石灰性土壤上供锌对不同基因型小麦生长及锌吸收的影响
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摘要
锌是动植物必需微量元素,锌通过植物进入食物链,直接或通过动物间接进入人类的膳食中,进而影响人体的营养平衡及身体健康。对以小麦为主食的人群来说,小麦籽粒锌是人体锌的最主要来源,因此小麦籽粒锌含量低极易导致敏感人群出现缺锌,而解决此问题最有效途径是提高小麦籽粒锌含量,其主要方法通过施用锌肥及锌高效基因型小麦筛选。本研究通过水培、盆栽及田间试验方法,探讨石灰性土壤中CaCO3对锌有效性的影响,施用锌肥对土壤中不同形态锌含量比例的影响,同时评价不同冬小麦基因型的锌效率及对锌肥后效的反应,以期提高小麦营养品质。本研究得到以下主要结论:
(1)通过营养液培养试验,研究了供Zn条件下添加CaCO3对3种基因型冬小麦(远丰998、中育6号、小偃22)幼苗生长及Zn吸收的影响。结果表明:供Zn和添加CaCO3对小麦幼苗生长量和根冠比均无显著性影响,3种基因型小麦间亦无显著性差异;添加CaCO3诱发了小麦叶片失绿黄化。无论供Zn还是不供Zn,添加CaCO3对3种基因型小麦根、茎、叶各部分的Zn含量及累积量均无显著性影响;与不供Zn处理相比,供Zn会大幅度提高根、茎、叶的Zn含量和累积量,供Zn使远丰998、中育6号、小偃22小麦植株Zn含量分别增加80.0%、104.8%和139.6%,缺Zn敏感型小麦远丰998植株Zn含量和累积量的增加幅度远小于不敏感型小麦中育6号和小偃22。供Zn和添加CaCO3对小麦幼苗根、茎、叶中P含量均无显著性影响,但远丰998小麦根、茎、叶3部分的P含量均明显低于其它两种非敏感型小麦。供Zn使小麦根、茎、叶3部分的P/Zn大幅度降低,添加CaCO3也使P/Zn呈现降低的趋势。不供Zn条件下添加CaCO3能诱发小麦失绿黄化但又不降低Zn吸收量。表明在水培条件下高含量CaCO3对小麦Zn吸收并未产生明显的抑制作用。
(2)土培试验设置了由低到高的土壤CaCO3含量梯度,以探讨CaCO3对土壤有效Zn含量、两种基因型小麦(远丰998,中育6号)生长发育及Zn、Fe营养的影响。结果表明:与不施Zn相比,施Zn使土壤有效Zn含量增加了3.22倍;高含量CaCO3使土壤有效Zn含量有所降低,新施入的有效态Zn仅有1.3%被小麦吸收,大部分则转化为无效态Zn;CaCO3含量达到111.8 g/kg时,可明显抑制小麦对Fe的吸收,进一步提高CaCO3含量抑制作用有所减弱;两种小麦基因型生长存在明显的差异,中育6号的根冠比和分蘖数都显著高于远丰998;施Zn可显著增加小麦Zn含量和吸收量,CaCO3含量达到111.8 g/kg可显著降低小麦根的Zn含量,对其他部分影响不明显;此外,施Zn可增加叶片的Fe含量和转运率。
(3)选择10种当地主要小麦基因型,在石灰性土壤上进行连续两年的田间试验,仅在第一年施入锌肥,两年内均施氮肥,测定第二季小麦种植前、灌浆期及收获期不同土层中不同形态锌含量。结果表明,在有效锌含量低的石灰性土壤施用锌肥可增加小麦籽粒锌含量;种植小麦可以使土壤表层有效锌含量增加;施入土壤中的锌增加了交换态、松结合态、碳酸盐结合态、氧化锰结合态锌含量,可为第二季小麦生长提供锌源,而第二季小麦生长后,这些形态锌含量下降,与不施锌肥处理相比,施锌处理仅矿物态锌含量高于不施锌处理,从而锌的潜在供应源减少,肥效降低,但有效锌含量仍保持较高水平,能否供下一季小麦生长所需还有待进一步研究;施氮肥增加了小麦对锌的吸收,但土壤自身能补充这部分消耗;氮锌配施增加了小麦对锌的吸收,同时也降低了施锌肥增加的有效锌含量。
(4)大田试验结果还表明,石灰性土壤上施锌取得了一定的增产效果,但不同基因型增产效果不同,其中武农148、新麦13、陕优253为锌高效基因型,西杂1号、陕优225、小偃22为锌低效基因型;施锌肥可增加锌低效基因型小麦籽粒锌含量,而锌高效基因型对锌肥的后效反应较差。施氮肥主要有利于叶片生长,可以提高分蘖数和叶绿素SPAD值,并且施氮肥和锌肥可以减缓叶片黄化。10种小麦基因型籽粒锌转运率为46.3-64.8%,可见小麦吸收到地上部的锌只有一半被转移进入籽粒,而茎、叶、颖壳中还含有不少的锌。施锌肥增加了大部分基因型锌转运率,但增加幅度有限,少部分基因型转运率有所降低。
Zinc is an essential micronutrient for plant and animal, it enters into to the food chain through plant, and it enters the diets of human being directly or through animal indirectly, so the balance of nutrition and the health of buman being is affected by zinc. Zinc in wheat grain is the most important source of zinc nutrition to people whose stape food mainly depends on wheat food. So zinc deficiency of wheat grain can lead to zinc deficiency of sensitive people easily. And the most useful approach to solve this problem is to increase the zinc concentration of wheat grain, application of zinc fertilizer and screening the wheat genotypes with high zinc efficiency at present. This study was to investigate effects of calcacium carbonate on the availability of soil zinc in calcareous soil, and the effects of zinc fertilization on the effects of zinc concentration and bioavailability of different types, the wheat genotypes with high zinc efficiency or with better response on residual effect in the soil were expected to be screened to improve the nutrition qulity of wheat. The main results of this study are as follows:
