激光共聚焦显微技术在瓮安生物群中的应用
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摘要
激光共聚焦显微技术是一种以激光作为激发光源,通过特殊装置"针孔"来过滤离焦光线以提高光学分辨率和对比度的光学成像技术。由于大部分化石不能自发荧光,该技术在古生物学领域尚未实现大范围的应用。但若围岩能自发荧光而与化石之间具有一定衬度,或化石因含特殊成分能在特定波段激光照射下自发荧光而产生结构衬度,则可以运用激光共聚焦显微技术获得在普通光学显微镜及荧光显微镜下难以清晰观察到的信息。为推动激光共聚焦技术在古生物学领域中的应用,文中系统介绍了该技术的原理与使用方法,并以埃迪卡拉纪磷酸盐化特异埋藏的瓮安生物群微体化石为例,展示了该技术在化石成像中的若干优势。实验结果表明,瓮安生物群微体化石因富含磷灰石可自发荧光实现成像,使用激光共聚焦显微成像技术观察瓮安生物群化石薄片不仅可以获得较好衬度,而且还能提高成像的分辨率和清晰度。此外,在化石薄片的厚度范围内还可以实现化石结构三维重建。
Confocal Laser Scanning Microscopy(CLSM) is a fluorescence imaging technique using laser and pinhole to obtain images with higher resolution and better contrast compared to conventional optical microscopy. Although CLSM is a powerful tool applied in many fields such as biology, it has not been widely used in palaeontology yet, because not all fossils are auto-fluorescent. However, if fossil matrix could fluoresce excited by laser beam, sharp contrast between fossils and matrix, or between different parts of fossils may be observed. In these cases, researchers can image inside microstructures with CLSM which can't be clearly visualized by epifluorescent microscope. In this article, we introduced the principle and work flow of sample preparation and imaged phosphatized microfossils from the Ediacaran Weng'an biota with CLSM. Our results suggest that the CLSM can help to obtain images with higher spatial resolution and better contrast in several cases.
Confocal Laser Scanning Microscopy(CLSM) is a fluorescence imaging technique using laser and pinhole to obtain images with higher resolution and better contrast compared to conventional optical microscopy. Although CLSM is a powerful tool applied in many fields such as biology, it has not been widely used in palaeontology yet, because not all fossils are auto-fluorescent. However, if fossil matrix could fluoresce excited by laser beam, sharp contrast between fossils and matrix, or between different parts of fossils may be observed. In these cases, researchers can image inside microstructures with CLSM which can't be clearly visualized by epifluorescent microscope. In this article, we introduced the principle and work flow of sample preparation and imaged phosphatized microfossils from the Ediacaran Weng'an biota with CLSM. Our results suggest that the CLSM can help to obtain images with higher spatial resolution and better contrast in several cases.
引文
An Yu-ying (安毓英),2002.Chapter two:Common laser.Laser & Infrared (激光与红外),32(3):201-205 (in Chinese).
Birkmann H,Lundin R F,1996.Confocal microscopy:Potential applications in micropaleontology.Journal of Paleontology,70(6):1084-1087.
Cai Chen-yang,Huang Di-ying,2014.The oldest micropepline beetle from Cretaceous Burmese amber and its phylogenetic implications (Coleoptera:Staphylinidae).Naturwissenschaften,101:813-817.
Chen Jun-yuan (陈均远),2004.The Dawn of Animal World.Nanjing:Jiangsu Science and Technology Publishing House.1-366 (in Chinese).
Chen Jun-yuan,Bottjer D J,Davidson E H,Dornbos S Q,Gao Xiang,Yang Yong-hua,Li Chia-wei,Li Gang,Wang Xiu-qiang,Xian Ding-chang,Wu Hung-jen.Hwu Yeu-kuang,Tafforeau P,2006.Phosphatized polar lobe-forming embryos from the Precambrian of southwest China.Science,312(5780):1644-1646.
