메뉴 건너뛰기




Volumn 59, Issue 9, 2008, Pages 2529-2543

IAA stimulates pollen tube growth and mediates the modification of its wall composition and structure in Torenia fournieri

Author keywords

AFM; Cellulose microfibrils; FTIR; IAA; PM H+ ATPase; Pollen tube; Secretory vesicles; Torenia fournieri

Indexed keywords

TORENIA FOURNIERI;

EID: 48649084435     PISSN: 00220957     EISSN: 14602431     Source Type: Journal    
DOI: 10.1093/jxb/ern119     Document Type: Article
Times cited : (68)

References (58)
  • 1
    • 49749125794 scopus 로고
    • Pollen germination and pollen tube growth, as influenced by pure growth substances
    • Addicott FT. 1943. Pollen germination and pollen tube growth, as influenced by pure growth substances. Plant Physiology 18, 270-279.
    • (1943) Plant Physiology , vol.18 , pp. 270-279
    • Addicott, F.T.1
  • 2
    • 29444446200 scopus 로고    scopus 로고
    • Role of auxin in regulating Arabidopsis flower development
    • Aloni R, Aloni E, Langhans M, Ullrich CI. 2006. Role of auxin in regulating Arabidopsis flower development. Planta 223, 315-328.
    • (2006) Planta , vol.223 , pp. 315-328
    • Aloni, R.1    Aloni, E.2    Langhans, M.3    Ullrich, C.I.4
  • 3
    • 15444366847 scopus 로고    scopus 로고
    • FTIR imaging of wheat endosperm cell walls in situ reveals compositional and architectural heterogeneity related to grain hardness
    • Barron C, Parker ML, Mills EN, Rouau X, Wilson RH. 2005. FTIR imaging of wheat endosperm cell walls in situ reveals compositional and architectural heterogeneity related to grain hardness. Planta 220, 667-677.
    • (2005) Planta , vol.220 , pp. 667-677
    • Barron, C.1    Parker, M.L.2    Mills, E.N.3    Rouau, X.4    Wilson, R.H.5
  • 4
    • 0035050721 scopus 로고    scopus 로고
    • On the alignment of cellulose microfibrils by cortical microtubules: A review and a model
    • Baskin TI. 2001. On the alignment of cellulose microfibrils by cortical microtubules: a review and a model. Protoplasma 215, 150-171.
    • (2001) Protoplasma , vol.215 , pp. 150-171
    • Baskin, T.I.1
  • 5
    • 33644832083 scopus 로고    scopus 로고
    • Pectin methylesterases and pectin dynamics in pollen tubes
    • Bosch M, Hepler PK. 2005. Pectin methylesterases and pectin dynamics in pollen tubes. The Plant Cell 17, 3219-3226.
    • (2005) The Plant Cell , vol.17 , pp. 3219-3226
    • Bosch, M.1    Hepler, P.K.2
  • 6
    • 0031691822 scopus 로고    scopus 로고
    • Effects of radiation and growth hormones on pollen germination, pollen tube growth and modulation of radiation responses of Pinus kesiya Royle ex Gord
    • Chauhan YS, Katiyar SR. 1998. Effects of radiation and growth hormones on pollen germination, pollen tube growth and modulation of radiation responses of Pinus kesiya Royle ex Gord. Cytologia 63, 341-348.
    • (1998) Cytologia , vol.63 , pp. 341-348
    • Chauhan, Y.S.1    Katiyar, S.R.2
  • 7
  • 8
    • 0003273105 scopus 로고    scopus 로고
    • Pollen tubes: Cellular organization and control of growth
    • Clement C, Pacini E, Audran JC, eds, Berlin: Springer
    • Derksen J, van Amstel ANM, Rutten ALM, Knuiman B, Li YQ, Pierson ES. 1999. Pollen tubes: cellular organization and control of growth. In: Clement C, Pacini E, Audran JC, eds. Anther and pollen. Berlin: Springer, 161-174.
    • (1999) Anther and pollen , pp. 161-174
    • Derksen, J.1    van Amstel, A.N.M.2    Rutten, A.L.M.3    Knuiman, B.4    Li, Y.Q.5    Pierson, E.S.6
  • 9
    • 33244486953 scopus 로고    scopus 로고
    • The maize primary cell wall microfibril: A new model derived from direct visualization
    • Ding SY, Himmel ME. 2006. The maize primary cell wall microfibril: a new model derived from direct visualization. Journal of Agriculture and Food Chemistry 54, 597-606.
