巨龙竹笋-幼竹期秆形发育的组织解剖学研究
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摘要
巨龙竹(Dendrocalamus sinicus Chia et J. L. Sun)是云南特有的珍稀木本丛生竹,其秆形分为通直型和弯曲型两种变型。为了揭示巨龙竹不同秆形的组织结构特征,本文通过定点观测0～49 d的巨龙竹笋和幼竹生长发育状况,并采用石蜡切片技术对笋期0～45 d内样品的组织结构进行比较解剖学研究。结果显示:(1)巨龙竹在笋-幼竹(0～49 d)发育期,秆高生长呈"慢-快"的趋势,21～35 d时弯曲型茎秆开始显现,易于辨别;(2) 5～30 d时,弯曲型茎秆中维管束的发育早于通直型;对比弯曲型茎秆内外两侧维管束,内侧维管束导管内径较小,但纤维鞘中的纤维细胞层数更多;(3)弯曲型茎秆中薄壁细胞的分化早于通直型,20 d后弯曲型茎秆中的薄壁细胞出现明显的可被番红-固绿染色的细胞核,并呈有规律的排列;弯曲型茎秆内侧薄壁细胞稍小于外侧薄壁细胞,但内侧被染色细胞核的薄壁细胞多于外侧。研究结果表明巨龙竹弯曲型茎秆性状在笋期第21～35 d内即可通过茎秆形态判别,弯曲型茎秆中维管束的发育以及薄壁细胞分化均早于通直型茎秆,同一时期弯曲型茎秆内侧细胞分裂较外侧旺盛、维管束木质化程度更高。
Dendrocalamus sinicus Chia et J. L. Sun,a rare bamboo species endemic to Yunnan Province in China,is one of the strongest woody bamboo species in the world. It occurs as two variants,i.e.,straight-culmed variant( SC) and bent-culmed variant( BC). To reveal the anatomical basis of SC and BC, we collected data on culm height and morphological characteristics of shoots and young culms( emerged from ground for 0-49 d) from six clumps of D. sinicus. In addition,shoot and young culm development( 0-45 d) were observed using comparative anatomy based on paraffin sectioning. Results showed that culmheight growth exhibited a "slow-fast"pattern,and the BC morphological character clearly appeared at 21-35 d,which was distinct from that of SC. The development of vascular bundles occurred earlier in BC( 5-30 d) than in SC. Within BC,compared with the outer vascular bundles, the inner vascular bundles possessed smaller diameters and more fibrocytes in the fibrous sheath. Parenchyma cell differentiation occurred earlier in BC than in SC. Nuclei dyed with Safranine and Fast Green appeared in the parenchyma cells,which became regularly arranged from 20 d in BC. The parenchyma cells in the inner culm of BC were slightly smaller than those in the outer culm,whereas there was a greater number of parenchyma cells with dyed nuclei in the inner culm than in the outer culm. These results showed that the bent culm shape in BC can be identified by culm morphology during 21-35 d.The development of vascular bundles and differentiation of parenchyma cells occurred earlier in BC than in SC. Furthermore,compared with the outer culm in BC at the corresponding period,cell divisions in the inner culm were more vigorous and the degree of vascular bundle lignification was also higher.
Dendrocalamus sinicus Chia et J. L. Sun,a rare bamboo species endemic to Yunnan Province in China,is one of the strongest woody bamboo species in the world. It occurs as two variants,i.e.,straight-culmed variant( SC) and bent-culmed variant( BC). To reveal the anatomical basis of SC and BC, we collected data on culm height and morphological characteristics of shoots and young culms( emerged from ground for 0-49 d) from six clumps of D. sinicus. In addition,shoot and young culm development( 0-45 d) were observed using comparative anatomy based on paraffin sectioning. Results showed that culmheight growth exhibited a "slow-fast"pattern,and the BC morphological character clearly appeared at 21-35 d,which was distinct from that of SC. The development of vascular bundles occurred earlier in BC( 5-30 d) than in SC. Within BC,compared with the outer vascular bundles, the inner vascular bundles possessed smaller diameters and more fibrocytes in the fibrous sheath. Parenchyma cell differentiation occurred earlier in BC than in SC. Nuclei dyed with Safranine and Fast Green appeared in the parenchyma cells,which became regularly arranged from 20 d in BC. The parenchyma cells in the inner culm of BC were slightly smaller than those in the outer culm,whereas there was a greater number of parenchyma cells with dyed nuclei in the inner culm than in the outer culm. These results showed that the bent culm shape in BC can be identified by culm morphology during 21-35 d.The development of vascular bundles and differentiation of parenchyma cells occurred earlier in BC than in SC. Furthermore,compared with the outer culm in BC at the corresponding period,cell divisions in the inner culm were more vigorous and the degree of vascular bundle lignification was also higher.
