二倍体菘蓝开花习性及传粉特性研究
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摘要
菘蓝是药材板蓝根和大青叶的基原植物,依赖种子繁殖。通过田间观测和人工授粉实验,对民乐县栽培菘蓝开花时序和传粉生物学系统进行研究,旨在为其种子繁育和杂种优势利用提供科学和技术依据。研究表明,菘蓝为无限花序,单株表现异向开花习性,花序从植株顶部向下,单花从花序基部向上依次开放。花两性,雄蕊6,4枚长,2枚短,花萼和花瓣均为4,花萼绿色,花瓣黄色,子房上位,1室。单花寿命6～7 d,异常气候可缩短至3～4 d,群体开花期30 d左右。即将开放的花蕾柱头已成熟,花瓣刚露出花萼后开花当天,花粉鲜黄而散,末花期子房膨大,花粉干枯。短角果结种子0～2粒。自然结籽率95.0%,不去雄和去雄套纸袋均不结实,同株异花去雄授粉结籽率仅4.0%,去雄套纸袋异株异花授粉结籽率72.0%,去雄不套袋结籽率53.3%,自交不亲和指数为0.96～1.00。不去雄套网袋结籽率5.0%,去雄后提高至13.7%,访花者主要有食蚜蝇、黑蜂等。说明菘蓝具有较强的自交不亲和性,不存在无融合生殖现象,属典型的异花授粉类型,需传粉者,传粉媒介以虫媒为主,风媒为辅。
Isatis indigotica is the primary ingredient of traditional Chinese drugs ‘Banlangen' and ‘Daqingye', and its cultivation relies on seed propagation. This research examined the flowering habit and pollination biology of I. indigotica cultivated in Minle county. The work included field observation and an artificial pollination experiment, in order to provide scientific and technical information to support seed production and heterosis utilization in this crop. It was found that the plant had an infinite inflorescence consisting of an upright main stem bearing raceme-like branches with new branches above as older matured branches below and 10-20 individual flowers per branch opening in an acropetal sequence over a flowering period of some 30 days. Flowers are bisexual with 6 stamens, four of which are long and two of which are short. There are 4 green calyces, 4 yellow petals with ovary superior, and 1 locule. The flower lifespan was 6-7 days but was shortened to 3-4 days under adverse climate conditions. The stigma matures just prior to the flowers opening. When the petals are just emerging from the calyx tube, bright fresh yellow pollen grains become visible and are dispersed by the flowering day. Anthers then begin to wither, and the ovary expands at the end of the flowering stage, growing a short pod with 0-2 seeds. Seed set was 95.0% for plants under natural conditions. Under covered conditions with paper bags, seed set was very low(approximately 4%) both with and without castration, but was 72.0% when plants were cross-pollinated after castration. When plants were cross-pollinated after castration, but the inflorescence was unbagged, seed set was 53.3%. The self-incompatibility index was assessed as 0.96-1.00. For bagged racemes, seed set was 5.0% without castration and increased to 13.7% after castration. The most common flower pollinator visitors were Episyrphus balteatus and Apis mellifer but various other insect species also visited. These results indicate that I. indigotica has strong self-incompatibility without apomixis, and can be considered a typical cross-pollinating plants needing pollinators, pollinated mainly by insects but supplemented by wind.
Isatis indigotica is the primary ingredient of traditional Chinese drugs ‘Banlangen' and ‘Daqingye', and its cultivation relies on seed propagation. This research examined the flowering habit and pollination biology of I. indigotica cultivated in Minle county. The work included field observation and an artificial pollination experiment, in order to provide scientific and technical information to support seed production and heterosis utilization in this crop. It was found that the plant had an infinite inflorescence consisting of an upright main stem bearing raceme-like branches with new branches above as older matured branches below and 10-20 individual flowers per branch opening in an acropetal sequence over a flowering period of some 30 days. Flowers are bisexual with 6 stamens, four of which are long and two of which are short. There are 4 green calyces, 4 yellow petals with ovary superior, and 1 locule. The flower lifespan was 6-7 days but was shortened to 3-4 days under adverse climate conditions. The stigma matures just prior to the flowers opening. When the petals are just emerging from the calyx tube, bright fresh yellow pollen grains become visible and are dispersed by the flowering day. Anthers then begin to wither, and the ovary expands at the end of the flowering stage, growing a short pod with 0-2 seeds. Seed set was 95.0% for plants under natural conditions. Under covered conditions with paper bags, seed set was very low(approximately 4%) both with and without castration, but was 72.0% when plants were cross-pollinated after castration. When plants were cross-pollinated after castration, but the inflorescence was unbagged, seed set was 53.3%. The self-incompatibility index was assessed as 0.96-1.00. For bagged racemes, seed set was 5.0% without castration and increased to 13.7% after castration. The most common flower pollinator visitors were Episyrphus balteatus and Apis mellifer but various other insect species also visited. These results indicate that I. indigotica has strong self-incompatibility without apomixis, and can be considered a typical cross-pollinating plants needing pollinators, pollinated mainly by insects but supplemented by wind.
