种子引发对甜高粱角质层蜡质及其抗性的影响

doi:10.11686/cyxb2021487

摘要/Abstract

摘要：

种子引发是一种可有效提高植物抗性及产量的方法，但引发是否可以调节作物生长期角质层蜡质沉积，从而参与植物抗性，目前尚不清楚。以甜高粱为试验对象，其种子用15% 聚乙二醇（PEG）、150 mg·L^-1水杨酸（SA）、20 mg·L^-1 脱落酸（ABA）、5 mg·L^-1赤霉素（GA）或水进行引发处理，之后在田间种植（2020年），分析测定了不同生长阶段（苗期、拔节期、抽穗期、成熟期）叶片角质层蜡质含量、叶片失水率、叶绿素浸提率；收获的种子进行第二年种植（2021年），分析种子引发对植株叶片角质层蜡质的跨代效应。结果表明，PEG、SA和ABA引发分别显著增加苗期蜡质总量57.6%，50.8%和80.3%。引发也可增加不同生育期蜡质烷烃组分的含量，其中，ABA引发效果显著，分别显著增加苗期、抽穗期与成熟期烷烃含量58.7%、35.5%和36.5%。具有不同碳链长度的蜡质组分丰度同样受引发影响，特别是C₂₇、C₂₉、C₃₃烷，C₂₈~C₃₂醛与C₂₈醇相对丰度。与此同时，引发后高粱叶片相对含水量增加，叶片失水率、叶绿素浸提率均下降。综合分析认为，PEG、SA和ABA引发均可起到调节作物角质层蜡质沉积、提高角质层抗性的作用。此外，种子引发对蜡质合成的影响具有跨代“记忆”，可为生产中培育抗性品种提供新方法。

关键词: 甜高粱, 角质层蜡质, 种子引发, 相对水分含量, 失水率, 抗性

Abstract:

Seed priming is an efficient method to improve plant tolerance and yield. However， it is still not clear whether the deposition of plant cuticular wax is altered by seed priming thus improving plant resistance to some kinds of stress. In this study conducted in 2020， seeds of sweet sorghum （Sorghum bicolor ‘Dochna’） were primed with 15% polyethylene glycol （PEG）， 150 mg·L^-1 salicylic acid （SA）， 20 mg·L^-1 abscisic acid （ABA）， 5 mg·L^-1 gibberellin （GA） or water （control）， and subsequently sorghum plants at four growth stages （seedling， jointing， heading and maturity） were analyzed for total wax content， leaf water loss rate and chlorophyll extraction rate. The seeds were harvested and reseeded in 2021， and the transgenerational effects of seed priming on cuticular wax were measured. It was found that， compared to the control， the contents of total wax at different growth stages were significantly influenced， with 57.6%， 50.8% and 80.3% increases observed at the seedling stage for PEG-， SA- and ABA-priming. The contents of alkanes were increased by 58.7%， 35.5% and 36.5% （P<0.05 in each case） at seedling， heading， and maturity stages for ABA-priming. Furthermore， the relative abundances of C₂₇， C₂₉， and C₃₃ alkanes， C₂₈-C₃₂ aldehydes， and C₂₈ alcohols， were also improved. Meanwhile， seed priming increased the leaf relative water content but reduced the water loss rate and chlorophyll extraction rate. We concluded that PEG-， SA- and ABA-priming all influenced the deposition of cuticular wax， contributing to an improvement in cuticular resistance. Importantly， the effects of seed priming on wax synthesis had transgenerational “memory”， indicating a potential new method for production of “resistant” seed lines.

Key words: sweet sorghum, cuticular wax, seed priming, relative water content, water loss rate, resistance

姚露花, 綦才, 杨建峰, 郭彦军. 种子引发对甜高粱角质层蜡质及其抗性的影响[J]. 草业学报, 2022, 31(7): 185-196.

Lu-hua YAO, Cai QI, Jian-feng YANG, Yan-jun GUO. Effects of seed priming on cuticular wax and resistance of sweet sorghum[J]. Acta Prataculturae Sinica, 2022, 31(7): 185-196.

