.
吴全宇,张瑞军,郑宝昌,等.荷刺2号等4个刺槐优良无性系选择研究[J].山东林业科技,2002,(06)：1-6.
吴全宇,郑宝昌,张瑞军,等.刺槐优良无性系荷刺1号选择的研究[J].山东林业科技,1999,(01)：7-10.
荀守华,乔玉玲,张江涛,等.我国刺槐遗传育种现状及发展对策[J].山东林业科技,2009,180(1)：92-96.
原法宪.刺槐开花生物学特性观察[J].山西林业科技,1978,(03):1-10.
张敦伦,张振芬,王方泉.刺槐无性系选种的研究[J].山东林业科技,1990,(2):16-20.
张国君,李云,姜金仲,等.饲料性四倍体刺槐叶粉饲用价值的比较研究[J].草业科学,2007,24(1)：26-31.
周琴宝.刺槐花的形态与解剖结构相关性的观察[J].江苏林业科技,1982,(1):34-39.
朱延林.刺槐优良无性系选育研究[J].河南林业科技,1989,9(1):25-30.
Aizen MK, Searcy KB, Mulcahy DL. Among-and withinflower comparisons of pollen tube growth following self-and cross-pollinations in Dianthus chinensis (Caryophyltaceae) [J]. American Journal of Botany,1990,77:671-676.
Barrett RP, Mebrahtu T, Hanover JW. Black locust:a multi-purpose tree species for temperate climates Advances in new crops. Janick J, Simon JE, Portland, OR, Timber Press:1990, pp278-283.
Barrett S. Mating strategies in flowering plants:the outcrossing-selfing paradigm and beyond [J]. Philosophical Transactions of the Royal Society of London B,2003,358:991-1004.
Bawa K and Webb C. Flower, fruit, and seed abortion in tropical forest trees:implications for the evolution of paternal and maternal reproductive patterns [J]. American Journal of Botany, 1984, (71):736-751.
Bellusci FG, Pellegrino G, Musacchio A. Different levels of inbreeding depression between outcrossing and selfing Serapias species [J]. Biologia Plantarum,2009,53:175-178.
Bongarten BC. Huber DA, Apsley DK. Environmental and genetic influences on short-rotation biomass production of black locust (Robinia pseudoacacia L.) in the Georgia Piedmont [J]. Forest Ecology and Management,1992,55:315-331.
Brown SK, Iezzoni AF, Fogle HW. et al. (Eds.). Fruit Breeding, vol. I:Tree and Tropical Fruits. John Wiley & Sons, Inc.,NY, pp.1996:213-255.
Bush R and Smouse P. Evidence for the adaptive significance of allozymes in forest trees [J]. New Forests,1992,6:179-196.
Charlesworth D and Charlesworth B. Inbreeding depression and its evolutionary consequences [J]. Annual Review of Ecology and Systematics,1987,18:237-268.
Charnov E. Simultaneous hermaphroditism and sexual selection [J]. Proceedings of the National Academy of Sciences of the United States of America,1979,76:2480-2484.
Dafni A and Firmage D. Pollen viability and longevity:Practical, ecological and evolutionary implications [J]. Plant Systematics and Evolution,2000,222:113-132.
Davis LE, Stephenson AG, Winsor JA. Pollen competition improves performance and reproductive output of the common zucchini squash under field conditions [J]. Journal of the American Society for Horticultural Science,1987,112:712-716.
Diaz A and Macnair M. Pollen tube competition as a mechanism of prezygotic reproductive isolation between Mimulus nasutus and its presumed progenitor M. guttatus [J]. New Phytologist, 1999,144:471-478.
Dini-Papanastasi O and Aravanopoulos FA. Artificial hybridization between Robinia pseudoacacia L. and R pscudoacacia var. monophylla Carr [J]. Forestry,2008,81:91-101.
Franklin T and Franklin F. The different mechanisms of gametophytic self-incompatibility [J]. Philosophical Transactions of the Royal Society of London B,2003,358:1025-1032.