Chen Jun-yuan,Bottjer D J,Davidson E H,Li Gang,Gao Feng,Cameron R A,Hadfield M G,Xian Ding-chang,Tafforeau P,Jia Quan-jie,2009.Phase contrast synchrotron X-ray microtomography of Ediacaran (Doushantuo) metazoan microfossils:Phylogenetic diversity and evolutionary implications.Precambrian Research,173:191-200.
Chen Jun-yuan,Bottjer D J,Oliveri P,Dornbos S Q,Gao Feng,Ruffins S,Chi Hui-mei,Li Chia-wei,Davidson E H,2004.Small bilaterian fossils from 40 to 55 Million Years before the Cambrian.Science,305(5681):218-222.
Chen Jun-yuan,Schopf J W,Bottjer D J,Zhang Chen-yu,Kudryavtsev A B,Tripathi A B,Wang Xiu-qiang,Yang Yong-hua,Gao Xiang,Yang Ying,2007.Raman spectra of a Lower Cambrian ctenophore embryo from southwestern Shaanxi,China.Proceedings of the National Academy of Sciences of the United States of America,104(15):6289-6292.
Chen Meng-e (陈孟莪),Liu Kui-wu (刘魁梧),1986.The geological significance of newly discovered microfossils from the Upper Sinian (Doushantuo Age) Phosphorites.Scientia Geologica Sinica (地质科学),21(1):46-53 (in Chinese with English abstract).
Chen Shang-bin (陈尚斌),Zuo Zhao-xi (左兆喜),Zhu Yan-ming (朱炎铭),Fu Chang-qing (付常青),Zhang Han (张寒),2015.Applicability of the testing method for the maturity of organic matter in shale gas reservoirs.Natural Gas Geoscience (天然气地球科学),26(3):564-574 (in Chinese with English abstract).
Chen Yu-ming (陈育明),Liu Yang (刘洋),2011.Status and research advances of metal halide lamp.China Light & Lighting (中国照明电器),2011(4):1-5 (in Chinese with English abstract).
Chi Hui-mei,Feng Man,Xiao Zhong-dang,Lu Zu-hong,2008.Preservation and fluorescence of the microfossils from Neoproterozoic Doushantuo Formation.Microscopy Research and Technique,71:260-266.
Chi Hui-mei,Xiao Zhong-dang,Fu De-gang,Lu Zu-hong,2006.Analysis of fluorescence from algae fossils of the Neoproterozoic Doushantuo formation of China by confocal laser scanning microscope.Microscopy Research and Technique,69(4):253-259.
Davidovits P,Egger M D,1969.Scanning Laser Microscope.Nature,223(5208):831.
Dobretsov M,Hayar A,Kockara N T,Kozhemyakin M,Light K E,Patyal P,Pierce D R,Wight P A,2019.A transgenic mouse model to selectively identify α3 Na,K-ATPase expressing cells in the nervous system.Neuroscience,398:274-294.
García-Plazaolaa J I,Fernández-Marína B,Dukec S O,Hernándeza A,López-Arbeload F,Becerril J M,2015.Autofluorescence:Biological functions and technical applications.Plant Science,236:136-145.
Guo Zi-xiao,Peng Xiao-tong,Czaja A D,Chen Shun,Ta Kai-wen,2018.Cellular taphonomy of well-preserved Gaoyuzhuang microfossils:A window into the preservation of ancient cyanobacteria.Precambrian Research,304:88-98.
Hickey P C,Swift S R,Roca M G,Read N D,2004.Live-cell imaging of filamentous fungi using vital fluorescent dyes and confocal microscopy.Methods in Microbiology,34(4):63-87.
Hochuli P A,Feist-Burkhardt S,2004.A boreal early cradle of Angiosperms?Angiosperm-like pollen from the Middle Triassic of the Barents Sea (Norway).Journal of Micropalaeontology,23:97-104.
Huang Di-ying,Bechly G,Nel P,Engel M S,Prokop J,Azar D,Cai Chen-yang,Kamp T,Staniczek A H,Garrouste R,Krogmann L,Rolo T S,Baumbach T,Ohlhoff R,Shmakov A S,Bourgoin T,Nel A,2016.New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda:Acercaria).Scientific Reports,6(6):23004.
