绣球属(Hydrangea L.)及近缘属花粉形态的研究
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摘要
为深入研究绣球属植物花粉形态的分类学价值和系统学意义,厘清绣球属与近缘属之间的系统发育关系,该文利用扫描电子显微镜(SEM,scanning electron microscope)对国产绣球属及其近缘属41种绣球花科(Hydrangeaceae)植物的花粉形态以及表面纹饰进行了观察。结果表明:绣球属及其近缘属的花粉为三孔沟;形状多数为长圆体形或近球体形;赤道面观为椭圆形或圆形;极面观多为圆形,少数为三角形或圆三角形。花粉外壁纹饰可分为网状和孔穴状。网眼内的三级纹饰可分为光滑和具颗粒状突起。根据花粉形状和外壁纹饰类型将上述物种划分为4个组,即花粉的形状为长圆体形,表面纹饰为孔穴纹饰;花粉的形状为长圆体形,表面纹饰为网状纹饰;花粉的形状为近球体形,表面纹饰为孔穴纹饰;花粉的形状为近球体形,表面纹饰为网状纹饰。以上可进一步细分为8个类型。上述表明花粉形态证据可为绣球属及其近缘属的属下分类和种间界定提供重要佐证;但结合前人的系统发育重建分析该属植物花粉形态的系统学意义相对有限,如花粉形态证据对于该属及其近缘种属系统发育树上大支的界定难以提供有力的证据。
In order to study taxonomic value and systematic significance of Hydrangea L. as well as clarify phylogenetic relationships among the genus and its related ones,pollen samplings of 41 species of Hydrangea and its related species from China and surrounding areas were observed under scanning electron microscope( SEM). The results were as follows: the pollens of all these species were tricolporate; the pollen morphology were most subprolate and a few subspheroida; ellipse or circular in equatorial view; circular or sometimes triangular in polar view. The ornamentations on the exine can be reticulate ornamentations or perforate ornamentations. The ornamentation in the mesh could be smooth or granular bulged. The above-mentioned species could be divided into four groups according to the pollen morphology and ornamentations on the exine,the characters of each group were respectively pollen morphology subprolate,ornamentation on the exine reticulate; pollen morphology subprolate,ornamentation on the exine perforate; pollen morphology subspheroida,ornamentation on the exine reticulate; pollen morphology subspheroida,ornamentation on the exine perforate. according to the ratio of polar axis to equatorial axis,these species can be further divided into eight types. These indicate that the pollen characters can provide valuable evidence for infrageneric taxonomic treatments and species circumscription. However,this evidence shows limited value in diagnosing some major clades of phylogenetic trees reconstructed in previous studies. Further studies including broader sampling and new evidence of Hydrangea pollen are needed to explore its taxonomic and systematic values.
In order to study taxonomic value and systematic significance of Hydrangea L. as well as clarify phylogenetic relationships among the genus and its related ones,pollen samplings of 41 species of Hydrangea and its related species from China and surrounding areas were observed under scanning electron microscope( SEM). The results were as follows: the pollens of all these species were tricolporate; the pollen morphology were most subprolate and a few subspheroida; ellipse or circular in equatorial view; circular or sometimes triangular in polar view. The ornamentations on the exine can be reticulate ornamentations or perforate ornamentations. The ornamentation in the mesh could be smooth or granular bulged. The above-mentioned species could be divided into four groups according to the pollen morphology and ornamentations on the exine,the characters of each group were respectively pollen morphology subprolate,ornamentation on the exine reticulate; pollen morphology subprolate,ornamentation on the exine perforate; pollen morphology subspheroida,ornamentation on the exine reticulate; pollen morphology subspheroida,ornamentation on the exine perforate. according to the ratio of polar axis to equatorial axis,these species can be further divided into eight types. These indicate that the pollen characters can provide valuable evidence for infrageneric taxonomic treatments and species circumscription. However,this evidence shows limited value in diagnosing some major clades of phylogenetic trees reconstructed in previous studies. Further studies including broader sampling and new evidence of Hydrangea pollen are needed to explore its taxonomic and systematic values.
