热带作物学报

剑麻斑马纹病抗病新品种——热麻1号剑麻

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张燕梅 , 李俊峰 , 陆军迎 , 赵艳龙 , 鹿志伟 , 杨子平 , 柯智 , 周文钊 ^*

热带作物学报 | 种质资源与遗传育种 2024,45(7): 1371-1377

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热带作物学报 |种质资源与遗传育种 2024 , 45 (7) : 1371 -1377

剑麻斑马纹病抗病新品种——热麻1号剑麻

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张燕梅（1975—），女，博士，副研究员，研究方向：作物育种。

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tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276598009781686834, companyId=1276598323452703149, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国热带农业科学院南亚热带作物研究所/海南省热带作物营养重点实验室/湛江市热带作物遗传改良重点实验室，广东湛江　524091)])])]

张燕梅, 李俊峰, 陆军迎, 赵艳龙, 鹿志伟, 杨子平, 柯智, 周文钊^*

作者信息

中国热带农业科学院南亚热带作物研究所/海南省热带作物营养重点实验室/湛江市热带作物遗传改良重点实验室，广东湛江　524091

通讯作者:

^* 周文钊（ZHOU Wenzhao），E-mail：zwenzhao@163.com。

A New Variety Rema No. 1 of Anti—zero Disease

Yanmei ZHANG, Junfeng LI, Junying LU, Yanlong ZHAO, Zhiwei LU, Ziping YANG, Zhi KE, Wenzhao ZHOU^*

Affiliations

South Subtropical Crops Research Institute, Chinese Academy of Tropical Agricultural Sciences / Key Laboratory of Tropical Crops Nutrition of Hainan Province / Zhanjiang City Key Laboratory for Tropical Crops Genetic Improvement, Zhanjiang, Guangdong 524091, China

出版时间: 2024-07-25 doi: 10.3969/j.issn.1000-2561.2024.07.009

文章导航

摘要

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斑马纹病是剑麻的主要病害之一，严重影响了剑麻产业的发展，目前关于斑马纹病的致病机制尚不清楚。为了培育出剑麻斑马纹病抗病新品种，本研究分别以剑麻斑马纹病抗病品种粤西114和感病品种H.11648为父母本进行回交，获得回交后代热麻1号剑麻。利用常规方法对热麻1号剑麻叶形、叶色、叶缘刺有无、叶缘刺类型、叶顶刺类型、叶长、叶宽、叶基厚度、顶刺长、顶刺基部宽、株高、花序形状、花轴高、花梗数、花、果实、种子、年展叶数、周期展叶数、生命周期、纤维率等进行分析，采用壁低渗火焰干燥法、流式细胞术结合菌斑接种法对热麻1号剑麻倍性、基因组大小以及对烟草疫霉的抗性等进行鉴定。结果表明：热麻1号剑麻叶片剑形、肉质、刚硬、叶片灰绿，叶面平整有蜡粉，叶顶有1.2~2.0 cm锐刺，叶缘无刺，叶片绕茎呈螺旋状排列上升；叶长约130 cm，叶宽约16 cm，叶基厚度约3.5 cm。株高1.8~2.5 m，圆锥形花序，花轴高3~7 m，花梗数20~32个，完全花、花黄色、花瓣6枚；蒴果、长圆形、成熟时黑褐色，种子半月形，黑色、纸质；周期展叶360~400片；纤维率约3.5%，纤维长约100 cm；生命周期6~9 a；热麻1号剑麻为四倍体（2X=4n），染色体数目为104~118，2C DNA含量为11.69 pg，对烟草疫霉高度抗病或免疫，有望成为剑麻斑马纹病发病区的先锋品种。

