Microsoft Word 胡阳阳_new_.doc

Size: px

Start display at page:

Download "Microsoft Word 胡阳阳_new_.doc"

夜识唐
5 years ago
Views:

1 作物学报 ACTA AGRONOMICA SINICA 208, 44(4): ISSN ; CN -809/S; CODEN TSHPA9 DOI: /SP.J 灌浆期高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性及淀粉积累的影响胡阳阳褚莹莹,** 卢红芳王晨阳,2,*,** 刘卫星康娟马耕李莎莎, ; 2, :, 366,, (HT) (DS) (HT+DS) 0 d, ( 25 / 5 ) ( 32 / 22 ), ( 75%) ( 50%),, HT DS HT+DS (SSS)(GBSS), ; (SBE) ADPG (AGPase) (SS) ; HT DS HT+DS,,,,, HT DS,, SSS GBSS (P<0.0), AGPase SBE SS ( 22~26 d) (P<0.0), : ; ; ; ; ; Effects of High Temperature and Water Deficiency during Grain Filling on Activities of Key Starch Synthesis Enzymes and Starch Accumulation in Wheat HU Yang-Yang,**, LU Hong-Fang,**, LIU Wei-Xing, KANG Juan, MA Geng, LI Sha-Sha, CHU Ying-Ying, and WANG Chen-Yang,2,* Henan Agricultural University, Zhengzhou , Henan, China; 2 National Wheat Engineering Research Center, Zhengzhou , Henan, China Abstract: High temperature and drought, the major abiotic stresses during grain filling of winter wheat, have great influence on photosynthate accumulation in grains. In this study, we evaluated the effects of high temperature, drought and the compound stress at grain-filling stage on the activities of key starch synthesis enzymes and the contents of starch and its components in grains of winter wheat cultivar Zhengmai 366. The pot-grown plants were moved from field to the climate chamber at 0 days after anthesis and exposed to the stresses until maturity. The climate chamber was desired with the normal growth condition (25 C/5 C, day/night) and high temperature (HT) stress (32 C/22 C, day/night) treatments. Under each temperature treatment, the soil (207YFD ), (205BAD26B0) (S200-0-G07) This study was supported by the National Key Research and Development Program of China (207YFD ), the National Key Technologies R&D Program of China (205BAD26B0), and the earmarked fund for Modern Wheat Industrial Technology System of Henan Province (S200-0-G07). * (Corresponding author):, xmzxwang@63.com ** (Contributed equally to this work) : @63.com Received(): ; Accepted(): ; Published online(): URL:

2 moisture was adjusted to ~75% (normal) and ~50% (drought stress, DS), respectively. The normal temperature and soil moisture condition was taken as the control. Compared with the control, HT, DS, and HT+DS significantly increased activities of soluble starch synthase (SSS) and granule bound starch synthase (GBSS) at early period of stress and rapidly declined them subsequently. However, all stress treatments reduced the activities of starch branching enzymes (SBE), ADPG pyrophosphorylase (AGPase) and sucrose synthase (SS) during the whole grain-filling stage. The interaction between HT and DS was significant on all enzyme activities. Under HT, DS and HT+DS conditions, starch accumulation rate of grain, and the contents of amylose, amylopectin and total starch decreased and growth duration shortened, leading to reduced grain weight and yield at maturity. HT showed greater influence than DS, and the effect of compound stress was larger than that of each individual stress. Correlation analysis revealed that the activities of SSS and GBSS were positively correlated with the contents of amylose, amylopectin and total starch at most stages (P < 0.0), whereas, those of SBE, AGPase and SS at late grain-filling stage (22 26 days after anthesis) (P < 0.0). These results indicate that high temperature and drought stresses cause yield loss and degrade grain quality by affecting activities of key starch synthesis enzymes and starch accumulation in wheat grains. Keywords: winter wheat; compound stress; high temperature; drought; starch accumulation; starch synthesis enzymes,, (SS) ADPG (AGPase) (SSS)(GBSS) (SBE) [-2],, SBE SSS GBSS AGPase [3-4],,, [5-6],, 0%~30% [7], GBSS SSS SBE AGPase, ;, [8] SSS SBE AGPase,, [9-0] (35 ),,,,, [-4],, 35~45 d,, (2~4 d),,, /,, /, 材料与方法 , 366, 6%, 33%(pH 8.0), 8 g kg g kg 58 mg kg 68 mg kg 205 mg kg, 26% 27 cm, 24 cm, 0 kg;, ,.5 g P 2 O 5.35 g K 2 O.5 g;, 2 ;,.5 g, 0,, 8:00 8:00, 500 μmol m 2 s,,, 50%±0% (25 /5 ) (32 /22 ) /, ( 75%) ( 50% ), 4, (CK) (DS) (HT) (HT+DS), 5 7 d

