水稻黄叶少分蘖突变体yllt1的鉴定及基因定位

采用⁶⁰Co-γ射线诱变籼稻（Oryza sativa subsp.indica）保持系‘T98B’获得一份兼具黄叶和少分蘖表型的突变体yllt1（yellow leaf and less tillering 1）,利用色素含量测定、构建显隐性混池和基因表达量测定等方法从表型和遗传层面对其遗传特征进行分析。结果显示：yllt1苗期叶绿素a和叶绿素b含量为野生型水稻品种‘T98B’的77.78%和60.00%,叶绿体发育异常,缺乏功能性叶绿体类囊体片层;其分蘖盛期的单株分蘖数为野生型的21.43%。遗传分析发现,在突变体yllt1与‘T98B’的杂交F₂群体中,黄叶与少分蘖性状的重组率为0.00%,表明yllt1同时控制叶色与分蘖表型;yllt1呈隐性遗传,受一个细胞核基因独立控制。该研究进一步采用连锁分析法将yllt1精细定位到第11染色体上,经测序分析推断发生了突变的登录号为LOC_Os11g05552的基因是yllt1的目的基因;该基因编码叶绿体前体信号识别颗粒54 kD（cpSRP54）蛋白,其第1外显子的第29位碱基C发生了缺失,将造成其蛋白产物从N-端至C-端氨基酸组成的严重破坏。RT-qPCR分析结果显示,yllt1叶中叶绿素合成基因OsCAO1、OsCAO2与OsNOL等的表达量明显下调;茎中分蘖正向调控基因OsTAC1受到显著抑制,而负调控基因OsTB1与OsDLT的表达量明显增强。研究结果表明cpSRP54同时参与了水稻叶色和分蘖的调控。     

水稻黄叶少分蘖突变体yllt1的鉴定及基因定位

采用~(60)Co-γ射线诱变籼稻(Oryza sativa subsp.indica)保持系‘T98B’获得一份兼具黄叶和少分蘖表型的突变体yllt1(yellow leaf and less tillering 1),利用色素含量测定、构建显隐性混池和基因表达量测定等方法从表型和遗传层面对其遗传特征进行分析。结果显示:yllt1苗期叶绿素a和叶绿素b含量为野生型水稻品种‘T98B’的77.78%和60.00%,叶绿体发育异常,缺乏功能性叶绿体类囊体片层;其分蘖盛期的单株分蘖数为野生型的21.43%。遗传分析发现,在突变体yllt1与‘T98B’的杂交F_2群体中,黄叶与少分蘖性状的重组率为0.00%,表明yllt1同时控制叶色与分蘖表型; yllt1呈隐性遗传,受一个细胞核基因独立控制。该研究进一步采用连锁分析法将yllt1精细定位到第11染色体上,经测序分析推断发生了突变的登录号为LOC_Os11g05552的基因是yllt1的目的基因;该基因编码叶绿体前体信号识别颗粒54 kD(cpSRP54)蛋白,其第1外显子的第29位碱基C发生了缺失,将造成其蛋白产物从N-端至C-端氨基酸组成的严重破坏。RT-qPCR分析结果显示,yllt1叶中叶绿素合成基因OsCAO1、OsCAO2与OsNOL等的表达量明显下调;茎中分蘖正向调控基因OsTAC1受到显著抑制,而负调控基因OsTB1与OsDLT的表达量明显增强。研究结果表明cpSRP54同时参与了水稻叶色和分蘖的调控。     

水稻分蘖的分子机理研究进展

分蘖是水稻等禾谷类作物生产的关键农艺性状,也是单子叶植物一种特殊的分枝现象.水稻分蘖的形成是一个复杂的过程,其间受遗传、植物激素、栽培环境等因素的综合影响.近年来,对水稻分蘖数改变的突变体研究取得令人瞩目的研究成果,本综述总结水稻分蘖的调控机理的最新研究进展.     

