菰适应湿地环境的解剖和屏障结构特征研究
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摘要
菰(Zizania latifolia)是一种多年生挺水植物,为了探讨该植物根、茎和叶的解剖结构、组织化学及其质外体屏障的通透性生理。该文利用光学显微镜和荧光显微镜,对菰的根、茎、叶进行了解剖学和组织化学研究。结果表明:(1)菰不定根解剖结构由外而内分别为表皮、外皮层、单层细胞的厚壁机械组织层、皮层、内皮层和维管柱;茎结构由外而内分别为角质层、表皮、周缘厚壁机械组织层、皮层、具维管束的厚壁组织层和髓腔。叶鞘具有表皮和具维管束皮层,叶片具有表皮,叶肉和维管束。(2)不定根具有位于内侧的内皮层及其邻近栓质化细胞和外侧的外皮层组成的屏障结构;茎具内侧厚壁机械组织层,外侧的角质层和周缘厚壁机械组织层组成的屏障结构,屏障结构的细胞壁具凯氏带、木栓质和木质素沉积的组织化学特点,叶表面具有角质层。(3)菰通气组织包括根中通气组织,茎、叶皮层的通气组织和髓腔。(4)菰的屏障结构和解剖结构是其适应湿地环境的重要特征,但其茎周缘厚壁层和厚壁组织层较薄。由此推测,菰适应湿地环境,但在旱生环境中分布有一定的局限性。
Wild rice( Zizania latifolia) is a famous,perennial,emergent vegetable in China. The current work explores the anatomy and histochemistry of roots,stems and leaves and the permeability of apoplastic barriers of wild rice. The anatomy and histochemistry of Z. latifolia were studied by optical microscope and fluorescence microscope. Sections were stained with Sudan red 7 B for suberin lamellae,berberine hemisulfate-aniline blue for Casparian bands and lignified walls,and phloroglucinol-HCl( Pg) for lignin. The results were as follows:( 1) The adventitious roots in wild rice suberized and lignified endodermis and adjacent,thick-walled cortical layers and suberized and lignified hypodermis,composed of a uniseriate sclerenchyma layer underlying uniseriate exodermis; The stems of wild plants included stolons,rhizomes,and culm. Rhizomes,stolons,and culms had two rings of thickened,lignified cells,the peripheral mechanical ring and the sclerenchyma ring; The latter delimits the cortex from the CC( central cylinder) and was usually associated with vascular bundles; Stems had thick epidermal cuticle,a narrow peripheral mechanical ring,cortex,sclerenchyma ring with vascular bundles and pith cavity from the outside and inside. Leaf sheaths had epidermis and cortex with vascular bundles,and leaf blades had epidermis,mesophyll and vascular bundles.( 2) Apoplastic barriers were found in roots and stems. The apoplastic barriers consist of that adventitious roots had endodermis,adjacent suberized cells and exodermis; Stems have cuticle,suberized and lignified peripheral mechanical ring and sclerenchyma ring,and the cell wall of apoplastic barriers had Casparian band,lignin and suberin. Leaves had cuticles at surface.( 3) The air space consist of aerenchyma in roots,and pith cavities and aerenchyma in stems and leaves.( 4) The peripheral mechanical ring and sclerenchyma ring were thinner in wild rice stems that adapt to wetland environment,but limited distribution to drought environment.