3 4 : 593, TDR300,, 4 d, 5 ( ), min, 70, ( ), 80,.2 (, SS-2-Y GBSS-2A-Y SSS-S-Y AGP-2A-Y SBE-2-Y) SS GBSS SSS AGPase SBE, GBSS SSS AGPase NADPH (nmol), SS (μg), SBE 660 nm, %.3 [5], Cn 4 Cn 4 [] Rn 8, n, R n n, C n+4 C n 4 n+4 n 4.4 SPSS 2.0, Microsoft Excel 高温与干旱胁迫下 SSS 活性变化 SSS ( -B) CK DS HT HT+DS SSS 26 d 22 d 8 d 4 d, HT DS HT+DS SSS CK, 2 结果与分析高温与干旱胁迫下 GBSS 活性变化 GBSS, CK DS GBSS 26 d, HT HT+DS 8 d 4 d ( -A),, ( 4 d) GBSS CK, 8 d, HT HT+DS GBSS, CK, DS 30 d CK (P<0.05) 图灌浆期不同胁迫处理小麦籽粒淀粉合成关键酶活性的变化 Fig. Dynamics of starch synthesis key enzyme activities during grain filling under different stress treatments HT: ; DS: ; HT+DS: DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS.

4 ~22 d SSS, (P<0.05) 2..3 高温与干旱胁迫下 SBE 活性变化 4 d ( 4 d), HT SBE CK,, CK 23.3% ( ) 3.2% ( ); DS HT+DS SBE, 4 d, % 46.8%, % 4.2% ( -C) 2..4 高温与干旱胁迫下 AGPase 活性变化 AGPase GBSS, CK DS AGPase 22 d, HT HT+DS 8 d 4 d 8 d, HT AGPase, AGPase, HT+DS, HT, DS ( -D) AGPase, 2..5 高温与干旱胁迫下 SS 活性变化 SS, DS SS 4 d, 8 d, ( -E) HT HT+DS SS, CK 23.8% 32.%, % 29.5%; DS SS 0~8 d CK, CK 9.% ( ) 25.6% ( ) HT SS DS, HT+DS, SS 2.2 HT 4 d SBE AGPase, ; DS 4 d GBSS, ; ( ) 高温与干旱胁迫对小麦籽粒淀粉含量的影响 ( 2), HT DS HT+DS CK, DS HT HT+DS 5.9% 0.2% 9.7% ( ) 5.5% 9.5% 9.2% ( ), 7.2%.2% 8.2% ( Enzyme 表高温与干旱胁迫对小麦籽粒淀粉合成关键酶活性影响的互作效应方差分析 (F 值 ) Table Variance analysis on interaction of HT and DS on activity of key starch synthesis enzymes (F-value) Source 4DAA 8DAA 22DAA 26DAA 4DAA 8DAA 22DAA 26DAA GBSS SSS SBE AGPase SS HT 22.8 ** 2.7 ** ** ** 2.6 ** 9.8 ** ** ** DS ** 39.3 ** ** 78.8 ** 40.5 ** DS HT 8.3 * 30.4 ** 99.3 ** 34. **.4 ** 63. ** 58. ** 47.6 ** HT 39. ** 23.3 ** 26.0 ** 344. ** 3.7 ** 50.0 ** ** ** DS 9. * 9.6 ** 42.9 ** 49.2 ** 0.0 * 72.5 ** ** DS HT ** 50.4 ** 0.8 * ** 42.7 **.6 ** HT ** 53.3 ** 6.6 ** ** 74.8 ** 62.4 ** DS 46.5 ** 4.3 ** ** 84.4 ** 7.9 ** 27.6 ** 86.2 ** ** DS HT 20.9 ** 3.2 ** 99.8 ** ** 7.6 *.5 ** 9.2 * HT * ** ** * ** 390. ** DS 6.0 * 98.0 ** 26.2 ** ** ** 94.6 ** ** DS HT 5.0 ** 20.7 ** ** 7. * 20.7 ** ** HT 74.0 ** 40. ** ** ** 54.8 ** 52.9 ** 66.0 ** ** DS 3.3 ** ** 5.0 ** 23.6 ** 9.6 * 34.4 ** 46.4 ** DS HT 43.5 ** 5.6 * ** 99.2 ** 9.0 *..5 ** * ** ()DAA: ; DS HT: * Significant difference at the 0.05 probability level; ** significant difference at the 0.0 probability level. DAA: days after anthesis; DS: drought stress; HT: high temperature stress; DS HT: interaction of HT and DS.