独脚金内酯调控水稻分蘖的研究进展

水稻(Oryza sativa)作为世界上最主要的粮食作物之一, 对其主要农艺性状调控机理的研究具有重要意义。分蘖是水稻生长发育过程中一种特殊的分枝, 它不仅是与水稻产量密切相关的重要农艺性状, 也是揭示高等植物侧枝生长发育机制的理想模型。独脚金内酯(strigolactone, SL)是一类新型植物激素, 能够抑制植物分枝的生长发育。近年来, 关于SL合成与信号在调控水稻分蘖方面的研究取得了重要进展, 但对其信号转导的下游组分的研究还相对匮乏。该文综述了SL合成途径、信号途径及下游靶基因调控水稻分蘖的研究进展, 并与在拟南芥(Arabidopsis thaliana)、豌豆(Pisum sativum)和矮牵牛(Petunia hybrida)中的研究进行了比较, 同时还对如何挖掘SL途径的新组分进行了讨论。     

表观遗传调控植物分枝/分蘖研究进展

分蘖是禾本科植物特有的分枝类型, 是影响作物产量的关键因素之一。分枝/分蘖数由叶腋处侧生分生组织的数量和侧芽的活性共同决定。表观遗传修饰调控植物生长发育的各个方面, 但是如何调控植物的分枝/分蘖数还未见系统报道。该综述归纳了表观遗传调控侧生分生组织的形成和侧芽向外生长两个方面, 并展望了表观遗传在调控植物分枝/分蘖中的研究方向, 以期为通过表观遗传修饰改良作物品种的育种途径提供理论指导。     

长雄蕊野生稻分蘖角度QTL定位及遗传分析

分蘖角度是水稻重要农艺性状之一,在调控水稻产量方面具有极其重要的作用。利用长雄蕊野生稻跟粳稻品种中花11杂交构建F_2作图群体,考察各单株分蘖角度,用均匀覆盖水稻基因组的125对多态性InDel标记,构建了长雄蕊野生稻分蘖角度InDel分子连锁图谱。共检测到3个分蘖角度相关QTLs(qTA-1-1,qTA-3-1,qTA-7-1),分别位于水稻1号,3号,7号染色体上,共解释21.40%的表型变异。其中,qTA-1-1可以解释6.41%的表型变异,是一个新的分蘖角度QTL位点;qTA-7-1解释了7.84%的表型变异,包含了一个调控普通野生稻分蘖角度的PROG1基因,对亲本进行比较测序分析,发现PROG1编码区中有一个18 bp的InDel、两个3bp的InDel和5个SNP,该差异造成长雄蕊野生稻和中花11中PROG1编码的蛋白不同。本研究构建了第一张高密度长雄蕊野生稻In Del标记遗传图谱,发掘分蘖角度新QTL位点,为育种家利用分子标记辅助选择培育水稻理想株型新品种打下基础。     

调亏灌水和分蘖干扰对冬小麦生长的补偿效应

为探索抑制个体功能的生长冗余以实现群体性能优化并挖掘作物高产潜力的途径,通过桶栽试验,选择分蘖能力中等的小偃22号和分蘖能力较强的郑麦7698,对比研究2种灌水模式（全生育期充分灌水和分生育期调亏灌水）和3种分蘖干扰（从拔节期开始去除所有小分蘖,仅保留主茎和1个大分蘖;抽穗期去除所有无效分蘖;以不作任何干扰为对照）,来模拟不同水分供应和不可预测干扰对冬小麦生理生长、产量和水分利用效率的补偿机制.结果表明： 2个冬小麦品种均存在生长冗余.与小偃22号相比,郑麦7698有效分蘖数较高,但穗部性状较差.调亏灌水和抽穗期去除无效分蘖均可减少生长冗余,弱化竞争能力,改变源 库关系,提高资源分配.但冗余消除过度（拔节期干扰）则会破坏植株固有的根冠平衡和功能结构,导致生长的不足补偿.与对照相比,调亏灌水联合抽穗期去除无效分蘖可在时空尺度上充分开发和利用作物自身调控潜力实现补偿生长,在不显著影响籽粒产量的同时可提高水分利用效率20.4%～25.4%,是适宜的减冗增效措施.     