Jackson F D,Horner J R,Varricchio D J,2010.A study of a Troodon egg containing embryonic remains using epifluorescence microscopy and other techniques.Cretaceous Research,31:255-262.
Leszczuk A,Szczuka E,2018.A study of a Troodon egg containing embryonic remains using epifluorescence microscopy and other techniques.Plant Physiology and Biochemistry,123:24-33.
Li Yong,Guo Jun-feng,Zhang Xing-liang,Zhang Wan-qian,Liu Yun-huan,Yang Wei-xian,Li Yuan-yuan,Liu Qian-qun,Shu De-gan,2008.Vase-shaped microfossils from the Ediacaran Weng’an biota,Guizhou,South China.Gondwana Research,14(1):263-268.
Lin Rui,Davis A,1988.A fluorogeochemical model for coal macerals.Organic Geochemistry,12(4):363-374.
Liu Peng-ju,Xiao Shu-hai,Yin Cong-yu,Zhou Chuan-ming,Gao Lin-zhi,Tang Feng,2010.Systematic description and phylogenetic affinity of tubular microfossils from the Ediacaran Doushantuo Formation at Weng’an,South China.Palaeontology,51(2):339-366.
Minsky M,1988.Memoir on inventing the confocal scanning microscope.Scanning,10(4):128-138.
Nix T,Feist-Burkhardt S,2003.New methods applied to the microstructure analysis of Messel oil shale:confocal laser scanning microscopy (CLSM) and environmental scanning electron microscopy (ESEM).Geological Magazine,140(4):469-478.
Pradier B,Bertrand P,Martinez L,Laggoun-Defagre F,1991.Fluorescence of organic matter and thermal maturity assessment.Organic Geochemistry,17(4):511-524.
Schopf J W,Kudryavtsev A B,Tripathi A B,Czaja A D,2010.Three-Dimensional Morphological (CLSM) and Chemical (Raman) Imagery of Cellularly Mineralized Fossils.Taphonomy:process and bias through time.In:Allison P A,Bottjer D J (ed.),Aims & Scope Topics in Geobiology Book.Springer:Heidelberg.Springer:Heidelberg.457-486.
Schopf J W,Sergeev V N,Kudryavtsev A B,2015.A new approach to ancient microorganisms:Taxonomy,paleoecology,and biostratigraphy of the Lower Cambrian Berkuta and Chulaktau microbiotas of South Kazakhstan.Journal of Paleontology,89(5):695-729.
Schopf J W,Tewari V C,Kudryavtsev A B,2008.Discovery of a new chert-permineralized microbiota in the Proterozoic Buxa Formation of the Ranjit Window,Sikkim,Northeast India,and its astobiological implications.Astrobiology,8(4):735-746.
Schopf J W,Tripathi A B,Kudryavtsev A B,2006.Three-dimensional confocal optical imagery of Precambrian Microscopic Organisms.Astrobiology,6(1):1-16.
Scott A C,Hemsley A R,1991.A comparison of new microscopical techniques for the study of fossils spore wall ultrastructure.Review of Palaeobotany and Palynology,67:133-139.
Talyzina N M,1998.Fluorescence intensity in Early Cambrian acritarchs from Estonia.Review of Palaeobotany and Palynology,100:99-108.
Wang Jin-xing (王金星),2004.Applications of a new method of confocal microsystem to geological studies.Geological Review (地质论评),50(2):215-217 (in Chinese with English abstract).
Xiao Shu-hai,Knoll A H,Yuan Xun-lai,Pueschel C M,2004.Phosphatized multicellular algae in the Neoproterozoic Doushantuo Formation,China,and the early evolution of florideophyte red algae.American Journal of Botany,91(2):214-227.
Xiao Shu-hai,Zhang Yun,Knoll A H,1998.Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorites.Nature,391(6667):553-558.
