花后干旱对西藏青稞叶片水势、光合生理、籽粒表型和产量影响

doi:10.3864/j.issn.0578-1752.2018.14.005

摘要/Abstract

摘要： 【目的】探索高原环境下青稞花后干旱胁迫响应模型及青稞受旱程度的快速、有效的检测方法，为青稞节水高产栽培提供理论依据和技术参考。【方法】利用干旱棚进行青稞盆栽模拟花后干旱，设轻度（对照灌水量的75%，LD）、中度（对照灌水量的50%，MD）和重度（对照灌水量的25%，HD）干旱胁迫处理，采用WP4C水势仪、LI-6400XT和OS5P便携式脉冲调制叶绿素荧光仪，分别测定叶水势（LWP）、叶蒸发冷却值（ΔT）、光合气体交换参数和叶绿素荧光变量；成熟后获取产量数据；利用数字图像法对籽粒表型进行定量分析，并依粒二维面积大小将籽粒划分大、中、小3个粒级。【结果】干旱水平与叶水势呈线性正相关，而与叶蒸发冷却值（ΔT）呈线性显著负相关（P＜0.05），两者均能灵敏反映青稞受旱程度。LD、MD和HD处理与CK相比，干旱胁迫导致旗叶净光合速率（P_n）、气孔导度（g_s）、胞间CO₂浓度（C_i）、蒸腾速率（T_r）、最大荧光（F_m）、PS II的最大量子效率（F_v/F_m）、PS II的实际光量子产量（ΦPSII）、光化学淬灭（qP）、光合电子传递的相对速率（ETR）呈降低趋势，而气孔限制（L_s）、初始荧光（F_o）、非光化学淬灭（NPQ）呈上升趋势，且MD和HD处理较CK对以上参数差异明显。随着干旱胁迫的加重，青稞千粒重、籽粒产量、单株粒重、干物质积累量和经济系数降低趋势愈明显。相关分析表明，干旱胁迫诱导g_s降低，直接导致?T上升，间接引起FWP下降，使得F_o、NPQ上升，F_m、F_v/F_m、ΦPSII、ETR、qP和P_n降低，进而引起粒二维面积、粒周长、粒长和粒宽减小，而粒圆度值增大；小粒占比明显增加，而大粒占比明显下降。【结论】LWP和?T对青稞花后干旱胁迫反映灵敏，可作为评价其受旱的指标。随着干旱胁迫加重，青稞旗叶光合和叶绿素荧光参数的变化加大，并造成5个籽粒表型性状值及粒级逐渐减小，最终导致千粒重、单穗粒重、籽粒产量、干物质积累量和经济系数下降。

关键词: 青稞, 花后干旱, 水势, 叶绿素荧光, 产量

Abstract: 【Objective】 Study on a drought stress response model of highland barley after anthesis in the plateau environment was established and a rapid, and effective detection method for the drought degree of highland barley was developed, so as to provide theoretical basis and technical reference for water-saving and high-yield cultivation of highland barley.【Method】The pot experiment of highland barley was carried out in drought shed at after anthesis, and light drought stress treatment (75% of controlled irrigation amount, LD), moderate drought stress treatment (50% of controlled irrigation amount, MD) and heavy drought stress treatment (25% of controlled irrigation amount, HD) were conducted for 13 days. WP4C PotentiaMeter, LI-6400XT and OS5P portable pulse modulated chlorophyll fluorometer were used to measure the leaf water potential (LWP), leaf evaporative cooling value (?T), photosynthetic gas exchange parameter and chlorophyll fluorescence variable, respectively. After maturity, plant laboratory test was conducted to obtain production data. Moreover, precise quantitative analysis of grain phenotypes of different treatments was conducted by digital image analysis method and grains were divided into three grain levels: large, medium and small according to the two-dimensional grain size.【Result】Drought level was positively correlated with leaf water potential and negatively correlated with leaf evaporative cooling value (?T) (P<0.05). Both of them were sensitive to reflect the drought degree of highland barley. Compared with CK, for LD, MD and HD treatments, the net photosynthetic rate of flag leaves (Pn), stomatal conductance (g_s), intercellular CO₂ concentration (Ci), transpiration rate (Tr), maximum fluorescence (Fm), maximum quantum efficiency of PS II (Fv/Fm), actual light quantum yield of PS II (ΦPSII), photochemical quenching (qP), and relative rate of photosynthetic electron transport (ETR) decreased due to drought stress, while stomatal limitation (Ls), original fluorescence (Fo) and non-photochemical quenching (NPQ) were on the rise. The effects of MD and HD on the above parameters were more significant compared with CK. The plant laboratory test data showed that with the increase of drought stress, the decreasing trend of 1000-grain weight, grain yield, grain weight per plant, dry matter accumulation, and economic coefficient of highland barley was more obvious. The correlation analysis showed that the decrease of g_s induced by drought stress directly led to the increase of ?T and indirectly led to the decrease of FWP, which resulted in the increase of Fo, NPQ and the decrease of Fm, Fv/Fm, ΦPSII, ETR, qP and Pn, thus causing the decrease of two-dimensional grain size, grain diameter, grain perimeter, grain length and grain width and the increase of grain roundness. The proportion of small grains increased obviously, while the proportion of large grains decreased obviously.【Conclusion】LWP and ?T were sensitive to the drought stress of highland barley after anthesis, which could be used as an effective method to evaluate the drought degree. With the aggravation of drought stress, the changes of photosynthetic and chlorophyll fluorescence parameters in flag leaves of highland barley increased, causing the gradual decrease in phenotypic characters and grain levels of five grains, thus resulting in the decrease of 1000-grain weight, grain weight per plant, grain yield, dry matter accumulation and economic coefficient.