(1) High concentration calcium carbonate in calcareous soils usually limits plant Zn uptake. In this paper, a solution culture experimentwas conducted to study the effects of CaCO3 addition with or no Zn supply on the growth and Zn up take of three genotypes winter wheat seedlings. The results showed that Zn supply or CaCO3 addition had no significant effects on seedlings biomass and root/canopy ratio, and there was no significant difference among the three genotypes. CaCO3 addition led to leaf chlorosis. No matter with or no Zn supply, CaCO3 addition had no significant effects on the Zn concentration and accumulation in wheat root, stem, and leaf. Zn supply increased the Zn concentration in Yuanfeng 998, Zhongyu 6, and Xiaoyan 22 by 80.0%, 104.8% and 139.6%, respectively. For Yuanfeng 998, a sensitive genotype to Zn-deficiency, the increment of Zn concentration and accumulation was much lower than that of Zhongyu 6 and Xiaoyan 22, non-sensitive genotypes to Zn-deficiency. Zn supply or CaCO3 addition had no significant effects on the P content in root, stem, and leaf, but the corresponding P content of Yuanfeng 998 was obviously lower than that of the other two genotypes. Zn supply greatly decreased the P/Zn ratio in root, stem and leaf, and CaCO3 addition also showed the same tendency. Under Zn-deficiency, CaCO3 addition caused leaf chlorosis, but did not decrease wheat plant Zn up take. In conclusion, at least under solution culture condition, high concentration calcium carbonate had no obvious inhibitory effect on plant zinc uptake, and whether this situation was true in calcareous soils is worthy to be further investigated.
(2) High content of calcium carbonate in calcareous soil is one of major factors which cause zinc deficiency of crops. In this research, varied amounts of calcium carbonate was added artificially to the calcareous soil and different levels of calcium carbonate was formed to investigate the effect of calcium carbonate on available zinc content in soil, the growth situation as well as Zn, Fe nutrition of two different wheat genotypes (Yuanfeng 998, Zhongyu 6). The obtained results showed that, the available Zn content of soil increased over 3 times at harvest from the initial 0.65 mg/kg at beginning of culture when adding Zn fertilizer (40 mg/kg) to soil; high content of calcium carbonate decreased the availability of zinc, and only about 1.3% of zinc added was taken up by wheat plants during the culture period, and most of newly added available zinc was transformed into unavailable form. The uptake of Fe by wheat plants was inhibited significantly when calcium carbonate content was high as 111.8 g/kg, however, the inhibition would become weaker when adding more calcium carbonate to soil. There existed obvious differences for their growth between two tested wheat genotypes, the ratio of root/shoot and tillering number of Zhongyu 6 was higher significantly than that of Yuanfeng 998. The Zn concentration and content of wheat plants was greatly increased following Zn application, the Zn concentration of root was inhibited when calcium carbonate was added to 111.8 g/kg, whereas it was not obvious for other parts of wheat plants. In addition, the Fe concentration of leaf and translocation efficiency was increased with Zn application.
