干旱胁迫下持绿小麦碳氮同化物转运与分配的研究
摘要
为了阐明持绿小麦碳氮同化物的转运与分配规律及干旱对其影响,本文以持绿型小麦豫麦66(YM66),早衰型小麦温麦6号(WM6),对照品种小偃22(XY22)作为试验材料进行盆栽试验。以称重控水法进行干旱处理,利用~(14)C放射性同位素示踪技术和~(15)N非放射性同位素技术进行花前和花后标记,研究不同持绿型的小麦,在灌浆期碳氮同化物的转运分配规律以及干旱对其影响。以阐明持绿型小麦同化物的生产、转运特性,为挖掘持绿型小麦的潜能提供理论依据。所得主要结果如下:
1.花前~(14)C标记表明,小麦花前标记的~(14)C-贮备物在开花时80%-90%已储存于茎鞘和颖壳中,仅有约10%-20%残留在叶片中;开花后这些贮备物都向籽粒转运,成熟时籽粒中~(14)C分配率约为20-40%,茎杆与颖壳中含有约60%-80%,极少量残留在叶片中。持绿型小麦YM66的~(14)C-贮备物从叶片和茎杆中输出得快,成熟时籽粒含有40%的~(14)C-贮备物,高于对照品种XY22和早衰品种WM6。对于YM66和XY2,干旱处理促进了~(14)C-贮备物向籽粒中的转运。
2.小麦花后~(14)C标记表明,在标记后12h,~(14)C-同化物有40%-60%储存在茎鞘和颖壳中,20%-50%储存在籽粒中,仅有15%-25%残留在叶片中,不同持绿性小麦之间~(14)C分配率差异较大;此后,叶片和茎鞘中的~(14)C向籽粒转运,成熟时,籽粒中~(14)C同化物的分配率为40%-80%,YM66和XY22籽粒中的~(14)C-同化物高达70%-80%,而WM6却只有40%-50%,对于YM66和XY22。干旱处理下,籽粒中~(14)C-同化物的分配率要高于正常水分处理的,而WM6则相反。
3.花前~(15)N标记表明,小麦花前标记的~(15)N-贮备物在开花时45%-60%储存于茎鞘和颖壳中,约有40%-55%仍留在叶片中;开花后茎鞘,颖壳和叶片中的~(15)N都向籽粒转运,成熟时籽粒中~(15)N分配率约为45-65%,茎鞘与颖壳中含有约15%-25%,有20%-40%残留在叶片中。成熟时YM66籽粒中~(15)N分配率最低,XY22最高。干旱胁迫使~(15)N-贮备物在籽粒中的分配率升高,说明干旱促进了花前N的转运。
4.花后~(15)N标记表明,在标记后12h, ~(15)N-同化物有50%-70%储存在茎鞘和颖壳中,有30%-50%留在叶片中;在12h-24h,叶片中~(15)N分配率升高,此后,叶片和茎鞘中~(15)N分配率持续下降;成熟时,籽粒中~(15)N同化物的分配率为50%-65%,茎鞘中为15%-25%,叶片残留30%-35%。干旱胁迫使YM66和XY22 ~(15)N-同化物在籽粒中的分配率有所增加。
5.开花后各个小麦品种叶片和茎鞘全氮含量呈下降趋势,籽粒全氮含量则先升后降。茎鞘中,持绿型品种YM66的全氮含量高于对照,WM6的最低,在叶片中则WM6和YM66的全氮含量较高,XY22的最低。籽粒中XY22的最高,YM66的最低。对于XY22和YM66,干旱降低了叶片和茎鞘中全氮含量,而增加了籽粒中的全氮含量。成熟时持绿小麦YM66总生物量和籽粒产量均高于非持绿品种,其中总生物量比对照品种提高了约10%。
Transportation and distribution of carbon and nitrogen assimilates in wheat under drought stress was studied by ~(14)C-labeling and ~(15)N-labeling.The materials are YM66 (stay-green), WM6(senescent) and XY22(control).The experiment was conducted by labelling leaf with ~(14)CO_2 before and after anthesis to elucidate the effect of drought on different kinds of stay-green wheat by control water.Also the another study was conducted with ~(15)N labeling leaf before and after anthesis to show transportation and distribution of nitrogen assimilates under drought stress.Our purpose was to elucidate the traits of production and distribution of carbon and nitrogen assimilates.The results are as follows:
1.The experiment of ~(14)C-labeling before anthesis showed that about 80%-90% of ~(14)C-reserves had been stored in the stems and rachises, and about 10%-20% remained in the leaves at anthesis;After anthesis, these reserves were remobilized and transported to the kernels. At maturity, ~(14)C-reserves in the leaves were exported almost entirely, but about 60%-80% ~(14)C-reserves remained in the stems and rachises, and about 20%-40% was imported into the kernels. The ~(14)C-reserves in stay-green wheat were transported more quickly from the leaves and stems, and at maturity ,the stay-green wheat YM66 has imported 40% of ~(14)C-reserves into the kernels, much more than XY22 and WM6.Drought stress promoted translocation and distribution of ~(14)C-reserves into the kernels of YM66 and XY22.
2.The experiment of ~(14)C-labeling after anthesis showed that about 40%-60% of ~(14)C-assimilates was stored in the stem and sheath after 24 hours, 20%-50% was transported into the kernels, and about 15%-25% remained in leaves.Then,these assimilates were remobilized and transported to the kernels.At maturity,about 40%-80%~(14)C was found in the kernels.It is up to70%-80% in YM66 and XY22 ,but 40%-50% in the WM6.For YM66 and XY22,the distribution of kernels under drought was higher than normal,but WM6 is opposite.