Wild rice( Zizania latifolia) is a famous,perennial,emergent vegetable in China. The current work explores the anatomy and histochemistry of roots,stems and leaves and the permeability of apoplastic barriers of wild rice. The anatomy and histochemistry of Z. latifolia were studied by optical microscope and fluorescence microscope. Sections were stained with Sudan red 7 B for suberin lamellae,berberine hemisulfate-aniline blue for Casparian bands and lignified walls,and phloroglucinol-HCl( Pg) for lignin. The results were as follows:( 1) The adventitious roots in wild rice suberized and lignified endodermis and adjacent,thick-walled cortical layers and suberized and lignified hypodermis,composed of a uniseriate sclerenchyma layer underlying uniseriate exodermis; The stems of wild plants included stolons,rhizomes,and culm. Rhizomes,stolons,and culms had two rings of thickened,lignified cells,the peripheral mechanical ring and the sclerenchyma ring; The latter delimits the cortex from the CC( central cylinder) and was usually associated with vascular bundles; Stems had thick epidermal cuticle,a narrow peripheral mechanical ring,cortex,sclerenchyma ring with vascular bundles and pith cavity from the outside and inside. Leaf sheaths had epidermis and cortex with vascular bundles,and leaf blades had epidermis,mesophyll and vascular bundles.( 2) Apoplastic barriers were found in roots and stems. The apoplastic barriers consist of that adventitious roots had endodermis,adjacent suberized cells and exodermis; Stems have cuticle,suberized and lignified peripheral mechanical ring and sclerenchyma ring,and the cell wall of apoplastic barriers had Casparian band,lignin and suberin. Leaves had cuticles at surface.( 3) The air space consist of aerenchyma in roots,and pith cavities and aerenchyma in stems and leaves.( 4) The peripheral mechanical ring and sclerenchyma ring were thinner in wild rice stems that adapt to wetland environment,but limited distribution to drought environment.
引文
ARMSTRONG W,COUSINS D,ARMSTRONG J,et al.,2000.Oxygen distribution in wetland plant roots and permeability barriers togas-exchange with the rhizosphere:A microelectrode and modelling study with Phragmites australis[J].Ann Bot,86(3):687-703.
ARMSTRONG J,JONES RE,ARMSTRONG W,2006.Rhizome phyllosphere oxygenation in Phragmites and other species in relation to redox potential,convective gas flow,submergence and aeration pathways[J].New Phytol,172(4):719-731.
BRUNDRETT MC,ENSTONE DE,PETERSON CA,1988.Aberberine-aniline blue fluorescent staining procedure for suberin,lignin,and callose in plant tissue[J].Protoplasma,146(2-3):133-142.
BRUNDRETT MC,KENDRICK B,PETERSON CA,1991.Efficient lipid staining in plant material with Sudan Red 7B or fluoral yellow 088 in polyethylene glycol-glycerol[J].Biotechnol Histochem,66(3):111-116.
COLMER TD,2003.Aerenchyma and an inducible barrier to radial oxygen loss facilitate root aeration in upland,paddy and deep-water rice(Oryza sativa L.)[J].Ann Bot,91(2):301-309.
ENSTONE DE,PETERSON CA,MA F,2002.Root endodermis and exodermis structure,function,and responses to the environment[J].J Plant Growth Regul,21(4):335-351.
EVERT RF,RUSSIN WA,BOSABALIDIS AM,1996.Anatomical and ultrastructural changes associated with sink-tosource transition in developing maize leaves[J].Int J Plant Sci,157(3):247-261.
FAHN A,1990.Plant Anatomy[M].Oxford,UK:Pergamon Press:1-544.
JENSEN WA,1962.Botanical histochemistry-principles and practice[M].W.H.Freeman,San Francisco,Calif,USA:1-408.
JORGENSON KD,LEE PE,KANAVILLIL K,2013.Ecological relationships of wild rice,Zizania spp.11.Electron microscopy study of iron plaques on the roots of northern wild rice(Zizania palustris)[J].Bot,91(3):189-201.
JUDZIEWICZ EJ,CLARK LG,2007.Classification and biogeography of New World Grasses:Anomochlooideae,Pharoideae,Ehrhartoideae,and Bambusoideae[J]//COLUMBUS JT,FRIAR EA,PORTER JM,et al.Monocots:Comparative Biology and Evolution Poales.Claremont CA.USA:Rancho Santa Ana Bot Gard,23(1):303-314.
JUNG J,LEE SC,CHO HK,2008.Anatomical patterns of aerenchyma in aquatic and wetland plants[J].J Plant Biol,51(6):428-439.