(3) 10 genotypes which were planted in Shaanxi Guanzhong Plain Area mainly were chosen in this study, a field experiment was carried out on calcaous soil for two years in succession, Zn fertilizer was applied in the first year, no zinc fertilization in the second year, but nitrogen fertilization was conducted in the continouns two years. The soil zinc contents of different forms in 0-20 cm and 20-40 cm soil depths was determined before wheat sowing, the filling stage and maturity stage, respectively. The results obtained showed that the Zn concentration in wheat grain could be increased by zinc fertilization on calcareous soil in which had low level of available zinc contend. The available Zn content in topsoil could be increased by planting wheat, the content of EX-Zn, WBO-Zn, CARB-Zn and OXMn-Zn were all increased by zinc fertilization, and these different forms of Zn could be utilized by wheat plants in the next year, furthermore, the contents of these various forms of Zn decreased after the wheat harvest, only the content of MIN-Zn was higher in soil at Zn fertilization compared to no zinc fertilization, so the potential Zn source in the soil reduced, and the zinc fertilizer response was decreased, but the available Zn content was still in high level, so it is worth further studying the residual effect of Zn fertilizer applied on the growth of wheat in the years that followed; application of nitrogen fertilizer increased the absorbtion of Zn from soil to wheat, but the soil can compensate the consumption by itself; To the combination of nitrogen and Zn fertilization, application of nitrogen fertilizer increased the absorbtion of Zn to wheat, but decreased the available zinc content in the soil which increased by zinc fertilization at the same time.
(4) Grain yield and Zn concentration of wheat grain were also investigated in the experiment to screen the genotypes with high Zn efficiency and with better response on residual effect in the soil. Zn fertilization increased grain yield to some extent, but there existed obvious genotypic difference of the increasing effects, among the 10 genotypes tested, Wunong148, Xinmai13 and Shanyou253 were three genotypes with high Zn efficiency; Xiza1, Shanyou225 and Xiaoyan22 were three genotypes with low Zn efficiency. The Zn concentration in grain of wheat geontypes with low Zn effiency could be increased by application of Zn fertilizer, but Zn fertilizer reponses of the genotypes with high Zn efficiency was not significant. Nitrogen fertilization was mainly beneficial for leaf growth, and it could increase the numbers of tillers and the chlorophyll SPAD values, the leaf chlorosis could be alleviated by application of nitrogen and zinc fertilizers. The result of this study showed that the range of translocation ratio of Zn to the grain was 46.3-64.8%, indicating that only about half of the total Zn taken up was translocated to the wheat grain, so a great deal of Zn remained in the shell, leaf and stem, reusing or increasing the translocation ratio could enhance the utilization of Zn fertilizer. The Zn translocation ratios of a majority of genotypes were increased by Zn fertilization, but the extent was not so large, and the zinc tranlocation ratios of a few of genotypes were decreased.
(1)通过营养液培养试验,研究了供Zn条件下添加CaCO3对3种基因型冬小麦(远丰998、中育6号、小偃22)幼苗生长及Zn吸收的影响。结果表明:供Zn和添加CaCO3对小麦幼苗生长量和根冠比均无显著性影响,3种基因型小麦间亦无显著性差异;添加CaCO3诱发了小麦叶片失绿黄化。无论供Zn还是不供Zn,添加CaCO3对3种基因型小麦根、茎、叶各部分的Zn含量及累积量均无显著性影响;与不供Zn处理相比,供Zn会大幅度提高根、茎、叶的Zn含量和累积量,供Zn使远丰998、中育6号、小偃22小麦植株Zn含量分别增加80.0%、104.8%和139.6%,缺Zn敏感型小麦远丰998植株Zn含量和累积量的增加幅度远小于不敏感型小麦中育6号和小偃22。供Zn和添加CaCO3对小麦幼苗根、茎、叶中P含量均无显著性影响,但远丰998小麦根、茎、叶3部分的P含量均明显低于其它两种非敏感型小麦。供Zn使小麦根、茎、叶3部分的P/Zn大幅度降低,添加CaCO3也使P/Zn呈现降低的趋势。不供Zn条件下添加CaCO3能诱发小麦失绿黄化但又不降低Zn吸收量。表明在水培条件下高含量CaCO3对小麦Zn吸收并未产生明显的抑制作用。