引文
[1]Chen S D.Genetic diversity and quality evaluation of Isatis indigotica.Nanjing:Nanjing Agricultural University,2014.陈苏丹.菘蓝遗传多样性及其药材品质评价研究.南京:南京农业大学,2014.
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[4]Yang C W.Advances in pharmacological research of Radix Isatidis.Chinese Journal of Modern Drug Application,2016,10(9):282-283.杨春望.板蓝根药理研究进展.中国现代药物应用,2016,10(9):282-283.
[5]Ye Q.Study on biological characteristics of Isatis indigotica.Yangling:Northwest A&F University,2006.叶青.菘蓝生物学特性的研究.杨凌:西北农林科技大学,2006.
[6]Gao Y H.Systematical research on self-incompatibility of Isatis indigotica.Changchun:Jilin Agricultural University,2008.高延辉.菘蓝自交不亲和特性的系统研究.长春:吉林农业大学,2008.
[7]Cao H L,Cao G F,Wei J H,et al.Effects of genetical and environmental factors on medicinal plant quality.Chinese Traditional&Herbal Drugs,2007,38(5):785-788.曹海禄,曹国番,魏建和,等.遗传和环境因子对药用植物品质的影响.中草药,2007,38(5):785-788.
[8]Zhang D Y.The life history evolution and reproductive ecology of plant.Beijing:Science Press,2004:97-279.张大勇.植物生活史进化与繁殖生态学.北京:科学出版社,2004:97-279.
[9]Wang E J,Chen Y,Han D H,et al.Correlation and path analysis of the yield folium isatidis and radix isatidis yield per plant and main agronomic traits in Isatis indigotica.Journal of Nuclear Agricultural Sciences,2018,32(2):399-406.王恩军,陈垣,韩多红,等.菘蓝农艺性状与药材产量的相关和通径分析.核农学报,2018,32(2):399-406.
[10]Dafni A,Maués M M.A rapid and simple procedure to determine stigma receptivity.Sexual Plant Reproduction,1998,11:177-180.
[11]Kunin W E,Shmida A.Plant reproductive traits as a function of local regional and global abundance.Conservation Biology,1997,11:183-192.
[12]Vogler D W,Kalisz S.Sex among the flowers:The distribution of plant mating systems.Evolution,2001,55(1):202-204.
[13]Ren M X.Floral syndrome and breeding system of Lobelia chinensis,aperennial herbs with stamen fusion.Journal of Plant Ecology,2009,33(2):361-368.任明迅.雄蕊合生植物半边莲的花部综合征与繁育系统.植物生态学报,2009,33(2):361-368.
[14]He Y P,Liu J Q.A review on recent advances in the studies of plant breeding system.Acta Phytoecologica Sinica,2003,(2):151-163.何亚平,刘建全.植物繁育系统研究的最新进展和评述.植物生态学报,2003,(2):151-163.
[15]Chen Y,Xu B Q,Guo F X,et al.Studies of the floral organ characteristic and sexual breeding system for Fritillaria unibracteata.Acta Prataculturae Sinica,2017,26(1):90-98.陈垣,徐博琼,郭凤霞,等.暗紫贝母花器官特征及有性繁育系统研究.草业学报,2017,26(1):90-98.
[16]Guo F X,Chen Y,Zhang J F,et al.Abloom phenology and pollination mode of endangered plant Fritillaria unibracteata.Acta Agrestia Sinica,2017,25(6):1324-1332.郭凤霞,陈垣,张荩凤,等.濒危植物暗紫贝母的开花物候和传粉方式.草地学报,2017,25(6):1324-1332.
[17]Gao Y H,Ren Y Y,Sun H M,et al.Identification of self-incompatibility of Isatis indigotica.Forest By-Product and Speciality in China,2007,(6):11-15.高延辉,任跃英,孙红梅,等.菘蓝自交不亲和性鉴定.中国林副特产,2007,(6):11-15.
[18]Gao Y H,Gao B,Wang Y H,et al.Screening of the chemicals to resolve the problem of self pollination failure of Isatis indigotica.Forest By-Product and Speciality in China,2008,(1):12-15.高延辉,高博,王云贺,等.克服菘蓝自交不亲和性的化学试剂筛选研究.中国林副特产,2008,(1):12-15.
[19]Wang E J,Han D H,Zhang Y,et al.Effect of harvested periods on the yield and quality dynamic for the Isatis indigotica root in Hexi Corridor.Journal of Chinese Medicinal Materials,2016,39(12):2686-2690.王恩军,韩多红,张勇,等.采收期对河西走廊产板蓝根产量和品质的影响.中药材,2016,39(12):2686-2690.