    • (2006) Journal of Agriculture and Food Chemistry , vol.54 , pp. 597-606
    • Ding, S.Y.1    Himmel, M.E.2
  • 10
    • 0033535048 scopus 로고    scopus 로고
    • Growing pollen tubes possess a constitutive alkaline band in the clear zone and a growth-dependent acidic tip
    • Feijo JA, Sainhas J, Hackett GR, Kunkel JG, Hepler PK. 1999. Growing pollen tubes possess a constitutive alkaline band in the clear zone and a growth-dependent acidic tip. Journal of Cell Biology 144, 483-496.
    • (1999) Journal of Cell Biology , vol.144 , pp. 483-496
    • Feijo, J.A.1    Sainhas, J.2    Hackett, G.R.3    Kunkel, J.G.4    Hepler, P.K.5
  • 11
    • 0031668983 scopus 로고    scopus 로고
    • Location of cellulose and callose in pollen tubes and grains of Nicotiana tabacum
    • Ferguson C, Teeri TT, Siika-aho M, Read SM, Basic A. 1998. Location of cellulose and callose in pollen tubes and grains of Nicotiana tabacum. Planta 206, 452-460.
    • (1998) Planta , vol.206 , pp. 452-460
    • Ferguson, C.1    Teeri, T.T.2    Siika-aho, M.3    Read, S.M.4    Basic, A.5
  • 13
    • 0030755821 scopus 로고    scopus 로고
    • pH gradients are not associated with tip growth in pollen tubes of Lilium longiflorum
    • Fricker MD, White NS, Obermeyer G. 1997. pH gradients are not associated with tip growth in pollen tubes of Lilium longiflorum. Journal of Cell Science 110, 1729-1740.
    • (1997) Journal of Cell Science , vol.110 , pp. 1729-1740
    • Fricker, M.D.1    White, N.S.2    Obermeyer, G.3
  • 14
    • 2142832702 scopus 로고    scopus 로고
    • The local cytomechanical properties of growing pollen tubes correspond to the axial distribution of structural cellular elements
    • Geitmann A, Parre E. 2004. The local cytomechanical properties of growing pollen tubes correspond to the axial distribution of structural cellular elements. Sexual Plant Reproduction 17, 9-16.
    • (2004) Sexual Plant Reproduction , vol.17 , pp. 9-16
    • Geitmann, A.1    Parre, E.2
  • 15
    • 33947095472 scopus 로고    scopus 로고
    • The architecture and properties of the pollen tube cell wall
    • Malhó R, ed, Berlin: Springer Verlag
    • Geitmann A, Steer M. 2006. The architecture and properties of the pollen tube cell wall. In: Malhó R, ed. The pollen tube. Plant Cell Monographs, Vol. 3. Berlin: Springer Verlag, 177-200.
    • (2006) The pollen tube. Plant Cell Monographs , vol.3 , pp. 177-200
    • Geitmann, A.1    Steer, M.2
  • 16
    • 0002874779 scopus 로고
    • Microtubule-mediated control of microfibril deposition: A re-examination of the hypothesis
    • Lloyd CW, ed, San Diego, CA: Academic Press
    • Giddings TH, Staehelin LA. 1991. Microtubule-mediated control of microfibril deposition: a re-examination of the hypothesis. In: Lloyd CW, ed. The cytoskeletal basis of plant growth and form. San Diego, CA: Academic Press, 85-100.
    • (1991) The cytoskeletal basis of plant growth and form , pp. 85-100
    • Giddings, T.H.1    Staehelin, L.A.2
  • 17
    • 34249984767 scopus 로고    scopus 로고
    • Pollen tube growth: Coping with mechanical obstacles involves the cytoskeleton
    • Gossot O, Geitmann A. 2007. Pollen tube growth: coping with mechanical obstacles involves the cytoskeleton. Planta 226, 405-416.
    • (2007) Planta , vol.226 , pp. 405-416
    • Gossot, O.1    Geitmann, A.2
  • 20
    • 0032303007 scopus 로고    scopus 로고
    • Guidance in vitro of the pollen tube to the naked embryo sac of Torenia fournieri
    • Higashiyama T, Haruko Kuroiwa H, Kawano S, Kuroiwa T. 1998. Guidance in vitro of the pollen tube to the naked embryo sac of Torenia fournieri. The Plant Cell 10, 2019-2031.