3.The experiment of ~(15)N-labeling before anthesis showed that about 45%-60% of ~(15)N-reserves assimilated before anthesis had been stored in the stems and rachises, and about 40%-55% remained in the leaves at anthesis;After anthesis, these reserves were remobilized and transported to the kernels, at maturity, about 15%-25% ~(15)N reserves remained in the stems and rachises, and about 45%-65% was imported into the kernels,20%-40% remained in the leaves.The distribution of YM66 in kernels is lower than others,but XY22 is higher. The drought promote the distribution in kernels.
4.The experiment of ~(15)N-labeling after anthesis showed that about 50%-70% of ~(15)N -assimilates was stored in the stem and sheath,about 30%-50% remained in the leaves.After 24 hours,distribution of leaves increasd first ,then got lower,and distribution of the stem and sheath always decreased.At maturity,about 50%-65%~(15)N was found in the kernels, 15%-25% in the stem and sheath and30%-35% in the leaves. The drought promote the distribution of YM66 and.XY22 in kernels.
5.After anthesis, The nitrogen content of leaves and stems has decreased,and the stay-green wheat YM66 had higher nitrogen level than XY22,and WM6 has lowest nitrogen content. The nitrogen content of kernels first increased and then decreased.In YM66 and XY22,the drought promote the nitrogen content in kernels and reduced the nitrogen content in leaves, stems and rachises. At maturity, Total Biomass and Seed Yield of YM66 is higher than XY22 and WM6.
1.花前~(14)C标记表明,小麦花前标记的~(14)C-贮备物在开花时80%-90%已储存于茎鞘和颖壳中,仅有约10%-20%残留在叶片中;开花后这些贮备物都向籽粒转运,成熟时籽粒中~(14)C分配率约为20-40%,茎杆与颖壳中含有约60%-80%,极少量残留在叶片中。持绿型小麦YM66的~(14)C-贮备物从叶片和茎杆中输出得快,成熟时籽粒含有40%的~(14)C-贮备物,高于对照品种XY22和早衰品种WM6。对于YM66和XY2,干旱处理促进了~(14)C-贮备物向籽粒中的转运。
2.小麦花后~(14)C标记表明,在标记后12h,~(14)C-同化物有40%-60%储存在茎鞘和颖壳中,20%-50%储存在籽粒中,仅有15%-25%残留在叶片中,不同持绿性小麦之间~(14)C分配率差异较大;此后,叶片和茎鞘中的~(14)C向籽粒转运,成熟时,籽粒中~(14)C同化物的分配率为40%-80%,YM66和XY22籽粒中的~(14)C-同化物高达70%-80%,而WM6却只有40%-50%,对于YM66和XY22。干旱处理下,籽粒中~(14)C-同化物的分配率要高于正常水分处理的,而WM6则相反。
3.花前~(15)N标记表明,小麦花前标记的~(15)N-贮备物在开花时45%-60%储存于茎鞘和颖壳中,约有40%-55%仍留在叶片中;开花后茎鞘,颖壳和叶片中的~(15)N都向籽粒转运,成熟时籽粒中~(15)N分配率约为45-65%,茎鞘与颖壳中含有约15%-25%,有20%-40%残留在叶片中。成熟时YM66籽粒中~(15)N分配率最低,XY22最高。干旱胁迫使~(15)N-贮备物在籽粒中的分配率升高,说明干旱促进了花前N的转运。
4.花后~(15)N标记表明,在标记后12h, ~(15)N-同化物有50%-70%储存在茎鞘和颖壳中,有30%-50%留在叶片中;在12h-24h,叶片中~(15)N分配率升高,此后,叶片和茎鞘中~(15)N分配率持续下降;成熟时,籽粒中~(15)N同化物的分配率为50%-65%,茎鞘中为15%-25%,叶片残留30%-35%。干旱胁迫使YM66和XY22 ~(15)N-同化物在籽粒中的分配率有所增加。
5.开花后各个小麦品种叶片和茎鞘全氮含量呈下降趋势,籽粒全氮含量则先升后降。茎鞘中,持绿型品种YM66的全氮含量高于对照,WM6的最低,在叶片中则WM6和YM66的全氮含量较高,XY22的最低。籽粒中XY22的最高,YM66的最低。对于XY22和YM66,干旱降低了叶片和茎鞘中全氮含量,而增加了籽粒中的全氮含量。成熟时持绿小麦YM66总生物量和籽粒产量均高于非持绿品种,其中总生物量比对照品种提高了约10%。
Transportation and distribution of carbon and nitrogen assimilates in wheat under drought stress was studied by ~(14)C-labeling and ~(15)N-labeling.The materials are YM66 (stay-green), WM6(senescent) and XY22(control).The experiment was conducted by labelling leaf with ~(14)CO_2 before and after anthesis to elucidate the effect of drought on different kinds of stay-green wheat by control water.Also the another study was conducted with ~(15)N labeling leaf before and after anthesis to show transportation and distribution of nitrogen assimilates under drought stress.Our purpose was to elucidate the traits of production and distribution of carbon and nitrogen assimilates.The results are as follows:
1.The experiment of ~(14)C-labeling before anthesis showed that about 80%-90% of ~(14)C-reserves had been stored in the stems and rachises, and about 10%-20% remained in the leaves at anthesis;After anthesis, these reserves were remobilized and transported to the kernels. At maturity, ~(14)C-reserves in the leaves were exported almost entirely, but about 60%-80% ~(14)C-reserves remained in the stems and rachises, and about 20%-40% was imported into the kernels. The ~(14)C-reserves in stay-green wheat were transported more quickly from the leaves and stems, and at maturity ,the stay-green wheat YM66 has imported 40% of ~(14)C-reserves into the kernels, much more than XY22 and WM6.Drought stress promoted translocation and distribution of ~(14)C-reserves into the kernels of YM66 and XY22.
