高粱组织培养技术研究进展
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摘要
高粱是全球第五大谷类作物,是食品、饲料和加工业的原料,也是一种潜在的能源作物,具有重要的应用价值。然而,通过基因工程技术对高粱进行遗传改良却十分困难,其主要原因是缺乏高效的组织培养体系。结合近年来的研究,从基因型、外植体、培养基、植物激素等影响高粱组织培养的主要因素进行综述,重点讨论了促进高粱再生的策略,并提出解决高粱组培再生困难的两大研究方向:一是深入研究植物再生的遗传及表观遗传机制;二是根据各基因型的代谢特点,进一步优化培养条件,旨为建立高效的高粱组织培养体系提供参考。
Sorghum(Sorghum bicolor L. Moench)is the fifth most widely planted cereal crop in the world. It is the raw material in food,fodder and processing industry. As well,it has been identified as a potential biofuel crop. Despite the important application value,the genetic improvement of sorghum via genetic engineering technology is very difficult,because there is no highly efficient sorghum tissue culture system.According to recent studies,we reviewed the main factors having great influences on sorghum tissue culture,such as genotypes,explants,media,plant hormones,etc. Importantly,we focused on the strategies which may promote the regeneration of sorghum. At the end,two directions for solving the difficulties in sorghum regeneration via tissue culture are given as :first,the genetic and epigenetical mechanisms of plant regeneration should be studied deeply;second,the culture conditions should be further optimized based on the metabolism characteristic of every genotype. These ideas will be references for the establishment of highly efficient sorghum tissue culture system.
Sorghum(Sorghum bicolor L. Moench)is the fifth most widely planted cereal crop in the world. It is the raw material in food,fodder and processing industry. As well,it has been identified as a potential biofuel crop. Despite the important application value,the genetic improvement of sorghum via genetic engineering technology is very difficult,because there is no highly efficient sorghum tissue culture system.According to recent studies,we reviewed the main factors having great influences on sorghum tissue culture,such as genotypes,explants,media,plant hormones,etc. Importantly,we focused on the strategies which may promote the regeneration of sorghum. At the end,two directions for solving the difficulties in sorghum regeneration via tissue culture are given as :first,the genetic and epigenetical mechanisms of plant regeneration should be studied deeply;second,the culture conditions should be further optimized based on the metabolism characteristic of every genotype. These ideas will be references for the establishment of highly efficient sorghum tissue culture system.
引文
[1]张丽敏,刘智全,陈冰嬬,等.我国能源甜高粱育种现状及应用前景[J].中国农业大学学报, 2012, 17(6):76-82.
[2]Dahlberg J, Berenji J, Sikora V, et al. Assessing sorghum[Sorghum bicolor(L)Moench]germplasm for new traits:food, fuels&unique uses[J]. Maydica, 2011, 56:85-92.
[3]Vermerris W. Survey of genomics approaches to improve bioenergytraits in maize, sorghum and sugarcane[J]. Journal of IntegrativePlant Biology, 2011, 53(2):105-119.
[4]刘公社,周庆源,宋松泉,等.能源植物甜高粱种质资源和分子生物学研究进展[J].植物学报, 2009, 44(3):253-261.
[5]Chandrashekar A, Satyanarayana KV. Disease and pest resistance ingrains of sorghum and millets[J]. Journal of Cereal Science, 2006,44(3):287-304.
[6]Nwanze KF, Seetharama N, Sharma HC, et al. Biotechnology inpestmanagement:Improving resistance in sorghum to insect pests[J].African Crop Science, 1995, 3(2):209-215.
[7]Chen M, Shelton A, Ye GY, et al. Insect-resistant geneticallymodified rice in China:From research to commercialization[J].Annual Review of Entomology, 2011, 56:81-101.
[8]Zhang BH. Transgenic cotton:From biotransformation methods toagricultural application[J]. Methods in Molecular Biology, 2013,958:3-15.