    • (1998) The Plant Cell , vol.10 , pp. 2019-2031
    • Higashiyama, T.1    Haruko Kuroiwa, H.2    Kawano, S.3    Kuroiwa, T.4
  • 21
    • 0037550211 scopus 로고
    • Relationship between polysaccharide synthesis and cell wall loosening in auxin-induced elongation of rice coleoptile segments
    • Hoson T, Masuda Y. 1992. Relationship between polysaccharide synthesis and cell wall loosening in auxin-induced elongation of rice coleoptile segments. Plant Science 83, 149-154.
    • (1992) Plant Science , vol.83 , pp. 149-154
    • Hoson, T.1    Masuda, Y.2
  • 22
    • 27744577989 scopus 로고    scopus 로고
    • VANGUARD1 encodes a pectin methylesterase that enhances pollen tube growth in the Arabidopsis style and transmitting tract
    • Jiang LX, Yang SL, Xie LF, Puah CS, Zhang XQ, Yang WC, Sundaresan V, Ye D. 2005. VANGUARD1 encodes a pectin methylesterase that enhances pollen tube growth in the Arabidopsis style and transmitting tract. The Plant Cell 17, 584-596.
    • (2005) The Plant Cell , vol.17 , pp. 584-596
    • Jiang, L.X.1    Yang, S.L.2    Xie, L.F.3    Puah, C.S.4    Zhang, X.Q.5    Yang, W.C.6    Sundaresan, V.7    Ye, D.8
  • 23
    • 18644384235 scopus 로고    scopus 로고
    • A conserved functional role of pectic polymers in stomatal guard cells from a range of plant species
    • Jones L, Milne JL, Ashford D, McCann MC, McQueen-Mason SJ. 2005. A conserved functional role of pectic polymers in stomatal guard cells from a range of plant species. Planta 221, 255-264.
    • (2005) Planta , vol.221 , pp. 255-264
    • Jones, L.1    Milne, J.L.2    Ashford, D.3    McCann, M.C.4    McQueen-Mason, S.J.5
  • 24
    • 0344961803 scopus 로고    scopus 로고
    • Hormonal status of the pollenpistil system at the progamic phase of fertilization after compatible and incompatible pollination in Petunia hybrida L
    • Kovaleva L, Zakharova E. 2003. Hormonal status of the pollenpistil system at the progamic phase of fertilization after compatible and incompatible pollination in Petunia hybrida L. Sexual Plant Reproduction 16, 191-196.
    • (2003) Sexual Plant Reproduction , vol.16 , pp. 191-196
    • Kovaleva, L.1    Zakharova, E.2
  • 26
    • 0037343729 scopus 로고    scopus 로고
    • Disruption of cellulose synthesis by isoxaben causes tip swelling and disorganizes cortical microtubules in elongating conifer pollen tubes
    • Lazzaro MD, Donohue JM, Soodavar FM. 2003. Disruption of cellulose synthesis by isoxaben causes tip swelling and disorganizes cortical microtubules in elongating conifer pollen tubes. Protoplasma 220, 201-207.
    • (2003) Protoplasma , vol.220 , pp. 201-207
    • Lazzaro, M.D.1    Donohue, J.M.2    Soodavar, F.M.3
  • 27
    • 26944464617 scopus 로고    scopus 로고
    • Identification of a Nicotiana plumbaginifolia plasma membrane H (+)-ATPase gene expressed in the pollen tube
    • Lefebvre B, Arango M, Oufattole M, Crouzet J, Purnelle B, Boutry M. 2005. Identification of a Nicotiana plumbaginifolia plasma membrane H (+)-ATPase gene expressed in the pollen tube. Plant Molecular Biology 58, 775-787.
    • (2005) Plant Molecular Biology , vol.58 , pp. 775-787
    • Lefebvre, B.1    Arango, M.2    Oufattole, M.3    Crouzet, J.4    Purnelle, B.5    Boutry, M.6
  • 28
    • 3042577084 scopus 로고    scopus 로고
    • Cell wall modification in grapevine cells in response to UV stress investigated by atomic force microscopy
    • Lesniewska E, Adrian M, Klinguer A, Pugin A. 2004. Cell wall modification in grapevine cells in response to UV stress investigated by atomic force microscopy. Ultramicroscopy 100, 171-178.
