热带作物学报

橡胶树育种技术研究进展

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张源源 , 李维国 , 张晓飞 , 高新生 , 王祥军 , 位明明 , 黄肖 , 周元元 , 黄华孙

热带作物学报 | 种质资源与遗传育种 2025,46(12): 2877-2889

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热带作物学报 |种质资源与遗传育种 2025 , 46 (12) : 2877 -2889

橡胶树育种技术研究进展

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张源源（ZHANG Yuanyuan），男，博士，副研究员，研究方向：橡胶树遗传育种；E-mail：zhangyuanyuan@catas.cn。

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张源源（ZHANG Yuanyuan），男，博士，副研究员，研究方向：橡胶树遗传育种；E-mail：zhangyuanyuan@catas.cn。

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张源源, 李维国, 张晓飞, 高新生, 王祥军, 位明明, 黄肖, 周元元, 黄华孙

作者信息

中国热带农业科学院橡胶研究所/国家橡胶树育种中心/热带作物生物育种全国重点实验室/农业农村部橡胶树生物学与遗传资源利用重点实验室，海南海口　571101

Progress in Breeding Techniques of Rubber Tree

Yuanyuan ZHANG, Weiguo LI, Xiaofei ZHANG, Xinsheng GAO, Xiangjun WANG, Mingming WEI, Xiao HUANG, Yuanyuan ZHOU, Huasun HUANG

Affiliations

Rubber Research Institute, Chinese Academy of Tropical Agricultural Science / State Centre for Rubber Breeding / State Key Laboratory of Tropical Crop Breeding / Key Laboratory of Biology and Genetic Resources of Rubber Tree, Ministry of Agriculture and Rural Affairs, Haikou, Hainan 571101, China

出版时间: 2025-12-25 doi: 10.3969/j.issn.1000-2561.2025.12.006

文章导航

摘要

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橡胶树作为重要的经济树种，其特殊代谢产物——天然橡胶是我国的战略资源。自19世纪从南美洲引种至东南亚以来，通过持续遗传改良，橡胶树产量获得大幅度提高。本文全面综述橡胶树的植物学特性，回溯其引种驯化历程，剖析传统育种技术（包括杂交育种、多倍体育种、诱变育种等）与现代生物育种技术（包括分子标记辅助选择、基因组选择、转基因、基因编辑等）在橡胶树育种中的应用现状、取得的成果、面临的挑战及未来研究方向。旨在为橡胶树遗传育种领域研究者提供参考，助力该领域研究发展，推动天然橡胶产业可持续发展。

关键词

橡胶树 / 植物学特性 / 引种驯化 / 传统育种 / 生物技术育种

Abstract

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Rubber tree (Hevea brasiliensis), an economically significant species, produces natural rubber, a strategic resource for China. Since rubber tree introduced from South America to Southeast Asia in the 19th century, rubber tree productivity has been substantially improved through sustained genetic improvement. This article comprehensively reviewed the botanical characteristics of the rubber tree, traced its history of introduction and domestication, and analyzed the current applications, achievements, challenges, and future research directions of both conventional breeding techniques (including hybrid breeding, polyploid breeding, mutagenesis breeding, etc.) and modern biotechnological breeding techniques (including molecular marker-assisted selection, genomic selection, genetic modification, gene editing, etc.) in rubber tree improvement. The review aims to provide a reference for researchers in rubber tree genetics and breeding, facilitate progress in this field, and promote the sustainable development of the natural rubber industry.