引文
[1]辉朝茂,杨宇明,杜凡.珍稀竹种巨龙竹生态生物学特性及开发利用基础性研究[M].昆明:云南科技出版社,2006.
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[3]谷志佳,杨汉奇,孙茂盛,杨宇明.巨龙竹资源分布特点及其开花结实现象[J].林业科学研究,2012,25(1):1-5.Gu ZJ,Yang HQ,Sun MS,Yang YM. Distribution characteristics,flowering and seeding of Dendrocalamus sinicus in Yunnan,China[J]. Forest Research,2012,25(1):1-5.
[4] Chen LN,Cui YZ,Wong KM,Li DZ,Yang HQ. Breeding system and pollination of two closely related bamboo species[J]. Ao B Plants,2017,9(3):1-11.
[5]杨汉奇,孙茂盛,阮桢媛,董禹然,梁宁.云南4种典型热带丛生竹的种源分化[J].林业科学研究,2014,27(2):168-173.Yang HQ,Sun MS,Ruan ZY,Dong YR,Liang N. Study on provenance differentiation of four typical tropical clump bamboos in Yunnan,China[J]. Forest Research,2014,27(2):168-173.
[6]杨汉奇,阮桢媛,田波,杨宇明,孙茂盛.通直型巨龙竹不同地理种源遗传分化的ISSR分析[J].浙江林学院学报,2010,27(1):81-86.Yang HQ,Ruan ZY,Tian B,Yang YM,Sun MS. Genetic differentiation of four straight-culmed provenances of Dendrocalamus sinicus based on ISSRs[J]. Journal of Zhejiang Forestry College,2010,27(1):81-86.
[7] Dong YR,Zhang ZR,Yang HQ. Sixteen novel microsatellite markers developed for Dendrocalamus sinicus(Poaceae),the strongest woody bamboo in the world[J]. Am J Bot,2012,99(9):e347-e349.
[8] Yang JB,Dong YR,Wong KM,Gu ZJ,Yang HQ,Li DZ.Genetic structure and differentiation in Dendrocalamus sinicus(Poaceae:Bambusoideae)populations provide insight into evolutionary history and speciation of woody bamboos[J]. Sci Rep,2018,8:16933.
[9] Li L,Cheng ZC,Ma YJ,Bai QS,Li XY,Cao ZH,et al.The association of hormone signalling genes,transcription and changes in shoot anatomy during moso bamboo growth[J]. Plant Biotechnol J,2018,16:72-85.
[10]龙海艳,顾小平,袁娜,岳晋军,楼崇.开花红哺鸡竹叶片的显微结构与光响应参数研究[J].植物科学学报,2014,32(3):273-278.Long HY,Gu XP,Yuan N,Yue JJ,Lou C. Microstructure and light-response parameters of flowering bamboo(Phyllostachys iridescens)blades[J]. Plant Science Journal,2014,32(3):273-278.
[11]黄慧敏,董蓉,向运蓉,何丹妮,陈娟,张小晶,陶建平.不同群落冠层环境下紫耳箭竹笋期生长发育研究[J].植物科学学报,2018,36(5):696-704.Huang HM,Dong R,Xiang YR,He DN,Chen J,Zhang XJ,Tao JP. Study on shooting rhythm of Fargesia decurvata under different canopy conditions[J]. Plant Science Journal,2018,36(5):696-704.
[12]普晓兰,杜凡.巨龙竹纤维形态及变异规律的研究[J].云南林业科技,2003,23(1):1-4.Pu XL,Du F. Study of fiber morphology and its variation law of Dendrocalamus sinicus[J]. Yunnan Forestry Science and Technology,2003,23(1):1-4.
