假俭草种质资源综合评价及其抗寒性的改良与调控
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摘要
假俭草(Eremochloa ophiuroides (Munro.) Hack.)是禾本科蜈蚣草属多年生草本植物,也是蜈蚣草属中唯一可用作草坪草的物种。它具有强壮的匍匐茎、蔓延力强而迅速、茎杆斜伸、叶形优美、植株低矮、养护水平低、耐贫瘠、病虫害少等优点,广泛用于庭院草坪、休憩草坪以及水土保持草坪建设中。然而,由于假俭草抗寒性相对较弱且枯黄期长,限制了其在亚热带北部及温带地区的广泛应用。因此,提高优良假俭草品种的抗寒性对扩大其应用范围具有重要作用。
本文在对165份中国假俭草种质资源坪用价值观测和综合分析的基础上,以优良选系E-126为材料,一方面,在以种子和侧芽为外植体建立再生体系的基础上,在低温胁迫下筛选抗寒突变体,以对假俭草进行遗传改良；另一方面,以尿素等化肥和6-BA等生长调节剂处理假俭草,通过栽培措施以提高其抗寒性,延长青绿期。全文结果如下；
1.本文在对165份中国假俭草种质资源的坪用价值、花序密度及抗病性进行系统观测分析的基础上,筛选出优良野生种质资源18份,并进一步以'Common'和'TifBlair'为对照,对其坪用价值、花序密度、花叶病毒抗性以及综合草坪质量进行了比较分析,为我国假俭草种质资源的利用提供依据。结果表明：(ⅰ)假俭草花序密度平均值为3.68,变异系数达29.5%,为变异幅度最大的性状；假俭草对花叶病毒的平均抗性为6.83,变异系数(CV)为16.7%,其中,春季花叶病毒抗性变异幅度明显高于夏季；假俭草的坪用价值平均为4.85,其冷季坪用价值变异系数是暖季坪用价值变异系数的2倍；作为营养繁殖型的假俭草,综合草坪质量平均为22.7,种源间变异系数10.0%,作为种子繁殖型假俭草,其综合草坪质量平均为20.3,种源间变异系数为12.1%；(ⅱ)作为营养繁殖型的假俭草种源,E-126等12份种源的综合草坪质量均明显高于对照；而作为种子繁殖型的种源,只有E022的综合草坪质量明显高于对照。
2.用NaOH、H2SO4、KNO3、磨破种皮和低温处理等方法对假俭草种子进行萌发试验,结果表明：以5%的NaOH处理15min效果最好,发芽率、发芽势和发芽指数分别为97.5%、72.5%和45.4,分别比对照高77.5%、72.5%、43.1%,发芽时间较对照提前4d。以E-126假俭草的侧芽和种子分别为外植体建立高效的再生体系,试验结果显示：适宜于侧芽生长、愈伤组织诱导、绿苗分化、试管苗生长和生根的培养基分别为MS+6-BA2.0mg/L+NAA0.8mg/L、MS+2,4-D1.0mg/L+6-BA0.1mg/L、MS+KT2.0mg/L、MS+6-BA2.0mg/L+NAA0.8mg/L和MS+NAA0.6mg/L;适宜于种子愈伤组织诱导、绿苗分化和生根的培养基分别为MS+2,4-D1.0mg/L+30g/L甘氨酸+椰子汁50ml/L、MS+KT2.0mg/L+椰子汁50ml/L和MS+NAA0.6mg/L+椰子汁50m1/L,本研究为开展假俭草体细胞突变体筛选和遗传转化提供基础和技术支持。
3.抗寒性弱是限制假俭草广泛应用的主要因素之一。本研究以E-126为材料,经低温诱导,筛选体细胞抗寒突变体。结果表明：愈伤组织在继代培养的过程中,经0℃的低温条件下培养26d,获得了2块存活的愈伤组织,该愈伤组织经过继代增殖后,进行分化、生根和移栽,获得株系1和株系2。苗期和成坪期外部形态观察和测量结果表明：假俭草低温诱导的株系1、株系2和对照在匍匐茎的节间长、节间宽、茎色、叶长、叶宽、叶色、叶毛上均没有显著性差异；株系1、株系2和对照的LTso分为-6.646℃、-6.546℃和-5.351℃,处理与对照之间达到显著性差异,而株系1和株系2之间无显著性差异；SRAP的分子鉴定表明：假俭草低温诱导的株系1和株系2在110bp、230bp以及240bp处均存在相同特征带,而对照没有这些条带,表明假俭草体细胞突变体植株的变异稳定且在分子水平与对照存在差异,但株系1和株系2无差异。因此,株系1和株系2可作为同一体细胞抗寒突变体株系加以利用。
4.用尿素(0、5、10、15和20g/L)、NaH2PO4(0、2、4、8和12g/L)、KN03(0、2.5、5、7.5和10g/L)、EDTA-Fe(0、1、5、10和15ml/L)、6-BA(0、5、10、15和20mg/L)和比久(B9)(0、1、3、5和7g/L)分别对盆栽E-126材料进行处理,研究其对假俭草的抗寒性和青绿期的影响。结果表明,在低温条件下,处理与对照相比,不仅降低了电导率,提高了叶绿素的含量、抑制了叶绿素的分解,而且提高了可溶性糖和脯胺酸的含量。效果最好的分别为尿素10g/L、NaH2PO48g/L、KNO37.5g/L、EDTA-Fe10ml/L、6-BA10mg/L和比久1g/L,均能显著提高假俭草的抗寒性和青绿期,其青绿期可分别延长11～12d、16d、12-13d、15d、16-17d和4～5d。在所有的处理中10mg/L6-BA效果最好,假俭草的青绿期可延长16-17d。
Centipedegrass (Eremochloa ophiuroides (Munro.) Hack), with strong stolons, beautiful leaves and dwarf swards, is the only one species of Eremochloa ssp. used as turfgrass (Bouton,1983; Hanna,1995). Due to low level of maintenance requirement and good adaptation to poor soil, centipedegrass has been widely used in the lawn of home yard, recreational turf and soil conservation turf from subtropical to tropical zones (Beard,1973; Hanson et al.,1969). However, the cold-resistance of centipedegrass is weaker than that of bermudagrass and zoysiagrass, which limits its wide application in the North of subtropical zone and temperate zone (Cardona et al.,1997; Fry et al.,1993; Fry et al.,1991; Philley et al.,1995). Therefore, the improvement and regulation of cold-resistance of centipedegrass is of great importance to its wide application.