图/表 9

图1 高粱种植期间重庆气温与降水情况

Fig.1 Temperature and precipitation in Chongqing during planting stage of sorghum

图2 种子引发对高粱叶片角质层蜡质总量的影响CK：对照Control；H2O：水引发Deionized water priming；PEG：聚乙二醇引发Polyethylene glycol priming；SA：水杨酸引发Salicylic acid priming；ABA：脱落酸引发Abscisic acid priming；GA：赤霉素引发Gibberellin priming. 不同小写字母表示不同处理间差异显著（P<0.05）。下同。Different lowercase letters indicate significant differences （P<0.05） among treatments. The same below.

Fig.2 Effects of different seed priming on total contents of leaf wax

图3 种子引发对高粱烷烃含量的影响

Fig.3 Effects of different seed priming on total contents of alkanes

图4 种子引发对高粱不同碳链烷烃相对丰度的影响

Fig.4 Effects of different seed priming on the relative abundance of alkanes with different carbon chain of alkanes

图5 种子引发对高粱蜡质醇组分含量及其碳链分布的影响

Fig.5 Effects of different seed priming on total contents of alcohols and carbon chain distribution

图6 种子引发对高粱蜡质醛组分及其碳链分布的影响

Fig.6 Effects of different seed priming on total contents of aldehydes and carbon chain distribution