关键词

热麻1号剑麻 / 斑马纹病 / 抗性

Abstract

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The sisal zebra disease, caused by fungal pathogen Phytophthora nicotianae Breda, is one of the main diseases in sisal, which seriously affects the development of sisal industry. The pathogenic mechanism is not clear up to now. In order to obtain new anti-zero disease sisal variety, Yuexi 114 and H.11648 were used as the parents to breed new offsprings. Rema No.1 is one of the backcross progenies of Yuexi 114 and H.11648. Morphological traits including leaf shape, leaf color, have/none marginal spine, apical spine type, leaf length, leaf width, thickness of leaf base, length of apical spine, and spine basal width, plant height, inflorescence shape and flower stalk height, number of pedical, flower, fruit and seed characters, number of leaf, fiber ratio were examined. Ploidy level, genome size and resistance to P. nicotianae Breda was determined by an analysis of wall removal and low permeability flame drying method, flow cytometry, and inoculating P. nicotianae Breda in field and lab respectively. Rema No.1 is characterized by the presence of gray green, fleshier and rigid, sword-leaf with wax powder on the leaf surface, 1.2-2.0 cm apical spine, none marginal spine. Leaf length, leaf width and leaf base is about 130 cm, 16 cm, and 3.5 cm, respectively. The leaves rise in a spiral arrangement around the stem. Rema No.1 is also characterized by plant height 1.8-2.5 m, flower stalk 3-7 m, paniculate inflorescence, with 20-32 pedicals, complete flower, yellow flower with 6 petals, oblong capsule, dark brown to almost black upon maturity, lunate black seeds, marginate. leaves 360-400, leaf fiber ratio 3.5%, fiber length 100 cm. The life span is 6-9 years. Rema No1 is tetraploid (2X=4n), chromosome number is between 104-118, the genome size of Rema No. 1 is 2C=11.69 pg. Rema No.1 is high resistant to Phytophthora nicotianae Breda, and would be the main cultures in high incidence area of zero leaf disease in sisal planting region.

Key words

Rema No.1 / zero disease / resistance

引用本文

张燕梅, 李俊峰, 陆军迎, 赵艳龙, 鹿志伟, 杨子平, 柯智, 周文钊. 剑麻斑马纹病抗病新品种——热麻1号剑麻. 热带作物学报, 2024 , 45 (7) : 1371 -1377 . DOI: 10.3969/j.issn.1000-2561.2024.07.009

Yanmei ZHANG, Junfeng LI, Junying LU, Yanlong ZHAO, Zhiwei LU, Ziping YANG, Zhi KE, Wenzhao ZHOU. A New Variety Rema No. 1 of Anti—zero Disease[J]. Chinese Journal of Tropical Crops, 2024 , 45 (7) : 1371 -1377 . DOI: 10.3969/j.issn.1000-2561.2024.07.009

正文

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剑麻是重要的热带经济作物之一，其纤维质地坚韧、富于弹性、拉力强、抗撕裂、耐磨、防腐以及耐低温等，广泛应用于国防、渔业、交通运输以及冶金等领域^[1-2]，随着材料科学技术的飞速发展，通过添加剑麻纤维改进新型复合材料及其衍生材料的性能^[3]，扩大复合材料在军事以及多种民用制造业中的应用，对推动我国经济快速发展起到了非常重要的作用。同时剑麻在食品科学^[4-6]、医学^[7]、生物质能源^[8]以及作物遗传改良^[9]等方面应用广泛，因此，加快剑麻种质创新，对推动我国经济长期稳定发展具有重要的经济价值。

斑马纹病是剑麻的主要病害之一，主要由烟草疫霉（Phytophthora nicotianae Breda）引起，因斑马纹病导致大面积麻园被毁，纤维产量下降，严重影响了国内外剑麻产业的发展^[10-11]。国内外学者对该病进行了大量研究，目前已形成了一套完整的病原菌分离、鉴定和防治技术^[12]。同时，在剑麻种质资源抗性鉴定、抗斑马纹病生理生化机制、功能基因挖掘以及转基因等方面也开展了研究^[13-17]，取得了一定进展，但抗病机制和致病机理仍不清楚。由于长期大量使用化学药剂对环境和生物多样性破坏严重，因此，培育出抗斑马纹病新品种，仍然是育种学家们一直以来的首要目标。20世纪80年代初，育种学家们以主栽品种H.11648为母本，普通剑麻为父本，通过有性杂交培育出来F₁代杂交品种粤西114，粤西114的特点是抗斑马纹病，对茎腐病也有一定抗性，但该品种叶基厚、叶缘有刺、生命周期短，在后续试验中发现对斑马纹病抗性不稳定^[18]。为了稳定和提高粤西114的优良性状，本研究利用粤西114与H.11648回交，获得杂交F₂代，通过对杂交F₂代叶缘刺有无、叶长、叶宽、株高、叶基厚度、年产叶数、周期产叶数、纤维率、抗病性、抗寒性等多个性状综合评价，初步筛选出叶缘无刺、长势良好、抗性比亲本粤西114和H.11648强的优良单株、然后在广东、广西等剑麻主产区开展品比试验和区域性试验，进一步筛选出了综合长势良好，叶缘无刺、对斑马纹病高度抗病的剑麻新品种——热麻1号剑麻。该品种2015年获得广东省品种登记权。由于热麻1号剑麻生命周期比主栽品种H.11648短，周期产业数比主栽品种H.11648少，热麻1号剑麻目前常用于剑麻斑马纹病病区补植品种和优良杂交亲本，还未广泛推广应用。