[20]Hou G P,Zhen D S,Jiang Q L,et al.Discussion on environment of traditional Chinese medicine producing area in Minle county.Modern Agricultural Sciences and Technology,2017,(6):114-115.侯格平,甄东升,姜青龙,等.民乐县中药材产地环境探讨.现代农业科技,2017,(6):114-115.
[21]Wang E J,Chen Y,Han D H,et al.Effect of cropping pattern on agronomic characteristics,yield and quality of Radix isatidis.Chinese Journal of Eco-Agriculture,2017,25(11):1661-1670.王恩军,陈垣,韩多红,等.栽培方式对菘蓝农艺性状及产量和品质的影响.中国生态农业学报,2017,25(11):1661-1670.
[22]Wang X J,Zhu L Q.Method for determining self-incompatibility of Brassica oleracea L.Journal of Agricultural Biotechnology,1998,6(2):195-198.王晓佳,朱利泉.甘蓝自交不亲和性的测定方法.农业生物技术学报,1998,6(2):195-198.
[23]Zhou Y L.Plant biology.Beijing:Higher Education Press,2000:241-243.周云龙.植物生物学.北京:高等教育出版社,2000:241-243.
[24]Dafni A.Pollination ecology.New York:Oxford University Press,1992:25-45.
[25]Wang Y H,Guo F X,Chen Y,et al.Studies on blossoming habit and seed filling characteristics of wild Aconitum sinomontanum.Acta Prataculturae Sinica,2013,22(4):76-82.王永辉,郭凤霞,陈垣,等.野生高乌头开花习性及种子灌浆特性研究.草业学报,2013,22(4):76-82.
[26]Shoichi K,Masuda J,Hayashi K.Life-history monographs of Japanese plants.10:Fritillaria koidzumiana,Ohwi(Liliaceae).Plant Species Biology,2008,23(1):51-57.
[27]Ren Y Y.Breeding of medicinal plants.Beijing:China Agriculture Press,2011:17-18.任跃英.药用植物育种学.北京:中国农业出版社,2011:17-18.
[28]Hu C,Liu Z J,Wu G Q,et al.Floral characteristic and breeding system of Anemone obtusiloba.Acta Agrestia Sinica,2013,21(4):783-788.胡春,刘左军,伍国强,等.钝裂银莲花花部综合特征及其繁育系统.草地学报,2013,21(4):783-788.
[2]Chinese Pharmacopoeia Commission.The Chinese pharmacopoeia(a).China Medical Science Press,2015:21-22,205-206.国家药典委员会.中国药典(一部).中国医药科技出版社,2015:21-22,205-206.
[3]Cheng W L,Fuu J T,Tsa C H,et al.Anti-SARS coronavirus 3C-like protease effects of Isatis indigotica root and plant-derived phenolic compounds.Antiviral Research,2005,68(1):36-42.
[4]Yang C W.Advances in pharmacological research of Radix Isatidis.Chinese Journal of Modern Drug Application,2016,10(9):282-283.杨春望.板蓝根药理研究进展.中国现代药物应用,2016,10(9):282-283.
[5]Ye Q.Study on biological characteristics of Isatis indigotica.Yangling:Northwest A&F University,2006.叶青.菘蓝生物学特性的研究.杨凌:西北农林科技大学,2006.
[6]Gao Y H.Systematical research on self-incompatibility of Isatis indigotica.Changchun:Jilin Agricultural University,2008.高延辉.菘蓝自交不亲和特性的系统研究.长春:吉林农业大学,2008.
[7]Cao H L,Cao G F,Wei J H,et al.Effects of genetical and environmental factors on medicinal plant quality.Chinese Traditional&Herbal Drugs,2007,38(5):785-788.曹海禄,曹国番,魏建和,等.遗传和环境因子对药用植物品质的影响.中草药,2007,38(5):785-788.
[8]Zhang D Y.The life history evolution and reproductive ecology of plant.Beijing:Science Press,2004:97-279.张大勇.植物生活史进化与繁殖生态学.北京:科学出版社,2004:97-279.
[9]Wang E J,Chen Y,Han D H,et al.Correlation and path analysis of the yield folium isatidis and radix isatidis yield per plant and main agronomic traits in Isatis indigotica.Journal of Nuclear Agricultural Sciences,2018,32(2):399-406.王恩军,陈垣,韩多红,等.菘蓝农艺性状与药材产量的相关和通径分析.核农学报,2018,32(2):399-406.
[10]Dafni A,Maués M M.A rapid and simple procedure to determine stigma receptivity.Sexual Plant Reproduction,1998,11:177-180.
[11]Kunin W E,Shmida A.Plant reproductive traits as a function of local regional and global abundance.Conservation Biology,1997,11:183-192.