图7 种子引发对高粱蜡质三萜类组分的影响

Fig.7 Effects of different seed priming on total contents of triterpenoids

图8 引发处理对叶片相对含水量及失水率的影响

Fig.8 Effects on relative water content and water loss rate of seed priming

图9 引发处理对高粱叶片叶绿素含量及叶绿素浸提率的影响

Fig.9 Effects on chlorophyll content and extraction rate of seed priming

参考文献 49

1	Rosa L， Chiarelli D D， Rulli M C， et al. Global agricultural economic water scarcity. Science Advances， 2020， 6（18）： eaaz6031.
2	Vogel E， Donat M G， Alexander L V， et al. The effects of climate extremes on global agricultural yields. Environmental Research Letters， 2019， 14（5）： 054010.
3	Akinseye F M， Ajeigbe H A， Traore P C S， et al. Improving sorghum productivity under changing climatic conditions： A modelling approach. Field Crops Research， 2020， 246： 107685.
4	Lal B， Gautam P， Nayak A K， et al. Tolerant varieties and exogenous application of nutrients can effectively manage drought stress in rice. Archives of Agronomy and Soil Science， 2020， 66（1）： 13-32.
5	Ferrante A， Mariani L. Agronomic management for enhancing plant tolerance to abiotic stresses： High and low values of temperature， light intensity， and relative humidity. Horticulturae， 2018， 4（3）： 21.
6	Ierna A， Mauromicale G. Potato growth， yield and water productivity response to different irrigation and fertilization regimes. Agricultural Water Management， 2018， 201： 21-26.
7	Grant C A， Flaten D N， Tomasiewicz D J， et al. The importance of early season phosphorus nutrition. Canadian Journal of Plant Science， 2001， 81（2）： 211-224.
8	Johnson R， Puthurb J T. Seed priming as a cost effective technique for developing plants with cross tolerance to salinity stress. Plant Physiology and Biochemistry， 2021， 162： 247-257.
9	Lei C， Bagavathiannan M， Wang H， et al. Osmopriming with polyethylene glycol （PEG） for abiotic stress tolerance in germinating crop seeds： A review. Agronomy， 2021， 11（11）： 2194.
10	Zhang F， Yu J L， Johnston C R， et al. Seed priming with polyethylene glycol induces physiological changes in sorghum （Sorghum bicolor L. Moench） seedlings under suboptimal soil moisture environments. PLoS One， 2015， 10（10）： e0140620.
11	Rhaman M S， Imran S， Rauf F， et al. Seed priming with phytohormones： An effective approach for the mitigation of abiotic stress. Plants-Basel， 2021， 10（1）： 37.
12	Fricke W， Akhiyarova G， Veselov D， et al. Rapid and tissue-specific changes in ABA and in growth rate in response to salinity in barley leaves. Journal of Experimental Botany， 2004， 55（399）： 1115-1123.
13	Iqbal M， Ashraf M. Seed treatment with auxins modulates growth and ion partitioning in salt-stressed wheat plants. Journal of Integrative Plant Biology， 2007， 49（7）： 1003-1015.
14	Yeats T H. The formation and function of plant cuticles. Plant Physiology， 2013， 163（1）： 5-20.
15	Ding F， Wang G， Wang M， et al. Exogenous melatonin improves tolerance to water deficit by promoting cuticle formation in tomato plants. Molecules， 2018， 23（7）： 1605.
16	Wang Y， Wang M， Sun Y， et al. FAR5， a fatty acyl-coenzyme A reductase， is involved in primary alcohol biosynthesis of the leaf blade cuticular wax in wheat （Triticum aestivum L.）. Journal of Experimental Botany， 2015， 66（5）： 1165-1178.
17	Xiao Y， Li X T， Yao L H， et al. Chemical profiles of cuticular waxes on various organs of Sorghum bicolor and their antifungal activities. Plant Physiology and Biochemistry， 2020， 155： 596-604.
18	Li H， Guo Y L， Cui Q， et al. Alkanes （C₂₉ and C₃₁）-mediated intracuticular wax accumulation contributes to melatonin- and ABA-induced drought tolerance in watermelon. Journal of Plant Growth Regulation， 2020， 39（4）： 1441-1450.
19	Martin B B， Romero P， Fich E A， et al. Cuticle biosynthesis in tomato leaves is developmentally regulated by abscisic acid. Plant Physiology， 2017， 174（3）： 1384-1398.
20	Oliveira A F M， Meirelles S T， Salatino A. Epicuticular waxes from caatinga and cerrado species and their efficiency against water loss. Anais Da Academia Brasileira De Ciencias， 2003， 75（4）： 431-439.
21	Weng H， Molina I， Shockey J， et al. Organ fusion and defective cuticle function in a lacs1 lacs2 double mutant of Arabidopsis. Planta， 2010， 231（5）： 1089-1100.
22	Bourdenx B， Bernard A， Domergue F， et al. Overexpression of Arabidopsis ECERIFERUM1 promotes wax very-long-chain alkane biosynthesis and influences plant response to biotic and abiotic stresses. Plant Physiology， 2011， 156（1）： 29-45.
23	Tomasi P， Wang H， Lohrey G T， et al. Characterization of leaf cuticular waxes and cutin monomers of Camelina sativa and closely-related Camelina species. Industrial Crops and Products， 2017， 98： 130-138.
24	Singh V K， Singh R， Tripathi S， et al. Seed priming： State of the art and new perspectives in the era of climate change//Climate change and soil interactions. Amsterdam， Netherlands： Elsevier， 2020： 143-170.