Glaettli M, Pescatore L, Goudet J. Proximity-dependent pollen performance in Silene vulgaris [J]. Annals of Botany,2006,98:431-437.
Goodwillie C, Kalisz S, Eckert G. The evolutionary enigma of mixed mating systems in plants: occurrence, theoretical explanations, and empirical evidence [J]. Annual Review of Ecology, Evolution and Systematics,2005,36:47-79.
Guerra ME, WUnsch A, Margarita LC, Rodrigo J. Flower Emasculation as the Cause for Lack of Fruit Set in Japanese Plum Crosses [J]. Journal of the American Society for Horticultural Science,2010,135:556-562.
Haileslassie T, Mollel M, Skogsmyr I. Effects of nutrient level on maternal choice and siring success in Cucumis sativus (Cucurbitaceae) [J]. Evolution Ecology,2005,19:275-288.
Harder LD and Barrett SCH. Mating cost of large floral displays in hermaphrodite plants [J]. Nature. 1995,373:512-515.
Harlow WM. Harrar ES, White FM. Text book of dendrology. New York. McGraw Hill,1979.
Harmon-Threatt AN, Burns JH, Shemyakina LA, Knight TM. Breeding system and pollination ecology of introduced plants compared to their native relatives [J]. American Journal of Botany.2009,96:1544-1550.
Hedhly A, Hormaza JI, Herrero, M. Flower emasculation accelerates ovule degeneration and reduces fruit set in sweet cherry [J]. Science Hortic-Amsterdam,2009,119: 455-457.
Hedrick P and Kalinowski S. Inbreeding depression in conservation biology [J]. Annual Review of Ecology Evolution and Systematics,2000,31:139-162.
Hufford KM and Hamrick JL. Viability selection at three early life stages of the tropical tree, Platypodium elegans (Fabaceae, Papilionoideae) [J]. Evolution 2003,57:518-526.
Husband BC and Schemske DW. Magnitude and timing of inbreeding depression in a diploid population of Epilobium angustifolium (Onagraceae) [J]. Heredity,1995,75:206-215.
Husband BC and Schemske DW. Evolution of the magnitude and timing of inbreeding depression in plants [J]. Evolution; international journal of organic evolution,1996,50:54-70.
Ishida KH, Yoshimaru H, Ito H. Effects of geitonogamy on the seed set of Magnolia obovata Thumb. (Magnoliaceae) [J]. International Journal of Plant Sciences,2003,164:729-735.
Jarne P and Charlesworth D. The evolution of the selfing rate in functionally hermaphrodite plants and animals [J]. Annual Review of Ecology and Systematics,1993,24:441-466.
Johnston MO and Schoen DJ. On the measurement of inbreeding depression [J]. Evolution,1994,48: 1735-1741.
Keresztesi B. Breeding and cultivation of black locust, Robinia pseudoacacia, in Hungary [J]. Forest Ecology and Management,1983,6:217-244.
Kittelson PM and Maron JL. Outcrossing rate and inbreeding depression in the perennial yellow bush lupine, Lupinus arboreus (Fabaceae) [J]. American Journal of Botany,2000,87(5): 652-660.
Kittelson PM and Maron JL. Outcrossing rate and inbreeding depression in the perennial yellow bush lupine. Lupinus arboreus (Fabaceae) [J]. American Journal of Botany,2000,87: 652-660.
Korbecka G, Klinkhamer PGL, Vrieling K. Selective embryo abortion hypothesis revisited-a molecular approach [J]. Plant biology,2002,4:298-310.
Kruszewski L and Galloway L. Explaining outcrossing rate in Campanulastrum americanum (Campanulaceae):geitonogamy and cryptic self-incompatibility [J]. International Journal of Plant Science,2006,167:455-461.
Lande R, Schemske DW. The evolution of self-fertilization and inbreeding depression in plants. I: Genetic models [J]. Evolution,1985,39:24-40.
Lee CB, Page LE, McClure BA, Holtsford. Post-pollination hybridization barriers in Nicotiana section Alatae [J]. Sexual Plant Reproduction,2008,21:183-195.