[9]Calla B, Blahut-Beatty L, Koziol L, et al. Genomic evaluation ofoxalate-degrading transgenic soybean in response to sclerotiniasclerotiorum infection[J]. Molecular Plant Pathology, 2014, 15(6):563-575.
[10]Polumahanthi S, Manin S, Pola S, et al. Tissue culture, molecularand genetic approaches to sorghum crop improvement[J]. IndianJournal of Plant Sciences, 2015, 4(2):97-113.
[11]Liu GQ, Gilding EK, Godwin ID. A robust tissue culture system forsorghum[Sorghum bicolor(L.)Moench][J]. South AfricanJournal of Botany, 2015, 98:157-160.
[12]Liu GQ, Godwin ID. Highly efficient sorghum transformation[J].Plant Cell Report, 2012, 31(6):999-1007.
[13]Wu E, Lenderts B, Glassman K, et al. Optimized Agrobacteriummediated sorghum transformation protocol and molecular data oftransgenic sorghum plants[J]. In Vitro Cell Dev Biol Plant,2014, 50(1):9-18.
[14]DoPT,LeeH,MookkanM,etal.RapidandefficientAgrobacterium-mediated transformati on of sorghum(Sorghum bicolor)employing standard binary vectors and bar gene as aselectable marker[J]. Plant Cell Report, 2016, 35(10):2065-2076.
[15]Hiei Y, Ishida Y, Komari. Progress of cereal transformationtechnology mediated by Agrobacterium tumefaciens[J]. Frontiersin Plant Science, 2014, 5:1-11.
[16]Li JQ, Wang LH, Zhan QW, et al. Development of a simple andefficient method for Agrobacterium-mediated transformation insorghum[J]. International Journal of Agriculture&Biology,2015, 18(1):134-138.
[17]Lowe K, Wu E, Wang N, et al. Morphogenic regulators Baby boomand Wuschel improve monocot transformation[J]. Plant Cell,2016, 28(9):1998-2015.
[18]Cai T, Butler L. Plant regeneration from embryogenic callusinitiated from immature inflorescences of several high-tanninsorghums[J]. Plant Cell, Tissue and Organ Culture, 1990, 20(2):101-110.
[19]Assem SK, Zamzam M, Hussein BA, et al. Evaluation of somaticembryogenesis and plant regeneration in tissue culture of tensorghum(Sorghum bicolor L.)genotype[J]. African Journal ofBiotechnology, 2014, 13(36):3672-3681.
[20]Gamhorg OL, Shyluk JP, Brar DS, et al. Morphogenesis and plantregeneration from callus of immature embryos of sorghum[J].Plant Science Letters, 1977, 10(1):67-74.
[21]Dustan DI, Short KC, Dhaliwal H, et al. Further studies on plantletproduction from cultured tissues of sorghum[J]. Protoplasma,1979, 101(4):355-361.
[22]Ma HT, Gu MH, Liang GH. Plant regeneration from culturedimmature embryos of Sorghum bicolor(L.)Moench[J]. TheorlAppl Genet, 1987, 73(3):389-394.
[23]马鸿图, Liang GH.高粱幼胚培养及再生植株变异的研究[J].遗传学报, 1985, 12(5):350-357.
[24]韩福光,张颍.高粱不同外植体愈伤组织诱导的研究[J].辽宁农业科学, 1993,(1):45-48.
[25]Brettell RIS, Wernicke W, Thomas E. Embryogenesis from culturedimmature inflorescences of Sorghum bicolor[J]. Protoplasma,1980, 104(1-2):141-148.
[26]Jogeswar G, Ranadheer D, Anjaiah V, et al. High frequency somaticembryogenesis and regeneration in different genotypes of Sorghum bicolor(L.)Moench from immature inflorescence explants[J].In Vitro Cell Dev Biol Plant, 2007, 43(2):159-166.