KAWAI M,SAMARAJEEWA PK,BARRERO RA,1998.Cellular dissection of the degradation pattern of cortical cell death during aerenchyma formation of rice roots[J].Planta,204(3):277-287.
KOTULA L,RANTHUNGRE K,SCHREIBER L,et al.,2009.Functional and chemical comparison of apoplastic barriers to radial oxygen loss in roots of rice(Oryza sativa L.)grown in aerated or deoxygenated solution[J].J Exp Bot,60(7):2155-2167.
KRISHNAMURTHY P,RANATHUNGE K,NAYAK S,et al.,2011.Root apoplastic barriers block Na+transport to shoots in rice(Oryza sativa L.)[J].J Exp Bot,62(12):4215-4228.
MCMANUS HA,SEAGO JR JL,MARSH LC,2002.Epifluorescent and histochemical aspects of shoot anatomy of Typha latifolia L.,Typha angustifolia L.and Typha glauca Godr.[J].Ann Bot,90(4):489-493.
METCALFE CR,1960.Anatomy of the Monocotyledons.I.Gramineae[M].London,UK:Oxford University Press:1-731.
MEYER CJ,PETERSON CA,2011.Casparian bands occur in the periderm of Pelargonium hortorum stem and root[J].Ann Bot,107(4):591-598.
MEYER CJ,SEAGO JR JL,PETERSON CA,2009.Environmental effects on the maturation of the endodermis and multiseriate exodermis of Iris germanica roots[J].Ann Bot,103(5):687-702.
RANTHUNGRE K,LIN J,STEUDLE E,et al.,2011.Stagnant deoxygenated growth enhances root suberization and lignifications,but differentially affects water and Na Cl permeabilities in rice(Oryza sativa L.)roots[J].Plant Cell Environ,34(8):1223-1240.
SANGSTER AG,1985.Silicon distribution and anatomy of the grass rhizome,with special reference to Miscanthus sacchariflorus(Maxim.)Hackel[J].Ann Bot,55(5):621-634.
SCHREIBER L,FRANKE RB,2011.Endodermis and exodermis in roots[J].Chichester,UK:John Wiley&Sons,Ltd.http://www.els.net[doi:10.1002/9780470015902.a0002086.pub2.]
SCULTHORPE CD,1967.The biology of aquatic vascular plants[M].London,UK:Edward Arnold Ltd:1-610.
SEAGO JR JL,MARSH LC,STEVENS KJ,et al.,2005.A reexamination of the root cortex in wetland flowering plants with respect to aerenchyma[J].Ann Bot,96(4):565-579.
SEAGO JR JL,PETERSON CA,ENSTONE DE,et al.,1999.Development of the endodermis and hypodermis of Typha glauca Godr.and T.angustifolia L.roots[J].Can JBot,77(1):122-134.
SEAGO JR JL,PETERSON CA,ENSTONE DE,2000.Cortical development in roots of the aquatic plant Pontederia cordata(Pontederiaceae)[J].Am J Bot,87(8):1116-1127.
SIMONE OD,HAASE K,MLLER E,et al.,2003.Apoplastic barriers and oxygen transport properties of hypodermal cell walls in roots from four Amazonian tree species[J].Plant Physiol,132(1):206-217.
SOUKUP A,ARMSTRONG W,SCHREIBER L,et al.,2007.Apoplastic barriers to radial oxygen loss and solute penetration:A chemical and functional comparison of the exodermis of two wetland species,Phragmites australis and Glyceria maxima[J].New Phytol,173(2):264-278.
STOVER EL,1928.The roots of wild rice Zizania aquatica L.[J].Ohio J Sci,28(1):43-49.
STOVER EL,1951.An introduction to the anatomy of seed plants[J].Aibs Bull,1(2):10.