2.The experiment of ~(14)C-labeling after anthesis showed that about 40%-60% of ~(14)C-assimilates was stored in the stem and sheath after 24 hours, 20%-50% was transported into the kernels, and about 15%-25% remained in leaves.Then,these assimilates were remobilized and transported to the kernels.At maturity,about 40%-80%~(14)C was found in the kernels.It is up to70%-80% in YM66 and XY22 ,but 40%-50% in the WM6.For YM66 and XY22,the distribution of kernels under drought was higher than normal,but WM6 is opposite.
3.The experiment of ~(15)N-labeling before anthesis showed that about 45%-60% of ~(15)N-reserves assimilated before anthesis had been stored in the stems and rachises, and about 40%-55% remained in the leaves at anthesis;After anthesis, these reserves were remobilized and transported to the kernels, at maturity, about 15%-25% ~(15)N reserves remained in the stems and rachises, and about 45%-65% was imported into the kernels,20%-40% remained in the leaves.The distribution of YM66 in kernels is lower than others,but XY22 is higher. The drought promote the distribution in kernels.
4.The experiment of ~(15)N-labeling after anthesis showed that about 50%-70% of ~(15)N -assimilates was stored in the stem and sheath,about 30%-50% remained in the leaves.After 24 hours,distribution of leaves increasd first ,then got lower,and distribution of the stem and sheath always decreased.At maturity,about 50%-65%~(15)N was found in the kernels, 15%-25% in the stem and sheath and30%-35% in the leaves. The drought promote the distribution of YM66 and.XY22 in kernels.
5.After anthesis, The nitrogen content of leaves and stems has decreased,and the stay-green wheat YM66 had higher nitrogen level than XY22,and WM6 has lowest nitrogen content. The nitrogen content of kernels first increased and then decreased.In YM66 and XY22,the drought promote the nitrogen content in kernels and reduced the nitrogen content in leaves, stems and rachises. At maturity, Total Biomass and Seed Yield of YM66 is higher than XY22 and WM6.
引文
[1] G.Spano,N.Di Fonzo and C.Perrotta.Physiological characterization of stay green mutants in durum wheat[J].Journal of experimental botany.2003,54(386):1415-1420
[2]胡廷积,杨永光,马元喜等.小麦生态与生产技术[M].河南:河南科学技术出版社,1986. 19-23
[3]董树亭,王空军,胡昌浩等.玉米品种更替过程中群体光合特性的演变[J].作物学报,2000,26 (2):200-204
[4]何萍,金继运.保绿型玉米的营养生理研究进展[J].玉米科学.2000, 8 (4): 41-44
[5]何萍,金继运.氮素营养对不同衰老类型玉米碳氮化合物解积的影响(收录入《青年学者论土壤与植物营养科学》),中国农业科技出版社,2000
[6] Xu, W. W., D.T. Rosenow, and H.T, Nguyen. Stay green trait in grain sorghum:Visual rating and objective measuring[J]. Plant Breeding,2000,119:365-367
[7]赵松岭.集水农业引论[M].西安:陕西科学技术出版社,1996.15-16.
[8] Howard Thomas, Catherine J. Howarth.Five ways to stay green[J]. Journal and experimental botany; 2002, 51(1):329-337
[9] Thomas H . Crops that stay green[J] Annals of applied biolpgy ,1993,123(1):193-219.
[10] Bekavac G.Path analysis of stay-green trait in maize[J].Cereal research communications 1998. 26(2): 1661-1671
[11] Guiamet JJ, Schwartz E, Pichersky E, Nooden LD (1991)Characterization of cytoplasmic and nuclear mutations affectingchlorophyll and chlorophyll-binding proteins during senescencein soybean. Plant Physiol 1996:227–231
[12] Gentinetta E, Ceppi D, Lepori C. A major gene for delayed senescence in maize. Pattern of photo- synthates accumulation and inheritance. Plant Breed1997:193–203
[13] Richard S .Inheritance of the Stay Green Trait in Sorghum[J].Crop Science,1994,34;970-972
[14] L M Dwyer and M Tollenaar.Genetic Improvement in Photosyn2thetic Response of Hybrid Maize Cultivars , 1959 to 1989[J] Plant Science ,1989 ,69 :81-91
[15] Guiamet JJ, Giannibelli MC . Nuclear and cytoplasmic“stay green”Mutations of soybean alter the loss of leaf soluble proteins during senescence. Physiol Plant 1996:665–661
[16] Rawson H M, Hindmarsh J H, Fischer R A, et al. Changes in leaf photosynthesis with plant ontogeny and relationships with yield per ear in whera cultivars and 120 progeny[J]. Plant Physiol, 1983, 10, 503- 514
[17] Mccaig T N, Clarke J M. Seasonal changes in nonstructural carbohydrate levels of wheat and oats grown in semiarid environment[J].Crop Science,1982,22:963-970
[18] Nooden L D, Gniamet J J, John I. Senescence mechanisms[J]. Physiol Plant, 1997,101:746-753
[19] Ntanos D A, Koutroubas S D. Dry matter anf N accumulation and translocation for Indica and Japonica rice Mediterranean conditions[J]. Field Crops,2002,74:93-101
[20] Palta J A, Kobata T, Turner N C,et al. Remobilization of carbon of carbon and nitrogen in wheat as influenced by postanthesis water deficits. Crop Science,1994,34:118-124
[21] Papakosta D K, Gagianas A A. Nitrogen and dry matter accumulation, remobilization and losses fronMdeiterranean wheat during grain filling[J]. Agron J, 1991,83:864-870
[22] Pheloung P C, Siddique K HM. Contributrion of stem dry matter to grain yield in wheat cultivars[J]. Journal of Plant Physilogy,1991,18:53-64
[23]郭文善,严六零,封超年等.小麦源库协调栽培途径的研究[J].江苏农学院学报,1995,16(1):33-37
[24]余松烈,元新华.小麦生育后期植株不同部位的光合同化量及其运转、贮藏与群体大小关系的研究[J].山东农学院学报,1892, 2:75-86
[25]张重义.高产小麦产量形成的特点[J].国外农学一麦类作物,1991(6): 47-50.