Based comprehensive turf quality evaluation of165native accessions of centipedegrass, the high quality accession E126was selected to establish the regeneration system with axillary buds and seeds as the explant for its cold resistance improvement. The callus of E-126were screened at low temperature, and the survival callus were regererated and the cold resistance and difference of molecular marker of the variant and check were tested; at the same time, E126were treated with fertilizers and regulators to improve the cold resistance and extend the green period. The main result is stated as followings:
The turf quality, inflorescence density and the mosiac virus resistance of165accessions of Eremochloa ophiuroides are observed, and the turf quality, inflorescence densitythe, mosiac virus resistance and comprehensive turf quality of18good performance accessions are compared with'Common'and'TifBlair'as the checks to provide the experimental basis for the Chinese germplasm utilization of Eremochloa ophiuroides. The result shows:(i) Average inflorescence density of Eremochloa ophiuroides is6.2with CV as29.5%, which are ranked as the greatest variation characteristics; average mosiac virus resistance is6.83with CV as16.7%, in which CV in spring is greater than CV in summer; the average turf quality is4.85, the CV of turf quality during cool season is twice of the CV of turf quality during warm season; the average comprehensive turf quality is22.7with the CV as10.0%when the accessions are used as vegetative propagation, whereas the average comprehensive turf quality is20.3with the CV as12.1%when the accessions are used as seed propagation.(ii) the comprehensive turf quality of12accessions including E126is significantly higher than that of checks when used as vegetative propagation, whereas only the comprehensive turf quality of one accession E022is obvious higher
Experiments of centipedegrass seeds treated with NaOH, H2SO4, KNO3, chafe or low temperature were progressed in laboratory. Results indicated that the seeds germinated best when they were treated with5%NaOH for15min,germination rate, germination potentialand Germination index of seed respectively were97.5%,72.5%and45.36, which respectively increased by77.5%,72.5%and43.10compared with checks. The plant regeneration system was established with the axillary bud and seed of Chinese selection E-126centipedegrass as explant. The optimum medium of growing axillary bud, callus induction, callus differentiation, shoot growth, plantlets rooting respectively were MS medium with6-BA2.0mg/L+NAA0.8mg/L,2,4-D1.0mg/L+6-BA0.1mg/L, KT2.0mg/L,6-BA2.0mg/L+NAA0.8mg/L and NAA0.6mg/L; the optimum medium of seed callus induction, callus differentiation, plantlets rooting respectively were MS medium with2,4-D1.0mg/L+mannitol30g/L+coconut milk50ml/L, KT2.0mg/L+coconut milk50ml/L and NAA0.6mg/L+coconut milk50ml/L. In conclusion, the plant regeneration system was successfully developed in this study, hopefully providing basic reference for screening of somatic variants and genetic transformation of centipedegrass.