[27]Murashige T, Skoog F. A revised medium for rapid growth andbioassays with tobacco tissue cultures[J]. Physiologia Plantarum,1962, 15:473-497.
[28]Gupta S, Khanna VK, Singh R, et al. Strategies for overcominggenotypic limitations of in vitro regeneration and determinationof genetic components of variability of plant regeneration traits insorghum[J]. Plant Cell Tiss Organ Cult, 2006, 86(3):379-388.
[29]徐丹,陈立余,徐子勤.甜高粱离体再生体系的建立和组织结构变化的观察[J].植物生理学通讯, 2009, 45(8):771-774.
[30]赵利铭.甜高粱遗传背景对其再生能力的影响及再生体系的建立[D].合肥:中国科学技术大学, 2010.
[31]Pola S, Saradamani N, Ramana T. Mature embryos as a sourcematerial for efficient regeneration response in sorghum(Sorghum bicolor L. Moench)[J]. Sjemenarstvo, 2009, 26:3-4.
[32]Wernicke W, Potrykus I, Thomas E. Morphogenesis from culturedleaf tissue of Sorghum bicolor—The morphogenetic pathway[J].ProtopIasma, 1982, 111(1):53-62.
[33]Pola S. Leaf discs as a source material for plant tissue culturestudies of Sorghum bicolor(L.)Moench[J]. Not Sci Biol, 2011,3(1):70-78.
[34]Pola SR, Mani NS. Somatic embryogenesis and plantlet regenerationin Sorghum bicolor(L.)Moench from leaf segments[J]. Journalof Cell and Molecular Biology, 2006, 5:99-107.
[35]Wen FS, Sorensen EL, Barnett F L, et al. Callus induction andplant regeneration from anther and inflorescence culture ofSorghum[J]. Euphytica, 1991, 52(3):177-181.
[36]Nirwan RS, Kothari SL. High frequency shoot organogenesis inSorghum bicolor(L)Moench[J]. J. Plant Biochemistry&Biotechnology, 2004, 13(2):149-152.
[37]Kishore NS, Visarada K, Lakshmi YA, et al. In vitro culturemethods in sorghum with shoot tip as the explant material[J].Plant Cell Rep, 2006, 25(3):174-182.
[38]刘宣雨,刘树君,宋松泉.建立甜高粱高频、高效再生体系的研究[J].中国农业科学, 2010, 43(23):4963-4969.
[39]Baskaran P, Rajeswari BR, Jayabalan N. Development of an in vitroregeneration system in Sorghum[Sorghum bicolor(L.)Moench]using root transverse thin cell layers(tTCLs)[J]. Turk J Bot,2006, 30:1-9.
[40]Sticklen MB, Oraby H. Shoot apical meristem:a sustainableexplant for genetic transformation of cereal crops[J]. In VitroCell Dev Bio Plant, 2005, 41(3):187-200.
[41]Maekinnon C, Gunderson G, Nabors MW. Plant regeneration bysomatic embryogenesis from callus cultures of sweet sorghum[J].Plant Cell Reports, 1986, 5(5):349-351.
[42]Linsmaier EM, Skoog F. Organic growth factor requirements oftobacco tissue cultures[J]. Physiologia Plantarum, 1965, 18:100-127.
[43]Goh E J, Seong E S, Yoo J H, et al. Effect of plant growth regulatorsand media on regeneration of Sorghum bicolor(L.)Moench[J].Korean J. Plant Res., 2011, 24(2):168-173.
[44]Chu CC, Wang CC, Sun CS, et al. Establishment of an efficientmedium for anther culture of rice through comparative experimentson the nitrogen sources[J]. Scientia Sinica, 1975, 18(5):659-668.
[45]Gamborg OL, Miller RA, Ojima K. Nutrient requments ofsuspension cultures of soybean root cells[J]. Expremental CellResearch, 1968, 50:151-158.