Yang Wei-ping (杨伟平),Zhang Hai-chun (张海春),Wang Bing (王冰),Xu Fang-ming (徐放鸣),1996.A new technique for microfossil study by Laser Scanning Confocal Imaging System.Acta Palaeontologica Sinica (古生物学报),35(6):730-733 (in Chinese with English abstract).
Yang Wei-ping (杨伟平),Zhang Hai-chun (张海春),Xu Fang-ming (徐放鸣),1998.Advanced research of ultra micro-structure of Middle Permian Acritarch Micrhystridium,West of Yunnan Province.Chinese Science Bulletin (科学通报),43(2):206-208 (in Chinese).
Yin Lei-ming (尹磊明),Bian Li-zeng (边立曾),Yuan Xun-lai (袁训来),2003.The discovery of branched tubules and microcircular spiral thickened tubules of the Ruyang Group in Middle Proterozoic,Shanxi Province.Scientia Sinica (中国科学),33(8):769-774 (in Chinese).
Yin Zong-jun,Zhu Mao-yan,Bottjer D J,Zhao Fang-chen,Tafforeau P,2016.Meroblastic cleavage identifies some Ediacaran Doushantuo (China) embryo-like fossils as metazoans.Geology,44(9):735-738.
Yin Zong-jun,Zhu Mao-yan,Davidson E H,Bottjer D J,Zhao Fang-chen,Tafforeau P,2015.Sponge grade body fossil with cellular resolution dating 60 Myr before the Cambrian.Proceedings of the National Academy of Sciences of the United States of America,112(12):E1453.
Yin Zong-jun,Zhu Mao-yan,Tafforeau P,Chen Jun-yuan,Liu Peng-ju,Li Gang,2013.Early embryogenesis of potential bilaterian animals with polar lobe formation from the Ediacaran Weng’an Biota,South China.Precambrian Research,225(1):44-57.
Yuan Xun-lai,Xiao Shu-hai,Taylor T N,2005.Lichen-like symbiosis 600 million years ago.Science,308(5724):1017-1020.
Yuan Xun-lai (袁训来),Xiao Shu-hai (肖书海),Yin Lei-ming (尹磊明),Knoll A H,Zhou Chuan-ming (周传明),Mu Xi-nan (穆西南),2002.Doushantuo Fossils:Life on the Eve of Animal Radiation.Hefei:University of Science and Technology of China Press.1-93.
Zhang Yun,YuanXun-lai,Yin Lei-ming,1998.Interpreting Late Precambrian Microfossils.Science,282(5395):1783.
Zhou Chuan-ming,Li Xian-hua,Xiao Shu-hai,Lan Zhong-wu,Ouyang Qing,Guan Cheng-guo,Chen Zhe,2017.A new SIMS zircon U–Pb date from the Ediacaran Doushantuo Formation:age constraint on the Weng'an biota.Geological Magazine,154(6):1193-1207.
Zhu Mao-yan,Zhang Jun-ming,Yang Ai-hua,2007.Integrated Ediacaran (Sinian) chronostratigraphy of South China.Palaeogeography,Palaeoclimatology,Palaeoecology,254:7-61.
Zhuo Er-jun (卓二军),Tang Ling-yu (唐领余),Zhang Hai-chun (张海春),2006.Application of Laser Scanning Confocal Microscope in fossil spore pollen observation.Acta Palaeontologica Sinica (古生物学报),45(3):430-436 (in Chinese with English abstract).
Birkmann H,Lundin R F,1996.Confocal microscopy:Potential applications in micropaleontology.Journal of Paleontology,70(6):1084-1087.
Cai Chen-yang,Huang Di-ying,2014.The oldest micropepline beetle from Cretaceous Burmese amber and its phylogenetic implications (Coleoptera:Staphylinidae).Naturwissenschaften,101:813-817.
Chen Jun-yuan (陈均远),2004.The Dawn of Animal World.Nanjing:Jiangsu Science and Technology Publishing House.1-366 (in Chinese).