TAN SD,ZHU MY,DANG HS,et al.,2009.Physiological responses of bermudagrass(Cynodon dactylon L.Pers.)to deep submergence stress in the Three Gorges Reservoir Area[J].Acta Ecol Sin,29(7):3685-3692.[谭淑端,朱明勇,党海山,等,2009.三峡库区狗牙根对深淹胁迫的生理响应[J].生态学报,29(7):3685-3692.]
TATEOKA T,1969.Root anatomy in grass systematic[J].Bull Nat Sci Museum,12(3):643-651.
VARTAPETIAN BB,JACKSON MB,1997.Plant adaptations to anaerobic stress[J].Ann Bot,79(Suppl.1):3-20.
WANG HF,ZENG B,LI Y,et al.,2008.Effects of long-term submergence on survival and recovery growth of four riparian plant species in Three Gorges Reservoir region[J].Chin JPlant Ecol,32(5):977-984.[王海锋,曾波,李娅,等,2008.长期完全水淹对4种三峡库区岸生植物存活及恢复生长的影响[J].植物生态学报,32(5):977-984.]
WATANABE H,SAIGUSA M,MORITA S,2006.Identification of casparian bands in the mesocotyl and lower internodes of rice(Oryza sativa L.)seedlings using fluorescence microscopy[J].Plant Prod Sci,9(4):390-394.
YANG CD,LI SF,DENG SM,et al.,2015.Study of the anatomy and apoplastic barrier characteristics of Imperata cylindrica[J].Acta Pratac Sin,24(3):213-218.[杨朝东,李守峰,邓仕明,2015.白茅解剖结构和屏障结构特征研究[J].草业学报,24(3):213-218.]
YANG CD,ZHANG X,2013.The permeability and supplement structures in stems for Paspalum distichum[J].Bull Bot Res,33(5):564-568.[杨朝东,张霞,2013.双穗雀稗(Paspalum distichum)通透性生理和茎解剖结构补充研究[J].植物研究,33(5):564-568.]
YANG CD,ZHANG X,LIU GF,et al.,2013.Progress on the structure and phisiological function of apoplastic barriers in root[J].Bull Bot Res,33(1):114-119.[杨朝东,张霞,刘国锋,等,2013.植物根中质外体屏障结构和生理功能研究进展[J].植物研究,33(1):114-119.]
YANG CD,ZHANG X,ZHOU CY,2011.Root and stem anatomy and histochemistry of four grasses from the Jianghan Floodplain along the Yangtze River,China[J].Flora,206(7):653-661.
ZHANG X,YANG CD,NING GG,2013.The comparison development of apoplastic barriers in Cynodon dactylon and Paspalum distichum roots[J].Hubei Agric Sci,52(20):4991-4994.[张霞,杨朝东,宁国贵,2013.狗牙根和双穗雀稗根中质外体屏障结构发育过程的比较研究[J].湖北农业科学,52(20):4991-4994.]
ARMSTRONG J,JONES RE,ARMSTRONG W,2006.Rhizome phyllosphere oxygenation in Phragmites and other species in relation to redox potential,convective gas flow,submergence and aeration pathways[J].New Phytol,172(4):719-731.
BRUNDRETT MC,ENSTONE DE,PETERSON CA,1988.Aberberine-aniline blue fluorescent staining procedure for suberin,lignin,and callose in plant tissue[J].Protoplasma,146(2-3):133-142.
BRUNDRETT MC,KENDRICK B,PETERSON CA,1991.Efficient lipid staining in plant material with Sudan Red 7B or fluoral yellow 088 in polyethylene glycol-glycerol[J].Biotechnol Histochem,66(3):111-116.
COLMER TD,2003.Aerenchyma and an inducible barrier to radial oxygen loss facilitate root aeration in upland,paddy and deep-water rice(Oryza sativa L.)[J].Ann Bot,91(2):301-309.
ENSTONE DE,PETERSON CA,MA F,2002.Root endodermis and exodermis structure,function,and responses to the environment[J].J Plant Growth Regul,21(4):335-351.