[26] Darwinkle A. Patterns of tillering and grain production of winter wheat at a wide range of plant densities[J]. Net. J. Agric. Sci., 1978,26:383-389
[27]杨兆生,许红霞,梁文科,等.紧凑大穗型小麦新品种3中6(37)的选育及亩产600 kg栽培技术研究[J]作物学报,1998,2 4(6):9 71-977
[28]张立言,刘树欣,李振国,等.高产麦田开花后干物质的积累、运转、分配与产量形成田[J].北京农学院学报,1988,3 ( 2):76-82.
[29] Gent M P N. Photosynthate reserves during grain filling in winter wheat[J].Agronomy Journal,1994,86:159-167
[30]山仑.春小麦对有限灌水的生理生态反应.见许越先等,农业用水有效性研究,科学出版社, 1992
[31] Austin R B, Morgan C L, Ford M A, et al. Contributions to grain yield fron pre-anthesis assimilation in tall and dwarf barley phenotypes in two contrasting seasons[J].Ann,Bot,1980b,45:309-319
[32] Kobata T, Palta J A, Turnwe N C. Rate of development of post-anthesis water deficits and grain filling of spring wheat[J].Crop Science,1992,32:1238-1242
[33] Zhang J, Sui X, Li B, et al. An improved water-use efficiency for winter wheat grown under reduced irrigation[J]. Field Crop Res,1998,59:91-98
[34] Brown J H, Paliyath G, Thompson J E. Physiological mechanisms of plant senescence, In F.C.Steward. Plant[J]. Physilogy,1991,227-275
[35] Palta J A, Kobata T, Turner N C,et al. Remobilization of carbon of carbon and nitrogen in wheat as influenced by postanthesis water deficits. Crop Science,1994,34:118-124
[36] Nicolas M E, Lambers H, Simpson R J, et al. Effect of drought on metabolism and partitioning of carbon in two wheat varieties differing in drought-tolerance[J]. Ann. Bot,1985,55:727-747
[37]杨建昌,王志琴,朱庆森.不同土壤水分状况下氮素营养对水稻产量的影响及其生理机制的研究[J]中国农业科学,1996,13(1):16-20
[38]朱庆森,张祖建,杨建昌.亚种间杂交稻产量源库关系的研究[J].中国农业科学,1997,30(3) : 52-59
[39] Evers A D, Lindley J. The particle size distribution in the. Wheat endosperm starch[J].Sci Food Agric,1977,28:98-102
[40] Weegels P I, Pijpekamp A M, Van de Graveland A. Depolymerisation and repolymerisation of wheat glutenin during macrooplymer content and quality parameters[J]. Cereal Sci,1996,23:103-114
[41] Ball S, Guan H P, James M. From glycogen to amylopectin: a model explaining the biogenesis of the p1ant starch granu1e[J].Cell,1996,86:349-352
[42] Przulj N, Momcilovic V.Genetic variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley[J]. Agron, 2001,15:255-265
[43]刘开昌,王庆成,张海松,冯凯。玉米叶片保绿性生理机理及遗传特性研究进展[J]山东农业科学,2003,2(2)48-50
[44]王之杰,王纪华,赵春江,等.用叶面标记态~(15)N研究冬小麦不同叶位氮素的运转[J ] .华北农学报,2004 ,19 (2) :71-75
[45]茹德平,赵彩霞,李习军,等.用~(15)N示踪技术研究高产小麦、玉米的施氮规律[J ] .核农学报,2005 ,19 (2) :151-154.
[46]李生秀,李宗让,田霄鸿,等.植物地上部分氮素的挥发损失[J ] .植物营养与肥料学报, 1995 ,2 :18-25.
[47]王百群,张卫,余存祖.用~(15)N示踪法研究不同土壤水分条件下小麦对氮的吸收利用[J ]核农学报,1999 ,13 (6) 3622367.
[48]尚兴甲,王梅芳,付宝余.运用同位素~(15)N研究冬小麦不同时期追施尿素的效果及氮肥的利用率[J]土壤肥料,2001 , (6) :9-11.
[49]沈其荣,徐国华.小麦和玉米叶面标记尿素态~(15)N的吸收和运输[J]土壤学报,2001 ,38(1) (1) :67-74.
[50]华珞,韦东普,白玲玉,等.氮锌硒肥配合施用对白三叶草的固氮作用与氮素转移的影响[J]生态学报, 2001 ,21 (4) : 5882592.
[51]王福钧,彭根元,兰林旺,等.应用~(15)N示踪技术研究水稻吸收肥料氮的动态及不同时期追施氮肥的作用[J]中国农业科学,1981 ,14 (4) :66271.
[52]刘强,罗泽民,荣湘民,等.不同时期不同施氮量对糙米蛋白质积累的影响初探[J ] .土壤学报,2000 ,37 (4) :529-535.
[53]黄见良,邹应斌,彭少兵,等.水稻对氮素的吸收、分配及其在组织中的挥发损失[J ] .植物营养与肥料学报,2004 ,10 (6) :579-581
[54]周珊庆,陈开铗,李合松,等.应用~(15)N示踪技术研究水稻对氮素的吸收利用[J ] .湖南农学院学报,1991 ,17 (4) 665-669
[55]王万里,林芝萍等.灌浆-成熟期间土壤干旱对小麦籽粒充实和物质转运的影响[J ]植物生理学报,1982, (1)67-68
[56] Spiertz J H J, Ellen J. Effects of nitrogen on crop development and grain growth of winter in relation to assimilation and utilization of assimilates and nutrients[J]. Agric.Sci,1978,26:210-231
[57] Van Sanford D A, Mackown C T. Cultivars differences in nitrogen remobilization during grain filling in soft red winter wheat[J]. Crop Science,1987,27:295-300
[58]张立言,刘树欣,李振国,等.高产麦田开花后干物质的积累、运转、分配与产量形成田[J].北京农学院学报,1988,3 ( 2):76-82.