Key words

Hevea brasiliensis / botanical characteristics / domestication / conventional breeding / biotechnological breeding

引用本文

张源源, 李维国, 张晓飞, 高新生, 王祥军, 位明明, 黄肖, 周元元, 黄华孙. 橡胶树育种技术研究进展. 热带作物学报, 2025 , 46 (12) : 2877 -2889 . DOI: 10.3969/j.issn.1000-2561.2025.12.006

Yuanyuan ZHANG, Weiguo LI, Xiaofei ZHANG, Xinsheng GAO, Xiangjun WANG, Mingming WEI, Xiao HUANG, Yuanyuan ZHOU, Huasun HUANG. Progress in Breeding Techniques of Rubber Tree[J]. Chinese Journal of Tropical Crops, 2025 , 46 (12) : 2877 -2889 . DOI: 10.3969/j.issn.1000-2561.2025.12.006

正文

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橡胶树[Hevea brasiliensis (Willd. ex A. Juss.) Muell. Arg.]的特殊代谢产物天然橡胶，是一种顺式-1,4-异戊二烯多聚物，具有独特的弹性形变恢复能力、耐高强度冲击能力，以及良好的耐磨性、热耗散性、低温延展性^[1]，在军事、航空航天、海洋工程、轨道交通等领域具有不可替代的作用，是我国的战略资源。天然橡胶还广泛用于生产轮胎、医疗器械（如乳胶手套、避孕套）、工程构件（如密封圈、减震元件）及日用品^[2]。天然橡胶除用于橡胶制品生产外，其原料胶乳还有潜在的药用价值^[3-5]。同时，橡胶树的木材^[6-8]和种子也具有重要的经济价值^[9-15]。

橡胶树自1876年从南美洲引种以来，其引种驯化历史已经超过120年。随着世界经济社会发展对天然橡胶需求的增长，橡胶树现已在亚洲、非洲、美洲、大洋洲的60多个国家和地区广泛栽培种植^[16]。期间，经过几代橡胶育种工作者的不懈努力，橡胶树的育种研究取得了巨大的进步。本文旨在综述橡胶树的植物学特性，梳理其引种驯化的历史，总结橡胶树的育种技术，取得的成果和存在的问题，以期为橡胶树遗传育种及相关领域研究工作者提供参考，助力橡胶树研究的进一步发展。

1　橡胶树植物学简介

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橡胶树又称巴西橡胶树、三叶橡胶树，为大戟科（Euphorbiaceae）橡胶属（Hevea）植物，染色体数为2n=2x=36，基因组约1.47~1.58 Gb^[17-20]。橡胶树属内有10个种和4个变种，其中巴西橡胶树是橡胶属内唯一大规模商业种植的种^[21]。橡胶树为高大乔木，株高可达30 m；叶片为掌状三出复叶，小叶椭圆形，网状叶脉；叶柄顶端具2~4枚腺体。橡胶树一年可开花2~3次，主花期（春花）为3~4月，夏花和秋花多发生于春花未有效结实时。橡胶树雌雄同株异花，花序腋生，有限圆锥花序；雄花占小花的90%以上，具卵状披针形花萼裂片；雄蕊10枚，排成2轮，花药2室，纵裂；雌花多着生于花序侧枝的顶端及下部，花萼形态同雄花但较大，花托绿色；子房2~6室，以3室为主，每室单胚珠，花柱短缩，柱头数目与子房室数一致；果实为蒴果，椭圆状，春花果实发育期为5—10月，每果含2~6粒种子。种子卵圆状，淡灰褐色，具斑纹。种子发育存在显著年龄效应，幼树童期约4~6年，初果期结实率低且种子饱满度普遍不足，至10龄后种子质量显著提升。

2　橡胶树的引种和驯化

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2.1　橡胶树的引种

橡胶树原产于南美洲亚马逊河流域，主要分布于巴西、委内瑞拉和圭亚那等地区。1876年，英国植物学家亨利·亚历山大·魏克汉（Henry Alexander Wickham）从亚马逊河中下游及塔帕若斯河（Tapajós River）流域采集70 000粒橡胶树种子，在英国皇家植物园成功培育出2899株实生苗，并于1877年将存活的1900余株转移至锡兰（现斯里兰卡），后逐步引种至新加坡、马来亚（现马来西亚）及印度尼西亚等东南亚地区^[22-24]。从这一批母树杂交选育获得的初生代、次生代及多生代无性系，构成了全球主要栽培品种的遗传基础，统称为魏克汉种质（Wickham germplasm）。为拓展橡胶树种质遗传多样性，国际橡胶研究与发展委员会（International Rubber Research and Development Board, IRRDB）于1981年组织多国专家重返亚马逊河流域开展野生种质资源系统性采集工作，从194株高产母树上共采取64 736粒种子及1522米芽条。这些种质材料分部分保存在巴西（50%）、马来西亚（37.5%）、科特迪瓦（12.5%）等国，该批资源被称为1981’IRRDB种质^[25]。我国通过国际合作从马来西亚引种并保存约6000份^[21]。此外，1995年马来西亚独自重返巴西进行了第二次种质采集，获得50 000余株实生苗^[26-27]，但未向国际社会共享该批资源。

