燕麦产量形成生理机制研究进展
Research Progress of Physiological Mechanism of Yield Formation in Oats
收稿日期: 2020-07-8 修回日期: 2020-09-10 网络出版日期: 2021-05-14
基金资助: |
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Received: 2020-07-8 Revised: 2020-09-10 Online: 2021-05-14
作者简介 About authors
赵宝平,主要从事作物生态生理研究,E-mail:
燕麦是我国北方重要的粮饲兼用作物。低产是制约燕麦产业健康稳定发展的主要问题。本文从燕麦小穗多花多粒特性、小穗不孕性和抗倒伏性能等角度分析了燕麦产量形成的主要特征;并从产量构成因素、光合生产性能和源库关系等方面系统梳理和总结了燕麦产量形成的生理机制研究进展,比较了皮燕麦和裸燕麦产量形成的生理学差异,提出了提高燕麦产量的技术途径。
关键词:
Oat is an important grain and forage crop in Northern China. Lower grain yield is a main problem restricting the healthy and stable development of the oat industry. We analyzed the yield formation characteristics from the aspects of spikelet fertility, multiflorous characteristics, panicle infertility and lodging. We also summarized the physiological mechanism of oats yield formation from the yield components and photosynthetic production performance and source-sink relationship, and emphatically compared the differences of yield formation between the hulled and the naked oats. We proposed forward future research directions and emphases on oat yield improvement.
Keywords:
本文引用格式
赵宝平, 刘景辉, 任长忠.
Zhao Baoping, Liu Jinghui, Ren Changzhong.
我国是裸燕麦(莜麦)起源地,具有悠久的种植历史和饮食文化传统,裸燕麦也是我国主要的燕麦栽培品种类型[1, 4]。然而,由于长期以来对燕麦的研究和重视程度不够,当前世界及我国燕麦籽粒产量平均水平远远低于小麦和大麦等麦类作物[5]。而裸燕麦的平均单产水平又低于国外主要种植的皮燕麦产量[6,7]。此外,国外燕麦主要种植在土壤肥沃和水分条件好的地区,栽培管理水平和机械化程度相对较高;而我国燕麦主要在偏远、干旱及贫瘠等土壤地区种植[8],水分、土壤养分及管理技术较为落后,干旱、土壤瘠薄等非生物胁迫及不合理的栽培管理措施严重影响了燕麦单产水平,低产低效问题也严重影响燕麦种植效益和产业健康发展。因此,本研究通过分析燕麦籽粒产量形成过程的主要特征,从产量构成因素、光合生产性能和源库关系等方面系统地梳理和总结国内外燕麦产量形成的生理机制研究进展,比较分析皮燕麦和裸燕麦产量形成的生理学差异,提出未来燕麦产量潜力提升的主要研究方向和重点,以期为我国燕麦高产栽培和品种改良提供参考。
1 燕麦籽粒产量形成的主要特征
1.1 燕麦穗及花序形态学特征与产量形成