1　材料与方法

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1.1　材料

H.11648、粤西114和热麻1号剑麻保存在中国热带农业科学院南亚热带作物研究所剑麻种质圃。Zea mays cv. CE777玉米种子由Doležel友情提供。

1.2　方法

1.2.1　亲本选择及杂交

以斑马纹病抗病品种粤西114为母本，以主栽品种H.11648为父本进行人工杂交，待果实成熟后取出种子并播种育苗，植株生长到30 cm左右，根据植株的表型特征如叶缘有/无刺、叶长、叶宽、叶片数、株型、叶基厚度等，结合育种目标初步筛选出候选材料，作为进一步鉴定的优株。

1.2.2　形态学特征观测

随机选取10株成龄剑麻，每株3片叶片，对热麻1号剑麻株型、叶片伸展模式、叶形、叶色、叶缘刺有无、叶缘刺类型、叶顶刺类型、叶片有无蜡粉、叶长、叶宽、叶基厚度、顶刺长、顶刺基部宽、株高、花轴高、花梗数、花、果实、种子等性状进行观测，其中质量性状用肉眼观测，数量性状用卷尺进行测定。叶长为叶基部到顶刺的距离（包括顶刺），叶宽为叶片最宽处的宽度，株高为植株基部到叶片顶端的高度，株高、花轴高、花梗数、花色、花序形状等在植株开花后测量，果实以及种子等数据在果实成熟时收集。所有数据用Excel软件进行分析统计。

1.2.3　纤维特性测定

待叶片成熟后，收割叶片，称量鲜叶重量，然后加工收获纤维，待纤维晾干后称取干纤维重量，计算纤维率，同时用拉力测量仪测量纤维拉力大小。纤维率=干纤维重/鲜叶重×100%。

1.2.4　斑马纹病抗性鉴定

大田抗病性测定：以热麻1号剑麻为试验对象，以H.11648和粤西114为对照，在雨季来临前种植于斑马纹病重病发病区，雨季过后调查斑马纹病发病情况，具体参考赵艳龙等^[12]的方法进行。室内抗病性鉴定：采用盆栽和离体叶片2种接种方式，取斑马纹病病菌接种于提前准备好的H.11648、粤西114和热麻1号剑麻叶片上，用棉花保湿，分别在接种24、48、72 h和7 d测定病班大小，具体参照张燕梅等^[2]的方法进行。

1.2.5　根尖染色体检测

分别取热麻1号剑麻、H.11648和粤西114根尖约1 cm，先用0.002 mol/L浓度的8-羟基喹啉预固定2 h，然后用卡诺固定液（无水乙醇∶冰乙酸=3:1）固定过夜，用5%纤维素酶和果胶酶（纤维素酶∶果胶酶=1:1）37 ℃孵育1 h后快速涂片并火焰干燥，最后用5%Giemsa染色10~15 min，在显微镜（ZEISS Axiolmager M2）下观察染色体数目，并拍照保存。

1.2.6　基因组大小测定

取热麻1号、H.11648和粤西114组培苗叶片，以玉米（Zea mays cv. CE777）为标准样品（standard plant），用流式细胞仪（Partec）测定基因组大小，具体参照PALOMINO等^[19]的流式细胞仪方法测定，待测样品2C DNA含量（pg）=待测样品峰图位置对应的荧光强度/标准样品峰图位置对应的荧光强度值×标准样品2C DNA含量。