EVERT RF,RUSSIN WA,BOSABALIDIS AM,1996.Anatomical and ultrastructural changes associated with sink-tosource transition in developing maize leaves[J].Int J Plant Sci,157(3):247-261.
FAHN A,1990.Plant Anatomy[M].Oxford,UK:Pergamon Press:1-544.
JENSEN WA,1962.Botanical histochemistry-principles and practice[M].W.H.Freeman,San Francisco,Calif,USA:1-408.
JORGENSON KD,LEE PE,KANAVILLIL K,2013.Ecological relationships of wild rice,Zizania spp.11.Electron microscopy study of iron plaques on the roots of northern wild rice(Zizania palustris)[J].Bot,91(3):189-201.
JUDZIEWICZ EJ,CLARK LG,2007.Classification and biogeography of New World Grasses:Anomochlooideae,Pharoideae,Ehrhartoideae,and Bambusoideae[J]//COLUMBUS JT,FRIAR EA,PORTER JM,et al.Monocots:Comparative Biology and Evolution Poales.Claremont CA.USA:Rancho Santa Ana Bot Gard,23(1):303-314.
JUNG J,LEE SC,CHO HK,2008.Anatomical patterns of aerenchyma in aquatic and wetland plants[J].J Plant Biol,51(6):428-439.
KAWAI M,SAMARAJEEWA PK,BARRERO RA,1998.Cellular dissection of the degradation pattern of cortical cell death during aerenchyma formation of rice roots[J].Planta,204(3):277-287.
KOTULA L,RANTHUNGRE K,SCHREIBER L,et al.,2009.Functional and chemical comparison of apoplastic barriers to radial oxygen loss in roots of rice(Oryza sativa L.)grown in aerated or deoxygenated solution[J].J Exp Bot,60(7):2155-2167.
KRISHNAMURTHY P,RANATHUNGE K,NAYAK S,et al.,2011.Root apoplastic barriers block Na+transport to shoots in rice(Oryza sativa L.)[J].J Exp Bot,62(12):4215-4228.
MCMANUS HA,SEAGO JR JL,MARSH LC,2002.Epifluorescent and histochemical aspects of shoot anatomy of Typha latifolia L.,Typha angustifolia L.and Typha glauca Godr.[J].Ann Bot,90(4):489-493.
METCALFE CR,1960.Anatomy of the Monocotyledons.I.Gramineae[M].London,UK:Oxford University Press:1-731.
MEYER CJ,PETERSON CA,2011.Casparian bands occur in the periderm of Pelargonium hortorum stem and root[J].Ann Bot,107(4):591-598.
MEYER CJ,SEAGO JR JL,PETERSON CA,2009.Environmental effects on the maturation of the endodermis and multiseriate exodermis of Iris germanica roots[J].Ann Bot,103(5):687-702.
RANTHUNGRE K,LIN J,STEUDLE E,et al.,2011.Stagnant deoxygenated growth enhances root suberization and lignifications,but differentially affects water and Na Cl permeabilities in rice(Oryza sativa L.)roots[J].Plant Cell Environ,34(8):1223-1240.
SANGSTER AG,1985.Silicon distribution and anatomy of the grass rhizome,with special reference to Miscanthus sacchariflorus(Maxim.)Hackel[J].Ann Bot,55(5):621-634.
SCHREIBER L,FRANKE RB,2011.Endodermis and exodermis in roots[J].Chichester,UK:John Wiley&Sons,Ltd.http://www.els.net[doi:10.1002/9780470015902.a0002086.pub2.]
SCULTHORPE CD,1967.The biology of aquatic vascular plants[M].London,UK:Edward Arnold Ltd:1-610.
SEAGO JR JL,MARSH LC,STEVENS KJ,et al.,2005.A reexamination of the root cortex in wetland flowering plants with respect to aerenchyma[J].Ann Bot,96(4):565-579.
SEAGO JR JL,PETERSON CA,ENSTONE DE,et al.,1999.Development of the endodermis and hypodermis of Typha glauca Godr.and T.angustifolia L.roots[J].Can JBot,77(1):122-134.