水稻OsTB1基因的结构及其表达分析

TCP基因是一类植物中新发现的、可能具有转录因子活性的基因家族,成员包括金鱼草的Cyclodiea (Cyc)、玉米的Teosinte Branched1 (TB1)以及水稻中的PCF1、PCF2等.玉米的TB1基因有维持玉米顶端优势的作用,与分蘖的发生密切相关;水稻和玉米同属禾本科,在发育的过程中都有分蘖的发生.通过筛选水稻的基因组文库,得到了水稻中的一个TB1同源基因Oryza sativa Teosinte Branched1 (OsTB1).该基因不含内含子,基因编码一个长度为388个氨基酸的蛋白,在氨基酸水平上与TB1的同源性为70%,含有保守的TCP区和R区,是属于TCP基因家族的一个成员.RT-PCR和mRNA原位杂交分析结果表明,OsTB1在水稻的侧芽中有很强的表达,在花序中有较弱的表达.以上结果显示该基因可能在水稻侧芽和花序的起始和发育过程中起重要作用.     

水稻分蘖氮响应调控机理研究进展

氮肥是作物产量增加最主要的驱动因素,然而氮肥滥用会造成生态环境的严重破坏。因此,提高作物氮素利用效率(nitrogen use efficiency,NUE)对未来农业可持续发展至关重要。产量性状对氮素的敏感性是衡量作物氮素利用效率的重要指标。禾本科作物的分蘖数、穗粒数和粒重是产量的直接决定因子,虽然影响三者本身的分子机制已有大量研究,但氮素对这些性状的调控机理仍知之甚少。分蘖数是对氮素响应最为敏感的性状之一,也是氮肥促进作物增产的关键要素。因此,研究氮素如何调控水稻的分蘖发育对于提高作物产量尤为重要。本文总结了水稻氮素利用效率的影响因素和分蘖发育的调控机理,聚焦氮素如何调控水稻分蘖发育的机制,并对该领域未来研究工作进行了展望,以期为作物氮高效精准改良提供参考。     

全基因组关联分析鉴定调控水稻分蘖角度的基因TAC3

水稻是主要的粮食作物,随着人口的不断增加,需要不断提高作物的产量来满足人们日益增长的粮食需求。株型是决定水稻群体产量的重要性状,提高水稻的种植密度是增加水稻产量的有效方法。水稻的分蘖角(Tiller angle, TA)是指侧生分蘖和主茎穗之间的夹角,它作为塑造理想株型的主要性状之一,决定植株的单位面积种植密度及作物产量。理想的分蘖角度既能避免角度过小导致田间湿度过高而诱发植物病害,又能避免因匍匐生长导致的光合作用效率降低和单位面积种植密度降低导致的产量下降,因此在水稻的长期驯化和遗传改良过程中分蘖角度受到了自然和人类的双重选择。PROG1(PROSTRATE GROWTH 1)基因由林鸿宣教授团队和孙传清教授团队分别揭示具有控制野生稻匍匐生长的功能,而其在栽培稻中的变异导致植株的直立生长,是驯化相关的基因(图1)。TAC1(Tiller Angle Control 1)基因由孙传清教授团队证明其在栽培稻籼粳亚群间的自然变异导致两个亚群间分蘖角度的差异(图1)。此外,研究人员利用反向遗传学等方法证明了一些基因具有控制水稻分蘖角度的功能。然而,关于水稻分蘖角度遗传分子机制的研究尤其是基于水稻自然变异的研究仍然相对有限。     

Effects of Plant Growth Regulators and Nutrient Supply on Tiller Bud Outgrowth in Barley (Hordeum distichum L.)

Seedlings of spring barley were raised in 100 and 20% nutrientsolution and treated with a foliar application of Terpal, Cerone,TIBA, GA₃ or BAP. The growth of individual tiller buds and tillers,the main shoot and the root system was recorded over the following15 d. Terpal and Cerone stimulated tiller bud elongation within5 d at both nutrient levels and after 15 d the number of emergedtillers was increased at the higher nutrient level. Terpal characteristicallypromoted the growth of secondary tiller buds whereas Ceronepromoted the emergence of the coleoptile tiller; both thesePGRs also retarded the development of the main shoot. TIBA increasedthe number of elongating tiller buds but this did not resultin greater tillering. GA₃ reduced the number of elongating tillerbuds and restricted their growth, especially in the high nutrientregime; this was accompanied by an increase in main shoot elongation.The growth and development of tiller buds was reduced by BAPand the number of emerged tillers was reduced at 15 d in bothnutrient levels; main shoot dry weight and root elongation werealso reduced. The results are considered in relation to theoverall influence of hormonal factors and mineral supply ontiller bud outgrowth. Hordeum distichum, spring barley, tiller bud outgrowth, plant growth regulators, Terpal, Cerone, GA₃, BAP, nutrient supply, apical dominance, TIBA     