Chen Jun-yuan,Bottjer D J,Davidson E H,Dornbos S Q,Gao Xiang,Yang Yong-hua,Li Chia-wei,Li Gang,Wang Xiu-qiang,Xian Ding-chang,Wu Hung-jen.Hwu Yeu-kuang,Tafforeau P,2006.Phosphatized polar lobe-forming embryos from the Precambrian of southwest China.Science,312(5780):1644-1646.
Chen Jun-yuan,Bottjer D J,Davidson E H,Li Gang,Gao Feng,Cameron R A,Hadfield M G,Xian Ding-chang,Tafforeau P,Jia Quan-jie,2009.Phase contrast synchrotron X-ray microtomography of Ediacaran (Doushantuo) metazoan microfossils:Phylogenetic diversity and evolutionary implications.Precambrian Research,173:191-200.
Chen Jun-yuan,Bottjer D J,Oliveri P,Dornbos S Q,Gao Feng,Ruffins S,Chi Hui-mei,Li Chia-wei,Davidson E H,2004.Small bilaterian fossils from 40 to 55 Million Years before the Cambrian.Science,305(5681):218-222.
Chen Jun-yuan,Schopf J W,Bottjer D J,Zhang Chen-yu,Kudryavtsev A B,Tripathi A B,Wang Xiu-qiang,Yang Yong-hua,Gao Xiang,Yang Ying,2007.Raman spectra of a Lower Cambrian ctenophore embryo from southwestern Shaanxi,China.Proceedings of the National Academy of Sciences of the United States of America,104(15):6289-6292.
Chen Meng-e (陈孟莪),Liu Kui-wu (刘魁梧),1986.The geological significance of newly discovered microfossils from the Upper Sinian (Doushantuo Age) Phosphorites.Scientia Geologica Sinica (地质科学),21(1):46-53 (in Chinese with English abstract).
Chen Shang-bin (陈尚斌),Zuo Zhao-xi (左兆喜),Zhu Yan-ming (朱炎铭),Fu Chang-qing (付常青),Zhang Han (张寒),2015.Applicability of the testing method for the maturity of organic matter in shale gas reservoirs.Natural Gas Geoscience (天然气地球科学),26(3):564-574 (in Chinese with English abstract).
Chen Yu-ming (陈育明),Liu Yang (刘洋),2011.Status and research advances of metal halide lamp.China Light & Lighting (中国照明电器),2011(4):1-5 (in Chinese with English abstract).
Chi Hui-mei,Feng Man,Xiao Zhong-dang,Lu Zu-hong,2008.Preservation and fluorescence of the microfossils from Neoproterozoic Doushantuo Formation.Microscopy Research and Technique,71:260-266.
Chi Hui-mei,Xiao Zhong-dang,Fu De-gang,Lu Zu-hong,2006.Analysis of fluorescence from algae fossils of the Neoproterozoic Doushantuo formation of China by confocal laser scanning microscope.Microscopy Research and Technique,69(4):253-259.
Davidovits P,Egger M D,1969.Scanning Laser Microscope.Nature,223(5208):831.
Dobretsov M,Hayar A,Kockara N T,Kozhemyakin M,Light K E,Patyal P,Pierce D R,Wight P A,2019.A transgenic mouse model to selectively identify α3 Na,K-ATPase expressing cells in the nervous system.Neuroscience,398:274-294.
García-Plazaolaa J I,Fernández-Marína B,Dukec S O,Hernándeza A,López-Arbeload F,Becerril J M,2015.Autofluorescence:Biological functions and technical applications.Plant Science,236:136-145.
Guo Zi-xiao,Peng Xiao-tong,Czaja A D,Chen Shun,Ta Kai-wen,2018.Cellular taphonomy of well-preserved Gaoyuzhuang microfossils:A window into the preservation of ancient cyanobacteria.Precambrian Research,304:88-98.
Hickey P C,Swift S R,Roca M G,Read N D,2004.Live-cell imaging of filamentous fungi using vital fluorescent dyes and confocal microscopy.Methods in Microbiology,34(4):63-87.