Lee T. Patterns of fruit and seed production. In:Lovett Doust J, Lovett Doust L, eds. Plant reproductive ecology:patterns and strategies. New York:Oxford University Press:1988, 179-202.
Lee T and Bazzaz F. Regulation of fruit and seed production in an annual legume, Cassia fasciculata [J]. Ecology,1982a,63:1363-1373.
Lee T and Bazzaz F. Regulation of fruit maturation pattern in an annual legume, Cassia fasciculata [J]. Ecology,1982b,63:1374-1388.
Lian C and Hogetsu T. Development of microsatellite markers in black locust (Robinia pseudoacacia) using a dual-suppression-PCR technique [J]. Molecular Ecology Notes.2002,2:211-213.
Lian C, Oishi R, Miyashita, Hogetsu T. High somatic instability of a microsatellite locus in a clonal tree, Robinia pseudoacacia [J]. Theoretical and Applied Genetics,2004,108:836-841.
Lloyd D. Self-and cross-fertilization in plants.Ⅱ. The selection of self-fertilization [J]. International Journal of Plant Science,1992,153:370-380.
Lyons EE, Waser NM, Price MV, Antonovics J, Motten AF. Sources of variation in plant reproductive success and implications for concepts of sexual selection [J]. American Naturalist,1989,134:409-433.
Marshall D and Diggle P. Mechanisms of differential pollen donor performance in wild radish, Raphanus sativus (Brassicaceae) [J]. American Journal of Botany.2001,88:242-257.
Marshall D and Folsom M. Mate choice in plants:an anatomical to population perspective [J]. Annual Review of Ecology and Systematics,1991,22:37-63.
Marshall D, Shaner MGM, Oliva JP. Effects of pollen load size on seed paternity in wild radish:the roles of pollen competition and mate choice [J]. Evolution,2007,61:1925-1937.
Melser C and Klinkhamer P. Selective seed abortion increases offspring survival in Cynoglossum officinale (Boraginaceae) [J]. American Journal of Botany,2001,88:1033-1040.
Menaali J and Rocha O. Effect of ovule position within the pod on the probability of seed production in Bauhinia ungulata [J]. Annals of Botany,2005,95:449-455.
Mishima K, Hirao T, Urano S, Watanabe. Isolation and characterization of microsatellite markers from Robinia pseudoacacia L [J]. Theoretical and Applied Genetics,2008,108:836-841.
Mulcahy D. The rise of the angiosperms:a genecological factor [J]. Science,1974,206:20-23.
Mulcahy D. Rise of the angiosperms:a genecological factor [J]. Science,1979,206:20-23.
Mulugeta D. Maxwell BD, Fay PK, Dyer WE. Kochia (Kochia scoparia) pollen dispersion, viability and germination [J]. Weed Science,1994,42:548-552.
Murawski D and Hamrick J. The effect of the density of flowering individuals on the mating systems of nine tropical tree species [J]. Heredity,1991,67:167-174.
Murawski DA, Hamrick JL, Hubbell SP, Foster RB. Mating systems of two Bombacaceous trees of a neotropical moist forest [J]. Oecologia,1990,82:501-506.
Nettancourt Dde. Incompatibility in angiosperms [J]. Sexual Plant Reproduction,1997,10:185-199.
Obaysshi K, Tsumura Y, Iharaujino T, Niiyama K, Tanouchi H, Suyama Y, Washitani I, Lee CT, Lee SL, Norwati M. Genetic diversity and outcrossing rate between undisturbed and selectively logged forests of Shorea curtisii (Dipterocarpaceae)using microsattellite DNA analysis [J]. International Journal of Plant Science,2002,163:151-158.
Okie WR, Weinberger JH. Plums. In:Janick, J, Moore, JN (Eds) Fruit Breeding, vol. I: Tree and Tropical Fruits. John Wiley & Sons. Inc., NY,1996, pp 213-255.
Queller D. Sexual selection in a hermaphroditic plant [J]. Nature,1983,305:706-707.