我国橡胶树引种始于20世纪初。1904年，云南傣族土司刀安仁从新加坡引进8000株橡胶树，定植于今云南盈江县，成为我国橡胶引种的开端。随后，台湾地区（1905年）和海南岛（1906年华侨何麟书引种）相继引种并试种橡胶树成功，为我国植胶业发展打下了初步基础^[28]。

2.2　橡胶树的驯化

橡胶树引种到东南亚之初主要采集母树的种子进行繁殖，至1914年，东南亚的橡胶园已达472 000 hm²^[29]。随着天然橡胶需求持续增长，而东南亚宜植胶土地开垦殆尽，单位面积产量的提高日益受到重视。为此，东南亚、非洲、美洲等地国家和公司启动了多个育种计划，通过定向选择高产亲本杂交、芽接无性系鉴定等技术，选育出RRIM^[1]、PB、RRIT、IRR、RRII、RRIC、IAC、IAN等系列无性系^[30-35]，使橡胶树的单位面积产量从最初魏克汉种质的300~400 kg/(hm²·a)提高到第一、二代杂种无性系的900~1600 kg/(hm²·a)，以及第三、四代杂种无性系的2000~2500 kg/(hm²·a)^[25-36]。

新中国成立后，我国于20世纪50年代建立专业的研究机构，开始系统研究橡胶树的种植和育种^[37]。早期，通过大规模的种子采集、高产单株繁殖鉴定等，筛选出一大批抗寒无性系，如93114、南华1、天任31-45等，但产量普遍不高。同一时期，国内开始规模引进国外优良无性系开展区域试种工作，并于20世纪80年代筛选出适宜我国特殊植胶环境的RRIM600、PR107、GT1、IAN873等优良无性系^[38]，成为我国早期的主栽品种。但早期筛选出的品种普遍抗性不足、产量抗性不协同，遭受严重风害、寒害时常造成严重的产量损失甚至胶园损毁。针对这些问题，橡胶育种单位通过选配优良亲本组合、人工授粉、早期鉴定等，选育出以热研73397、海垦2、文昌217、南俸37、大丰95、保亭235、大岭68-35、93114、红星1、化59-2、云研77-4等为代表的中规模推广级以上品种^[37]。随后经过多年的系统鉴定，至今仍有热研73397、热研917（热研72059）、海垦2、文昌11、文昌217、大丰95、大丰99等高产抗风品种^[39-43]，以及云研77-4、云研77-2、云研73-46、云研80-1983、湛试32713等高产抗寒品种^[44-46]，以及超高产品种热研879^[36]，成为当前我国自主培育的橡胶树主推品种。此外，通过引进国外一批胶木兼优品种在国内开展联合区域试验鉴定，选育出热垦628、热垦525等胶木兼优品种^[47-48]。以上品种为我国天然橡胶产业的发展提供了有力支撑。

3　橡胶树育种技术研究进展

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橡胶树引入东南亚初期，推广种植的主要是母树的实生苗，但因为基因型各异，实生苗的产量高低不一，严重影响橡胶的产量。1916年，荷兰人赫尔顿在印尼爪哇岛发明了橡胶树芽接技术^[49]，使得生产中的高产单株可以快速增殖，并以无性系的形式推广种植，推动橡胶树育种进入了无性系育种阶段。随后“雄蕊塞入法”授粉技术的发明^[50]，为橡胶树的人工杂交育种提供了技术条件。此后，橡胶树育种借鉴其他作物育种技术，逐渐发展出了多样化的育种技术，以满足橡胶产业发展的需求。