[46]Elkonin LA, Pakhomova NV. Influence of nitrogen and phosphoruson induction embryogenic callus of sorghum[J]. Plant Cell,Tissue and Organ Culture, 2000, 61(2):115-123.
[47]Kumar V, Campbell LM, Rathore KS. Rapid recovery andcharacterization of transformants following Agrobacterium-mediatedT-DNA transfer to sorghum[J]. Plant Cell Tiss Organ Cult, 2011,104(2):137-146.
[48]Purnhauser L. Stimulation of shoot and root regeneration in wheat(Triticum aestivum)callus cultures by copper[J]. CerealResearch Communication, 1991, 19(4):419-423.
[49]Purnhauser L, Gyulai G. Effect of copper on shoot and rootregenerationinwheat,triticale,rapeandtobaccotissuecultures[J]. Plant Cell Tissue and Organ Culture, 1993, 35(2):131-139.
[50]Dahleen L. Improved plant regeneration from barley callus culturesby increased copper levels[J]. Plant Cell Tissue Organ Culture,1995, 43(3):267-269.
[51]Nirwan RS, Kothar SL. High copper levels improve callus inductionand plant regeneration in Sorghum bicolor(L.)Moench In VitroCell Dev Biol Plant, 2003, 39(2):161-164.
[52]Liu GQ, Gilding EK, Godwin ID. Additive effects of three auxinsand copper on sorghum in vitro root induction[J]. In Vitro CellDev Biol Plant, 2013, 49(2):19-197.
[53]Lidon FC, Barreiro MG, Henriquest F. Interactions betweenbiomass production and ethylene biosynthesis in copper treatedrice[J]. Journal of Plant Nutrition, 1995, 18(6):1301-1314.
[54]Rao AM, Sree KP, Kishor PBK. Enhanced plant regenerationingrainandsweetsorghumbyasparagine,prolineandcefotaxime[J]. Plant Cell Reports, 1995, 15(1-2):72-75.
[55]Elkonin LA, Lopushanskaya RF, Pakhomova NV. Initiation andmaintenance of friable, embryogenetic callus of sorghum(Sorghum bicolor(L.)Moench)by amino acids[J]. Maydica, 1995, 40(2):153-157.
[56]Hagio T. Adventitious shoot regeneration from immature embryos ofsorghum[J]. Plant Cell, Tissue and Organ Culture, 2002, 68(1):65-72.
[57]Nguyen TV, Thu TT, Claeys M, et al. Agrobacterium mediatedtransformation of sorghum(Sorghum bicolor(L.)Moench)usingan improved in vitro regeneration system[J]. Plant Cell Tiss andOrgan Cult, 2007, 91(2):155-164.
[58]Lu L, Wu XG, Yin XY, et al. Development of marker-freetransgenic sorghum[Sorghum bicolor(L.)Moench]usingstandard binary vectors with bar as a selectable marker[J]. Plantcell Tiss Organ Cult, 2009, 99(1):97-108.
[59]Che P, Anand A, Wu E, et al. Developing a flexible, high-efficiencyAgrobacterium-mediated sorghum transformation system with broadapplication[J]. Plant Biotechnology Journal, 2018:1-8.
[60]KumarV,StadenJV.Newinsightsintoplantsomaticembryogenesis:an epigenetic view[J]. Acta Physiol Plant,2017, 39:1-17.
[2]Dahlberg J, Berenji J, Sikora V, et al. Assessing sorghum[Sorghum bicolor(L)Moench]germplasm for new traits:food, fuels&unique uses[J]. Maydica, 2011, 56:85-92.
[3]Vermerris W. Survey of genomics approaches to improve bioenergytraits in maize, sorghum and sugarcane[J]. Journal of IntegrativePlant Biology, 2011, 53(2):105-119.
[4]刘公社,周庆源,宋松泉,等.能源植物甜高粱种质资源和分子生物学研究进展[J].植物学报, 2009, 44(3):253-261.