1.1.1 燕麦小穗多花多粒特性 燕麦小穗第3位粒结实和籽粒败育机制是燕麦区别于其他麦类作物的2个独特的特征[11]。通常,皮燕麦的小穗中通常有2个小花(也有3或4个)。与皮燕麦相比,裸燕麦每个小穗的小花数和结实粒数有3~8个,甚至12个,呈现出更加明显的多花多粒特性[5, 12-13],并且具有更长的小穗轴[14]。多花多粒特性是裸燕麦区别于皮燕麦的最大特点,一些学者推测裸燕麦多花的特性会增加小穗粒数使得其籽粒产量潜力高于皮燕麦[5]。然而,很多研究发现,裸燕麦的产量潜力低于皮燕麦[7, 15]。王茜等[16]通过比较13个皮燕麦和7个裸燕麦品种的籽粒产量发现,皮燕麦产量(带壳)接近裸燕麦的2倍,具有明显的产量优势。Peltonen-Sainio[6]认为裸燕麦籽粒低产主要是由于其单穗小穗数少而小穗粒数多造成的。尤其是裸燕麦小穗中的第3位粒及之后的小花常常会退化或结实率降低,灌浆速率下降,有超过70%的小花败育,远大于小麦的50%[10],并且导致粒重(弱势粒)显著下降,经济产量降低以及籽粒大小不均匀[17,18],影响燕麦加工利用[18]。
燕麦小穗不孕特性存在基因型差异。不同地区的燕麦种质资源的小穗不孕率存在很大差异,其中山西省的品种资源不孕率为4%,而内蒙古的种质资源不孕率达到14%;皮燕麦与裸燕麦之间的小穗不孕率差异不显著[23]。此外,抗旱、耐高温程度不同的品种其小穗不孕率也存在差异,抗性强的品种小穗不孕率低[21]。裸燕麦小穗不孕主要发生在穗的下部,中部次之,上部极少,这主要与其幼穗分化顺序和花序分枝特点有关[13, 24],即每小穗平均维管束数量由上部一次枝梗的8个下降到底端分枝的2个,使得不孕小穗主要发生在穗的下部,说明小穗不孕与维管束系统的同化物运输能力关系密切[25]。在田间试验中也发现燕麦小穗数与其维管束数量和面积关系密切,并且不同环境条件对维管束形态学特征的影响存在基因型差异[26]。
1.2 燕麦抗倒伏性能
倒伏的原因主要与燕麦株高有关[30,31]。裸燕麦与皮燕麦的株高一般均在100cm以上,是燕麦更易倒伏的主要原因[3]。Ma等[15]和Zhou等[32]通过对裸燕麦、皮燕麦与小麦比较发现,增施氮肥使裸燕麦和皮燕麦均出现比小麦更严重的倒伏现象,并限制了灌浆期光合产物和干物质积累,导致籽粒产量更低。此外,燕麦的易倒伏性也限制了氮肥投入的增加以获得更高籽粒产量。燕麦倒伏性还与株型、茎粗、茎秆强度和弹性等特性有关[30]。拥有直立叶片的株型和坚实根系的燕麦品种具有较强的抗倒性[27]。Wu等[33]研究发现,叶鞘持绿性好的燕麦品种可提高茎秆强度和硬度,对茎杆抗折倒的贡献率更高。Ma等[34]研究得出燕麦茎秆中全磷含量低于13.6kg/hm2时,倒伏很少发生,推测茎秆中氮和磷含量增加可能降低了茎秆强度而导致倒伏。关于皮、裸燕麦抗倒伏性差异,研究发现控制裸粒性的N1基因位点通过调控木质素向小穗或茎中沉积,使茎中木质素含量发生变化,进而影响倒伏[31]。
2 燕麦产量形成的生理机制研究
2.1 籽粒产量与产量构成因素
由以上分析可知,穗粒数是决定燕麦产量的主要因素,说明小穗和小花的发育状况对燕麦产量潜力具有决定性作用[35,36]。在生产实践和研究中发现,影响燕麦穗粒数的原因主要包括2个方面:一是品种差异。燕麦每穗小穗数与穗长、轮层数和每轮层第1分枝数密切相关,因此选择穗轴节点数较多的品种可增加其小穗数[24]。Wang等[8]发现,近年育成的裸燕麦品种相对于早期地方品种而言,在干旱下拥有更高的光合产物向穗部分配能力和灌浆速率,能够更快转化为籽粒产量,其穗粒数和粒重高于早期地方品种。Peltonen-Sainio等[10]研究发现,开花前生育日数短的品种的小花数会减少,但籽粒产量未下降。二是同化物的竞争。果穗在有限资源的情况下对同化物的竞争能力影响了小花的成粒数[9]。在研究中发现穗粒数与茎秆中可溶性碳水化合物含量呈显著负相关[40],从而减少穗发育早期籽粒的败育[43]。研究[44]发现小花发育速度与植株体内碳素水平的高低密切相关,在保持一定氮素水平供应条件下,植株体内碳水化合物含量越高,越有利于可孕小花的发育。外源激素也会影响燕麦小穗小花发育。随着喷施外源细胞分裂素苄氨基嘌呤(6-BA)浓度增加,燕麦不孕小穗数也相应增加,只有在1×10-8和1×10-3g/L处理下可育小穗和小花数增加,但不孕小穗和小花的比例没有减少[9]。以上研究说明不同基因型品种穗粒数形成与光合产物向穗部的分配能力以及生理变化等密切相关,但不同穗型、株型燕麦穗粒数的变化、小穗数或小穗粒数之间的关系以及其形成过程与生理学途径尚不明确。