Growth-Regulating Substances and the Growth of Tiller Buds in Barley; Effects of IAA and GA3

Application of cytokinin to barley seedlings grown without mineralnutrients leads to rapid growth of coleoptile and first leaftiller buds. IAA and GA₃ cannot substitute for cytokinin inthis effect and applications of TIBA and CCC are also ineffectiveon bud growth. However, when bud growth was promoted, eitherby application of cytokinins or by supplying plants with mineralnutrients, IAA and GA₃ applications caused enhanced tiller growthindicating that these compounds can stimulate growth of activelygrowing buds. The results are compatible with an interpretationwhich stresses the importance of cytokinin availability in determiningtiller bud growth.     

Effects of external ABA,GA<Subscript>3</Subscript> and NAA on the tiller bud outgrowth of rice is related to changes in endogenous hormones

Hormones play an important role in regulating the growth of rice tiller buds. However, little is known about the hormonal changes that occur during tiller bud growth and the mechanism of hormonal regulation of tiller bud growth. Here, two rice cultivars, Yangdao 6 (Indica) and Nanjing 44 (Japonica), were used to investigate the changes in plant hormones during tiller bud growth and the mechanism that underlies the hormonal regulation of tiller bud growth. In the present study, panicles were removed after heading to stimulate the growth of dormant tiller buds located at the elongated upper internodes. At the same time, external abscisic acid (ABA), gibberellic acid (GA₃) and α-naphthalene acetic acid (NAA) were applied. The results demonstrated that auxin and cytokinin (CTK) play important and different roles in the regulation of tiller bud growth. Auxin in the nodes inhibits tiller bud growth, while CTK is transferred to the tiller buds to promote growth. The inhibitory effects of GA₃ and NAA on tiller bud growth are mainly due to the control of the indole-3-acetic acid (IAA) or CTK contents in plants. As opposed to auxin and CTK, the ABA contents in nodes and tiller buds remained unchanged before tiller bud growth after panicle removal. Meanwhile, external ABA application only slightly slowed the growth of the tiller buds, suggesting that ABA may not be a key regulator of tiller bud growth. These results indicate that auxin, CTK and ABA together likely play roles in the regulation of tiller bud growth.     

Growth of Tiller Buds in Barley: Effects of Shade Treatment and Mineral Nutrition

Examination of the stem apex of Proctor barley showed that thebud of the coleoptile tiller, Tc, is probably present in thedry grain and that the bud, TI, carried in the axil of the firstleaf is present at or soon after 24 h from planting. Subsequentlytiller buds are initiated with a plastochron of about 4 days,this being rather longer than that for the foliar primordia.During the initial phase of bud growth vascular connectionsare established with the leaf above, but not to the subtendingleaf. At some time after these vascular connections are formedand when it has a dry weight of 4–7 µg the bud entersa phase of rapid, exponential growth in dry weight. Shading the first leaf delays the onset of rapid growth forboth Tc and Ti, but after a lag period rapid growth commences;this is coincident with development of the second leaf as anorgan exporting assimilated carbon. The phase of rapid growth of tiller buds is delayed when applicationof either nitrogenous or nonnitrogenous minerals is delayed.Ammonium was found to be less satisfactory as a nitrogen sourcethan nitrate, probably because of toxicity effects. Slight growthof Tc and T1 occurs in presence of nonnitrogenous minerals andabsence of nitrogen but growth is greater when nitrogen is suppliedin absence of the other minerals, although such growth is substantiallyless than that found when all nutrients are supplied. The interactionbetween nitrogen and non-nitrogenous minerals which controlsbud growth was not found to affect growth of the parent plantwhich is, as previously shown, controlled by timing of the nitrogensupply. AnotheT distinction is that higher concentrations ofnitrogen and the other minerals are required for maximum growthof the bud than for that of the plant. Tiller bud growth is interpreted as occurring in two phases.In the first, initiation, phase there is a close associationwith the subtending leaf, and nutritionally bud and leaf arelinked. This phase is followed by one in which the bud is directlyconnected by vascular traces to the leaf above, which becauseof this controls bud growth by modulating supply of assimilatedcarbon and nitrogen, and other minerals to it.     