[5]Chandrashekar A, Satyanarayana KV. Disease and pest resistance ingrains of sorghum and millets[J]. Journal of Cereal Science, 2006,44(3):287-304.
[6]Nwanze KF, Seetharama N, Sharma HC, et al. Biotechnology inpestmanagement:Improving resistance in sorghum to insect pests[J].African Crop Science, 1995, 3(2):209-215.
[7]Chen M, Shelton A, Ye GY, et al. Insect-resistant geneticallymodified rice in China:From research to commercialization[J].Annual Review of Entomology, 2011, 56:81-101.
[8]Zhang BH. Transgenic cotton:From biotransformation methods toagricultural application[J]. Methods in Molecular Biology, 2013,958:3-15.
[9]Calla B, Blahut-Beatty L, Koziol L, et al. Genomic evaluation ofoxalate-degrading transgenic soybean in response to sclerotiniasclerotiorum infection[J]. Molecular Plant Pathology, 2014, 15(6):563-575.
[10]Polumahanthi S, Manin S, Pola S, et al. Tissue culture, molecularand genetic approaches to sorghum crop improvement[J]. IndianJournal of Plant Sciences, 2015, 4(2):97-113.
[11]Liu GQ, Gilding EK, Godwin ID. A robust tissue culture system forsorghum[Sorghum bicolor(L.)Moench][J]. South AfricanJournal of Botany, 2015, 98:157-160.
[12]Liu GQ, Godwin ID. Highly efficient sorghum transformation[J].Plant Cell Report, 2012, 31(6):999-1007.
[13]Wu E, Lenderts B, Glassman K, et al. Optimized Agrobacteriummediated sorghum transformation protocol and molecular data oftransgenic sorghum plants[J]. In Vitro Cell Dev Biol Plant,2014, 50(1):9-18.
[14]DoPT,LeeH,MookkanM,etal.RapidandefficientAgrobacterium-mediated transformati on of sorghum(Sorghum bicolor)employing standard binary vectors and bar gene as aselectable marker[J]. Plant Cell Report, 2016, 35(10):2065-2076.
[15]Hiei Y, Ishida Y, Komari. Progress of cereal transformationtechnology mediated by Agrobacterium tumefaciens[J]. Frontiersin Plant Science, 2014, 5:1-11.
[16]Li JQ, Wang LH, Zhan QW, et al. Development of a simple andefficient method for Agrobacterium-mediated transformation insorghum[J]. International Journal of Agriculture&Biology,2015, 18(1):134-138.
[17]Lowe K, Wu E, Wang N, et al. Morphogenic regulators Baby boomand Wuschel improve monocot transformation[J]. Plant Cell,2016, 28(9):1998-2015.
[18]Cai T, Butler L. Plant regeneration from embryogenic callusinitiated from immature inflorescences of several high-tanninsorghums[J]. Plant Cell, Tissue and Organ Culture, 1990, 20(2):101-110.
[19]Assem SK, Zamzam M, Hussein BA, et al. Evaluation of somaticembryogenesis and plant regeneration in tissue culture of tensorghum(Sorghum bicolor L.)genotype[J]. African Journal ofBiotechnology, 2014, 13(36):3672-3681.
[20]Gamhorg OL, Shyluk JP, Brar DS, et al. Morphogenesis and plantregeneration from callus of immature embryos of sorghum[J].Plant Science Letters, 1977, 10(1):67-74.
[21]Dustan DI, Short KC, Dhaliwal H, et al. Further studies on plantletproduction from cultured tissues of sorghum[J]. Protoplasma,1979, 101(4):355-361.
[22]Ma HT, Gu MH, Liang GH. Plant regeneration from culturedimmature embryos of Sorghum bicolor(L.)Moench[J]. TheorlAppl Genet, 1987, 73(3):389-394.
[23]马鸿图, Liang GH.高粱幼胚培养及再生植株变异的研究[J].遗传学报, 1985, 12(5):350-357.