[59]马溶慧.高产小麦群体质量指标及其与产量关系的研究[D].河南农业大学硕士学位论文,2005
[60] Yang J,Zhang J,Wang Z et al.Water deficitinduced senescence and its relationship to the remobilization of pre-stored carbon in wheat during grain filling.Agronomy Journal,2001,93:196～206.
[61] Yang J,Peng S,Visperas R M et al.Grain filling pattern and cytokinin cotent in the grains and roots of rice plants.Plant Growth Relation,2000,30(3):261～270.
[62] Yang J Zhang J,Ye Y et al Involvement of abscisic acid and ethylene in the responses of rice grains to water stress during filling.Plant,Cell and Environment,2004,27:1055～1064.
[63] Xue QW,Zhu Z X,Jack TM et al.Physiological mechanisms contributing to the increased water-use efficiency in winter wheat under deficit irrigation,Journal of Plant Physiology,2006,163(2):154～164.
[64] Ruuska S A,Rebetzke G J,van Herwaarden A et al.Genotypic variation in water-soluble carbohydrate accumulation in wheat.Functional Plant Biology,2006,33:799～809 .[65]王维,蔡一霞,张建华等。适度干旱对贪青小麦茎贮藏碳水化合物向籽粒运转的调节.作物学报,2005,31(3):289～296.
[66] Yang J,Zhang J,Wang Z et al.Activities of starch hydrolytic enzymes and sucrose-phosphate synthase in the stems of rice subjected to water stress during grain filling.Journal of Experimental Botany 2001,52:2169～2179.
[67] Lunn J E, Furbank R T. Sucrose biosynthesis in C4 Plants[J]. New Phytol, 1999,143 :221-223
[68] Leloir L F, Cardini C E. The biosynthesis of sucrose phosphate[J]. J Biol Chem, 1955,214:157-165
[69] Gallagher J N,Bisoe P N,Hunter B.Effects of drought ongraingrowth.Nature,1976,64:541-542.
[70] Austin R B C,Morgan L,Ford M A et al.Contribution to grain yield from pre-anthesis assimilation in tall and dwarf barley phenotype in two contrasting season.Annals of Botany Botany Botany,1980,45:309-319.
[71] Bidinger F R,Musgrav R B,Fischer R A.Contribution of stored pre-anthesis assimilates to grain yield in wheat and barley.Nature,1977,270:431-433.。
[72]吕金印,山仑,高俊凤.土壤干湿交替对小麦花前碳同化物分配的影响.西北植物学报,2004,24:1565-1569.
[73] Plauta Z,Butow B J,Blumenthalb C S et al.Transport of dry matter into developing wheat kernels and its contribution to grain yield under post-anthesis water deficit and elevated temperature.Field Crops Research,2004,86:185-198
[74]徐凤.小麦品种源库生态规律的研究[J].安徽农学院学报.1985 (1):1-12
[75]徐凤.小麦品种源库流的辩证关系及高产品种结构型问题[M].安徽农业科技,1978,:65-92
[76]严威凯.不同小麦品种源库状况的初步研究[J].西北农业大学学报,1988,39-44.
[77]曹显祖,朱庆森.水稻品种的源库特征及其类型划分的研究[J].作物学报,1987,13(4):265-272.
[78]任明全.小麦品种的产量潜力及其遗传改良[J].国外农学-麦类作物,1993,(2):41-46.
[79]彭永欣,郭文善,封超年等.小麦源库机理与高产栽培配套技术[M].稻麦研究新进展,东南大学出版社,1991: 146
[81]王甫同,郭文善,朱新开.小麦高粒叶比群体源形成特性[M].江苏农学院学报,1996, 17:85-90.
[81]彭永欣,封超年,郭文善等.小麦潜在源库与产量关系的研究[M].稻麦研究新进展,东南大学出版社,1991: 142.
[82]王振林,尹燕枰,贺明荣等.小麦源库比与产盘形成期同化物分配及结实性的研究[J].山东农大学报,1995, 26 (2):144-149
[83]郭文善,凌启鸿,彭永欣等.小麦群体粒叶比与高产栽培途径的研究.稻麦研究新进展[M],东南大学出版社,1991: 134-141
[84]李金才,陈峰.播种密度对小麦茎秆大维管束系统和穗部发育影响的研究[M].江苏农学院学报,1996, 17:123-127
[85]李国强,朱云集,郭天财等.硫氮配施对强筋小麦豫麦34籽粒灌浆特性的影响[J].麦类作物学报2006, 26 (2) : 98-102
[86]李秀君,潘宗东.不同粒重小麦品种子粒灌浆特性研究[J].中国农业科技导报,2005,,7:(1)
[87]沈学善,朱云集,郭天财等.施硫对‘豫麦50’籽粒灌浆特性及产量的影响[J].西北植物学报,2007 ,27 (6) :1265– 1269
[88]蔡庆生.小麦子粒生长各阶段干物质积累量与粒重的关系[J].南京农业大学学报,1993,16 (1) :27～32
[89]段国辉,高海涛,温红霞等.黄淮麦区不同习性小麦籽粒灌浆特性分析[J].植物生理科学,2007,12:(23),224-228.
[90]张晓龙.小麦品种籽粒灌浆的研究[J].作物学报,1982,8(2):87-92.
[91]刘丰明,陈明灿,郭香风.高产小麦粒重形成的灌浆特性分析[J].麦类作物学报, 1997,17(6):38-41.
[92] Blade S F, R J Baker. Kernel weight response to source-sink changes in spring wheat[J]. Crop Sci. 1991, 31(5):1117-1120.
[93] Perez P R, Martinez-carrasco, I M Martin Del Molino,et al. Nitrogen uptake and accumulation in grains of three winter wheat varieties with altered source-sink ratios[J]. Exper. Bot.,1989,40(215):707- 710.