25	Wang Y， Chang J， Cai D， et al. Repeated-batch fermentation of L-lactic acid from acid hydrolysate of sweet sorghum juice using mixed neutralizing agent under unsterilized conditions. Journal of Chemical Technology and Biotechnology， 2017， 92（7）： 1848-1854.
26	Zhang C， Wen H， Zheng J， et al. A combination of evaporation and chemical preservation for long-term storage of fresh sweet sorghum juice and subsequent bioethanol production. Journal of Food Processing and Preservation， 2018， 42（12）： e13825.
27	Patil J V， Reddy P S， Prabhakar， et al. History of post-rainy season sorghum research in India and strategies for breaking the yield plateau. Indian Journal of Genetics and Plant Breeding， 2014， 74（3）： 271-285.
28	Zhang X， Ni Y， Xu D X， et al. Integrative analysis of the cuticular lipidome and transcriptome of Sorghum bicolor reveals cultivar differences in drought tolerance. Plant Physiology and Biochemistry， 2021， 163： 285-295.
29	Bao S D. Agricultural chemical analysis of soil. Beijing： China Agriculture Press， 2005.
	鲍士旦. 土壤农化分析. 北京：中国农业出版社， 2005.
30	Sheteiwy M S， Fu Y， Hu Q， et al. Seed priming with polyethylene glycol induces antioxidative defense and metabolic regulation of rice under nano-ZnO stress. Environmental Science and Pollution Research， 2016， 23（19）： 19989-20002.
31	Buschhaus C， Jetter R. Composition and physiological function of the wax layers coating Arabidopsis leaves： Beta-amyrin negatively affects the intracuticular water barrier. Plant Physiology， 2012， 160（2）： 1120-1129.
32	Yao L H， Guo N， He Y J， et al.Variations of soil organic matters and plant cuticular waxes along an altitude gradient in Qinghai-Tibet Plateau. Plant and Soil， 2019， 458（1/2）： 41-58.
33	Weatherley H B A P. A re-examination of the relative turgidity technique for estimating water deficits in leaves. Australian Journal of Biological Sciences， 1962， 15（3）： 413-428.
34	Burkhardt J， Pariyar S. Particulate pollutants are capable to ‘degrade’ epicuticular waxes and to decrease the drought tolerance of Scots pine （Pinus sylvestris L.）. Environmental Pollution， 2014， 184： 659-667.
35	Chen H M， Xie Z W， Liu P， et al. Impact of water stress on leaf epicuticular wax accumulation of rice. Genomics and Applied Biology， 2013， 32（6）： 777-781.
	陈红妙，谢振文，刘平，等. 干旱胁迫对水稻叶表面蜡质积累的影响. 基因组学与应用生物学， 2013， 32（6）： 777-781.
36	Lutts S， Benincasa P， Wojtyla L， et al. Seed priming： New comprehensive approaches for an old empirical technique//New challenges in seed biology-basic and translational research driving seed technology. Rijeka， Croatia： InTech， 2016.
37	Chen K， Arora R. Priming memory invokes seed stress-tolerance. Environmental and Experimental Botany， 2013， 94： 33-45.
38	Ibrahim E A. Seed priming to alleviate salinity stress in germinating seeds. Journal of Plant Physiology， 2016， 192： 38-46.
39	Shaheenuzzamn M， Shi S， Sohail K， et al. Regulation of cuticular wax biosynthesis in plants under abiotic stress. Plant Biotechnology Reports， 2021， 15： 1-12.
40	Romero P， Lafuente M T. Relative humidity regimes modify epicuticular wax metabolism and fruit properties during Navelate orange conservation in an ABA-dependent manner. Food Chemistry， 2022， 369： 130946.
41	Chaudhary K， Geeta R， Panjabi P. Origin and diversification of ECERIFERUM1 （CER1） and ECERIFERUM3 （CER3） genes in land plants and phylogenetic evidence that the ancestral CER1/3 gene resulted from the fusion of pre-existing domains. Molecular Phylogenetics and Evolution， 2021， 159： 107101.
42	Li D T， Dai Y T， Chen X， et al. Ten fatty acyl-CoA reductase family genes were essential for the survival of the destructive rice pest， Nilaparvata lugens. Pest Management Science， 2020， 76（7）： 2304-2315.
43	Shepherd T， Griffiths D W. The effects of stress on plant cuticular waxes. New Phytologist， 2006， 171（3）： 469-499.
44	Wang D J， Ni Y， Liao L X， et al. Poa pratensis ECERIFERUM1 （PpCER1） is involved in wax alkane biosynthesis and plant drought tolerance. Plant Physiology and Biochemistry， 2021， 159： 312-321.
45	Lewandowska M， Keyl A， Feussner I. Wax biosynthesis in response to danger： Its regulation upon abiotic and biotic stress. New Phytologist， 2020， 227（3）： 698-713.
46	Leide J， Hildebrandt U， Reussing K， et al. The developmental pattern of tomato fruit wax accumulation and its impact on cuticular transpiration barrier properties： Effects of a deficiency in a beta-ketoacyl-coenzyme A synthase （LeCER6）. Plant Physiology， 2007， 144（3）： 1667-1679.
47	Seo P J， Lee S B， Suh M C， et al. The MYB96 transcription factor regulates cuticular wax biosynthesis under drought conditions in Arabidopsis. Plant Cell， 2011， 23（3）： 1138-1152.
48	Jahan M S， Shu S， Wang Y， et al. Melatonin alleviates heat-induced damage of tomato seedlings by balancing redox homeostasis and modulating polyamine and nitric oxide biosynthesis. BMC Plant Biology， 2019， 19（1）： 414.
49	Yuan Z， Jiang Y， Liu Y， et al. Exogenous hormones influence Brassica napus leaf cuticular wax deposition and cuticle function. PeerJ， 2020， 8： e9264.