通过栽培管理调节穗内部结构可改善运输系统输送光合产物的能力[45]。研究[46]发现施用氮肥可有效增加有效分蘖数和穗粒数,因此单位面积粒数增加。在开花前增施氮肥显著增加皮燕麦的小穗数和结实率,从而增加了穗粒数,对小穗粒数影响不大[10];在孕穗前减少氮素供应其小穗数显著减少[47]。Peltonen-Sainio[6]研究发现,增加播种量和施氮量对小穗粒数没有显著影响,却显著减少了燕麦小穗数和穗粒数。此外,燕麦的穗粒数与光合有效辐射(特别是开花前及开花期)的截获密切相关,通过合理氮肥管理可以实现光合有效辐射利用最大化,从而提高燕麦穗粒数[48]。说明氮素对燕麦小穗小花发育及穗粒数形成影响显著,但不同养分管理调控措施对燕麦小穗和小花败育的生理机制影响尚不明确。
2.2 燕麦光合性能
绿色革命以来主要作物增产原因分析表明,作物总光合能力的增加主要体现在光合面积和光合持续时间的增加上,而单位叶面积光合速率并没有增加,甚至出现下降[49]。在燕麦的早期研究中发现,燕麦籽粒产量与植株干物质积累量呈正相关关系[50,51]。然而,Ma等[15]研究发现在高氮处理下,虽然皮燕麦或裸燕麦的生物量与小麦相似,但其籽粒产量显著低于小麦,主要原因是皮燕麦或裸燕麦的收获指数较低且开花后更易倒伏。此外,研究[7, 52]发现裸燕麦的生物产量高于皮燕麦,然而其籽粒产量却低于皮燕麦,可能是裸燕麦在开花期叶面积指数(LAI)较高而收获指数较低导致的[15]。Ma等[15]还进一步探讨了LAI和干物质积累量的关系,随着施氮量增加,LAI和干物质积累量呈显著正相关关系,但在高氮处理下燕麦的干物质积累量并没有增加,可能是由于LAI与光合辐射截获量之间并不是线性关系,即当LAI大于3时,光合有效辐射截获率达到90%以上,LAI进一步增加时光截获量不再增加,并且易引起倒伏。
在单叶光合能力和叶面积持续期研究方面,Hisir等[53]研究得出,燕麦籽粒产量与叶片叶绿素含量和叶面积持续期呈显著正相关关系。研究还发现新育成的燕麦品种开花后叶面积衰老速度低于老品种,有利于籽粒灌浆[54,55]。而Peltonen-Sainio[56]在生长期较短的芬兰高纬度地区研究发现,籽粒产量与叶面积持续期并不存在相关关系。此外,增加追氮用量可延缓开花后叶片衰老,提高光合能力[57]。Sadras等[40]通过对29个燕麦品种的研究得出,燕麦籽粒产量与灌浆期叶片叶绿素相对值呈显著正相关。Sadras等[58]进一步比较了16个燕麦品种后发现,可溶性糖含量低的燕麦品种可增加叶面积持续期和单位面积粒数进而提高产量,并提出水溶性碳水化合物含量低的性状可作为高产燕麦品种选择的重要标准。说明开花后叶片持绿性好和较长的叶面积持续期是燕麦获得高产的保证。
2.3 燕麦产量源库关系
作物产量可能会受到源活性、库活性或者源库关系2个方面的限制[43]。与小麦、大麦等其他麦类作物的穗状花序不同,燕麦的圆锥状花序绿色面积大且分散,有利于截获太阳辐射,提高开花后花序光合作用对产量的贡献,因此在源库关系方面与其他麦类作物存在许多差异[11, 17]。在源库限制方面,在开花盛期由于光合产物供需不平衡导致完全发育小花之间竞争加剧出现籽粒败育,导致库容改变,并且单位面积籽粒数多的群体中籽粒败育率高[11]。不同时期外部环境胁迫会导致籽粒败育情况发生。Doehlert等[61]发现春季干旱导致光合产物供应减少,小穗籽粒败育率上升,超过30%小穗出现籽粒败育,并提出利用第3位粒结实可提高燕麦库容大小。第3位粒灌浆结实能力可塑性取决于灌浆期光合产物的供给状况。Doehlert等[61]研究发现9.5%的小穗有第3位粒结实,而Browne等[11]认为只有不超过5%的第3位粒结实。这意味着在关键生育阶段,由于外界环境胁迫导致光合能力下降(源限制),进而影响小穗籽粒败育或第3位粒结实(库容变化)以及维管束系统发育(流限制),最终影响燕麦籽粒结实和灌浆。