Key words: hulless barley, post-anthesis drought, water potential, chlorophyll fluorescence, yield

侯维海，王建林，胡单，冯西博. 花后干旱对西藏青稞叶片水势、光合生理、籽粒表型和产量影响[J]. 中国农业科学, 2018, 51(14): 2675-2688.

HOU WeiHai, WANG JianLin, HU Dan, FENG XiBo. Effects of Drought in Post-Flowering on Leaf Water Potential, Photosynthetic Physiology, Seed Phenotype and Yield of Hulless Barley in Tibet Plateau[J]. Scientia Agricultura Sinica, 2018, 51(14): 2675-2688.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2018.14.005

https://www.chinaagrisci.com/CN/Y2018/V51/I14/2675

参考文献

[1] Kebbas S, Lutts S, Aid F. Effect of drought stress on the photosynthesis of Acacia tortilis subsp. raddiana at the young seedling stage. Photosynthetica, 2015, 53(2): 288-298.

[2] MAKSUP S, ROYYTSKULS, SUPAIBULWATANA K. Physiological and comparative proteomic analyses of Thai jasmine rice and two chick cultivars in response to drought stress. Journal of Plant Interactions, 2014, 9(1): 43-55.

[3] AMINIAN R, MOHAMMADI S, HOSHMAND S, KHODOMBASHI M. Chromosomal analysis of photosynthesis rate and stomatal conductance and their relationships with grain yield in wheat (Triticum aestivum L.) under water-stressed and well-watered conditions. Acta Physiologiae Plantarum, 2011, 33(3): 755-764.

[4] BISWAL A K, KOHLI A. Cereal flag leaf adaptations for grain yield under drought: knowledge status and gaps. Molecular Breeding, 2013, 31(4): 749-766.

[5] 关卫星. 浅谈西藏高寒农区发展旱作农业生产存在的问题及建议. 西藏农业科技, 2017, 39(2): 1-2.

GUAN W X. Discussion on problems and suggestions of developing dry farming in alpine agricultural area of Tibet. Tibet journal of agricultural sciences, 2017, 39(2): 1-2. (in Chinese)

[6] CECCARELLI S. Drought and drought resistance. Encyclopedia Biotechnol. Agric Food, 2010, 1(1): 205-207.

[7] SHANMUGAM S, KJAER K H, OTTOSEN C O, ROSENQVIST E, SHARMA D K, WOLLENWEBER B. The alleviating effect of elevated CO₂on heat stress susceptibility of two wheat (Triticum aestivum L.) cultivars. Journal of Agronomy and Science, 2013, 1995(5): 340-350.

[8] 王艺陶, 周宇飞, 李丰先, 苏仲. 干旱胁迫下高粱叶温与叶片水分状况的关系. 干旱地区农业研究, 2013, 31(6): 146-151.

WANG Y T, ZHOU Y F, LI F X, SU Z. Relationship between leaf temperature and water status in sorghum under drought stress. Agricultural Research in the Arid Areas, 2013, 31(6): 146-151. (in Chinese)

[9] ROMANO G, ZIA S, SPREER W, SANCHEZ C, CAIRNS J, ARAUS J L, MULLER J. Use of thermography for high throughput phenotyping of tropical maize adaptation in water stress. Computers and electronics in agriculture, 2011, 79(1): 67-74.

[10] KALAJI M H, SCHANSKER G, LADLE R J, GOLTSEV V, BOSA K, ALLAKHVERDIEV I, BRESTIC M, BUSSOTTI F. Frequently asked questions about in vivo chlorophyll fluorescence: practical issues. Photosynthesis Research, 2014, 122(2): 121-158.