引文
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ENGLER A,1930.Hydrangea in Engler and Prantl[M].Die Natürlichen Pflanzenfamilien,2(18a):201-207.
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GUO H,YAN RY,ZHANG M,et al.,2016.Taxonomic significance of the pollen morphology of sect.Racemosae(Impatiens)[J].Guihaia,36(1):83-95.[郭慧,闫瑞亚,张梅,等,2016.凤仙花属总状花序组的花粉形态及其分类学意义[J].广西植物,36(1):83-95.]
HAO G,HU QM,1996.A study of pollen morphology of Hydrangeoideae(Hydrangeaceae)[J].J Trop Subtrop Bot,4(3):26-31.[郝刚,胡启明,1996.绣球亚科花粉形态的研究[J].热带亚热带植物学报,4(3):26-31.]
HUFFORD L,MOODY ML,SOLTIS DE,2001.A phylogenetic analysis of Hydrangeaceae based on sequences of the plastid gene mat K and their combination with rbc L and morphological data[J].Int J Plant Sci,162(4):835-846.
HUFFORD L,1997.A phylogenetic analysis of Hydrangeaceae based on morphological data[J].Int J Plant Sci,158(5):652-672.
JACOBS SJ,2010.Flag flower morphology and phylogeny of Hydrangeaceae tribe Hydrangeeae[D].Washington:Washington State University School of Biological Sciences.
LI GQ,SUN L,LIU Y,2005.Pollen features of 6 species in sect.Theopsis of genus Camellia[J].J Trop Subtrop Bot,1:40-44.[李广清,孙立,刘燕,2005.山茶属连蕊茶组6种植物花粉形态特征研究[J].热带亚热带植物学报,1:40-44.]
LI TQ,CAO HJ,KANG MS,et al.,2010.Pollen flora of China woody plant by SEM[M].Beijing:Science Press.[李天庆,曹慧娟,康木生,等,2010.中国木本植物花粉电镜扫描图志[M].北京:科学出版社.]
MAXIMOWICZ CJ,1867.Revisio Hydrangeearum asiae orientalis[J].Memoires de l'Academie Imperiale des Sci de St.-Petersbourg,10(16):6-18.
MCCLINTOCK E,1957.A monograph of the genus Hydrangea[J].Proc Calif Acad Sci,29:147-256.
SAMAIN MS,WANKE S,GOETGHEBEUR P,2010.Unraveling extensive paraphyly in the genus Hydrangea s.l.with implications for the systematics of tribe Hydrangeaceae[J].Syst Bot,35(3):593-600.
SOLTIS DE,SOLTIS PS,CHASE MW,et al.,2000.Angiosperm phylogeny inferred from a combined data set of 18SrDNA,rbc L,and atp B sequences[J].Bot J Linn Soc,133:381-461.
SOLTIS DE,SOLTIS PS,1997.Phylogenetic relationships in Saxifragaceae sensu lato:A comparison of topologies based on 18Sr DNA and rbc L sequences[J].Am J Bot,84:504-522.
SOLTIS DE,XIANG QY,HUFFORD L,1995.Relationships and evolution of Hydrangeaceae based on rbc L sequence data[J].Am J Bot,82(4):504-514.
WANG CY,LIU XL,2017.Pollen morphology of 13 subgenus Yulania species[J].J NE For Univ,45(11):54-59.[王晨宇,刘秀丽,2017.13种玉兰亚属植物的花粉形态扫描电镜观察[J].东北林业大学学报,45(11):54-59.]
WANG FX,YU CH,1954.Research on morphology of pollens1.Terminology and research methods[J].Acta Bot Sin,3(1):81-104.[王伏雄,喻诚鸿,1954.花粉形态的研究1.术语及研究方法[J].植物学报,3(1):81-104.]
WANG KF,WANG XZ,1983.Introduction to palynology[M].Beijing:Peking University Press:21-32.[王开发,王宪曾,1983.孢粉学概论[M].北京:北京大学出版社:21-32.]