[13]普晓兰,杜凡.巨龙竹竹材结构及其变异的解剖学研究[J].西南林学院学报,2003,24(1):1-5.Pu XL,Du F. Anatomical Studies on the culm and variation of Dendrocalamus sinicus[J]. Journal of Southwest Forestry College,2003,24(1):1-5.
[14]崔凯.毛竹茎秆快速生长的机理研究[D].北京:中国林业科学研究院,2011.
[15]叶祖云,阮少江.绿竹笋的石蜡切片制作[J].宁德师专学报(自然科学版),2005,17(2):140-141.Ye ZY,Ruan SJ. Manufaction of paraffin section on green bamboo shoot[J]. Journal of Ningde Teachers College(Natural Science),2005,17(2):140-141.
[16]李龙.毛竹笋生长空间变化规律和生长素相关基因分析[D].北京:中国林业科学研究院,2016.
[17]李霞,郭振华.麻竹节间伸长过程的初步研究[J].植物分类与资源学报,2014,36(1):22-28.Li X,Guo ZH. A pilot study on internode elongation in a paleotropical bamboo,Dendrocalamus latiflorus(Poaceae:Bambusoideae)[J]. Plant Diversity and Resources,2014,36(1):22-28.
[18]辉朝茂,张国学,李在留,谷中明,杨宇明.珍稀竹种巨龙竹种群特性及其保护研究[J].竹子研究汇刊,2004,23(4):4-9.Hui ZM,Zhang GX,Li ZL,Gu ZM,Yang YM. A study on the population characteristics of the valuable and rare bamboo species of Dendrocalamus sinicus and its conservation in Yunnan, China[J]. Journal of Bamboo Research,2004,23(4):4-9.
[19]徐有明,郝培应,费本华.竹笋形态发育构建过程中解剖结构及化学成分的动态变化[J].东北林业大学学报,2008,36(4):8-11.Xu YM,Hao PY,Fei BH. Dynamic change of anatomical structure and chemical constitutes of bamboo shoots for Phyllostachys pubescence at development stage[J].Journal of Northeast Forestry University,2008,36(4):8-12.
[20]金亮,薛庆中,肖建富,张宪银.不同倍性水稻植株茎解剖结构比较研究[J].浙江大学学报(农业与生命科学版),2009,35(5):489-496.Jin L,Xue QZ,Xiao JF,Zhang XY. Comparative studies on stem anatomical structure of rice plants with various ploidy level[J]. Journal of Zhejiang University(Agriculture&Life Science),2009,35(5):489-496.
[21]李素坤,张秋芝,郝玉兰,南张杰,朱瑾,陈丽,潘金豹.玉米成熟期茎秆石蜡切片方法的研究[J].安徽农业科学,2010,38(8):3935-3937.Li SK,Zhang QZ,Hao YL,Nan ZJ,Zhu J,Chen L,Pan JB. Research on the method of paraffin section of the corn stalk in its maturity stage[J]. Journal of Anhui Agricultural Sciences,2010,38(8):3935-3937.
[22]李兴会,罗蓓,何蕊.青皮竹和慈竹不同发育期竹材解剖特征研究[J].世界竹藤通讯,2017,15(4):9-12.Li XH,Luo B,He R. Anatomical characteristics of Bambusa texlilis McClure and Bambusa emeiensis in different growth periods[J]. World Bamboo and Rattan,2017,15(4):9-12.
[23] Gritsch CS,Murphy RJ. Ultrastructure of fibre and parenchyma cell walls during early stages of culm development in Dendrodalamus asper[J]. Ann Bot,2005,95(4):619-629.
[24]杨艳华,朱镇,张亚东,赵庆勇,周丽慧,王才林.水稻茎秆解剖结构与抗倒伏能力关系的研究[J].广西植物,2012,32(6):834-839.Yang YH,Zhu Z,Zhang YD,Zhao QY,Zhou LH,Wang CL. Relationship between anatomic structure of the stem and lodging resistance of rice[J]. Guihaia,2012,32(6):834-839.