[1]	杨德智, 王晨, 侯明杰, 王虎成. 饲用甜高粱和全株玉米青贮对肉羊前胃微生态的影响[J]. 草业学报, 2022, 31(4): 145-154.
[2]	王志恒, 魏玉清, 赵延蓉, 王悦娟. 基于转录组学比较研究甜高粱幼苗响应干旱和盐胁迫的生理特征[J]. 草业学报, 2022, 31(3): 71-84.
[3]	李彦忠, 喻军强, 李明. 48个苜蓿品种对3种病害抗性的初步评价[J]. 草业学报, 2021, 30(9): 62-75.
[4]	刘建新, 刘瑞瑞, 贾海燕, 卜婷, 李娜. NaHS引发提高裸燕麦种子活力的生理机制[J]. 草业学报, 2021, 30(2): 135-142.
[5]	常丹丹, 王旭, 田新会, 杜文华. 甘肃中部地区秋播小黑麦套作式复种甜高粱的效应及品质研究[J]. 草业学报, 2021, 30(11): 212-220.
[6]	再吐尼古丽·库尔班, 吐尔逊·吐尔洪, 涂振东, 王卉, 山其米克, 艾克拜尔·伊拉洪. 长期不同施肥处理对连作高粱生长规律及产量的影响研究[J]. 草业学报, 2020, 29(8): 81-92.
[7]	周恩光, 王虎成, 尚占环. 甜高粱的饲用价值及其绵羊体外瘤胃发酵产气性能研究[J]. 草业学报, 2020, 29(5): 43-49.
[8]	高丽敏, 田倩, 苏晶, 沈益新. 施氮水平对甜高粱干物质产量及氮肥利用率的影响[J]. 草业学报, 2020, 29(4): 192-198.
[9]	王婷, 王虎成, 苟娜娜, 冯强, 贺春贵, 尚占环. 甜高粱饲草及葡萄籽对小尾寒羊生产性能及血液生理参数的影响[J]. 草业学报, 2019, 28(9): 155-163.
[10]	姜红岩, 滕珂, 檀鹏辉, 尹淑霞. 日本结缕草ZjZFN1基因对拟南芥的转化及其耐旱性分析[J]. 草业学报, 2019, 28(4): 129-138.
[11]	郭志鹏, 冯长松, 张靖雪, 王苗利, 曲根, 刘建宇, 管永卓, 张晓婷, 郭玉霞, 严学兵. 30个紫花苜蓿品种对苜蓿花叶病毒病的田间抗性初步研究[J]. 草业学报, 2019, 28(4): 157-167.
[12]	孙铭, 王思琪, 艾尔肯·达吾提, 毛培胜. 抗氧化剂引发对无芒雀麦老化种子发芽及幼苗生长的影响[J]. 草业学报, 2019, 28(11): 105-113.
[13]	王亚麒, 袁玲. 甜高粱、高丹草和拉巴豆对难溶性磷的活化与吸收[J]. 草业学报, 2019, 28(10): 33-43.
[14]	李晓婷, 赵晓, 王登科, 黄蕾, 姚露花, 王党军, 和玉吉, 郭彦军. 天然草地植物叶角质层蜡质的化学组成及其对自由放牧的响应[J]. 草业学报, 2018, 27(6): 137-147.
[15]	赵春旭, 姜寒玉, 董文科, 陈红, 方彦霞, 谢丽萍, 马晖玲. 外源复合激素对匍匐翦股颖抗褐斑病的诱导抗性[J]. 草业学报, 2018, 27(11): 120-130.