3.1　传统育种技术

3.1.1　杂交育种

橡胶树的杂交育种程序包括杂交授粉、子代有性系比较试验、无性系比较试验、品种比较试验以及区域试验等环节^[51]。杂交授粉包括人工授粉和自然授粉2种。人工授粉采用“雄蕊塞入法”，即在雌花成熟（花瓣伸长变黄）且未开放时，将雄蕊整个取出并塞入雌花中，然后取树枝上的胶乳将雌花瓣粘合起来，避免外来花粉的侵入。自然授粉则是指选择优良杂交组合建设种子园，在橡胶树开花传粉阶段不进行人工干预，通过虫媒或风媒完成授粉。子代有性系比较试验是在果实成熟阶段收集种子，进行育苗、定植于大田，以早期阶段评估生长速度、产量等性状，筛选优良单株。无性系比较试验是将筛选出的优良单株通过芽接等手段繁殖成无性系，小规模种植以再次鉴定其目的性状，筛选优良无性系。筛选出的优良无性系，再较大规模繁殖，并按照生产标准种植和管理，经多年生产性鉴定进一步筛选出优异无性系，为品种比较试验。优异无性系品种在全国不同的生态类型区进行生产适应性种植鉴定，即为品种区域试验。

杂交育种的关键是选择杂交组合，不同品种的一般配合力和特殊配合力存在差异，导致从不同杂交组合子代中选育出无性系优良品种的概率不同。通过对国内选育品种的分子标记结合亲本系谱分析发现，RRIM600、PR107、GT1和PB86为我国橡胶树育种的骨干亲本，从这些骨干亲本及衍生后代中选育出热研系列、云研系列、大丰系列、文昌系列等众多优良品种^[52-53]，但也导致我国育成品种遗传基础狭窄^[54]。同时，长期以单一或少数性状为目标的育种工作，使当前品种存在“高产不抗逆，抗逆产量低”的问题。例如，在海南地区以高产为主要目标的育种工作中，高产基因不断聚合，生长、抗逆等副性状基因逐渐丢失，选育出的品种产量较高，但抗风抗寒能力一般，如热研88-13、热研8-333、热研879、热研106等^[55-57]；抗风抗寒综合能力较好的品种，产量却并非最高，如热研73397、热研917等^{[39, 41]}。在云南、广东以抗寒高产为主要目标的育种工作中，抗寒基因不断聚合，高产基因没有同步聚合，选育出的品种抗寒能力强，产量却较低，如云研77-4、湛试32713等^[58-59]。为解决遗传基础狭窄、基因型高度杂合、多性状难以聚合等问题，育种工作者尝试了其他育种技术和方法。

3.1.2　多倍体育种

多倍体育种作为一种有效的育种手段，能够利用杂种优势和基因剂量效应^[60-61]，是林木良种选育的重要方法之一。多倍体育种在杨树^[62-64]、桑树^[65]等树种中开始较早，并取得了显著成效。橡胶树作为较早开展育种的树种之一，其多倍体育种研究也有较长的历史^[66]。20世纪60至80年代，世界主要的橡胶树育种国家均开展了橡胶树多倍体研究。当时各国普遍采用秋水仙碱处理生长点诱导四倍体的方法，通过处理橡胶树的茎尖、芽点、种子等，获得了大量的多倍体植株^[67-71]。中国在橡胶树多倍体育种研究中取得显著进展，形成了从诱变处理，分离筛选，到细胞学鉴定和形态、组织学鉴定的有效方法，能够在1.5~2年内获得需要诱导的优良品系的稳定多倍体^[72]。然而，这些多倍体材料在后续产量比较试验中并未表现出预期的高产性状^[73]。近几年，国内外以愈伤组织、体胚等为诱变材料再次进行了四倍体诱导^[74-75]，但获得的四倍体植株的农艺性状有待进一步观察。