[25] Cui L,Peng WX,Sun ZJ,Shang LL,Chen GN. Weibull statistical analysis of tensile strength of vascular bundle in inner layer of Moso bamboo culm in molecular parasitology and vector biology[J]. Pak J Pharm Sci,2014,27(S4):1083-1087.
[2]刘世男,辉朝茂.珍稀竹种巨龙竹的研究现状和展望[J].世界竹藤通讯,2011,9(5):26-30.Liu SN,Hui ZM. Research status and outlook of rare bamboo species of Dendrocalamus sinicus[J]. World Bamboo and Rattan,2011,9(5):26-30.
[3]谷志佳,杨汉奇,孙茂盛,杨宇明.巨龙竹资源分布特点及其开花结实现象[J].林业科学研究,2012,25(1):1-5.Gu ZJ,Yang HQ,Sun MS,Yang YM. Distribution characteristics,flowering and seeding of Dendrocalamus sinicus in Yunnan,China[J]. Forest Research,2012,25(1):1-5.
[4] Chen LN,Cui YZ,Wong KM,Li DZ,Yang HQ. Breeding system and pollination of two closely related bamboo species[J]. Ao B Plants,2017,9(3):1-11.
[5]杨汉奇,孙茂盛,阮桢媛,董禹然,梁宁.云南4种典型热带丛生竹的种源分化[J].林业科学研究,2014,27(2):168-173.Yang HQ,Sun MS,Ruan ZY,Dong YR,Liang N. Study on provenance differentiation of four typical tropical clump bamboos in Yunnan,China[J]. Forest Research,2014,27(2):168-173.
[6]杨汉奇,阮桢媛,田波,杨宇明,孙茂盛.通直型巨龙竹不同地理种源遗传分化的ISSR分析[J].浙江林学院学报,2010,27(1):81-86.Yang HQ,Ruan ZY,Tian B,Yang YM,Sun MS. Genetic differentiation of four straight-culmed provenances of Dendrocalamus sinicus based on ISSRs[J]. Journal of Zhejiang Forestry College,2010,27(1):81-86.
[7] Dong YR,Zhang ZR,Yang HQ. Sixteen novel microsatellite markers developed for Dendrocalamus sinicus(Poaceae),the strongest woody bamboo in the world[J]. Am J Bot,2012,99(9):e347-e349.
[8] Yang JB,Dong YR,Wong KM,Gu ZJ,Yang HQ,Li DZ.Genetic structure and differentiation in Dendrocalamus sinicus(Poaceae:Bambusoideae)populations provide insight into evolutionary history and speciation of woody bamboos[J]. Sci Rep,2018,8:16933.
[9] Li L,Cheng ZC,Ma YJ,Bai QS,Li XY,Cao ZH,et al.The association of hormone signalling genes,transcription and changes in shoot anatomy during moso bamboo growth[J]. Plant Biotechnol J,2018,16:72-85.
[10]龙海艳,顾小平,袁娜,岳晋军,楼崇.开花红哺鸡竹叶片的显微结构与光响应参数研究[J].植物科学学报,2014,32(3):273-278.Long HY,Gu XP,Yuan N,Yue JJ,Lou C. Microstructure and light-response parameters of flowering bamboo(Phyllostachys iridescens)blades[J]. Plant Science Journal,2014,32(3):273-278.
[11]黄慧敏,董蓉,向运蓉,何丹妮,陈娟,张小晶,陶建平.不同群落冠层环境下紫耳箭竹笋期生长发育研究[J].植物科学学报,2018,36(5):696-704.Huang HM,Dong R,Xiang YR,He DN,Chen J,Zhang XJ,Tao JP. Study on shooting rhythm of Fargesia decurvata under different canopy conditions[J]. Plant Science Journal,2018,36(5):696-704.
[12]普晓兰,杜凡.巨龙竹纤维形态及变异规律的研究[J].云南林业科技,2003,23(1):1-4.Pu XL,Du F. Study of fiber morphology and its variation law of Dendrocalamus sinicus[J]. Yunnan Forestry Science and Technology,2003,23(1):1-4.