[12]Vogler D W,Kalisz S.Sex among the flowers:The distribution of plant mating systems.Evolution,2001,55(1):202-204.
[13]Ren M X.Floral syndrome and breeding system of Lobelia chinensis,aperennial herbs with stamen fusion.Journal of Plant Ecology,2009,33(2):361-368.任明迅.雄蕊合生植物半边莲的花部综合征与繁育系统.植物生态学报,2009,33(2):361-368.
[14]He Y P,Liu J Q.A review on recent advances in the studies of plant breeding system.Acta Phytoecologica Sinica,2003,(2):151-163.何亚平,刘建全.植物繁育系统研究的最新进展和评述.植物生态学报,2003,(2):151-163.
[15]Chen Y,Xu B Q,Guo F X,et al.Studies of the floral organ characteristic and sexual breeding system for Fritillaria unibracteata.Acta Prataculturae Sinica,2017,26(1):90-98.陈垣,徐博琼,郭凤霞,等.暗紫贝母花器官特征及有性繁育系统研究.草业学报,2017,26(1):90-98.
[16]Guo F X,Chen Y,Zhang J F,et al.Abloom phenology and pollination mode of endangered plant Fritillaria unibracteata.Acta Agrestia Sinica,2017,25(6):1324-1332.郭凤霞,陈垣,张荩凤,等.濒危植物暗紫贝母的开花物候和传粉方式.草地学报,2017,25(6):1324-1332.
[17]Gao Y H,Ren Y Y,Sun H M,et al.Identification of self-incompatibility of Isatis indigotica.Forest By-Product and Speciality in China,2007,(6):11-15.高延辉,任跃英,孙红梅,等.菘蓝自交不亲和性鉴定.中国林副特产,2007,(6):11-15.
[18]Gao Y H,Gao B,Wang Y H,et al.Screening of the chemicals to resolve the problem of self pollination failure of Isatis indigotica.Forest By-Product and Speciality in China,2008,(1):12-15.高延辉,高博,王云贺,等.克服菘蓝自交不亲和性的化学试剂筛选研究.中国林副特产,2008,(1):12-15.
[19]Wang E J,Han D H,Zhang Y,et al.Effect of harvested periods on the yield and quality dynamic for the Isatis indigotica root in Hexi Corridor.Journal of Chinese Medicinal Materials,2016,39(12):2686-2690.王恩军,韩多红,张勇,等.采收期对河西走廊产板蓝根产量和品质的影响.中药材,2016,39(12):2686-2690.
[20]Hou G P,Zhen D S,Jiang Q L,et al.Discussion on environment of traditional Chinese medicine producing area in Minle county.Modern Agricultural Sciences and Technology,2017,(6):114-115.侯格平,甄东升,姜青龙,等.民乐县中药材产地环境探讨.现代农业科技,2017,(6):114-115.
[21]Wang E J,Chen Y,Han D H,et al.Effect of cropping pattern on agronomic characteristics,yield and quality of Radix isatidis.Chinese Journal of Eco-Agriculture,2017,25(11):1661-1670.王恩军,陈垣,韩多红,等.栽培方式对菘蓝农艺性状及产量和品质的影响.中国生态农业学报,2017,25(11):1661-1670.
[22]Wang X J,Zhu L Q.Method for determining self-incompatibility of Brassica oleracea L.Journal of Agricultural Biotechnology,1998,6(2):195-198.王晓佳,朱利泉.甘蓝自交不亲和性的测定方法.农业生物技术学报,1998,6(2):195-198.
[23]Zhou Y L.Plant biology.Beijing:Higher Education Press,2000:241-243.周云龙.植物生物学.北京:高等教育出版社,2000:241-243.
[24]Dafni A.Pollination ecology.New York:Oxford University Press,1992:25-45.
[25]Wang Y H,Guo F X,Chen Y,et al.Studies on blossoming habit and seed filling characteristics of wild Aconitum sinomontanum.Acta Prataculturae Sinica,2013,22(4):76-82.王永辉,郭凤霞,陈垣,等.野生高乌头开花习性及种子灌浆特性研究.草业学报,2013,22(4):76-82.
[26]Shoichi K,Masuda J,Hayashi K.Life-history monographs of Japanese plants.10:Fritillaria koidzumiana,Ohwi(Liliaceae).Plant Species Biology,2008,23(1):51-57.
[27]Ren Y Y.Breeding of medicinal plants.Beijing:China Agriculture Press,2011:17-18.任跃英.药用植物育种学.北京:中国农业出版社,2011:17-18.
[28]Hu C,Liu Z J,Wu G Q,et al.Floral characteristic and breeding system of Anemone obtusiloba.Acta Agrestia Sinica,2013,21(4):783-788.胡春,刘左军,伍国强,等.钝裂银莲花花部综合特征及其繁育系统.草地学报,2013,21(4):783-788.