5 4 : 595 ) 6.7%.2% 8.3% ( ), 6.9% 0.99% 8.6% ( ) 6.4% 0.8% 8.5% ( ), HT DS, HT+DS,, DS HT HT+DS CK 4 d 8 d 2 d 图 3 不同处理小麦籽粒淀粉积累速率的变化 Fig. 3 Changes of starch accumulating rate in wheat grains under different treatments HT: ; DS: ; HT+DS: DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS. 图 2 不同处理籽粒直链淀粉 (A) 支链淀粉(B) 和总淀粉含量 (C) 的变化 Fig. 2 Changes of amylose (A) amylopection (B), and total starch (C) contents under different treatments HT: ; DS: ; HT+DS: DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS 高温与干旱胁迫对小麦籽粒淀粉积累速率的影响, HT HT+DS 4 d, CK DS 8 d ( 3) HT DS HT+DS, CK 6.0%.4% 2.8% ( ) 8.4%.9% 26.4% ( ) 2.4 HT DS (HT+DS) ( 2) CK, HT DS HT+DS 0.6% 4.% 4.5% ( ) 0.2% 4.2% 4.9% ( ), 3.8% 6.9% 9.7% ( ) 3.4% 6.0% 8.7% ( ), 3.0% 6.2% 8.4% ( ) 2.7% 5.6% 7.8% ( ) HT DS HT+DS / CK 3.7% 3.5% 6.4% ( ) 3.7%.9% 4.6% ( ), 2.5 HT DS 0 d,,,, ( 3) HT+DS 33.4%~36.6% 33.5%~38.%; HT 7.6%~22.5% 20.5%~2.5%; DS, 3.7%~4.5%, 8.%~8.3% 2.6, GBSS SSS ( 22 d ); ( 22~26 d), SBE AGPase ; SS

6 表 2 不同处理成熟期小麦籽粒淀粉及其组分含量 Table 2 Starch content and its components in wheat grain under different treatments at maturity Treatment Amylose Amylopectin Total starch / Amylose/ amylopectin Amylose Amylopectin Total starch / Amylose/ amylopectin CK 5.9 a 50.8 a 66.7 a 0.3 c 5.6 a 50.0 a 65.6 a 0.3 c DS 5.3 b 47. b 62.4 b 0.32 b 5.0 b 46.9 b 6.9 b 0.32 bc HT 4.2 c 43.8 c 58. c 0.33 b 4.0 c 43.2 c 57.3 c 0.32 ab HT+DS 3.6 d 40.8 d 54.4 d 0.33 a 3.3 d 40.6 d 53.9 d 0.33 a (P<0.05) HT+DS: Values within a column followed by different letters are significantly different among treatments at P < DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS. 表 3 不同处理对成熟期小麦籽粒产量及其构成因素的影响 Table 3 Effects of different treatments on grain yield and its components of wheat Treatment SN (pot ) GNP TGW (g) GY (g pot ) SN (pot ) GNP TGW (g) GY (g pot ) CK 35.7 a 45.5 a 48. a 72.6 a 43.0 a 46.0 a 42.9 a 67.7 a DS 37.0 a 43.3 a 4.5 b 59.3 b 42.3 a 46.6 a 36.7 b 55.4 b HT 38.0 a 45.8 a 37.3 c 56.9 b 39.3 a 43.5 a 35.4 c 53.8 b HT+DS 38.0 a 46.5 a 30.5 d 48.2 c 40.7 a 46. a 28.6 d 4.9 c (P<0.05) SN: ; GNS: ; TGW: ; GY: ; HT+DS: Values within a column followed by different letters are significantly different among treatments at P < SN: spike number; GNS: grain number per spike; TGW: thousand-grain weight; GY: grain yield; DS: drought stress; HT: high temperature stress; HT+DS: compound stress of HT and DS. ( 4 d), ( 22~26 d) ( 4), GBSS SSS SBE AGPase SS, 3 讨论 3., [6] AGPase SSS GBSS SBE SS [7-8],, [8,9-22], GBSS SSS SBE AGPase SS,, [23-25] [26] (35±3, 3 d), AGPase SSS SBE 7%~3% [9] ( 2~5 ), SS AGPase SSS 7.7% 0.0% 2.5%, 4.% 27.7% 27.9%,, SS AGPase SSS SBE.% 5.6% 3.6% 3.5%, GBSS SSS,,,,, SSS SBE AGPase SS, GBSS AGPase SS [2,0,27-28], SSS GBSS, SS AGPase SBE (22~26 d),