[13]普晓兰,杜凡.巨龙竹竹材结构及其变异的解剖学研究[J].西南林学院学报,2003,24(1):1-5.Pu XL,Du F. Anatomical Studies on the culm and variation of Dendrocalamus sinicus[J]. Journal of Southwest Forestry College,2003,24(1):1-5.
[14]崔凯.毛竹茎秆快速生长的机理研究[D].北京:中国林业科学研究院,2011.
[15]叶祖云,阮少江.绿竹笋的石蜡切片制作[J].宁德师专学报(自然科学版),2005,17(2):140-141.Ye ZY,Ruan SJ. Manufaction of paraffin section on green bamboo shoot[J]. Journal of Ningde Teachers College(Natural Science),2005,17(2):140-141.
[16]李龙.毛竹笋生长空间变化规律和生长素相关基因分析[D].北京:中国林业科学研究院,2016.
[17]李霞,郭振华.麻竹节间伸长过程的初步研究[J].植物分类与资源学报,2014,36(1):22-28.Li X,Guo ZH. A pilot study on internode elongation in a paleotropical bamboo,Dendrocalamus latiflorus(Poaceae:Bambusoideae)[J]. Plant Diversity and Resources,2014,36(1):22-28.
[18]辉朝茂,张国学,李在留,谷中明,杨宇明.珍稀竹种巨龙竹种群特性及其保护研究[J].竹子研究汇刊,2004,23(4):4-9.Hui ZM,Zhang GX,Li ZL,Gu ZM,Yang YM. A study on the population characteristics of the valuable and rare bamboo species of Dendrocalamus sinicus and its conservation in Yunnan, China[J]. Journal of Bamboo Research,2004,23(4):4-9.
[19]徐有明,郝培应,费本华.竹笋形态发育构建过程中解剖结构及化学成分的动态变化[J].东北林业大学学报,2008,36(4):8-11.Xu YM,Hao PY,Fei BH. Dynamic change of anatomical structure and chemical constitutes of bamboo shoots for Phyllostachys pubescence at development stage[J].Journal of Northeast Forestry University,2008,36(4):8-12.
[20]金亮,薛庆中,肖建富,张宪银.不同倍性水稻植株茎解剖结构比较研究[J].浙江大学学报(农业与生命科学版),2009,35(5):489-496.Jin L,Xue QZ,Xiao JF,Zhang XY. Comparative studies on stem anatomical structure of rice plants with various ploidy level[J]. Journal of Zhejiang University(Agriculture&Life Science),2009,35(5):489-496.
[21]李素坤,张秋芝,郝玉兰,南张杰,朱瑾,陈丽,潘金豹.玉米成熟期茎秆石蜡切片方法的研究[J].安徽农业科学,2010,38(8):3935-3937.Li SK,Zhang QZ,Hao YL,Nan ZJ,Zhu J,Chen L,Pan JB. Research on the method of paraffin section of the corn stalk in its maturity stage[J]. Journal of Anhui Agricultural Sciences,2010,38(8):3935-3937.
[22]李兴会,罗蓓,何蕊.青皮竹和慈竹不同发育期竹材解剖特征研究[J].世界竹藤通讯,2017,15(4):9-12.Li XH,Luo B,He R. Anatomical characteristics of Bambusa texlilis McClure and Bambusa emeiensis in different growth periods[J]. World Bamboo and Rattan,2017,15(4):9-12.
[23] Gritsch CS,Murphy RJ. Ultrastructure of fibre and parenchyma cell walls during early stages of culm development in Dendrodalamus asper[J]. Ann Bot,2005,95(4):619-629.
[24]杨艳华,朱镇,张亚东,赵庆勇,周丽慧,王才林.水稻茎秆解剖结构与抗倒伏能力关系的研究[J].广西植物,2012,32(6):834-839.Yang YH,Zhu Z,Zhang YD,Zhao QY,Zhou LH,Wang CL. Relationship between anatomic structure of the stem and lodging resistance of rice[J]. Guihaia,2012,32(6):834-839.
[25] Cui L,Peng WX,Sun ZJ,Shang LL,Chen GN. Weibull statistical analysis of tensile strength of vascular bundle in inner layer of Moso bamboo culm in molecular parasitology and vector biology[J]. Pak J Pharm Sci,2014,27(S4):1083-1087.
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