[24]韩福光,张颍.高粱不同外植体愈伤组织诱导的研究[J].辽宁农业科学, 1993,(1):45-48.
[25]Brettell RIS, Wernicke W, Thomas E. Embryogenesis from culturedimmature inflorescences of Sorghum bicolor[J]. Protoplasma,1980, 104(1-2):141-148.
[26]Jogeswar G, Ranadheer D, Anjaiah V, et al. High frequency somaticembryogenesis and regeneration in different genotypes of Sorghum bicolor(L.)Moench from immature inflorescence explants[J].In Vitro Cell Dev Biol Plant, 2007, 43(2):159-166.
[27]Murashige T, Skoog F. A revised medium for rapid growth andbioassays with tobacco tissue cultures[J]. Physiologia Plantarum,1962, 15:473-497.
[28]Gupta S, Khanna VK, Singh R, et al. Strategies for overcominggenotypic limitations of in vitro regeneration and determinationof genetic components of variability of plant regeneration traits insorghum[J]. Plant Cell Tiss Organ Cult, 2006, 86(3):379-388.
[29]徐丹,陈立余,徐子勤.甜高粱离体再生体系的建立和组织结构变化的观察[J].植物生理学通讯, 2009, 45(8):771-774.
[30]赵利铭.甜高粱遗传背景对其再生能力的影响及再生体系的建立[D].合肥:中国科学技术大学, 2010.
[31]Pola S, Saradamani N, Ramana T. Mature embryos as a sourcematerial for efficient regeneration response in sorghum(Sorghum bicolor L. Moench)[J]. Sjemenarstvo, 2009, 26:3-4.
[32]Wernicke W, Potrykus I, Thomas E. Morphogenesis from culturedleaf tissue of Sorghum bicolor—The morphogenetic pathway[J].ProtopIasma, 1982, 111(1):53-62.
[33]Pola S. Leaf discs as a source material for plant tissue culturestudies of Sorghum bicolor(L.)Moench[J]. Not Sci Biol, 2011,3(1):70-78.
[34]Pola SR, Mani NS. Somatic embryogenesis and plantlet regenerationin Sorghum bicolor(L.)Moench from leaf segments[J]. Journalof Cell and Molecular Biology, 2006, 5:99-107.
[35]Wen FS, Sorensen EL, Barnett F L, et al. Callus induction andplant regeneration from anther and inflorescence culture ofSorghum[J]. Euphytica, 1991, 52(3):177-181.
[36]Nirwan RS, Kothari SL. High frequency shoot organogenesis inSorghum bicolor(L)Moench[J]. J. Plant Biochemistry&Biotechnology, 2004, 13(2):149-152.
[37]Kishore NS, Visarada K, Lakshmi YA, et al. In vitro culturemethods in sorghum with shoot tip as the explant material[J].Plant Cell Rep, 2006, 25(3):174-182.
[38]刘宣雨,刘树君,宋松泉.建立甜高粱高频、高效再生体系的研究[J].中国农业科学, 2010, 43(23):4963-4969.
[39]Baskaran P, Rajeswari BR, Jayabalan N. Development of an in vitroregeneration system in Sorghum[Sorghum bicolor(L.)Moench]using root transverse thin cell layers(tTCLs)[J]. Turk J Bot,2006, 30:1-9.
[40]Sticklen MB, Oraby H. Shoot apical meristem:a sustainableexplant for genetic transformation of cereal crops[J]. In VitroCell Dev Bio Plant, 2005, 41(3):187-200.
[41]Maekinnon C, Gunderson G, Nabors MW. Plant regeneration bysomatic embryogenesis from callus cultures of sweet sorghum[J].Plant Cell Reports, 1986, 5(5):349-351.
[42]Linsmaier EM, Skoog F. Organic growth factor requirements oftobacco tissue cultures[J]. Physiologia Plantarum, 1965, 18:100-127.