[11] MATHOBO R, MARAISA D, STEYNA J M. The effect of drought stress on yield, leaf gaseous exchange and chlorophyll fluorescence of dry beans (Phaseolus vulgaris L.). Agricultural Water Management, 2017, 180: 118-125.

[12] LIAO J X, WANG G X. Effects of drought stress on leaf gas exchange and chlorophyll fluorescence of Glycyrrhiza uralensis. Russian Journal of Ecology, 2014, 45(6): 532-538.

[13] WU C W, LEE Y M, PENG C L. Chlorophyll fluorescence upper-to- lower-leaf ratio for determination of irrigation time for Pentas lanceolata. Photosynthetica, 2016, 54(2): 196-200.

[14] BRESTIC M, ZIVCAK M. PSII fluorescence techniques for measurement of drought and high temperature stress signal in crop plants: protocols and applications. Molecular Stress Physiology of Plants, 2013: 87-131.

[15] GUO S S, TIANS S, LIU W Q, WANG W Q, SUI N. Energy dissipation and antioxidant enzyme system protect photosystem II of sweet sorghum under drought stress. Photosynthetica, 2018, 56(3): 861-872.

[16] BEYEL V, BRÜGGEMANN W. Differential inhibition of photosynthesis during pre-flowering drought stress in Sorghum bicolor genotypes with different senescence traits. Physiologia Plantarum, 2010, 124(2): 249-259.

[17] 吴金芝, 王志敏, 李友军, 冯伟森, 田文忠, 高海涛. 不同冬小麦品种旗叶叶绿素荧光特性及其对干旱胁迫的响应. 麦类作物学报, 2015, 35(5): 699-706.

WU J Z, WANG Z M, LI Y J, FENG W S, TIAN W Z, GAO H T. Flag leaf chlorophyll fluorescence characteristics and its response to drought stress in different cultivars of winter wheat. Journal of triticeae crop, 2015, 35(5): 699-706. (in Chinese)

[18] 张仁和, 郑友军, 马国胜, 张兴华, 路海东, 史俊通, 薛吉全. 干旱胁迫对玉米苗期叶片光合作用和保护酶的影响[J]. 生态学报, 2011, 31(5): 1303-1311.

Zhang R H, Zheng Y J, Ma G S, Zhang X H, Lu H D, Shi J T, Xue J Q. Effects of drought stress on photosynthetic traits and protective enzyme activity in maize seeding. Acta Ecologica Sinica, 2011, 31(5): 1303-1311. (in Chinese)

[19] 何丽, 杜彦斌, 张金, 崔同霞. 干旱对胡麻现蕾期光合特性及产量的影响. 西北农林科技大学学报(自然科学版), 2017, 45(4): 59-64.

HE L, DU Y B, ZHANG J, CUI T X. Influence of drought stress on photosynthetic and fluorescence characteristics of benne at squaring stage. Journal of Northwest A＆F University(Natural Science Edition), 2017, 45(4): 59-64. (in Chinese)

[20] MENG L L, SONG J F, WEN J, ZHANG J, WEI J H. Effects of drought stress on fluorescence characteristics of photosystem II in leaves of Plectranthus scutellarioides. Photosynthetica, 2015, 54(3): 1-9.

[21] SHAH N H, PAULSEN G M. Interaction of drought and high temperature on photosynthesis and grain-filling of wheat. Plant and Soil, 2003, 257(1): 219-226.

[22] GUÓTH A, TARI I, GALLÉ A, CSISZÁR J, PÉCSVÁRADI A. Comparison of the drought stress responses of tolerant and sensitive wheat cultivars during grain filling: Changes in flag leaf photosynthetic activity, ABA levels, and grain yield. Plant Growth Regul, 2009, 28(2): 167-176.

[23] SÁNCHEZ-DÍAZ M, GARCÍA J L, ANTOLÍN M C, ARAUS J L. Effects of soil drought and atmospheric humidity on yield, gas exchange, and stable carbon isotope composition of barley. Photosythetica, 2002, 40(3): 415-421.

[24] THAMEUR A, LACHIHEB B, FERCHICHI A. Drought effect on growth, gas exchange and yield, in two strains of local barley Ardhaoui, under water deficit conditions in southern Tunisia. Journal of environmental management, 2012, 113(1): 495-500.

[25] 颉建明, 郁继华, 颉敏华. 青花菜叶片PSII光化学效率和光合对强光高温的响应. 兰州大学学报(自然科学版), 2008, 44(1): 51-55.