7 4 : 597 表 4 小麦籽粒淀粉合成关键酶活性与淀粉积累的相关系数 Table 4 Correlation coefficients of activities of key starch synthesis enzymes with contents of starch and its components Days after anthesis (DAA) Key starch synthesis 4 d 8 d 22 d 26 d Amylose enzymes 4 DAA 8 DAA 22 DAA 26 DAA GBSS ** ** ** Amylopectin content SSS ** * ** SBE ** 0.94 ** AGP ** ** SS ** ** ** GBSS 0.88 ** ** Amylose content SSS ** ** ** SBE ** AGP ** SS * ** GBSS ** ** ** Total starch content SSS ** 0.88 ** ** SBE ** ** AGP ** ** SS 0.78 ** ** ** GBSS 0.7 ** ** ** Amylopectin content SSS ** * * ** SBE ** ** AGP ** ** SS * ** ** GBSS ** ** ** Amylose content SSS ** ** ** SBE ** AGP ** SS ** GBSS ** ** ** Total starch content SSS ** 0.76 ** * ** SBE ** ** AGP ** 0.95 ** SS * ** ** * ** * and ** indicate significant correlation at the 0.05 and 0.0 probability levels, respectively. 3.2, [29-3] [32-34], 30, [22,35-37] [38], (38, 2~4 d) 3~5 d, d, ( ), , ,

8 598 44,,,,,,,,,,, [39] [33], [2] (38 ) (55%), 34.4% 7.6% 43.2%,, 0.9% 6.7% 8.6%, 20.2% 4.% 35.%, 20.% 8.2% 35.8%,, (), 4 结论,,,,, GBSS SSS SBE AGPase SS, References [],,,.., 200, 27: Li Y G, Yu Z W, Jiang D, Yu S L. Studies on the dynamic changes of the synthesis of sucrose in the flag leaf and starch in the grain and related enzymes of high-yielding wheat. Acta Agron Sin, 200, 27: (in Chinese with English [2],,,,,,.., 2007, 33: Yan S H, Wang Z L, Dai Z M, Li W Y, Fu G Z, He M R, Yin Y P. Activity of enzymes involved in starch synthesis and accumulation in grains of two wheat cultivars with a different amylose content. Acta Agron Sin, 2007, 33: (in Chinese with English [3] Wang Z B, Li W H, Qi J C, Shi P C, Yin Y A. Starch accumulation, activities of key enzyme and gene expression in starch synthesis of wheat endosperm with different starch contents. J Food Sci Technol, 204, 5: [4],,,,,. 9., 2005, 38: Li C Y, Feng C N, Zhang Y, Guo W S, Zhu X K, Peng Y X. Effects of the ratio between basal N and top dressing N on grain starch formation in weak gluten wheat variety Ningmai 9 and its enzymes activities. Sci Agric Sin, 2005, 38: (in Chinese with English [5],,,,,,.., 2004, 30: 88 9 Wang C Y, Guo T C, Yan Y L, Zhu Y J, Xia G J, Wang H C, Zhou J Z. Effects of short post-anthesis high temperature stress on leaf photosynthetic potential in winter wheat (Triticum aestivum L.). Acta Agron Sin, 2004, 30: 88 9 (in Chinese with English [6],,,,,,.., 2008, 27: Zhang H H, He M R, Liu Y H, Li F, Wu C P, Zhang B, Wang X Y. Effect of high temperature stress at late grain-filling stage and nitrogen and sulphur fertilization on wheat grain yield and quality. Chin J Ecol, 2008, 27: (in Chinese with English [7],,,,,,.., 2008, 28: Liu Z G, Sun J S, Zhang J Y, Wang J L, Meng Z J, Li X D, Liu X F. Effect of drought at different growing stages on yield and quality characteristics of strong-gluten wheat. J Trit Crops, 2008, 28: (in Chinese with English [8],,,.., 205, 35: Yang Y, Li Y, Kang J H, Liu P. Effect of heat stress after anthesis on starch synthesis in spring wheat. J Trit Crops, 205, 35: (in Chinese with English [9],,,,.., 206, 33: Li Y T, Kang J H, Wu H L, Li Y, Yao S. Effect of water stress after anthesis on starch formation and the activity of associated enzymes in spring wheat. Pratacult Sci, 206, 33: (in Chinese with English [0],,,,,.., 205, 33: 04 0 Zhao J J, Yao S, Li Y, Kang J H, Wu H L, Xu Q. Effects of drought on the starch contents and activities of key enzymes after anthesis in spring wheat grains. Agric Res Arid Areas, 205, 33: 04 0 (in Chinese with English [],,,,,..