SEAGO JR JL,PETERSON CA,ENSTONE DE,2000.Cortical development in roots of the aquatic plant Pontederia cordata(Pontederiaceae)[J].Am J Bot,87(8):1116-1127.
SIMONE OD,HAASE K,MLLER E,et al.,2003.Apoplastic barriers and oxygen transport properties of hypodermal cell walls in roots from four Amazonian tree species[J].Plant Physiol,132(1):206-217.
SOUKUP A,ARMSTRONG W,SCHREIBER L,et al.,2007.Apoplastic barriers to radial oxygen loss and solute penetration:A chemical and functional comparison of the exodermis of two wetland species,Phragmites australis and Glyceria maxima[J].New Phytol,173(2):264-278.
STOVER EL,1928.The roots of wild rice Zizania aquatica L.[J].Ohio J Sci,28(1):43-49.
STOVER EL,1951.An introduction to the anatomy of seed plants[J].Aibs Bull,1(2):10.
TAN SD,ZHU MY,DANG HS,et al.,2009.Physiological responses of bermudagrass(Cynodon dactylon L.Pers.)to deep submergence stress in the Three Gorges Reservoir Area[J].Acta Ecol Sin,29(7):3685-3692.[谭淑端,朱明勇,党海山,等,2009.三峡库区狗牙根对深淹胁迫的生理响应[J].生态学报,29(7):3685-3692.]
TATEOKA T,1969.Root anatomy in grass systematic[J].Bull Nat Sci Museum,12(3):643-651.
VARTAPETIAN BB,JACKSON MB,1997.Plant adaptations to anaerobic stress[J].Ann Bot,79(Suppl.1):3-20.
WANG HF,ZENG B,LI Y,et al.,2008.Effects of long-term submergence on survival and recovery growth of four riparian plant species in Three Gorges Reservoir region[J].Chin JPlant Ecol,32(5):977-984.[王海锋,曾波,李娅,等,2008.长期完全水淹对4种三峡库区岸生植物存活及恢复生长的影响[J].植物生态学报,32(5):977-984.]
WATANABE H,SAIGUSA M,MORITA S,2006.Identification of casparian bands in the mesocotyl and lower internodes of rice(Oryza sativa L.)seedlings using fluorescence microscopy[J].Plant Prod Sci,9(4):390-394.
YANG CD,LI SF,DENG SM,et al.,2015.Study of the anatomy and apoplastic barrier characteristics of Imperata cylindrica[J].Acta Pratac Sin,24(3):213-218.[杨朝东,李守峰,邓仕明,2015.白茅解剖结构和屏障结构特征研究[J].草业学报,24(3):213-218.]
YANG CD,ZHANG X,2013.The permeability and supplement structures in stems for Paspalum distichum[J].Bull Bot Res,33(5):564-568.[杨朝东,张霞,2013.双穗雀稗(Paspalum distichum)通透性生理和茎解剖结构补充研究[J].植物研究,33(5):564-568.]
YANG CD,ZHANG X,LIU GF,et al.,2013.Progress on the structure and phisiological function of apoplastic barriers in root[J].Bull Bot Res,33(1):114-119.[杨朝东,张霞,刘国锋,等,2013.植物根中质外体屏障结构和生理功能研究进展[J].植物研究,33(1):114-119.]
YANG CD,ZHANG X,ZHOU CY,2011.Root and stem anatomy and histochemistry of four grasses from the Jianghan Floodplain along the Yangtze River,China[J].Flora,206(7):653-661.
ZHANG X,YANG CD,NING GG,2013.The comparison development of apoplastic barriers in Cynodon dactylon and Paspalum distichum roots[J].Hubei Agric Sci,52(20):4991-4994.[张霞,杨朝东,宁国贵,2013.狗牙根和双穗雀稗根中质外体屏障结构发育过程的比较研究[J].湖北农业科学,52(20):4991-4994.]