    • (2004) Ultramicroscopy , vol.100 , pp. 171-178
    • Lesniewska, E.1    Adrian, M.2    Klinguer, A.3    Pugin, A.4
  • 29
    • 0347613014 scopus 로고    scopus 로고
    • Detection and localization of pectin methylesterase isoforms in pollen tubes of Nicotiana tabacum L
    • Li YQ, Mareck A, Faleri C, Moscatelli A, Liu Q, Cresti M. 2002. Detection and localization of pectin methylesterase isoforms in pollen tubes of Nicotiana tabacum L. Planta 214, 734-740.
    • (2002) Planta , vol.214 , pp. 734-740
    • Li, Y.Q.1    Mareck, A.2    Faleri, C.3    Moscatelli, A.4    Liu, Q.5    Cresti, M.6
  • 31
    • 0035212734 scopus 로고    scopus 로고
    • Pectin methylesterases: Cell wall enzymes with important roles in plant physiology
    • Micheli F. 2001. Pectin methylesterases: cell wall enzymes with important roles in plant physiology. Trends in Plant Science 6, 414-419.
    • (2001) Trends in Plant Science , vol.6 , pp. 414-419
    • Micheli, F.1
  • 32
    • 9244220189 scopus 로고    scopus 로고
    • Ethylene synthesis and auxin augmentation in pistil tissues are important for egg cell differentiation after pollination in maize
    • Mol R, Filek M, Machackova I, Matthys-Rochon E. 2004. Ethylene synthesis and auxin augmentation in pistil tissues are important for egg cell differentiation after pollination in maize. Plant and Cell Physiology 45, 1396-1405.
    • (2004) Plant and Cell Physiology , vol.45 , pp. 1396-1405
    • Mol, R.1    Filek, M.2    Machackova, I.3    Matthys-Rochon, E.4
  • 33
    • 0035781005 scopus 로고    scopus 로고
    • Plant mitochondria and oxidative stress: Electron transport, NADPH turnover, and metabolism of reactive oxygen species
    • Moller IM. 2001. Plant mitochondria and oxidative stress: electron transport, NADPH turnover, and metabolism of reactive oxygen species. Annual Review of Plant Physiology & Plant Molecular Biology 52, 561-591.
    • (2001) Annual Review of Plant Physiology & Plant Molecular Biology , vol.52 , pp. 561-591
    • Moller, I.M.1
  • 35
    • 0033792717 scopus 로고    scopus 로고
    • Auxin and ETTIN in Arabidopsis gynoecium morphogenesis
    • Nemhauser JL, Feldman LJ, Zambryski PC. 2000. Auxin and ETTIN in Arabidopsis gynoecium morphogenesis. Development 127, 3877-3888.
    • (2000) Development , vol.127 , pp. 3877-3888
    • Nemhauser, J.L.1    Feldman, L.J.2    Zambryski, P.C.3
  • 37
    • 14644408804 scopus 로고    scopus 로고
    • Pectin and the role of the physical properties of the cell wall in pollen tube growth of Solanum chacoense
    • Parre E, Geitmann A. 2005. Pectin and the role of the physical properties of the cell wall in pollen tube growth of Solanum chacoense. Planta 220, 582-592.
    • (2005) Planta , vol.220 , pp. 582-592
    • Parre, E.1    Geitmann, A.2
  • 39
    • 0034908310 scopus 로고    scopus 로고
    • Dynamics of the apical vesicle accumulation and the rate of growth are related in individual pollen tubes
    • Parton RM, Fischer-Parton S, Watahiki MK, Trewavas AJ. 2001. Dynamics of the apical vesicle accumulation and the rate of growth are related in individual pollen tubes. Journal of Cell Science 114, 2685-2695.
    • (2001) Journal of Cell Science , vol.114 , pp. 2685-2695
    • Parton, R.M.1    Fischer-Parton, S.2    Watahiki, M.K.3    Trewavas, A.J.4
  • 41
    • 49749115566 scopus 로고
    • On factors influencing fruit-set and sterility in arecanut (Areca catechu Linn.). II. Germination of pollen grains and growth of pollen tube under the influence of certain auxin, vitamins and trace elements
    • Raghavan V, Baruah HK. 1956. On factors influencing fruit-set and sterility in arecanut (Areca catechu Linn.). II. Germination of pollen grains and growth of pollen tube under the influence of certain auxin, vitamins and trace elements. Phyton 7, 77-82.