WANG PL,1997.China practical pollen[M].Chengdu:Sichuan Science and Technology Press:5-150.[王萍莉,1997.中国实用花粉[M].成都:四川科学技术出版社:5-150.]
WEI ZF,HUANG SM,1995.Hydrangeoideae.FRPS[M].Beijing:Science Press,35(1):201-258.[卫兆芬,黄淑美,1995.绣球花亚科Hydrangeoideae.中国植物志[M].北京:科学出版社,35(1):201-258.]
WEI ZF,1994.A revision of the genus Hydrangea in China[J].Guihaia,14(2):101-121.[卫兆芬,1994.中国绣球属植物的修订[J].广西植物,14(2):101-121.]
YU HX,GUO XL,ZHOU SD,et al.,2018.Pollen and seed micro-morphology comparison and taxonomic significance of the Chinese Allium sect.Daghestanica[J].Acta Bot BorealOccident Sin,38(1):61-67.[于换喜,郭先林,周颂东,等,2018.中国葱属Daghestanica组植物的花粉和种子微形态学研究及其分类学意义[J].西北植物学报,38(1):61-67.]
ZENG XM,TONG F,LIU YM,et al.,2017.Pollen and seed micro-morphology characteristics and taxonomic significance of genus Ypsilandra[J].Acta Bot Boreal-Occident Sin,37(4):695-704.[曾心美,童芬,刘艳梅,等,2017.丫蕊花属植物花粉和种子微形态特征比较及其分类学意义[J].西北植物学报,37(4):695-704.]
CHEN HY,1954.Research on the information of Hydrangeoideae[J].Acta Phytotax Sin,3(2):108-161.[陈焕镛,1954.绣球亚科资料的研究[J].植物分类学报,3(2):108-161.]
CHEN YL,AKIYAMA S,OHBA H,2013.Balsaminaceae through Saxifragaceae[M]//WU ZY,RAVEN PH.Flora of China.Beijing:Science Press;St.Louis:Missouri Botanical Garden Press,8:411-422.
ENGLER A,1930.Hydrangea in Engler and Prantl[M].Die Natürlichen Pflanzenfamilien,2(18a):201-207.
ERDTMAN G,1978.Palynological handbook[M].Translated by Paleontology Research Laboratory,Chinese Academy of Sciences.Beijing:Science Press:238.[ERDTMAN G,1978.孢粉学手册[M].中国科学院植物研究所古植物研究室孢粉组译.北京:科学出版社:238.]
GE LP,2003.Systematics of the tribe Hydrangeeae DC.(Hydrangeaceae)[D].Beijing:Institute of Botany,Chinese Academy of Sciences.[葛丽萍,2003.绣球族的系统学研究[D].北京:中国科学院植物研究所.]
GUO H,YAN RY,ZHANG M,et al.,2016.Taxonomic significance of the pollen morphology of sect.Racemosae(Impatiens)[J].Guihaia,36(1):83-95.[郭慧,闫瑞亚,张梅,等,2016.凤仙花属总状花序组的花粉形态及其分类学意义[J].广西植物,36(1):83-95.]
HAO G,HU QM,1996.A study of pollen morphology of Hydrangeoideae(Hydrangeaceae)[J].J Trop Subtrop Bot,4(3):26-31.[郝刚,胡启明,1996.绣球亚科花粉形态的研究[J].热带亚热带植物学报,4(3):26-31.]
HUFFORD L,MOODY ML,SOLTIS DE,2001.A phylogenetic analysis of Hydrangeaceae based on sequences of the plastid gene mat K and their combination with rbc L and morphological data[J].Int J Plant Sci,162(4):835-846.
HUFFORD L,1997.A phylogenetic analysis of Hydrangeaceae based on morphological data[J].Int J Plant Sci,158(5):652-672.
JACOBS SJ,2010.Flag flower morphology and phylogeny of Hydrangeaceae tribe Hydrangeeae[D].Washington:Washington State University School of Biological Sciences.