1.3　数据处理

病斑大小用直尺测量并用Excel软件分析数据和作图。

2　结果与分析

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2.1　表型特征及纤维特性

热麻1号剑麻植株高大，约1.9 m，叶片向上螺旋排列，灰绿色，肉质刚直，叶剑形，表面平整，有少量蜡粉，叶缘无刺，叶尖有1.6~2.0 cm黑褐色坚硬的锐刺，叶长120~135 cm，叶宽15~ 17 cm，叶基厚度约3.5 cm，年展叶45~55片，周期展叶数360~400片。圆锥花序，花轴高5.1~ 5.5 m，花梗数20~32个，完全花，花黄色，花被浅淡黄色，下部联合形成萼筒，上部分裂成6片花瓣，雌蕊1枚，柱头3裂，子房下位，自交结实。蒴果，长圆形，果实大小约5 cm×6 cm，早期为绿色，成熟后为黑褐色。每个果实里约300枚种子，种子半月形，成熟后为黑色，纸质扁平，有光泽。生命周期为6~9 a，叶片纤维含量为3.5%，纤维细而均匀，长度为110 cm，纤维拉力为66.2 kg、束纤维强力为649 N（图1）。

2.2　倍性和基因组大小测定

根尖染色体观测表明，热麻1号剑麻染色体数目为104~118，长染色体和中等长度的染色体共计20条，其余染色体为棒状或点状，为四倍体。H.11648染色体数目为48~60条，长染色体和中等长度的染色体共10条，为二倍体。粤西114染色体数目为101~112条，其中长染色体和中等长度的染色体共20条，为四倍体（图2）。流式细胞仪分析显示，热麻1号剑麻、H.11648和粤西114分别在荧光强度402、244和403处出现了峰图，分别为四倍体、二倍体和四倍体，基因组大小2C分别为11.69、7.07、11.74 pg（图3）。

2.3　斑马纹病抗性检测

大田抗病试验表明，H.11648、粤西114和热麻1号剑麻抗病指数分别为78.1、63.2和22.4，对斑马纹病菌表现为高度感病，中度感病和高度抗病^[12]。盆栽试验和离体叶片接种后发现，H.11648病斑最大，2019—2021年连续3 a接种烟草疫霉，每年接种1周后观测发现病斑大小均超过15 cm，粤西114次之，2019年约10 cm，2021年约15 cm，2020年约2 cm，抗性极不稳定，热麻1号剑麻除伤口处有与伤口相同大小的约0.6 cm褐斑外，随着接种时间延长，病斑没有进一步扩展，检测结果与大田试验一致，且2019、2020和2021年接种结果均不变，即热麻1号剑麻对烟草疫霉的抗性远超过亲本H.11648和粤西114，对斑马纹病菌表现为高度抗病（图4）。

3　讨论

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培育抗病高产剑麻新品种一直是剑麻育种的首要目标，采取的方式方法也多种多样，比如人工杂交、化学诱变、细胞融合、单倍体育种、基因工程以及基因编辑等，其中人工杂交是较为常用的方法之一，在此过程中，亲本的选择尤为重要。H.11648是目前剑麻产业中唯一的主栽品种，平均年长叶60~70片，周期长叶560~650片，纤维率5%以上，生长周期10 a以上，综合性状优良，常常作为重要的亲本材料广泛应用于剑麻育种，但H.11648易感斑马纹病。粤西114是H.11648与普通剑麻的杂交F₁代，通过长期试验观察发现，该品种对斑马纹病菌表现为较强的抗病性，但抗性不稳定，因此，利用粤西114和H.11648为亲本进行杂交，有望获得抗病高产的新品种。热麻1号剑麻是粤西114和H.11648回交子代，对斑马纹病菌高度抗病甚至免疫，并且抗性稳定，此外，热麻1号剑麻耐旱耐寒，速生粗长，本研究室前期在云南元谋干热河谷区的品种试验发现，热麻1号剑麻在云南干热河谷区生长较其他品种要好，因此，可以广泛应用于云南干热河谷区生态恢复。由于热麻1号剑麻生产周期短，纤维含量比H.11648低，目前常作为抗性杂交亲本使用，暂时还未推广应用。随着育种工作的不断推进，目前已从热麻1号剑麻的自交子代和杂交后代中相继筛选出部分优良单株，进一步的品种区域试验正在进行，有望获得产量高、抗性强的剑麻优良品种，因此，有一定的应用前景。