进入21世纪，随着在云南大面积推广种植的品种云研77-2和云研77-4被鉴定为三倍体^[76]，橡胶树多倍体育种再次引起国内橡胶树育种工作者的关注。北京林业大学联合云南省热带作物科学研究所开展了人工诱导橡胶树2n雌雄配子的研究，通过秋水仙碱、高温等处理手段，获得了人工诱导的2n花粉、2n卵细胞，并经过人工授粉杂交，获得了三倍体植株^[77-79]。同时，中国热带农业科学院橡胶研究所和云南省热带作物科学研究所通过连续多年多点对多品种的子代进行倍性鉴定，筛选出大量多倍体植株，并发现花发育期的温度与橡胶树GT1子代三倍体的发生频率相关，温度越低，三倍体频率越高^[80]。目前，除了与云研77-2和云研77-4同批选出的云研73-46^[81]外，尚未有新的多倍体品种推广应用。多倍体育种在橡胶树育种中的应用仍处于探索阶段，但其潜力巨大，未来有望通过进一步的研究和技术改进，选育出更多符合生产需要的多倍体新品种。

3.1.3　诱变育种

诱变育种的本质是通过理化处理诱导染色体数量、片段改变或碱基突变。因此，广义的诱变育种包括染色体倍数的变化，即多倍体诱导。狭义的诱变育种则主要是指通过化学诱变剂处理或辐射等物理方法引起染色体片段或碱基的突变。早期，由于缺乏诱变所需的离体再生材料，橡胶树的化学诱变研究较少。橡胶树组织培养技术获得突破后，陈祖兴等^[82]尝试使用EMS（ethyl methane sulfonate）诱导橡胶树花药次生体细胞胚突变，确定了0.5%的EMS浸渍4 h为适宜的处理条件，并成功获得了再生植株。辐射诱变方面，郑学勤等^[72]使用3500伦琴伽马射线照射GT1自然授粉种子，播种育苗后进行产量早期预测，筛选出叶绿素缺乏的黄色斑块叶和小叶柄扭曲的表型变异单株。此外，辐射诱变还作为多倍体诱导的辅助手段^[83]，但其理论基础和实际作用尚不明确。随着我国航天技术的发展，利用体细胞胚可保存较长时间的优势，通过航天搭载开展空间环境诱变育种，有望在微重力与宇宙射线协同作用下诱发更丰富的遗传变异，为创制新种质提供新途径。诱变育种能够产生新的性状变异，对于拓宽橡胶树遗传基础和研究突变性状的分子机理有重要意义。

3.2　现代生物育种技术

采用常规杂交、多倍体诱导、诱变等第一、二代育种技术开展橡胶树多目标性状遗传改良，存在成本高、育种周期长、育种效率低下等问题，难以在短期内育成高产、速生、优质、抗逆等综合性状优良的新品种，逐渐无法满足橡胶树现代育种研究的要求，亟需以分子标记辅助选择、转基因、基因编辑、基因组选择等为代表的现代生物育种技术助力橡胶树育种研究，加快新品种的培育进程，相关技术在橡胶树研究中已取得显著进展。