Poor cold-resistance of centipedegrass is one of the major factors limiting its wide application. The objective of this study is to obtain cold-resistant somatic mutants from the seed of eminent Chinese native centipedegrass selection E-126by induction and screening at low temperature. The callus induced by seeds of centipedegrass was cultivated at the low temperature of0℃for twenty-six days during the process of subculture and two pieces of survival callus tissues were obtained. Strain1and Strain2were obtained after survival callus subculture proliferation, differentiation, rhizogenesis as well as transplantation. No significant differences were found in leaf color, lamellar hair, leaf length and leaf width among Strainl, Strain2and the control. LT50of Strainl and Strain2were-6.646℃and-6.546℃, respectively, which was significantly lower that of control (LT50=-5.351℃), no significant differences were demonstrated between LT50of Strain1and Strain2. The results of sequence related amplified polymorphism (SRAP) markers demonstrated that there were identical characteristic bands at110bp,230bp and240bp of centipedegrass Strain1and Strain2obtained through low temperature induction, however, no characteristic bands at110bp,230bp and240bp were found in control plant, which indicated that the somatic mutants of centipedegrass were genetically stable comparing with that of the control plant on molecular level. Strain1and Strain2could be utilized as the identical strain.
The new selection of E-126was used to respectively study the effect of carbamide (0,5,10,15and20g/L), NaH2PO4(0,2,4,8and12g/L), KNO3(0,2.5,5,7.5and10g/L), EDTA-Fe (0,1,5,10and15ml/L),6-BA(0,5,10,15and20mg/L) and B9(0,1,3,5and7g/L) on the coldness and green period of centipedegrass during the cold season. The result showed:(1) the relative electrical conductivity and chlorophyll decomposition of centipedegrass were reduced, whereas the content of soluble sugar and proline in centipedegrass leaf were obviously increased under the treatment compared with the check, which means that the treatment could improve the coldness and performance of centipedegrass during the cold season.;(2)10g/L carbamide,8g/L NaH2PO4,7.5g/L KNO3,10ml/L EDTA-Fe,10mg/L6-BA and1g/L B9respectively treated resulted in the best result and could respectively prolong the green period by11-12,16,12-13,15,16-17 and4-5days compared with the check. The result with10mg/L6-BA treatmented was the best in all treatments and prolong the green period by16-17days compared with the check.
本文在对165份中国假俭草种质资源坪用价值观测和综合分析的基础上,以优良选系E-126为材料,一方面,在以种子和侧芽为外植体建立再生体系的基础上,在低温胁迫下筛选抗寒突变体,以对假俭草进行遗传改良；另一方面,以尿素等化肥和6-BA等生长调节剂处理假俭草,通过栽培措施以提高其抗寒性,延长青绿期。全文结果如下；