(2)土培试验设置了由低到高的土壤CaCO3含量梯度,以探讨CaCO3对土壤有效Zn含量、两种基因型小麦(远丰998,中育6号)生长发育及Zn、Fe营养的影响。结果表明:与不施Zn相比,施Zn使土壤有效Zn含量增加了3.22倍;高含量CaCO3使土壤有效Zn含量有所降低,新施入的有效态Zn仅有1.3%被小麦吸收,大部分则转化为无效态Zn;CaCO3含量达到111.8 g/kg时,可明显抑制小麦对Fe的吸收,进一步提高CaCO3含量抑制作用有所减弱;两种小麦基因型生长存在明显的差异,中育6号的根冠比和分蘖数都显著高于远丰998;施Zn可显著增加小麦Zn含量和吸收量,CaCO3含量达到111.8 g/kg可显著降低小麦根的Zn含量,对其他部分影响不明显;此外,施Zn可增加叶片的Fe含量和转运率。
(3)选择10种当地主要小麦基因型,在石灰性土壤上进行连续两年的田间试验,仅在第一年施入锌肥,两年内均施氮肥,测定第二季小麦种植前、灌浆期及收获期不同土层中不同形态锌含量。结果表明,在有效锌含量低的石灰性土壤施用锌肥可增加小麦籽粒锌含量;种植小麦可以使土壤表层有效锌含量增加;施入土壤中的锌增加了交换态、松结合态、碳酸盐结合态、氧化锰结合态锌含量,可为第二季小麦生长提供锌源,而第二季小麦生长后,这些形态锌含量下降,与不施锌肥处理相比,施锌处理仅矿物态锌含量高于不施锌处理,从而锌的潜在供应源减少,肥效降低,但有效锌含量仍保持较高水平,能否供下一季小麦生长所需还有待进一步研究;施氮肥增加了小麦对锌的吸收,但土壤自身能补充这部分消耗;氮锌配施增加了小麦对锌的吸收,同时也降低了施锌肥增加的有效锌含量。
(4)大田试验结果还表明,石灰性土壤上施锌取得了一定的增产效果,但不同基因型增产效果不同,其中武农148、新麦13、陕优253为锌高效基因型,西杂1号、陕优225、小偃22为锌低效基因型;施锌肥可增加锌低效基因型小麦籽粒锌含量,而锌高效基因型对锌肥的后效反应较差。施氮肥主要有利于叶片生长,可以提高分蘖数和叶绿素SPAD值,并且施氮肥和锌肥可以减缓叶片黄化。10种小麦基因型籽粒锌转运率为46.3-64.8%,可见小麦吸收到地上部的锌只有一半被转移进入籽粒,而茎、叶、颖壳中还含有不少的锌。施锌肥增加了大部分基因型锌转运率,但增加幅度有限,少部分基因型转运率有所降低。
Zinc is an essential micronutrient for plant and animal, it enters into to the food chain through plant, and it enters the diets of human being directly or through animal indirectly, so the balance of nutrition and the health of buman being is affected by zinc. Zinc in wheat grain is the most important source of zinc nutrition to people whose stape food mainly depends on wheat food. So zinc deficiency of wheat grain can lead to zinc deficiency of sensitive people easily. And the most useful approach to solve this problem is to increase the zinc concentration of wheat grain, application of zinc fertilizer and screening the wheat genotypes with high zinc efficiency at present. This study was to investigate effects of calcacium carbonate on the availability of soil zinc in calcareous soil, and the effects of zinc fertilization on the effects of zinc concentration and bioavailability of different types, the wheat genotypes with high zinc efficiency or with better response on residual effect in the soil were expected to be screened to improve the nutrition qulity of wheat. The main results of this study are as follows:
(1) High concentration calcium carbonate in calcareous soils usually limits plant Zn uptake. In this paper, a solution culture experimentwas conducted to study the effects of CaCO3 addition with or no Zn supply on the growth and Zn up take of three genotypes winter wheat seedlings. The results showed that Zn supply or CaCO3 addition had no significant effects on seedlings biomass and root/canopy ratio, and there was no significant difference among the three genotypes. CaCO3 addition led to leaf chlorosis. No matter with or no Zn supply, CaCO3 addition had no significant effects on the Zn concentration and accumulation in wheat root, stem, and leaf. Zn supply increased the Zn concentration in Yuanfeng 998, Zhongyu 6, and Xiaoyan 22 by 80.0%, 104.8% and 139.6%, respectively. For Yuanfeng 998, a sensitive genotype to Zn-deficiency, the increment of Zn concentration and accumulation was much lower than that of Zhongyu 6 and Xiaoyan 22, non-sensitive genotypes to Zn-deficiency. Zn supply or CaCO3 addition had no significant effects on the P content in root, stem, and leaf, but the corresponding P content of Yuanfeng 998 was obviously lower than that of the other two genotypes. Zn supply greatly decreased the P/Zn ratio in root, stem and leaf, and CaCO3 addition also showed the same tendency. Under Zn-deficiency, CaCO3 addition caused leaf chlorosis, but did not decrease wheat plant Zn up take. In conclusion, at least under solution culture condition, high concentration calcium carbonate had no obvious inhibitory effect on plant zinc uptake, and whether this situation was true in calcareous soils is worthy to be further investigated.