3.2.1　分子标记辅助选择育种

与传统育种方法相比，分子标记辅助选择（marker-assisted selection, MAS）能够显著提高杂交后代选择的效率，降低育种成本，缩短育种周期^[84]。MAS通过利用与目标性状紧密连锁的分子标记，在幼苗阶段即可筛选出具有优良性状的个体，从而加速优良品种的培育^[85]。在橡胶树中，已经开发了多种分子标记，包括限制性片段长度多态性（restriction fragment length polymorphism, RFLP）、随机扩增多态性DNA（randomly amplified polymorphic DNA, RAPD）、扩增片段长度多态性（amplified fragment length polymorphism, AFLP）、简单重复序列标记（simple sequence repeats, SSR）、单核苷酸多态性（single nucleotide polymorphism, SNP）、内含子长度多态性（intron length polymorphic, ILP）、多核苷酸多态性（Multiple nucleotide polymorphism, MNP）等，但这些标记主要集中在种质资源的遗传多样性分析、无性系的遗传关系分析和品种指纹图谱构建研究上，而用于辅助选择育种的分子标记相对较少^[86]。陈守才等^[87]利用22个抗白粉病和感白粉病品种鉴定到一个与橡胶树抗白粉病表型密切相关的RAPD标记opv-390。VENKATACHALAM等^[88-89]分别鉴定到一个与橡胶树矮化表型连锁的和一个与橡胶树死皮相关的RAPD标记。BHUSUDSAWANG等^[90]使用26个与次生生长相关的候选基因筛选出115个ILP标记，通过关联分析检测到9个标记与胸径生长性状显著关联，其中COBL064_1与旱季胸径（GiD）和雨季胸径（GiW）关联最强，分别解释了14.0%和14.6%的表型变异，并在标记与有性系的关联验证中发现COBL064_1和PIN227_1与GiD和GiW显著关联。在COBL4基因中鉴定出128个SNP和178个In/Del，其中3'UTR中的SNP（CA5589）与胸径生长性状的关联最强，解释了11.1%的表型变异。CHAO等^[19]通过对335份材料的基于重测序的GWAS分析，鉴定到一个调控乳管列数的基因HbPSK5，并开发出相关的SNP标记Chr15:1200714，有望用于高产性状的分子标记辅助选择。然而，这些标记目前均未应用于育种实践中。MAS在橡胶树中的应用仍处于研究和开发阶段，但其在提高育种效率和精准度方面的潜力巨大，未来有望在橡胶树的遗传改良中发挥重要作用。

3.2.2　基因组选择育种

随着基因组测序技术和计算机科学的快速发展，通过高密度的分子标记准确预测作物产量等复杂性状成为可能。植物全基因组选择育种技术作为分子标记辅助选择技术的高阶版，通过收集训练群体的表型数据和基因型数据，使用特定的模型估计分子标记效应值或个体育种值，再根据待测群体的基因型数据和模型拟合结果对待测群体的表型值进行预测，并根据预测结果进行定向选择。全基因组选择育种可以减少育种工作量，显著缩短育种周期，提高育种效率^[91]。橡胶树基因组选择育种方面也有初步尝试。CROS等^[92]通过对189个和143个来自PB260×RRIM600杂交组合的子代无性系的表型鉴定，使用332个SSR标记进行基因分型，探讨了基因组预测方法、训练集大小和标记密度对基因组选择精度的影响。发现使用所有无性系和标记进行不同地点验证时，基因组选择精度为0.53。标记密度和训练集大小对精度有显著影响，但300个标记已足够，使用杂合度最高的125~200个标记时，跨地点的基因组选择精度可达0.56，而训练群体数量超过175个后对精度的提升有限。进一步通过模拟分析，利用基因组预选技术对3000个幼苗进行筛选，可以将橡胶产量的选择响应提高10.3%。SOUZA等^[93]比较了多种包含基因型（Genotype, G）×环境（Environment, E）相互作用的基因组预测模型，发现G×E相互作用会降低基因组选择的准确性，从而限制遗传增益的提升，进而认为开发能够有效管理G×E相互作用的预测模型对于优化橡胶树育种策略至关重要。AONO等^[94]开发了一种基于机器学习的新型方法，用于基于分子标记预测橡胶树茎围，先通过神经网络预判个体所属遗传群体，然后针对性训练亚群特异性模型，结合多方法联合标记筛选（L1-SVM/Pearson/梯度树提升），将橡胶树茎围预测精度从传统基因组选择模型的R=0.16提升至0.68，实现了群体结构下的高精度非线性建模，为复杂性状育种提供了新范式。WANG等^[95]使用靶点测序基因分型（genotyping by target sequencing, GBTS）技术开发了40K高密度育种芯片，并对223个橡胶树无性系进行基因分型，结果表明无性系间的遗传多样性较低，遗传基础有限。随后，通过全基因组关联（genome-wide association study, GWAS）分析鉴定了与茎围增长速度相关的标记。基于GWAS结果，使用7个基因组选择模型来评估不同标记数据集对茎围生长速度的预测准确性，其中贝叶斯最小绝对收缩和选择运算模型（bayesian least absolute shrinkage and selection operator, BayesLASSO）具有最高预测精度（0.59），被选为育种值预测的最佳基因组选择模型和基因型数据集，并依据该模型筛选出50个优良亲本组合。以上研究结果表明，基因组选择育种技术在提高橡胶树选育种效率方面具有巨大的应用潜力与价值。