[43]Goh E J, Seong E S, Yoo J H, et al. Effect of plant growth regulatorsand media on regeneration of Sorghum bicolor(L.)Moench[J].Korean J. Plant Res., 2011, 24(2):168-173.
[44]Chu CC, Wang CC, Sun CS, et al. Establishment of an efficientmedium for anther culture of rice through comparative experimentson the nitrogen sources[J]. Scientia Sinica, 1975, 18(5):659-668.
[45]Gamborg OL, Miller RA, Ojima K. Nutrient requments ofsuspension cultures of soybean root cells[J]. Expremental CellResearch, 1968, 50:151-158.
[46]Elkonin LA, Pakhomova NV. Influence of nitrogen and phosphoruson induction embryogenic callus of sorghum[J]. Plant Cell,Tissue and Organ Culture, 2000, 61(2):115-123.
[47]Kumar V, Campbell LM, Rathore KS. Rapid recovery andcharacterization of transformants following Agrobacterium-mediatedT-DNA transfer to sorghum[J]. Plant Cell Tiss Organ Cult, 2011,104(2):137-146.
[48]Purnhauser L. Stimulation of shoot and root regeneration in wheat(Triticum aestivum)callus cultures by copper[J]. CerealResearch Communication, 1991, 19(4):419-423.
[49]Purnhauser L, Gyulai G. Effect of copper on shoot and rootregenerationinwheat,triticale,rapeandtobaccotissuecultures[J]. Plant Cell Tissue and Organ Culture, 1993, 35(2):131-139.
[50]Dahleen L. Improved plant regeneration from barley callus culturesby increased copper levels[J]. Plant Cell Tissue Organ Culture,1995, 43(3):267-269.
[51]Nirwan RS, Kothar SL. High copper levels improve callus inductionand plant regeneration in Sorghum bicolor(L.)Moench In VitroCell Dev Biol Plant, 2003, 39(2):161-164.
[52]Liu GQ, Gilding EK, Godwin ID. Additive effects of three auxinsand copper on sorghum in vitro root induction[J]. In Vitro CellDev Biol Plant, 2013, 49(2):19-197.
[53]Lidon FC, Barreiro MG, Henriquest F. Interactions betweenbiomass production and ethylene biosynthesis in copper treatedrice[J]. Journal of Plant Nutrition, 1995, 18(6):1301-1314.
[54]Rao AM, Sree KP, Kishor PBK. Enhanced plant regenerationingrainandsweetsorghumbyasparagine,prolineandcefotaxime[J]. Plant Cell Reports, 1995, 15(1-2):72-75.
[55]Elkonin LA, Lopushanskaya RF, Pakhomova NV. Initiation andmaintenance of friable, embryogenetic callus of sorghum(Sorghum bicolor(L.)Moench)by amino acids[J]. Maydica, 1995, 40(2):153-157.
[56]Hagio T. Adventitious shoot regeneration from immature embryos ofsorghum[J]. Plant Cell, Tissue and Organ Culture, 2002, 68(1):65-72.
[57]Nguyen TV, Thu TT, Claeys M, et al. Agrobacterium mediatedtransformation of sorghum(Sorghum bicolor(L.)Moench)usingan improved in vitro regeneration system[J]. Plant Cell Tiss andOrgan Cult, 2007, 91(2):155-164.
[58]Lu L, Wu XG, Yin XY, et al. Development of marker-freetransgenic sorghum[Sorghum bicolor(L.)Moench]usingstandard binary vectors with bar as a selectable marker[J]. Plantcell Tiss Organ Cult, 2009, 99(1):97-108.
[59]Che P, Anand A, Wu E, et al. Developing a flexible, high-efficiencyAgrobacterium-mediated sorghum transformation system with broadapplication[J]. Plant Biotechnology Journal, 2018:1-8.
[60]KumarV,StadenJV.Newinsightsintoplantsomaticembryogenesis:an epigenetic view[J]. Acta Physiol Plant,2017, 39:1-17.