Hochuli P A,Feist-Burkhardt S,2004.A boreal early cradle of Angiosperms?Angiosperm-like pollen from the Middle Triassic of the Barents Sea (Norway).Journal of Micropalaeontology,23:97-104.
Huang Di-ying,Bechly G,Nel P,Engel M S,Prokop J,Azar D,Cai Chen-yang,Kamp T,Staniczek A H,Garrouste R,Krogmann L,Rolo T S,Baumbach T,Ohlhoff R,Shmakov A S,Bourgoin T,Nel A,2016.New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda:Acercaria).Scientific Reports,6(6):23004.
Jackson F D,Horner J R,Varricchio D J,2010.A study of a Troodon egg containing embryonic remains using epifluorescence microscopy and other techniques.Cretaceous Research,31:255-262.
Leszczuk A,Szczuka E,2018.A study of a Troodon egg containing embryonic remains using epifluorescence microscopy and other techniques.Plant Physiology and Biochemistry,123:24-33.
Li Yong,Guo Jun-feng,Zhang Xing-liang,Zhang Wan-qian,Liu Yun-huan,Yang Wei-xian,Li Yuan-yuan,Liu Qian-qun,Shu De-gan,2008.Vase-shaped microfossils from the Ediacaran Weng’an biota,Guizhou,South China.Gondwana Research,14(1):263-268.
Lin Rui,Davis A,1988.A fluorogeochemical model for coal macerals.Organic Geochemistry,12(4):363-374.
Liu Peng-ju,Xiao Shu-hai,Yin Cong-yu,Zhou Chuan-ming,Gao Lin-zhi,Tang Feng,2010.Systematic description and phylogenetic affinity of tubular microfossils from the Ediacaran Doushantuo Formation at Weng’an,South China.Palaeontology,51(2):339-366.
Minsky M,1988.Memoir on inventing the confocal scanning microscope.Scanning,10(4):128-138.
Nix T,Feist-Burkhardt S,2003.New methods applied to the microstructure analysis of Messel oil shale:confocal laser scanning microscopy (CLSM) and environmental scanning electron microscopy (ESEM).Geological Magazine,140(4):469-478.
Pradier B,Bertrand P,Martinez L,Laggoun-Defagre F,1991.Fluorescence of organic matter and thermal maturity assessment.Organic Geochemistry,17(4):511-524.
Schopf J W,Kudryavtsev A B,Tripathi A B,Czaja A D,2010.Three-Dimensional Morphological (CLSM) and Chemical (Raman) Imagery of Cellularly Mineralized Fossils.Taphonomy:process and bias through time.In:Allison P A,Bottjer D J (ed.),Aims & Scope Topics in Geobiology Book.Springer:Heidelberg.Springer:Heidelberg.457-486.
Schopf J W,Sergeev V N,Kudryavtsev A B,2015.A new approach to ancient microorganisms:Taxonomy,paleoecology,and biostratigraphy of the Lower Cambrian Berkuta and Chulaktau microbiotas of South Kazakhstan.Journal of Paleontology,89(5):695-729.
Schopf J W,Tewari V C,Kudryavtsev A B,2008.Discovery of a new chert-permineralized microbiota in the Proterozoic Buxa Formation of the Ranjit Window,Sikkim,Northeast India,and its astobiological implications.Astrobiology,8(4):735-746.
Schopf J W,Tripathi A B,Kudryavtsev A B,2006.Three-dimensional confocal optical imagery of Precambrian Microscopic Organisms.Astrobiology,6(1):1-16.
Scott A C,Hemsley A R,1991.A comparison of new microscopical techniques for the study of fossils spore wall ultrastructure.Review of Palaeobotany and Palynology,67:133-139.
Talyzina N M,1998.Fluorescence intensity in Early Cambrian acritarchs from Estonia.Review of Palaeobotany and Palynology,100:99-108.