3.2.3　转基因育种

转基因技术作为现代生物育种技术的代表之一，可以快速定向改变育种目标性状。自2003年印度首次报道在橡胶树品种RRII105中实现了农杆菌介导的转基因植株的遗传转化和再生^[96]以来，各国在转基因愈伤组织的筛选和再生、农杆菌介导的遗传转化、标记基因的优化、抗生素在转基因过程中的作用、体细胞胚胎发生效率提升等方面取得了显著进展。2006年，法国BLANC等^[97]以农杆菌EHA105为介导载体，pCAMBIA2301为质粒载体，建立了巴西橡胶树无性系PB260愈伤组织的高效转化方法。随后的研究发现，添加硝酸银能够提高农杆菌介导的巴西橡胶树叶片愈伤组织的遗传转化效率和体细胞胚胎发生效率^[98]，并开发了利用叶片和根作为农杆菌浸染外植体的遗传转化方法^[99]。此后，中国HUANG等^[100]报道了以体胚替代愈伤组织作为侵染对象的农杆菌介导的转基因方法。在基因转入与转化效率提升方面，印度JAYASHREE等^[101]于2018年将hmgr1基因转入橡胶树，显著提高了胶乳产量。2021年，HUANG等^[102]研究了花青素生物合成基因的转录激活基因R2R3-MYB转录因子HbAN2作为视觉选择标记，在橡胶树遗传转化中的应用，以提高转化效率。同时，他们还对农杆菌介导的橡胶树遗传转化过程中的抗生素浓度、预培养时间、农杆菌浓度、超声和共培养等条件进行优化，改良了橡胶树转基因方法^[103]。此后，LUO等^[104]通过过表达HbGRF4或HbGRF4-HbGIF1，显著提高了橡胶树热研73397的体细胞胚胎发生效率。这些研究不仅提高了橡胶树的遗传转化效率，还为橡胶树的遗传改良和分子育种提供了新的方法和策略。随着橡胶树功能基因研究的不断深入，转基因技术在橡胶树育种研究中的作用将愈发重要，有望为橡胶树产业的可持续发展提供更有力的技术支撑。

3.2.4　基因编辑育种

CRISPR/Cas9技术作为前沿生物技术之一，在植物抗病性改良、抗逆性增强、产量提高、品质改良以及育种材料创制等方面取得了重要进展^[105-107]。橡胶树作为重要的热带经济树种，在基因编辑领域也取得了显著进展。FAN等^[108]设计了5个针对橡胶树开花相关基因的单导向RNA（sgRNA），通过将Cas9蛋白与sgRNA预组装后直接递送至橡胶树原生质体，成功诱导了靶位点的突变。5个靶位点的突变频率介于3.74%~20.11%之间，并检测到+1 nt插入和缺失2种突变模式，展示了RNPs在橡胶树基因编辑中的潜力。随后，DAI等^[109]进一步优化了CRISPR/Cas9系统在橡胶树中的应用，通过鉴定和利用橡胶树内源U6启动子驱动sgRNA转录，建立了CRISPR/Cas9短暂编辑系统。研究发现，不同的U6启动子具有不同的编辑效率，从8.47%到24.92%不等。通过深入分析10个sgRNA针对5个开花时间相关基因的突变谱，检测到缺失、插入和碱基替换3种突变模式。此外，通过构建稳定转化编辑载体，针对橡胶树基因HbPDS在橡胶树愈伤组织中进行转化，成功获得了目标位点的“+1” bp纯合插入，表现为预期的白化表型。2024年，YANG等^[110]发现10 mg/L的潮霉素是适合胚胎筛选的浓度，而超过90%的T₀基因编辑胚是嵌合体，通过将1个T₀代胚切成33小块诱导次生胚，产生了T₁代胚，且50%的T₁代胚是纯合的，编辑效率提高到近90%。基于此发现，建立了针对HbPDS基因的CRISPR载体转化胚胎的方法，最终从29个T₁代胚中再生出8株植株，包括4株嵌合体和4株纯合白化植株，研究结果显著提高了橡胶树纯合编辑植株的获得效率，克服了嵌合体转化胚胎的限制。橡胶树基因编辑领域取得的以上进展，提高了基因编辑的效率和精确性，为橡胶树的遗传改良和分子育种提供了强有力的工具。