根尖分生组织细胞生长旺盛，常用于染色体观察，并且相对于流式细胞仪倍性分析而言，染色体观察能更直观地观察到染色体数目，形态特征等。本研究结果显示粤西114染色体数目为102~114条，根据中长染色体数目，结合流式细胞仪分析结果判定为四倍体，染色体形态正常，未检测到三倍体和丝状小染色体，这与LV等^[20]研究结果有一定差异，可能与剑麻复杂的倍性，以及自然条件下容易变异有关^[21-24]。建议扩大观测样本数量，防止实验结果出现偏差。此外，粤西114的母本H.11648为二倍体，父本普通剑麻为五倍体，有望获得三倍体和四倍体的子代粤西114。四倍体的粤西114和二倍体的H.11648回交获得四倍体热麻1号剑麻，一方面说明剑麻遗传的复杂性，同时也证实了剑麻多倍体和非整倍体广泛存在，变异频繁而且比例还很高的观点^[21-22]，因此，建议在杂交亲本选择时，尽量不选或少选多倍体。

流式细胞术作为一种高效快速的倍性鉴定和DNA含量分析方法，在剑麻种质资源鉴定与评价中应用广泛^[23-24]。该方法能快速直观地检测出目标样本的倍性，而且灵敏度高，尤其对于大样本，省时省力。由于流式细胞术无法直观展示染色体的核型特征，染色体是否丢失或结构发生变化等信息，建议在进行分析时，将流式检测技术与常规的染色体制片技术相结合，进一步分析染色体形态结构特征及倍性。

前人研究表明剑麻染色体基数为30，其中5条长染色体，25条相对小染色体，随着倍性增加，染色体条数成比例增加^[20]。热麻1号剑麻和粤西114染色体数目存在较大出入，一方面可能是实验操作过程中小染色体降解或丢失，也可能是不正常减数分裂，如丢失、交换等，或非整倍体存在等多种因素所致，同时这也会导致在加倍过程中基因组变小^[25]。ROBERT等^[21]以洋葱（Allium cepa cv. Ailsa Craig）为参考基因组，对H.11648等龙舌兰属剑麻7个品种进行基因组大小分析，结果显示H.11648基因组4C DNA为15.23 pg，本研究以玉米（Zea mays cv. CE777）为参考基因组，显示H.11648基因组2C DNA为7.07 pg，另外，从理论而言，热麻1号剑麻和粤西114为四倍体，2C DNA为H.11648的2倍，约14.14 pg，实际分析结果分别为11.69 pg和11.74 pg，出入较大，可能与参考基因组的选择有关。MORENOSALAZAR^[26]、PALOMINO^[27]和ROBERT等^[21]分别在对Agave angustifolia Haw进行基因组大小分析时，以玉米（Zea mays cv. CE777）为参考基因组，2C DNA为8.499 pg^[26]和8.477 pg^[27]，以洋葱（Allium cepa cv. Ailsa Craig）为参考基因组，2C DNA为7.45 pg^[20]。DOLEŽEL等^[24]认为，为了降低试验结果误差，应选取与研究对象基因组大小相近的物种作为参考基因组。

基金

收起

国家麻类产业技术体系项目(CARS-16)
广东省现代农业产业技术体系创新团队建设项目(2023KJ104)
中央级公益性科研院所基本科研业务费专项(1630062023004)

参考文献

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2024年第45卷第7期

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doi: 10.3969/j.issn.1000-2561.2024.07.009

接收时间：2023-08-01
首发时间：2026-06-24
出版时间：2024-07-25

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收稿日期：2023-08-01
修回日期：2023-09-07

基金

国家麻类产业技术体系项目(CARS-16)

广东省现代农业产业技术体系创新团队建设项目(2023KJ104)

中央级公益性科研院所基本科研业务费专项(1630062023004)

作者信息

中国热带农业科学院南亚热带作物研究所/海南省热带作物营养重点实验室/湛江市热带作物遗传改良重点实验室，广东湛江　524091

通讯作者:

^* 周文钊（ZHOU Wenzhao），E-mail：zwenzhao@163.com。

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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