Growth-Regulating Substances and the Growth of Tiller Buds in Barley; Effects of Cytokinins

An hypothesis was set up from which it was predicted that applicationof cytokinin to barley seedlings grown without mineral nutrientswould lead to rapid growth of the coleoptile and first leaftiller buds. Application of cytokinins to the leaves was ineffective,but supplying a number of known cytokinins by steeping the rootsof 4 d old seedlings in solution for 4 h led to significantgrowth of the coleoptile bud. Adenine and cytokinin analogueshad no effect. Supplying cytokinins through the roots also furtherenhanced the growth of buds of plants given mineral nutrients.Cytokinin treatment reduced root dry matter, with small reductionsin mean axis length and number of lateral roots. For plantsnot given mineral nutrients reduction in root weight was compensatedby an increase in weight of the aerial parts; however, for plantssupplied with mineral nutrients this was not so and the lowerroot weight resulted in a smaller total plant dry weight. An interpretation of tiller bud growth in terms of control byinteracting effects of mineral nutrition, assimilate supply,and cytokinin availability is proposed.     

Tiller Bud Suppression in Reproductive Plants of Lolium multiflorum Lam. Cv. Westerwoldicum

The control of tiller bud growth during reproductive developmentwas investigated in experimental plants ofLolium multiflorumLam. cv. Westerwoldicum that were reduced to a main axis havinga developing but unemerged ear, elongating stem internodes,a series of expanded leaves, slow-growing tiller buds and aroot system. Isolation of the ear by excision of its base, ordecapitation so as to remove the ear together with the upperleaves, promoted the movement of ¹⁴C-assimilates to tiller buds,decapitation being the more effective treatment. Applicationof 0.1 per cent indol–3yl-acetic acid (IAA) to cut tissuesof decapitated plants diverted ¹⁴C-assimilates to upper internodesbut did not reduce import by buds, whereas application of 1.0per cent IAA both diverted labelled assimilates to upper internodesand reduced bud import. Radioactivity from [¹⁴C] IAA appliedto the upper leaves or to the ear base was recovered from budsin very small amounts; larger amounts were recovered from budsfollowing the application of labelled IAA to an elongating internode,especially from the bud at the base of the treated internode.It is suggested that tiller bud suppression may be influencedby the movement of inhibitory levels of auxin into buds fromnearby elongating stem internodes, whose activity in turn maybe controlled by the developing inflorescence and upper leaves.     

Hormonal Regulation of Lateral Bud (Tiller) Release in Oats (Avena sativa L.)

Stem segments containing a single node and quiescent lateral bud (tiller) were excised from the bases of oat shoots (cv. `Victory') and used to study the effects of plant hormones on release of lateral buds and development of adventitious root primordia. Kinetin (10⁻⁵ and 10⁻⁶ molar) stimulates development of tillers and inhibits development of root primordia, whereas indoleacetic acid (IAA) (10⁻⁵ and 10⁻⁶ molar) causes the reverse effects. Abscisic acid strongly inhibits kinetin-induced tiller bud release and elon-gation and IAA-induced adventitious root development. IAA, in combination with kinetin, also inhibits kinetin-induced bud prophyll (outermost leaf of the axillary bud) elongation. The IAA oxidase cofactor p-coumaric acid stimulates lateral bud release; the auxin transport inhibitor 2,3,5-triiodo-benzoic acid and the antiauxin α (p-chlorophenoxy)-isobutyric acid inhibit IAA-induced adventitious root formation. Gibberellic acid is synergistic with kinetin in the elongation of the bud prophyll. In intact oat plants, tiller release is induced by shoot decapitation, geostimulation, or the emergence of the inflorescence. Results shown support the apical dominance theory, namely, that the cytokinin to auxin ratio plays a decisive role in determining whether tillers are released or adventitious roots develop. They also indicate that abscisic acid and possibly gibberellin may act as modulator hormones in this system.