对于裸燕麦来说,其多花多粒特性会表现出更明显的籽粒结实粒数可塑性[5]。然而,由于裸燕麦的小穗轴比皮燕麦更长以及特殊的小花形态[62],导致进入穗部的光合产物减缓,灌浆速率受到影响。因此,裸燕麦的多花多粒性没有使燕麦库容量增大,反而导致库活性(灌浆速率)下降,未能成为增加产量潜力的优势,而成为产量形成的劣势。Finnan等[35]通过系统总结发现,在单位面积粒数达到某一值(25 000粒/m2)之后,裸燕麦的籽粒产量会达到最高值(拐点),之后不再增长,在籽粒产量达到拐点之前表现出明显的库限制,达到拐点之后即表现为源限制,并且拐点位置还与品种和环境因素密切相关。为此进一步提出,在产量没有达到最高值之前应考虑增加投入来增加粒数,在产量达到最高之后可通过提高光合同化能力,促进籽粒灌浆,提高库强度。Zhao等[63]通过比较不同水分条件下皮燕麦、裸燕麦源库关系发现,裸燕麦在充分供水条件下的籽粒产量高于皮燕麦(脱壳后)的主要原因是裸燕麦具有更高的源活性和单穗小穗数。因此通过育种或栽培等调控途径增加每穗小穗数,而不是小穗粒数,可能是提高裸燕麦穗粒数(库容)的重要策略。
3 展望
从以上对燕麦产量形成生理机制的研究分析发现,燕麦产量潜力较低,既有品种本身遗传和生理特性的原因,又有环境胁迫和栽培条件不适宜的因素,还有对燕麦研究不重视和投入不够等问题。因此,为了提升我国燕麦籽粒产量潜力,需重点从以下4个方面加强研究或投入。
3.1 燕麦种质资源创新与新品种选育
燕麦产量是多基因控制的数量性状,随着二代测序技术的快速发展,利用全基因组关联分析(genome-wide association study)定位与产量性状相关的QTL,结合转录组测序,SNP-index分析鉴定候选基因,验证基因功能,挖掘优异基因资源,创制收获指数高、产量潜力大、抗倒性强以及抗旱、耐瘠薄、耐盐碱等抗性强的燕麦种质资源;利用现代分子设计育种与传统育种技术相结合等方法,通过降低株高、改变株型和提高收获指数等途径选育矮秆抗倒燕麦品种,解决倒伏问题并提高光合产物的同化利用效率;通过表型鉴定和分子标记辅助选择等手段选育抗旱、耐瘠薄和耐盐碱的燕麦品种,解决燕麦产量低且不稳的问题。
3.2 燕麦抗逆及高产生理机制研究
针对我国燕麦主产区干旱和土壤瘠薄等环境特点,开展燕麦对非生物胁迫适应机制、抗逆生理基础及调控机制,从转录组、蛋白组和代谢组等组学层面研究不同逆境胁迫下燕麦产量的形成规律及其与环境的关系。在产量生理方面,重点从皮燕麦和裸燕麦产量形成生理学差异角度,研究燕麦穗粒数的形成规律,并从基因表达、激素平衡和碳氮代谢等角度系统地开展燕麦小穗不孕与花粒败育的生理机制研究。此外,随着燕麦部分染色体基因组序列拼接的完成和部分转录组测序数据的释放,可通过高通量测序数据结合产量形成生理性状定位控制燕麦产量主要性状基因并解析调控产量形成分子机制,揭示提高燕麦产量的关键生理学途径。
3.3 燕麦高产、抗逆栽培技术研发
通过种植密度、水分和养分管理等栽培耕作调控措施来协调株型发育与籽粒形成、源库关系、地上部与地下部生长关系,提高燕麦抗逆性和籽粒产量。通过外源激素和水肥耦合等栽培调控手段协调植株衰老、光合作用与同化物向籽粒转运关系,促进同化物向籽粒转运和灌浆,挖掘裸燕麦多花多粒特性、增加穗粒数的潜力,克服粒重下降的弱势,提升燕麦产量潜力。
3.4 加大对燕麦产业的扶持力度
1960-2005年,全世界燕麦单产只增加了39%,而同期小麦和玉米单产分别增加了147%和143%[64]。其主要原因是在燕麦育种和农学等方面研究的投入较少。因此,要进一步加大对燕麦种质资源创新、品种选育和高产栽培生理等方面的研究力度,提升科技创新对燕麦产业发展的驱动和推动能力,鼓励和引导高校及科研院所建立协同创新平台和联合研发基地或实验室,以提高燕麦产量。
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