    • (1956) Phyton , vol.7 , pp. 77-82
    • Raghavan, V.1    Baruah, H.K.2
  • 42
    • 84980139514 scopus 로고
    • Effect of time factor on the stimulation of pollen germination and pollen tube growth by certain auxins, vitamins, and trace elements
    • Raghavan V, Baruah HK. 1959. Effect of time factor on the stimulation of pollen germination and pollen tube growth by certain auxins, vitamins, and trace elements. Physiologia Plantarum 12, 441-451.
    • (1959) Physiologia Plantarum , vol.12 , pp. 441-451
    • Raghavan, V.1    Baruah, H.K.2
  • 43
    • 0026911103 scopus 로고
    • The acid growth theory of auxininduced cell elongation is alive and well
    • Rayle DL, Cleland RE. 1992. The acid growth theory of auxininduced cell elongation is alive and well. Plant Physiology 99, 1271-1274.
    • (1992) Plant Physiology , vol.99 , pp. 1271-1274
    • Rayle, D.L.1    Cleland, R.E.2
  • 45
    • 37249067542 scopus 로고    scopus 로고
    • Elaborate spatial patterning of cell-wall PME and PMEI at the pollen tube tip involves PMEI endocytosis, and reflects the distribution of esterified and de-esterified pectins
    • Röckel N, Wolf S, Kost B, Rausch T, Greiner S. 2008. Elaborate spatial patterning of cell-wall PME and PMEI at the pollen tube tip involves PMEI endocytosis, and reflects the distribution of esterified and de-esterified pectins. The Plant Journal 53, 133-143.
    • (2008) The Plant Journal , vol.53 , pp. 133-143
    • Röckel, N.1    Wolf, S.2    Kost, B.3    Rausch, T.4    Greiner, S.5
  • 46
    • 28444480365 scopus 로고    scopus 로고
    • COBRA, an Arabidopsis extracellular glycosyl-phosphatidyl inositol-anchored protein, specifically controls highly anisotropic expansion through its involvement in cellulose microfibril orientation
    • Roudier F, Fernandez AG, Fujita M, Himmelspach R, Borner GHH, Schindelman G, Song S, Baskin TI, Dupree P, Wasteneys GO, Benfey PN. 2005. COBRA, an Arabidopsis extracellular glycosyl-phosphatidyl inositol-anchored protein, specifically controls highly anisotropic expansion through its involvement in cellulose microfibril orientation. The Plant Cell 17, 1749-1763.
    • (2005) The Plant Cell , vol.17 , pp. 1749-1763
    • Roudier, F.1    Fernandez, A.G.2    Fujita, M.3    Himmelspach, R.4    Borner, G.H.H.5    Schindelman, G.6    Song, S.7    Baskin, T.I.8    Dupree, P.9    Wasteneys, G.O.10    Benfey, P.N.11
  • 47
    • 84989998732 scopus 로고
    • Fine structure of Petunia pollen grain and pollen tube
    • Sassen MMA. 1964. Fine structure of Petunia pollen grain and pollen tube. Acta Botanica Neerlandica 13, 175-181.
    • (1964) Acta Botanica Neerlandica , vol.13 , pp. 175-181
    • Sassen, M.M.A.1
  • 48
    • 33747098886 scopus 로고    scopus 로고
    • Roles of the ubiquitin/proteasome pathway in pollen tube growth with emphasis on MG132-induced alterations in ultrastructure, cytoskeleton, and cell wall components
    • Sheng X, Hu Z, Lu H, Wang X, Baluska F, Samaj J, Lin J. 2006. Roles of the ubiquitin/proteasome pathway in pollen tube growth with emphasis on MG132-induced alterations in ultrastructure, cytoskeleton, and cell wall components. Plant Physiology 141, 1578-1590.