Wang Jin-xing (王金星),2004.Applications of a new method of confocal microsystem to geological studies.Geological Review (地质论评),50(2):215-217 (in Chinese with English abstract).
Xiao Shu-hai,Knoll A H,Yuan Xun-lai,Pueschel C M,2004.Phosphatized multicellular algae in the Neoproterozoic Doushantuo Formation,China,and the early evolution of florideophyte red algae.American Journal of Botany,91(2):214-227.
Xiao Shu-hai,Zhang Yun,Knoll A H,1998.Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorites.Nature,391(6667):553-558.
Yang Wei-ping (杨伟平),Zhang Hai-chun (张海春),Wang Bing (王冰),Xu Fang-ming (徐放鸣),1996.A new technique for microfossil study by Laser Scanning Confocal Imaging System.Acta Palaeontologica Sinica (古生物学报),35(6):730-733 (in Chinese with English abstract).
Yang Wei-ping (杨伟平),Zhang Hai-chun (张海春),Xu Fang-ming (徐放鸣),1998.Advanced research of ultra micro-structure of Middle Permian Acritarch Micrhystridium,West of Yunnan Province.Chinese Science Bulletin (科学通报),43(2):206-208 (in Chinese).
Yin Lei-ming (尹磊明),Bian Li-zeng (边立曾),Yuan Xun-lai (袁训来),2003.The discovery of branched tubules and microcircular spiral thickened tubules of the Ruyang Group in Middle Proterozoic,Shanxi Province.Scientia Sinica (中国科学),33(8):769-774 (in Chinese).
Yin Zong-jun,Zhu Mao-yan,Bottjer D J,Zhao Fang-chen,Tafforeau P,2016.Meroblastic cleavage identifies some Ediacaran Doushantuo (China) embryo-like fossils as metazoans.Geology,44(9):735-738.
Yin Zong-jun,Zhu Mao-yan,Davidson E H,Bottjer D J,Zhao Fang-chen,Tafforeau P,2015.Sponge grade body fossil with cellular resolution dating 60 Myr before the Cambrian.Proceedings of the National Academy of Sciences of the United States of America,112(12):E1453.
Yin Zong-jun,Zhu Mao-yan,Tafforeau P,Chen Jun-yuan,Liu Peng-ju,Li Gang,2013.Early embryogenesis of potential bilaterian animals with polar lobe formation from the Ediacaran Weng’an Biota,South China.Precambrian Research,225(1):44-57.
Yuan Xun-lai,Xiao Shu-hai,Taylor T N,2005.Lichen-like symbiosis 600 million years ago.Science,308(5724):1017-1020.
Yuan Xun-lai (袁训来),Xiao Shu-hai (肖书海),Yin Lei-ming (尹磊明),Knoll A H,Zhou Chuan-ming (周传明),Mu Xi-nan (穆西南),2002.Doushantuo Fossils:Life on the Eve of Animal Radiation.Hefei:University of Science and Technology of China Press.1-93.
Zhang Yun,YuanXun-lai,Yin Lei-ming,1998.Interpreting Late Precambrian Microfossils.Science,282(5395):1783.
Zhou Chuan-ming,Li Xian-hua,Xiao Shu-hai,Lan Zhong-wu,Ouyang Qing,Guan Cheng-guo,Chen Zhe,2017.A new SIMS zircon U–Pb date from the Ediacaran Doushantuo Formation:age constraint on the Weng'an biota.Geological Magazine,154(6):1193-1207.
Zhu Mao-yan,Zhang Jun-ming,Yang Ai-hua,2007.Integrated Ediacaran (Sinian) chronostratigraphy of South China.Palaeogeography,Palaeoclimatology,Palaeoecology,254:7-61.
Zhuo Er-jun (卓二军),Tang Ling-yu (唐领余),Zhang Hai-chun (张海春),2006.Application of Laser Scanning Confocal Microscope in fossil spore pollen observation.Acta Palaeontologica Sinica (古生物学报),45(3):430-436 (in Chinese with English abstract).
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