[94] Martra N P, Singha Roy and S P Sen. Source-sink relationship: control of photosynthesis in the flag leaf by the panicle in wheat[J]. Plant Cell Physiol,1987,28(5):911-917.
[95] Koshkin E I, Tararina V V. Yield and source/sink relations of spring wheat cultivars[J]. Field Crop Research,1989,(22):294-306
[96] D J Miralles and G A Slafer. Individual grain weight responses to genetic reduction in culm length in wheat as affected by source-sink manipulations[J]. Field Crop Search,1995,(43):55-66.
[97] Evans L T ed. Crop Physiology. Cambridge University Press,London.1975: 101-149.
[98] Jenner C F, Ugalde T D, Aspinal D. The physiology of starch and protein deposition in the endosperm of wheat. Australian Journal ofPlant Physiollogy, 1991, 18: 211-216.
[2]胡廷积,杨永光,马元喜等.小麦生态与生产技术[M].河南:河南科学技术出版社,1986. 19-23
[3]董树亭,王空军,胡昌浩等.玉米品种更替过程中群体光合特性的演变[J].作物学报,2000,26 (2):200-204
[4]何萍,金继运.保绿型玉米的营养生理研究进展[J].玉米科学.2000, 8 (4): 41-44
[5]何萍,金继运.氮素营养对不同衰老类型玉米碳氮化合物解积的影响(收录入《青年学者论土壤与植物营养科学》),中国农业科技出版社,2000
[6] Xu, W. W., D.T. Rosenow, and H.T, Nguyen. Stay green trait in grain sorghum:Visual rating and objective measuring[J]. Plant Breeding,2000,119:365-367
[7]赵松岭.集水农业引论[M].西安:陕西科学技术出版社,1996.15-16.
[8] Howard Thomas, Catherine J. Howarth.Five ways to stay green[J]. Journal and experimental botany; 2002, 51(1):329-337
[9] Thomas H . Crops that stay green[J] Annals of applied biolpgy ,1993,123(1):193-219.
[10] Bekavac G.Path analysis of stay-green trait in maize[J].Cereal research communications 1998. 26(2): 1661-1671
[11] Guiamet JJ, Schwartz E, Pichersky E, Nooden LD (1991)Characterization of cytoplasmic and nuclear mutations affectingchlorophyll and chlorophyll-binding proteins during senescencein soybean. Plant Physiol 1996:227–231
[12] Gentinetta E, Ceppi D, Lepori C. A major gene for delayed senescence in maize. Pattern of photo- synthates accumulation and inheritance. Plant Breed1997:193–203
[13] Richard S .Inheritance of the Stay Green Trait in Sorghum[J].Crop Science,1994,34;970-972
[14] L M Dwyer and M Tollenaar.Genetic Improvement in Photosyn2thetic Response of Hybrid Maize Cultivars , 1959 to 1989[J] Plant Science ,1989 ,69 :81-91
[15] Guiamet JJ, Giannibelli MC . Nuclear and cytoplasmic“stay green”Mutations of soybean alter the loss of leaf soluble proteins during senescence. Physiol Plant 1996:665–661
[16] Rawson H M, Hindmarsh J H, Fischer R A, et al. Changes in leaf photosynthesis with plant ontogeny and relationships with yield per ear in whera cultivars and 120 progeny[J]. Plant Physiol, 1983, 10, 503- 514
[17] Mccaig T N, Clarke J M. Seasonal changes in nonstructural carbohydrate levels of wheat and oats grown in semiarid environment[J].Crop Science,1982,22:963-970
[18] Nooden L D, Gniamet J J, John I. Senescence mechanisms[J]. Physiol Plant, 1997,101:746-753
[19] Ntanos D A, Koutroubas S D. Dry matter anf N accumulation and translocation for Indica and Japonica rice Mediterranean conditions[J]. Field Crops,2002,74:93-101
[20] Palta J A, Kobata T, Turner N C,et al. Remobilization of carbon of carbon and nitrogen in wheat as influenced by postanthesis water deficits. Crop Science,1994,34:118-124
[21] Papakosta D K, Gagianas A A. Nitrogen and dry matter accumulation, remobilization and losses fronMdeiterranean wheat during grain filling[J]. Agron J, 1991,83:864-870
[22] Pheloung P C, Siddique K HM. Contributrion of stem dry matter to grain yield in wheat cultivars[J]. Journal of Plant Physilogy,1991,18:53-64
[23]郭文善,严六零,封超年等.小麦源库协调栽培途径的研究[J].江苏农学院学报,1995,16(1):33-37
[24]余松烈,元新华.小麦生育后期植株不同部位的光合同化量及其运转、贮藏与群体大小关系的研究[J].山东农学院学报,1892, 2:75-86
[25]张重义.高产小麦产量形成的特点[J].国外农学一麦类作物,1991(6): 47-50.
[26] Darwinkle A. Patterns of tillering and grain production of winter wheat at a wide range of plant densities[J]. Net. J. Agric. Sci., 1978,26:383-389
[27]杨兆生,许红霞,梁文科,等.紧凑大穗型小麦新品种3中6(37)的选育及亩产600 kg栽培技术研究[J]作物学报,1998,2 4(6):9 71-977
[28]张立言,刘树欣,李振国,等.高产麦田开花后干物质的积累、运转、分配与产量形成田[J].北京农学院学报,1988,3 ( 2):76-82.