    • (2006) Plant Physiology , vol.141 , pp. 1578-1590
    • Sheng, X.1    Hu, Z.2    Lu, H.3    Wang, X.4    Baluska, F.5    Samaj, J.6    Lin, J.7
  • 49
    • 0002153560 scopus 로고
    • Microtubules and the regulation of cell morphogenesis by plant hormones
    • Lloyd CW, ed, London: Academic Press
    • Shibaoka H. 1991. Microtubules and the regulation of cell morphogenesis by plant hormones. In: Lloyd CW, ed. The cytoskeletal basis of plant growth and form. London: Academic Press, 159-168.
    • (1991) The cytoskeletal basis of plant growth and form , pp. 159-168
    • Shibaoka, H.1
  • 50
    • 0342748430 scopus 로고    scopus 로고
    • Localization of pectins in the pollen tube wall of Ornithogalum virens: Does the pattern of pectin distribution depend on the growth rate of the pollen tube?
    • Stepka M, Ciampolimi F, Cresti M. 2000. Localization of pectins in the pollen tube wall of Ornithogalum virens: does the pattern of pectin distribution depend on the growth rate of the pollen tube? Planta 210, 630-635.
    • (2000) Planta , vol.210 , pp. 630-635
    • Stepka, M.1    Ciampolimi, F.2    Cresti, M.3
  • 51
    • 0034522561 scopus 로고    scopus 로고
    • New techniques enable comparative analysis of microtubule orientation, wall texture and growth rate in intact roots of Arabidopsis
    • Sugimoto K, Williamson RE, Wasteneys GO. 2000. New techniques enable comparative analysis of microtubule orientation, wall texture and growth rate in intact roots of Arabidopsis. Plant Physiology 124, 1493-1506.
    • (2000) Plant Physiology , vol.124 , pp. 1493-1506
    • Sugimoto, K.1    Williamson, R.E.2    Wasteneys, G.O.3
  • 53
    • 0001641624 scopus 로고
    • Effects of auxin on wall polysaccharide composition and enzyme activity during extension-growth of Pellia (Bryophyta)
    • Thomas RJ, Behringer FJ, Lombard CS, Sparkowski JJ. 1984. Effects of auxin on wall polysaccharide composition and enzyme activity during extension-growth of Pellia (Bryophyta). Physiologia Plantarum 60, 502-506.
    • (1984) Physiologia Plantarum , vol.60 , pp. 502-506
    • Thomas, R.J.1    Behringer, F.J.2    Lombard, C.S.3    Sparkowski, J.J.4
  • 56
    • 43149096258 scopus 로고    scopus 로고
    • Pollen tube growth is affected by exogenous hormones and correlated with hormone changes in styles before and after pollination in Torenia fournieri L
    • DOI: 10.1007/s10725-008-9268-5
    • Wu JZ, Qin Y, Zhao J. 2008. Pollen tube growth is affected by exogenous hormones and correlated with hormone changes in styles before and after pollination in Torenia fournieri L. Plant Growth Regulation DOI: 10.1007/s10725-008-9268-5.
    • (2008) Plant Growth Regulation
    • Wu, J.Z.1    Qin, Y.2    Zhao, J.3
  • 57
    • 26444580703 scopus 로고    scopus 로고
    • Heterotrimeric G protein α-subunit is localized in the plasma membrane of Pinus bungeana pollen tubes
    • Zhang LY, Fang KF, Lin JX. 2005. Heterotrimeric G protein α-subunit is localized in the plasma membrane of Pinus bungeana pollen tubes. Plant Science 169, 1066-1073.
    • (2005) Plant Science , vol.169 , pp. 1066-1073
    • Zhang, L.Y.1    Fang, K.F.2    Lin, J.X.3
  • 58
    • 0035733658 scopus 로고    scopus 로고
    • Changes of calcium distribution in egg cells, zygotes and two-celled proembryos of rice (Oryza sativa L.)
    • Zhao J, Yu FL, Liang SP, Zhou C, Yang HY. 2002. Changes of calcium distribution in egg cells, zygotes and two-celled proembryos of rice (Oryza sativa L.). Sexual Plant Reproduction 14, 331-337.
    • (2002) Sexual Plant Reproduction , vol.14 , pp. 331-337
    • Zhao, J.1    Yu, F.L.2    Liang, S.P.3    Zhou, C.4    Yang, H.Y.5


* 이 정보는 Elsevier사의 SCOPUS DB에서 KISTI가 분석하여 추출한 것입니다.