4　问题和展望

收起

在未来很长一段时间内，橡胶树同其他林木一样，以杂交育种为代表的传统育种方法仍将是获得新品种的最有效方式^[111]，而橡胶树传统育种至今存在结实率低、选择效率低下、育种周期长、遗传基础狭窄等问题。针对以上问题，提出以下解决途径：（1）研发胚挽救技术，在坐果期拯救面临败育风险的未成熟胚胎，提高人工杂交授粉的结实率。（2）营建高世代无性系种子园，直接收集自然授粉种子，省去人工杂交授粉的繁琐过程，提高杂交子代获得效率。（3）完善基于早期试割、乳管列数、生理指标、橡胶合成相关基因表达量等检测手段的综合产量早期精准鉴定技术，研发基于关键基因多态性的高通量早期产量选择技术，以提高早期鉴定准确度，缩短早期鉴定时间，提高选择效率。（4）学习胶木兼优选育的“以空间换时间”的成功经验，合并无性系比较试验和品种比较试验程序，并在不同植胶类型区建立无性系比较试验区替代品种区域试验，增加无性系数量，扩大无性系规模，延长测定年限，进一步缩短育种周期。（5）持续开展诱变、多倍体诱导等成熟育种技术的应用研究，创制育种中间材料。

目前，现代育种技术用于橡胶树新品种选育面临诸多挑战，如缺乏可靠实用的分子标记用于辅助选择、基因组选择育种研究仍处于起步阶段、转基因效率有待提高、基因编辑技术的稳定性需进一步验证、多目标性状的关键基因鉴定不足等，这些因素限制了现代育种技术在橡胶树中的应用。因此，尽管橡胶树现代育种技术取得了显著进展，但离实际应用仍存在一定距离。未来，有必要持续加大投入，深化现代育种技术在橡胶树研究中的应用。具体包括：（1）充分利用基因组重测序等高通量测序技术，通过全基因组关联分析、数量性状位点定位等正向遗传学手段，系统鉴定调控产量、抗逆性、品质等多目标性状的关键基因，分析其优势单倍型，为转基因和基因编辑育种提供遗传靶标。（2）基于关键基因鉴定结果，开发高效、可靠的分子标记，支撑种子园杂交子代群体的高通量分子标记辅助筛选。（3）提升转基因效率和基因编辑技术的精准性与稳定性，为关键基因的功能验证和性状定向改良提供技术支撑。（4）利用人工智能、深度学习及大语言模型等前沿技术，赋能橡胶树育种全过程，具体包括：①整合基因组、转录组、表型组、环境等多维异构大数据，挖掘复杂性状的遗传调控网络；②利用人工智能/机器学习构建基因型-表型预测模型，加速优异种质筛选和杂交组合设计；③应用人工智能辅助设计更高效、脱靶率更低的基因编辑方案；④利用大语言模型等工具高效挖掘和梳理海量文献与育种数据中的隐性知识，辅助育种决策。

通过传统育种与现代育种技术的协同创新与融合发展，将系统性提升橡胶树育种的精准度、效率和成功率，加速培育出高产、优质、多抗且适应未来需求的新品种，为天然橡胶产业可持续发展提供核心种源保障。

基金

收起

国家重点研发计划项目(2022YDF2301205)
国家农业产业技术体系专项(CARS-33-YZ2)

参考文献

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文献

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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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