1.本文在对165份中国假俭草种质资源的坪用价值、花序密度及抗病性进行系统观测分析的基础上,筛选出优良野生种质资源18份,并进一步以'Common'和'TifBlair'为对照,对其坪用价值、花序密度、花叶病毒抗性以及综合草坪质量进行了比较分析,为我国假俭草种质资源的利用提供依据。结果表明：(ⅰ)假俭草花序密度平均值为3.68,变异系数达29.5%,为变异幅度最大的性状；假俭草对花叶病毒的平均抗性为6.83,变异系数(CV)为16.7%,其中,春季花叶病毒抗性变异幅度明显高于夏季；假俭草的坪用价值平均为4.85,其冷季坪用价值变异系数是暖季坪用价值变异系数的2倍；作为营养繁殖型的假俭草,综合草坪质量平均为22.7,种源间变异系数10.0%,作为种子繁殖型假俭草,其综合草坪质量平均为20.3,种源间变异系数为12.1%；(ⅱ)作为营养繁殖型的假俭草种源,E-126等12份种源的综合草坪质量均明显高于对照；而作为种子繁殖型的种源,只有E022的综合草坪质量明显高于对照。
2.用NaOH、H2SO4、KNO3、磨破种皮和低温处理等方法对假俭草种子进行萌发试验,结果表明：以5%的NaOH处理15min效果最好,发芽率、发芽势和发芽指数分别为97.5%、72.5%和45.4,分别比对照高77.5%、72.5%、43.1%,发芽时间较对照提前4d。以E-126假俭草的侧芽和种子分别为外植体建立高效的再生体系,试验结果显示：适宜于侧芽生长、愈伤组织诱导、绿苗分化、试管苗生长和生根的培养基分别为MS+6-BA2.0mg/L+NAA0.8mg/L、MS+2,4-D1.0mg/L+6-BA0.1mg/L、MS+KT2.0mg/L、MS+6-BA2.0mg/L+NAA0.8mg/L和MS+NAA0.6mg/L;适宜于种子愈伤组织诱导、绿苗分化和生根的培养基分别为MS+2,4-D1.0mg/L+30g/L甘氨酸+椰子汁50ml/L、MS+KT2.0mg/L+椰子汁50ml/L和MS+NAA0.6mg/L+椰子汁50m1/L,本研究为开展假俭草体细胞突变体筛选和遗传转化提供基础和技术支持。
3.抗寒性弱是限制假俭草广泛应用的主要因素之一。本研究以E-126为材料,经低温诱导,筛选体细胞抗寒突变体。结果表明：愈伤组织在继代培养的过程中,经0℃的低温条件下培养26d,获得了2块存活的愈伤组织,该愈伤组织经过继代增殖后,进行分化、生根和移栽,获得株系1和株系2。苗期和成坪期外部形态观察和测量结果表明：假俭草低温诱导的株系1、株系2和对照在匍匐茎的节间长、节间宽、茎色、叶长、叶宽、叶色、叶毛上均没有显著性差异；株系1、株系2和对照的LTso分为-6.646℃、-6.546℃和-5.351℃,处理与对照之间达到显著性差异,而株系1和株系2之间无显著性差异；SRAP的分子鉴定表明：假俭草低温诱导的株系1和株系2在110bp、230bp以及240bp处均存在相同特征带,而对照没有这些条带,表明假俭草体细胞突变体植株的变异稳定且在分子水平与对照存在差异,但株系1和株系2无差异。因此,株系1和株系2可作为同一体细胞抗寒突变体株系加以利用。
4.用尿素(0、5、10、15和20g/L)、NaH2PO4(0、2、4、8和12g/L)、KN03(0、2.5、5、7.5和10g/L)、EDTA-Fe(0、1、5、10和15ml/L)、6-BA(0、5、10、15和20mg/L)和比久(B9)(0、1、3、5和7g/L)分别对盆栽E-126材料进行处理,研究其对假俭草的抗寒性和青绿期的影响。结果表明,在低温条件下,处理与对照相比,不仅降低了电导率,提高了叶绿素的含量、抑制了叶绿素的分解,而且提高了可溶性糖和脯胺酸的含量。效果最好的分别为尿素10g/L、NaH2PO48g/L、KNO37.5g/L、EDTA-Fe10ml/L、6-BA10mg/L和比久1g/L,均能显著提高假俭草的抗寒性和青绿期,其青绿期可分别延长11～12d、16d、12-13d、15d、16-17d和4～5d。在所有的处理中10mg/L6-BA效果最好,假俭草的青绿期可延长16-17d。
Centipedegrass (Eremochloa ophiuroides (Munro.) Hack), with strong stolons, beautiful leaves and dwarf swards, is the only one species of Eremochloa ssp. used as turfgrass (Bouton,1983; Hanna,1995). Due to low level of maintenance requirement and good adaptation to poor soil, centipedegrass has been widely used in the lawn of home yard, recreational turf and soil conservation turf from subtropical to tropical zones (Beard,1973; Hanson et al.,1969). However, the cold-resistance of centipedegrass is weaker than that of bermudagrass and zoysiagrass, which limits its wide application in the North of subtropical zone and temperate zone (Cardona et al.,1997; Fry et al.,1993; Fry et al.,1991; Philley et al.,1995). Therefore, the improvement and regulation of cold-resistance of centipedegrass is of great importance to its wide application.