(2) High content of calcium carbonate in calcareous soil is one of major factors which cause zinc deficiency of crops. In this research, varied amounts of calcium carbonate was added artificially to the calcareous soil and different levels of calcium carbonate was formed to investigate the effect of calcium carbonate on available zinc content in soil, the growth situation as well as Zn, Fe nutrition of two different wheat genotypes (Yuanfeng 998, Zhongyu 6). The obtained results showed that, the available Zn content of soil increased over 3 times at harvest from the initial 0.65 mg/kg at beginning of culture when adding Zn fertilizer (40 mg/kg) to soil; high content of calcium carbonate decreased the availability of zinc, and only about 1.3% of zinc added was taken up by wheat plants during the culture period, and most of newly added available zinc was transformed into unavailable form. The uptake of Fe by wheat plants was inhibited significantly when calcium carbonate content was high as 111.8 g/kg, however, the inhibition would become weaker when adding more calcium carbonate to soil. There existed obvious differences for their growth between two tested wheat genotypes, the ratio of root/shoot and tillering number of Zhongyu 6 was higher significantly than that of Yuanfeng 998. The Zn concentration and content of wheat plants was greatly increased following Zn application, the Zn concentration of root was inhibited when calcium carbonate was added to 111.8 g/kg, whereas it was not obvious for other parts of wheat plants. In addition, the Fe concentration of leaf and translocation efficiency was increased with Zn application.
(3) 10 genotypes which were planted in Shaanxi Guanzhong Plain Area mainly were chosen in this study, a field experiment was carried out on calcaous soil for two years in succession, Zn fertilizer was applied in the first year, no zinc fertilization in the second year, but nitrogen fertilization was conducted in the continouns two years. The soil zinc contents of different forms in 0-20 cm and 20-40 cm soil depths was determined before wheat sowing, the filling stage and maturity stage, respectively. The results obtained showed that the Zn concentration in wheat grain could be increased by zinc fertilization on calcareous soil in which had low level of available zinc contend. The available Zn content in topsoil could be increased by planting wheat, the content of EX-Zn, WBO-Zn, CARB-Zn and OXMn-Zn were all increased by zinc fertilization, and these different forms of Zn could be utilized by wheat plants in the next year, furthermore, the contents of these various forms of Zn decreased after the wheat harvest, only the content of MIN-Zn was higher in soil at Zn fertilization compared to no zinc fertilization, so the potential Zn source in the soil reduced, and the zinc fertilizer response was decreased, but the available Zn content was still in high level, so it is worth further studying the residual effect of Zn fertilizer applied on the growth of wheat in the years that followed; application of nitrogen fertilizer increased the absorbtion of Zn from soil to wheat, but the soil can compensate the consumption by itself; To the combination of nitrogen and Zn fertilization, application of nitrogen fertilizer increased the absorbtion of Zn to wheat, but decreased the available zinc content in the soil which increased by zinc fertilization at the same time.
(4) Grain yield and Zn concentration of wheat grain were also investigated in the experiment to screen the genotypes with high Zn efficiency and with better response on residual effect in the soil. Zn fertilization increased grain yield to some extent, but there existed obvious genotypic difference of the increasing effects, among the 10 genotypes tested, Wunong148, Xinmai13 and Shanyou253 were three genotypes with high Zn efficiency; Xiza1, Shanyou225 and Xiaoyan22 were three genotypes with low Zn efficiency. The Zn concentration in grain of wheat geontypes with low Zn effiency could be increased by application of Zn fertilizer, but Zn fertilizer reponses of the genotypes with high Zn efficiency was not significant. Nitrogen fertilization was mainly beneficial for leaf growth, and it could increase the numbers of tillers and the chlorophyll SPAD values, the leaf chlorosis could be alleviated by application of nitrogen and zinc fertilizers. The result of this study showed that the range of translocation ratio of Zn to the grain was 46.3-64.8%, indicating that only about half of the total Zn taken up was translocated to the wheat grain, so a great deal of Zn remained in the shell, leaf and stem, reusing or increasing the translocation ratio could enhance the utilization of Zn fertilizer. The Zn translocation ratios of a majority of genotypes were increased by Zn fertilization, but the extent was not so large, and the zinc tranlocation ratios of a few of genotypes were decreased.
引文
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