Real D. Rizza MD, Quesenberry, Echenique M. Reproductive and molecular evidence for allogamy in Lotononis bainesii Baker [J]. Crop Science,2004,44:394-400.
Redei K. Black Locust (Robinia pseudoacacia L.) Growing in Hungary. Budapest, Publications of the Hungarian Forest Research Institute,1998.
Richards AJ. Plant breeding systems. London, UK, George Allen & Unwin,1986.
Rocha O and Stephenson A. Effects of ovule position on seed production, seed weight, and progeny performance in Phaseolus coccineus L [J]. American Journal of Botany,1990,77: 1320-1329.
Rocha O and Stephenson A. Effects of nonrandom seed abortion on progeny performance in Phaseolus coccineus L [J]. Evolution,1991,45:1198-1208.
Rocha O and Stephenson A. Order of fertilization within the ovary in Phaseolus coccineus L. (Leguminosae) [J]. Sexual Plant Reproduction,1991.9:273-278.
Schemske DW and Lande R. The evolution of self-fertilization and inbreeding depression in plants. Ⅱ. Empirical observations [J]. Evolution,1985,39:41-52.
Seavey S and Bawa K. Late-acting self-incompatibility in angiosperms [J]. The Botanical Review, 1986,52:195-219.
Skogsmyr I and Lankinen A. Sexual selection:an evolutionary force in plants? [J]. Biological Reviews,2002,77:537-562.
Snow A. Postpollination selection and male fitness in plants [J]. American Naturalist,1994,144: 69-83.
Souto CP, Aizen MA, Premoli AC. Effects of crossing distance and genetic relatedness on pollen performance in Alstroemeria aurea (Alstroemeriaceae) [J]. American Journal of Botany, 2002,89:427-432.
Stephenson AG. Flower and fruit abortion:proximate causes and ultimate functions [J]. Annual Review of Ecology and Systematics,1981,12:253-279.
Surles S, Hamrick JL, Bongarten, BC. Mating systems in open-pollinated families of black locust (Robinia pseudoacacia L.) [J]. Silvae Genet,1990,39:35-40.
Susko DJ. Effect of ovule position on patterns of seed maturation and abortion in Robinia pseudoacacia (Fabaceae) [J]. Canadian Journal of Botany,2006,84:1259-1265.
Torres C, Eynard MC, Aizen MA, Galetto L. Selective fruit maturation and seedling performance in Acacia caven (Fabaceae) [J]. International Journal of Plant Sciences,2002,163:809-813.
Travers S and Mazer S. The absence of cryptic self-incompatibility in Clarkia unguiculata (Onagraceae) [J]. American Journal of Botany,2000,87:191-196.
Waser N and Price M. Crossing distance effects on prezygotic performance in plants:an argument for female choice [J]. Oikos,1993,68:303-308.
Weir B. Genetic data analysis Ⅱ. Sinauer Associates. Sunderland, MA,1996.
Wiens D, Calvin CL, Wilson CA, Davern CI, Frank D, Seavy SR. Reproductive success, spontaneous embryo abortion, and genetic load in flowering plants [J]. Oecologia,1987,71: 501-509.
Wilcock C and Neiland R. Pollination failure in plants:why it happens and when it matters [J]. Trends in Plant Science,2002,7:270-277.
Williams E and Rouse J. Relationships of pollen size, pistil length and pollen tube growth rates in Rhododendron and their influence on hybridization [J]. Sexual Plant Reproduction,1990.3: 7-17.
Willson M. Sexual selection in plants and animals [J]. Trends in Ecology Evolution,1979,5: 210-214.
Willson M. Sexual selection and dicliny in angiosperms [J]. American Naturalist,1982,119: 579-583.
Winsor JA, Peretz S, Stephenson AG. Pollen competition in a natural population of Cucurbita foetidissima (Cucurbitaceae) [J]. American Journal of Botany,2000,87:527-532.
Wright S. Systems of mating. Ⅱ. The effects of inbreeding on the genetic composition of a population [J]. Genetics,1921,6:124-143.