[29] Gent M P N. Photosynthate reserves during grain filling in winter wheat[J].Agronomy Journal,1994,86:159-167
[30]山仑.春小麦对有限灌水的生理生态反应.见许越先等,农业用水有效性研究,科学出版社, 1992
[31] Austin R B, Morgan C L, Ford M A, et al. Contributions to grain yield fron pre-anthesis assimilation in tall and dwarf barley phenotypes in two contrasting seasons[J].Ann,Bot,1980b,45:309-319
[32] Kobata T, Palta J A, Turnwe N C. Rate of development of post-anthesis water deficits and grain filling of spring wheat[J].Crop Science,1992,32:1238-1242
[33] Zhang J, Sui X, Li B, et al. An improved water-use efficiency for winter wheat grown under reduced irrigation[J]. Field Crop Res,1998,59:91-98
[34] Brown J H, Paliyath G, Thompson J E. Physiological mechanisms of plant senescence, In F.C.Steward. Plant[J]. Physilogy,1991,227-275
[35] Palta J A, Kobata T, Turner N C,et al. Remobilization of carbon of carbon and nitrogen in wheat as influenced by postanthesis water deficits. Crop Science,1994,34:118-124
[36] Nicolas M E, Lambers H, Simpson R J, et al. Effect of drought on metabolism and partitioning of carbon in two wheat varieties differing in drought-tolerance[J]. Ann. Bot,1985,55:727-747
[37]杨建昌,王志琴,朱庆森.不同土壤水分状况下氮素营养对水稻产量的影响及其生理机制的研究[J]中国农业科学,1996,13(1):16-20
[38]朱庆森,张祖建,杨建昌.亚种间杂交稻产量源库关系的研究[J].中国农业科学,1997,30(3) : 52-59
[39] Evers A D, Lindley J. The particle size distribution in the. Wheat endosperm starch[J].Sci Food Agric,1977,28:98-102
[40] Weegels P I, Pijpekamp A M, Van de Graveland A. Depolymerisation and repolymerisation of wheat glutenin during macrooplymer content and quality parameters[J]. Cereal Sci,1996,23:103-114
[41] Ball S, Guan H P, James M. From glycogen to amylopectin: a model explaining the biogenesis of the p1ant starch granu1e[J].Cell,1996,86:349-352
[42] Przulj N, Momcilovic V.Genetic variation for dry matter and nitrogen accumulation and translocation in two-rowed spring barley[J]. Agron, 2001,15:255-265
[43]刘开昌,王庆成,张海松,冯凯。玉米叶片保绿性生理机理及遗传特性研究进展[J]山东农业科学,2003,2(2)48-50
[44]王之杰,王纪华,赵春江,等.用叶面标记态~(15)N研究冬小麦不同叶位氮素的运转[J ] .华北农学报,2004 ,19 (2) :71-75
[45]茹德平,赵彩霞,李习军,等.用~(15)N示踪技术研究高产小麦、玉米的施氮规律[J ] .核农学报,2005 ,19 (2) :151-154.
[46]李生秀,李宗让,田霄鸿,等.植物地上部分氮素的挥发损失[J ] .植物营养与肥料学报, 1995 ,2 :18-25.
[47]王百群,张卫,余存祖.用~(15)N示踪法研究不同土壤水分条件下小麦对氮的吸收利用[J ]核农学报,1999 ,13 (6) 3622367.
[48]尚兴甲,王梅芳,付宝余.运用同位素~(15)N研究冬小麦不同时期追施尿素的效果及氮肥的利用率[J]土壤肥料,2001 , (6) :9-11.
[49]沈其荣,徐国华.小麦和玉米叶面标记尿素态~(15)N的吸收和运输[J]土壤学报,2001 ,38(1) (1) :67-74.
[50]华珞,韦东普,白玲玉,等.氮锌硒肥配合施用对白三叶草的固氮作用与氮素转移的影响[J]生态学报, 2001 ,21 (4) : 5882592.
[51]王福钧,彭根元,兰林旺,等.应用~(15)N示踪技术研究水稻吸收肥料氮的动态及不同时期追施氮肥的作用[J]中国农业科学,1981 ,14 (4) :66271.
[52]刘强,罗泽民,荣湘民,等.不同时期不同施氮量对糙米蛋白质积累的影响初探[J ] .土壤学报,2000 ,37 (4) :529-535.
[53]黄见良,邹应斌,彭少兵,等.水稻对氮素的吸收、分配及其在组织中的挥发损失[J ] .植物营养与肥料学报,2004 ,10 (6) :579-581
[54]周珊庆,陈开铗,李合松,等.应用~(15)N示踪技术研究水稻对氮素的吸收利用[J ] .湖南农学院学报,1991 ,17 (4) 665-669
[55]王万里,林芝萍等.灌浆-成熟期间土壤干旱对小麦籽粒充实和物质转运的影响[J ]植物生理学报,1982, (1)67-68
[56] Spiertz J H J, Ellen J. Effects of nitrogen on crop development and grain growth of winter in relation to assimilation and utilization of assimilates and nutrients[J]. Agric.Sci,1978,26:210-231
[57] Van Sanford D A, Mackown C T. Cultivars differences in nitrogen remobilization during grain filling in soft red winter wheat[J]. Crop Science,1987,27:295-300
[58]张立言,刘树欣,李振国,等.高产麦田开花后干物质的积累、运转、分配与产量形成田[J].北京农学院学报,1988,3 ( 2):76-82.
[59]马溶慧.高产小麦群体质量指标及其与产量关系的研究[D].河南农业大学硕士学位论文,2005
[60] Yang J,Zhang J,Wang Z et al.Water deficitinduced senescence and its relationship to the remobilization of pre-stored carbon in wheat during grain filling.Agronomy Journal,2001,93:196～206.
[61] Yang J,Peng S,Visperas R M et al.Grain filling pattern and cytokinin cotent in the grains and roots of rice plants.Plant Growth Relation,2000,30(3):261～270.
[62] Yang J Zhang J,Ye Y et al Involvement of abscisic acid and ethylene in the responses of rice grains to water stress during filling.Plant,Cell and Environment,2004,27:1055～1064.