LI GQ,SUN L,LIU Y,2005.Pollen features of 6 species in sect.Theopsis of genus Camellia[J].J Trop Subtrop Bot,1:40-44.[李广清,孙立,刘燕,2005.山茶属连蕊茶组6种植物花粉形态特征研究[J].热带亚热带植物学报,1:40-44.]
LI TQ,CAO HJ,KANG MS,et al.,2010.Pollen flora of China woody plant by SEM[M].Beijing:Science Press.[李天庆,曹慧娟,康木生,等,2010.中国木本植物花粉电镜扫描图志[M].北京:科学出版社.]
MAXIMOWICZ CJ,1867.Revisio Hydrangeearum asiae orientalis[J].Memoires de l'Academie Imperiale des Sci de St.-Petersbourg,10(16):6-18.
MCCLINTOCK E,1957.A monograph of the genus Hydrangea[J].Proc Calif Acad Sci,29:147-256.
SAMAIN MS,WANKE S,GOETGHEBEUR P,2010.Unraveling extensive paraphyly in the genus Hydrangea s.l.with implications for the systematics of tribe Hydrangeaceae[J].Syst Bot,35(3):593-600.
SOLTIS DE,SOLTIS PS,CHASE MW,et al.,2000.Angiosperm phylogeny inferred from a combined data set of 18SrDNA,rbc L,and atp B sequences[J].Bot J Linn Soc,133:381-461.
SOLTIS DE,SOLTIS PS,1997.Phylogenetic relationships in Saxifragaceae sensu lato:A comparison of topologies based on 18Sr DNA and rbc L sequences[J].Am J Bot,84:504-522.
SOLTIS DE,XIANG QY,HUFFORD L,1995.Relationships and evolution of Hydrangeaceae based on rbc L sequence data[J].Am J Bot,82(4):504-514.
WANG CY,LIU XL,2017.Pollen morphology of 13 subgenus Yulania species[J].J NE For Univ,45(11):54-59.[王晨宇,刘秀丽,2017.13种玉兰亚属植物的花粉形态扫描电镜观察[J].东北林业大学学报,45(11):54-59.]
WANG FX,YU CH,1954.Research on morphology of pollens1.Terminology and research methods[J].Acta Bot Sin,3(1):81-104.[王伏雄,喻诚鸿,1954.花粉形态的研究1.术语及研究方法[J].植物学报,3(1):81-104.]
WANG KF,WANG XZ,1983.Introduction to palynology[M].Beijing:Peking University Press:21-32.[王开发,王宪曾,1983.孢粉学概论[M].北京:北京大学出版社:21-32.]
WANG PL,1997.China practical pollen[M].Chengdu:Sichuan Science and Technology Press:5-150.[王萍莉,1997.中国实用花粉[M].成都:四川科学技术出版社:5-150.]
WEI ZF,HUANG SM,1995.Hydrangeoideae.FRPS[M].Beijing:Science Press,35(1):201-258.[卫兆芬,黄淑美,1995.绣球花亚科Hydrangeoideae.中国植物志[M].北京:科学出版社,35(1):201-258.]
WEI ZF,1994.A revision of the genus Hydrangea in China[J].Guihaia,14(2):101-121.[卫兆芬,1994.中国绣球属植物的修订[J].广西植物,14(2):101-121.]
YU HX,GUO XL,ZHOU SD,et al.,2018.Pollen and seed micro-morphology comparison and taxonomic significance of the Chinese Allium sect.Daghestanica[J].Acta Bot BorealOccident Sin,38(1):61-67.[于换喜,郭先林,周颂东,等,2018.中国葱属Daghestanica组植物的花粉和种子微形态学研究及其分类学意义[J].西北植物学报,38(1):61-67.]
ZENG XM,TONG F,LIU YM,et al.,2017.Pollen and seed micro-morphology characteristics and taxonomic significance of genus Ypsilandra[J].Acta Bot Boreal-Occident Sin,37(4):695-704.[曾心美,童芬,刘艳梅,等,2017.丫蕊花属植物花粉和种子微形态特征比较及其分类学意义[J].西北植物学报,37(4):695-704.]