9 4 : 599, 2008, 28: Miao J L, Wang C Y, Guo T C, Ma D Y, Hu J B, Feng H. Effects of post-anthesis interactions of high temperature and drought stress on content and composition of grain starch in two wheat cultivars with different gluten strength. J Trit Crops, 2008, 28: (in Chinese with English [2],,,,,,.., 206, 36: Yue P L, Wang C Y, Lu H F, Liu W X, Ma G, Wang Q, Hu Y Y. Effect of heat and drought stress on starch accumulation during grain filling stage. J Trit Crops, 206, 36: (in Chinese with English [3],,,.., 204, 34: Lu H F, Wang C Y, Guo T C, Yin Y X. Effects of hightemperature and drought stress on protein concentration and keys enzymes activities in relation to nitrogen metabolism in wheat grains during the early stage of grain filling. Acta Ecol Sin, 204, 34: (in Chinese with English [4],,,,,,.., 204, 34: Wang C Y, Miao J L, Zhang M W, Ma D Y, Feng W, Xie Y X, Guo T C. Effects of post-anthesis high temperature, drought stress and their interaction on the starch pasting properties of two wheat cultivars with different gluten strength. Acta Ecol Sin, 204, 34: (in Chinese with English [5].. :, 985. pp He Z F. Analysis Technique for Grain Quality in Cereals and Oils. Beijing: China Agriculture Press, 985. pp (in Chinese) [6] Rahman S, Regina A, Li Z Y, Mukai Y, Yamamoto M, Kosar- Hashemi B, Abrahams S, Morell M K. Comparison of starch branching enzyme genes reveals evolutionary relationships among isoform. Characterization of a gene for starch-branching enzymes IIa from the wheat D genome donor Aegilops tauschii. Plant Physiol, 200, 25: [7] Nakamura Y, Yuki K, Park S Y, Ohya T. Carbohydrate metabolism in the developing endosperm of rice grains. Plant Cell Physiol, 989, 30: [8],,,,. /., 200, 30: Cao Y N, Hu W G, Wang G P, Liu L X, Wang C S. Dynamic changes of starch accumulation and enzymes relating to starch biosynthesis of kernel during grain filling in waxy and non-waxy winter wheat. Acta Bot Boreali-Occid Sin, 200, 30: (in Chinese with English [9],,.., 20, 35: Shi H Q, Gong Y H, Zhang D W. Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis. J Plant Ecol, 20, 35: (in Chinese with English [20] Dai Z M, Yin Y P, Wang Z L. Comparison of starch accumulation and enzyme activity in grains of wheat cultivars differing in kernel type. Plant Growth Regul, 2009, 57: [2] Ahmadi A, Baker D A. The effect of water stress on the activities of key regulatory enzymes of the sucrose to starch pathway in wheat. Plant Growth Regul, 200, 35: 8 9 [22],,,,.., 202, 28(2): Bao G, Tan Z H, Zhou Y, Bao Y H, Sa C L. Advances of evaluation of climate changes impact on crop yield. Chin Agric Sci Bull, 202, 28(2): (in Chinese with English [23],,,,,,.., 2006, 32: Zhao H, Dai T B, Jing Q, Jiang D, Cao W X, Lu W, Tian X W. Effects of high temperature during grain filling on key enzymes involved in starch synthesis in two wheat cultivars with different quality types. Acta Agron Sin, 2006, 32: (in Chinese with English [24],,,,,,,,,.., 2008, 34: Yan S H, Yin Y P, Li W Y, Liang T B, Li Y, Wu Y H, Wang P, Geng Q H, Dai Z M, Wang Z L. Effect of high temperature during grain filling on starch accumulation, starch granule distribution, and activities of related enzymes in wheat grains. Acta Agron Sin, 2008, 34: (in Chinese with English [25],,,,.., 20, 29: Wang Z H, Guo X P, Lyu S F, Li Y J, Deng X P. Effects of soil drought on starch accumulation of grain and activity of corresponding starch synthesis enzymes in different winter wheat varieties. Agric Res Arid Areas, 20, 29: (in Chinese with English [26],,,,,.., 205, 36: Sun L Y, Su W, Wu H L, Li Y, Wang X, Kang J H. Effects of the high temperature after the flowering period on starch formation in spring wheat. J Agric Sci, 205, 36: (in Chinese with English [27],,,.., 2003, 29: 75 8 Wang Y F, Yu Z W, Li S X, Yu S L. Activity of enzymes related to starch synthesis and their effect during the filling of winter wheat. Acta Agron Sin, 2003, 29: 75 8 (in Chinese with English [28],,,,,,,.., 2008, 34: Li J M, Wang Z L, Gao R Q, Li S F, Cai R G, Yan S H, Yu A L, Yin Y P. Activities of enzymes involved in sucrose and starch synthesis during grain filling and the relation to nitrogen metabo-