[63] Xue QW,Zhu Z X,Jack TM et al.Physiological mechanisms contributing to the increased water-use efficiency in winter wheat under deficit irrigation,Journal of Plant Physiology,2006,163(2):154～164.
[64] Ruuska S A,Rebetzke G J,van Herwaarden A et al.Genotypic variation in water-soluble carbohydrate accumulation in wheat.Functional Plant Biology,2006,33:799～809 .[65]王维,蔡一霞,张建华等。适度干旱对贪青小麦茎贮藏碳水化合物向籽粒运转的调节.作物学报,2005,31(3):289～296.
[66] Yang J,Zhang J,Wang Z et al.Activities of starch hydrolytic enzymes and sucrose-phosphate synthase in the stems of rice subjected to water stress during grain filling.Journal of Experimental Botany 2001,52:2169～2179.
[67] Lunn J E, Furbank R T. Sucrose biosynthesis in C4 Plants[J]. New Phytol, 1999,143 :221-223
[68] Leloir L F, Cardini C E. The biosynthesis of sucrose phosphate[J]. J Biol Chem, 1955,214:157-165
[69] Gallagher J N,Bisoe P N,Hunter B.Effects of drought ongraingrowth.Nature,1976,64:541-542.
[70] Austin R B C,Morgan L,Ford M A et al.Contribution to grain yield from pre-anthesis assimilation in tall and dwarf barley phenotype in two contrasting season.Annals of Botany Botany Botany,1980,45:309-319.
[71] Bidinger F R,Musgrav R B,Fischer R A.Contribution of stored pre-anthesis assimilates to grain yield in wheat and barley.Nature,1977,270:431-433.。
[72]吕金印,山仑,高俊凤.土壤干湿交替对小麦花前碳同化物分配的影响.西北植物学报,2004,24:1565-1569.
[73] Plauta Z,Butow B J,Blumenthalb C S et al.Transport of dry matter into developing wheat kernels and its contribution to grain yield under post-anthesis water deficit and elevated temperature.Field Crops Research,2004,86:185-198
[74]徐凤.小麦品种源库生态规律的研究[J].安徽农学院学报.1985 (1):1-12
[75]徐凤.小麦品种源库流的辩证关系及高产品种结构型问题[M].安徽农业科技,1978,:65-92
[76]严威凯.不同小麦品种源库状况的初步研究[J].西北农业大学学报,1988,39-44.
[77]曹显祖,朱庆森.水稻品种的源库特征及其类型划分的研究[J].作物学报,1987,13(4):265-272.
[78]任明全.小麦品种的产量潜力及其遗传改良[J].国外农学-麦类作物,1993,(2):41-46.
[79]彭永欣,郭文善,封超年等.小麦源库机理与高产栽培配套技术[M].稻麦研究新进展,东南大学出版社,1991: 146
[81]王甫同,郭文善,朱新开.小麦高粒叶比群体源形成特性[M].江苏农学院学报,1996, 17:85-90.
[81]彭永欣,封超年,郭文善等.小麦潜在源库与产量关系的研究[M].稻麦研究新进展,东南大学出版社,1991: 142.
[82]王振林,尹燕枰,贺明荣等.小麦源库比与产盘形成期同化物分配及结实性的研究[J].山东农大学报,1995, 26 (2):144-149
[83]郭文善,凌启鸿,彭永欣等.小麦群体粒叶比与高产栽培途径的研究.稻麦研究新进展[M],东南大学出版社,1991: 134-141
[84]李金才,陈峰.播种密度对小麦茎秆大维管束系统和穗部发育影响的研究[M].江苏农学院学报,1996, 17:123-127
[85]李国强,朱云集,郭天财等.硫氮配施对强筋小麦豫麦34籽粒灌浆特性的影响[J].麦类作物学报2006, 26 (2) : 98-102
[86]李秀君,潘宗东.不同粒重小麦品种子粒灌浆特性研究[J].中国农业科技导报,2005,,7:(1)
[87]沈学善,朱云集,郭天财等.施硫对‘豫麦50’籽粒灌浆特性及产量的影响[J].西北植物学报,2007 ,27 (6) :1265– 1269
[88]蔡庆生.小麦子粒生长各阶段干物质积累量与粒重的关系[J].南京农业大学学报,1993,16 (1) :27～32
[89]段国辉,高海涛,温红霞等.黄淮麦区不同习性小麦籽粒灌浆特性分析[J].植物生理科学,2007,12:(23),224-228.
[90]张晓龙.小麦品种籽粒灌浆的研究[J].作物学报,1982,8(2):87-92.
[91]刘丰明,陈明灿,郭香风.高产小麦粒重形成的灌浆特性分析[J].麦类作物学报, 1997,17(6):38-41.
[92] Blade S F, R J Baker. Kernel weight response to source-sink changes in spring wheat[J]. Crop Sci. 1991, 31(5):1117-1120.
[93] Perez P R, Martinez-carrasco, I M Martin Del Molino,et al. Nitrogen uptake and accumulation in grains of three winter wheat varieties with altered source-sink ratios[J]. Exper. Bot.,1989,40(215):707- 710.
[94] Martra N P, Singha Roy and S P Sen. Source-sink relationship: control of photosynthesis in the flag leaf by the panicle in wheat[J]. Plant Cell Physiol,1987,28(5):911-917.
[95] Koshkin E I, Tararina V V. Yield and source/sink relations of spring wheat cultivars[J]. Field Crop Research,1989,(22):294-306
[96] D J Miralles and G A Slafer. Individual grain weight responses to genetic reduction in culm length in wheat as affected by source-sink manipulations[J]. Field Crop Search,1995,(43):55-66.
[97] Evans L T ed. Crop Physiology. Cambridge University Press,London.1975: 101-149.
[98] Jenner C F, Ugalde T D, Aspinal D. The physiology of starch and protein deposition in the endosperm of wheat. Australian Journal ofPlant Physiollogy, 1991, 18: 211-216.