Based comprehensive turf quality evaluation of165native accessions of centipedegrass, the high quality accession E126was selected to establish the regeneration system with axillary buds and seeds as the explant for its cold resistance improvement. The callus of E-126were screened at low temperature, and the survival callus were regererated and the cold resistance and difference of molecular marker of the variant and check were tested; at the same time, E126were treated with fertilizers and regulators to improve the cold resistance and extend the green period. The main result is stated as followings:
The turf quality, inflorescence density and the mosiac virus resistance of165accessions of Eremochloa ophiuroides are observed, and the turf quality, inflorescence densitythe, mosiac virus resistance and comprehensive turf quality of18good performance accessions are compared with'Common'and'TifBlair'as the checks to provide the experimental basis for the Chinese germplasm utilization of Eremochloa ophiuroides. The result shows:(i) Average inflorescence density of Eremochloa ophiuroides is6.2with CV as29.5%, which are ranked as the greatest variation characteristics; average mosiac virus resistance is6.83with CV as16.7%, in which CV in spring is greater than CV in summer; the average turf quality is4.85, the CV of turf quality during cool season is twice of the CV of turf quality during warm season; the average comprehensive turf quality is22.7with the CV as10.0%when the accessions are used as vegetative propagation, whereas the average comprehensive turf quality is20.3with the CV as12.1%when the accessions are used as seed propagation.(ii) the comprehensive turf quality of12accessions including E126is significantly higher than that of checks when used as vegetative propagation, whereas only the comprehensive turf quality of one accession E022is obvious higher
Experiments of centipedegrass seeds treated with NaOH, H2SO4, KNO3, chafe or low temperature were progressed in laboratory. Results indicated that the seeds germinated best when they were treated with5%NaOH for15min,germination rate, germination potentialand Germination index of seed respectively were97.5%,72.5%and45.36, which respectively increased by77.5%,72.5%and43.10compared with checks. The plant regeneration system was established with the axillary bud and seed of Chinese selection E-126centipedegrass as explant. The optimum medium of growing axillary bud, callus induction, callus differentiation, shoot growth, plantlets rooting respectively were MS medium with6-BA2.0mg/L+NAA0.8mg/L,2,4-D1.0mg/L+6-BA0.1mg/L, KT2.0mg/L,6-BA2.0mg/L+NAA0.8mg/L and NAA0.6mg/L; the optimum medium of seed callus induction, callus differentiation, plantlets rooting respectively were MS medium with2,4-D1.0mg/L+mannitol30g/L+coconut milk50ml/L, KT2.0mg/L+coconut milk50ml/L and NAA0.6mg/L+coconut milk50ml/L. In conclusion, the plant regeneration system was successfully developed in this study, hopefully providing basic reference for screening of somatic variants and genetic transformation of centipedegrass.
Poor cold-resistance of centipedegrass is one of the major factors limiting its wide application. The objective of this study is to obtain cold-resistant somatic mutants from the seed of eminent Chinese native centipedegrass selection E-126by induction and screening at low temperature. The callus induced by seeds of centipedegrass was cultivated at the low temperature of0℃for twenty-six days during the process of subculture and two pieces of survival callus tissues were obtained. Strain1and Strain2were obtained after survival callus subculture proliferation, differentiation, rhizogenesis as well as transplantation. No significant differences were found in leaf color, lamellar hair, leaf length and leaf width among Strainl, Strain2and the control. LT50of Strainl and Strain2were-6.646℃and-6.546℃, respectively, which was significantly lower that of control (LT50=-5.351℃), no significant differences were demonstrated between LT50of Strain1and Strain2. The results of sequence related amplified polymorphism (SRAP) markers demonstrated that there were identical characteristic bands at110bp,230bp and240bp of centipedegrass Strain1and Strain2obtained through low temperature induction, however, no characteristic bands at110bp,230bp and240bp were found in control plant, which indicated that the somatic mutants of centipedegrass were genetically stable comparing with that of the control plant on molecular level. Strain1and Strain2could be utilized as the identical strain.
The new selection of E-126was used to respectively study the effect of carbamide (0,5,10,15and20g/L), NaH2PO4(0,2,4,8and12g/L), KNO3(0,2.5,5,7.5and10g/L), EDTA-Fe (0,1,5,10and15ml/L),6-BA(0,5,10,15and20mg/L) and B9(0,1,3,5and7g/L) on the coldness and green period of centipedegrass during the cold season. The result showed:(1) the relative electrical conductivity and chlorophyll decomposition of centipedegrass were reduced, whereas the content of soluble sugar and proline in centipedegrass leaf were obviously increased under the treatment compared with the check, which means that the treatment could improve the coldness and performance of centipedegrass during the cold season.;(2)10g/L carbamide,8g/L NaH2PO4,7.5g/L KNO3,10ml/L EDTA-Fe,10mg/L6-BA and1g/L B9respectively treated resulted in the best result and could respectively prolong the green period by11-12,16,12-13,15,16-17 and4-5days compared with the check. The result with10mg/L6-BA treatmented was the best in all treatments and prolong the green period by16-17days compared with the check.
引文
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