10 lism in strong-and weak-gluten wheat cultivars. Acta Agron Sin, 2008, 34: (in Chinese with English [29] Panozzo J F, Eagles H A. Cultivar and environmental effects on quality characters in wheat: I. Starch. Aust J Agric Res, 998, 49: [30] Morris C F, Shackley B J, King G E, Kidwell K K. Genotypicand environmental variation for flour swelling volume in wheat. Cereal Chem, 997, 74: 6 2 [3] Jenner C F. Starch synthesis in the kernel of wheat under high temperature conditions. Aust J Plant Physiol, 994, 2: [32],,.., 2003, 29: Xu Z Z, Yu Z W, Zhang Y L. The effects of soil moisture on grain starch synthesis and accumulation of winter wheat. Acta Agron Sin, 2003, 29: (in Chinese with English [33],,,,.., 205, 35: Yang X J, Ju H, Wang Z S, Guo A T, Yang Y M. Effect of high temperature and drought after anthesis on photosynthesis, antioxidant properties and grain weight in winter wheat. J Trit Crops, 205, 35: (in Chinese with English [34],,,,.., 205, 36: Su W, Sun L Y, Kang J H, Wu H L, Li Y. Effect of high temperature and drought treatment on photosynthetic characteristics and yield of different varieties of spring wheat. J Agric Sci, 205, 36: (in Chinese with English [35] Jenner C F. Effects of exposure of wheat ears to high temperature on dry matter accumulation and carbohydrate metabolism in the grain of two cultivars: II. Carry-over effects. Aust J Plant Physiol, 99, 8: [36] Rane J, Nagarajan S. High temperature index-for field evaluation of heat tolerance in wheat varieties. Agric Systems, 2004, 79: [37] Shah N H, Paulsen G M. Interaction of drought and high temperature on photosynthesis and grain-filling of wheat. Plant Soil, 2003, 257: [38],,,,,.., 2008, 42: Hu J B, Wang C Y, Guo T C, Miao J L, Zhu Y J, He D X. Effects of high temperature and drought stress on grain filling characteristics in wheat during grain filling period. J Henan Agric Univ, 2008, 42: (in Chinese with English [39],,,,,,,.., 205, 35: Li C Y, Fu K Y, Zhang R Q, Xu F F, Shi X Y, Zhu Y Q, Qin A X, Li C. Effect of high temperature post anthesis on the development of starch granules in winter wheat. J Trit Crops, 205, 35: (in Chinese with English

Microsoft Word - A200812-1992.doc

Microsoft Word - A200812-1992.doc 1 长期遮荫对小麦籽粒淀粉理化特性的影响张传辉, 姜东, 戴廷波, 荆奇, 曹卫星南京农业大学, 农业部作物生长调控重点开放实验室, 南京 (2195) E-mail: jiangd@njau.edu.cn 摘要 : 揭示了长期遮荫对小麦籽粒淀粉理化特性的影响选