XIE J M, YU J H, XIE J H. Responses of PSII photochemical efficiencies and photosynthesis in Brassica oleracea L. var Italicap leaves under strong light and high temperature. Journal of Lanzhou university(natural sciences), 2008, 44(1): 52-55. (in Chinese)

[26] Deb S K, Shukla M K, Mexal J G. Estimating midday leaf and stem water potentials of mature pecan trees from soil water content and climatic parameters. Hortscience A Publication of the American Society for Horticultural Science, 2012, 47(7): 907-916.

[27] 徐林娟, 徐正浩, 李舸, 何勇, 谢法权, 孙映宏, 袁侠凡. 不同土壤水分供给下水稻叶水势的变化规律. 核农学报, 2011, 25(3): 553-558.

XU L J, XU Z H, LI G, HE Y, XIE F Q, SUN Y H, YUAN X F. response of leaf water potential in rice to different soil water supply levels. Journal of Nuclear Agricultural Sciences, 2011, 25(3): 553-558. (in Chinese)

[28] JALEELC A, MANIVANNAN P, WAHID A, FAROOQ M, SOMASUNDARAM R, PANNEERSELVAM R. Drought stress in plants: a review on morphological characteristics and pigments composition. International journal of agriculture and Biology, 2009, 11(1): 100-105.

[29] GALLÉ A, FELLER U. Changes of photosynthetic traits in beech saplings (Fagussylvatica) under severe drought stress and during recovery. Physiologia Plantarum, 2007, 131(3): 412-421.

[30] LANG Y, WANG M, XIA J, ZHAO Q. Effects of soil drought stress on photosynthetic gas exchange traits and chlorophyll fluorescence in Forsythia suspensa. Journal of forestry research, 2017(4): 1-9.

[31] SHANMUGAM S, KJAER K H, OTTOSEN C O, ROSENQVIST E, SHARMA D K, WOLLENWEBER B. The alleviating effect of elevated CO₂on heat stress susceptibility of two wheat (Triticum aestivum L.) cultivars. Journal of Agronomy and Science, 2013, 1995(5): 340-350.

[32] LAMBERT J M. Expression of osmotic adjustment and dehydration tolerance in diverse rice lines. Field Crops Research, 1996, 48(2): 185-197.

[33] COLOM M R,VAZZANA C. Photosynthesis and PSII functionality of drought-resistant and drought-sensitive weeping lovegrass plants. Environmental and Experimental Botany, 2003, 49(2): 135-144.

[34] MCDOWELLN G. Mechanisms linking drought, hydraulics, carbon metabolism, and vegetation mortality. Plant Physiology, 2011, 155(3): 1051-1059.

[35] WOO N S, BADGER M R, POGSON B J. A rapid, non-invasive procedure for quantitative assessment of drought survival using chlorophyll fluorescence. Plant Methods, 2008, 4(1): 27.

[36] SPERDOULI I, MOUSTAKAS M. Spatio-temporal heterogeneity in Arabidopsis thaliana leaves under drought stress. Plant Biology, 2012, 14(1): 118-128.

[37] EHLERT B, HINCHA D K. Chlorophyll fluorescence imaging accurately quantifies freezing damage and cold acclimation responses in Arabidopsis leaves. Plant Methods, 2008, 4(1): 4-12.

[38] BRESSON J, VASSEUR F, DAUZAT M, LABADIE M, VAROQUAUX F, TOURAINE B, VILE D. Interact to survive: Phyllobacterium brassicacearum improves Arabidopsis tolerance to severe water deficit and growth recovery. Plos one, 2014, 9(9): e107607.

[39] MISHRA R K, SINGHAL G S. Function of photosynthetic apparatus of intact wheat leaves under high light and heat-stress and its relationship with peroxidation of thylakoid lipids. Plant Physiology, 1992, 98(1): 1-6.

[40] HURA T, GRZESIAK S, HURA K, THIEMT E, TOKARZ K, Wedzony M. Physiological and biochemical tools in drought-tolerance detection in genotypes of winter triticale: accumulation of ferulic acid correlates with drought tolerance. Annals of Botany, 2007, 100(4): 767-775.

[41] TARI I, CAMEN D, CORADINI G, CSISZA´R J, FEDIUC E, GE´MES K, LAZAR A, MADOSA E, MIHACEA S, POOR P, POSTELNICU S, STAICU M, SZEPESI A, NEDELEA G, ERDEI L. Changes in chlorophyll fluorescence parameters and oxidative stress responses of bush bean genotypes for selecting contrasting acclimation strategies under water stress. Acta Biologica Hungarica, 2008, 59(3): 335-345.

[42] WARE M A, BELGIO E, RUBAN A V. Photoprotective capacity of nonphotochemical quenching in plants acclimated to different light intensities. Photosynthesis Research, 2015, 126(2/3): 261-274.